Retracing the Aurochs: History, Morphology & Ecology of an Extinct Wild Ox

AUROCHS history, morphology and Cis van Vuure ecology of an extinct wild ox M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 4...

Author: Cis Van Vuure

31 downloads 1277 Views 22MB Size Report

This content was uploaded by our users and we assume good faith they have the permission to share this book. If you own the copyright to this book and it is wrongfully on our website, we offer a simple DMCA procedure to remove your content from our site. Start by pressing the button below!
Report copyright / DMCA form

DOWNLOAD PDF

AUROCHS history, morphology and Cis van Vuure ecology of an extinct wild ox

M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10

THE

M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10

RETRACING

M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10

M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10 M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10 M 80 60 40 20 10 Y 80 60 40 20 10 C 80 60 40 20 10 B 80 60 40 20 10

The book is addressed to specialists in mammalogy (interested in the archaeology, ecology, morphology and history of mammals), palaeoecology (because of the herbivore-impact debate concerning the former natural landscape), forest ecology and European nature conservation, to cattle breeders and other cattle lovers.

RETRACING THE AUROCHS

Directly related to the ecology of the aurochs is the ongoing discussion concerning the supposed impact large herbivores render on forest structure, as well as on the appearance of the natural landscapes they live in. Here, too, existing and new surprising data from various fields of science are compiled to elucidate this problem. In addition, due attention is paid to the origin, description and evaluation of the so-called bred-back aurochs (Heck cattle), used in several places for educational and nature conservation purposes. Only now can Heck cattle be properly compared with the original aurochs prototype.

Cis van Vuure

This book tells the comprehensive story of the extinct aurochs (Bos primigenius), the wild ancestor of our domestic cattle, and of what is still left of it. Not only until its extinction in 1627, the aurochs, because of its appearance and nature, had left a deep impression on people, but also afterwards this bovine species, due to scanty information and questionable interpretations, has been subject to discussion and mystification. In spite of a growing amount of individual scientific studies, no exhaustive overview on this subject has ever been made. After many years of research at Wageningen University, the existing information as well as numerous new or unused data from the fields of archaeozoology, history, philology, ecology and palaeoecology have been brought together by the author to build up a complete picture of the physical appearance, the way of life and the environment of the aurochs.

Black Cyan Page 1

CONTENTS


history morphology and ecology of an extinct wild ox

1

Black Cyan Page 2

2


Black Cyan Page 3

CONTENTS

RETRACING THE AUROCHS history, morphology and ecology of an extinct wild ox

by

Cis van Vuure

Sofia–Moscow 2005

3

Black Cyan Page 4

4


RETRACING THE AUROCHS history, morphology and ecology of an extinct wild ox © T. van Vuure Translation: K.H.M. van den Berg

First published 2005 ISBN 954-642-235-5

© PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner.

Pensoft Publishers [email protected] www.pensoft.net

Printed in Bulgaria, June 2005

Black Cyan Page 5

CONTENTS

“YOU

CAN BREED ANYTHING YOU LIKE, AS LONG AS YOU HAVE AT

YOUR DISPOSAL THE SEPARATE HEREDITARY FACTORS FOR THE DESIRED CHARACTERISTICS.”

Erwin Baur (genetics researcher, teacher of Lutz Heck). “WE

HAVE NO WORD IN OUR LANGUAGE THAT MEANS ‘WILDERNESS’,

AS ANYWHERE WE GO IS OUR HOME”.

George Barnaby (Dene Indians tribe (Central Canada), previously hunter-gatherers, 1999). “…AND

THE RESULT WAS ASTONISHING!

IN THE

ANTWERP ZOO,

A

GENUINE AUROCHS!

HE

IS

TOGETHER WITH HIS WIFE AND CHILD, AND HE

HAS THE SWEETEST DISPOSITION. IF YOU CALL HIM, HE WILL COME UP TO YOU AND PATIENTLY ALLOW YOU TO SCRATCH HIS PREHISTORIC HEAD”.

Yvonne Kroonenberg (author, 1995). “A

CLEAR AND COMPLETE LITERARY REFERENCE IS HALF THE BATTLE”.

5

Black Cyan Page 6

6


This page intentionally left blank

Black Cyan Page 7

CONTENTS

Contents Preface

11 Word of thanks 14

1. Introduction

17

2. Sources for aurochs research

25

3. Systematics and evolution 31 3.1. The scientific name 31 3.2. Evolution 32 3.3. Systematics 36

Summary 40 4. Distribution area

41 North Africa 42 Asia 44 Europe 48 4.1. Occurrence inside the outermost boundaries Summary 52 5. Decline and disappearance 53 5.1. Outside Europe 53 5.2. Inside Europe 55 5.2.1. The last living area 64 5.2.2. Causes of extinction 72

Summary 78 6. Language and symbolism 6.1. Etymology 79

79

50

7

Black Cyan Page 8

8


6.2. Toponyms 81 6.3. The aurochs-European bison confusion 6.4. Nature and symbolism 88

83

Summary 97 7. Characteristics of the aurochs 7.1. Size 101

100

Comparison of the Pleistocene and the Holocene aurochs 101 The Holocene aurochs 105 The height of the withers 109 Summary 119 7.2. Horns 120 Size 121 Colour 123 Shape 124 Position 131 Summary 135 7.3. The colour of the fur 135 Pictures and descriptions 137 Genetic aspects of the fur colour 147 Comparison with related bovine species 150 Reconstruction of the fur colour of the aurochs 152 Summary 155 7.4. Other physical characteristics 155 Hooves 155 Udder 158 Fur 160 Hide 164 Summary 164 7.5. Summarising table 165

Black Cyan Page 9

CONTENTS

8. Changes resulting from domestication

166

Summary 174 9. Ecology of the aurochs and other wild cattle 9.1. Habitat 175 9.1.1. The natural landscape of Europe 175

175

Introduction 175 Insects 177 Pollen research 181 Roman writers 192 Frontier forests 198 The last Central-European wilderness 200 Summary 212 9.1.2. Food 213 Reconstruction of the food of the aurochs in its original habitat 231 Summary 231 9.1.3. The habitat of the aurochs 232 Other bovine species 241 Reconstruction of the aurochs habitat 245 Seasonal migration 257 Summary 259 9.2. Predation 259 Summary 264 9.3. Social structure and reproduction 264 Summary 271 9.4. The impact of large herbivores on the forest growth Introduction 271 Areas comparable to Europe 276 Population density of wild cattle 280 The effects of feeding: forest elephants, European bison and others 286

271

9

Black Cyan Page 10

10


Wood Buffalo National Park: A case study 297 Discussion 302 Summary 319 10. The breeding-back experiment of the Heck brothers 10.1. Cattle breeds 323

323

Summary 329 10.2. The breeding-back experiment 330 Summary 344 10.3. Evaluation of the breeding-back experiment 345 Description of Heck cattle 350 Summary 358 11. Final remarks, conclusions and recommendations 11.1. The animal 360 11.2. Effects and landscape 369 11.3. The animal in the landscape 373

360

Summary of conclusions and recommendations 380 Appendix 383 References 387 Register 425

Black Cyan Page 11

PREFACE

PREFACE This research of the aurochs, its appearance, its way of living and habitat began in 1980, when a photograph of a so-called ‘bred-back aurochs’ in the Munich Zoo caught the attention of Cis van Vuure. It showed a head of Heck cattle, and it prompted Cis to conduct an extensive correspondence with breeders of Heck cattle, and to start his own research. In 1982, he came into contact with the Stichting Kritisch Bosbeheer (SKB, see Ch. 1) and its ideas regarding natural processes in forests and the original fauna in those forests. These ideas caused a fresh breeze to blow through Dutch nature management, and even Staatsbosbeheer (the Dutch Forestry Commission) hazarded experiments that diverged from the policy carried out up to that point. One of those experiments was the plan to distribute Heck cattle in the Oostvaardersplassen reserve (Central Netherlands) to keep the landscape in this area open for geese. In the autumn of 1982, a tour of a number of Heck cattle breeding farms in Germany and Austria made clear that the assertion that “Heck cattle are a lot like aurochs” is not necessarily correct. Vague preconceptions and personal preferences have played important parts in the breeding of these animals. The choice for Heck cattle as large grazers, their introduction in the Netherlands in 1983 and the ideas behind this stimulated Cis’s search for the facts behind the extinct aurochs and its alleged effects on the landscape. In 1985, he felt he had gathered sufficient material to provide an adequate description of the appearance and ecology of the aurochs. At the time, attempts to gather funds through the SKB to commit all his information to paper failed. Clearly, the time was not right. In retrospect this was fortunate, since later on, the puzzle turned out to consist of far more pieces, the collection and fitting together of which took more time and more research. In the course of 1995, after several years’ interruption, Cis was once again able to resume his work. Aurochs research still consisted of many fragmented constituent studies, and a comprehensive investigation had not been

11

Black Cyan Page 12

12


realised yet. The lack of this, in combination with new factual material, inspired a second, this time successful, attempt to procure funds. Both the Wetenschapswinkel (Science Shop) of Wageningen University and the Ministry of the Flemish Community realised the importance of this research. Another contribution was made by the World Wildlife Fund. Thus, after years of personal research, Cis could be appointed to the staff of the Sub-department of Nature Conservation of Wageningen University in April 1998. For over a year, with great commitment and enthusiasm, he arranged his data and carried out additional research in this sub-department. We are glad to be able to share in the benefits of all his efforts, made partly at the sacrifice of a lot of personal time. His findings are represented in this book. That this was a very topical issue became clear when a supervisory committee had to be formed. It was not at all difficult to find people willing to participate in this, from all over the country as well as from various disciplines. In the course of this research, this supervisory committee got together quite regularly; without exception, the discussions during those gatherings were lively, inspiring and stimulating. The following people were members of the committee: Ruud Lardinois and Hans van der Lans (Stichting Kritisch Bosbeheer), Beatrijs Van der Aa, Koen De Smet and Daniël Josten (Ministry of the Flemish Community, Nature Division), Han Olff and Sip van Wieren (Tropical Nature Conservation and Vertebrate Ecology Group of Wageningen UR), Peter Bergström (formerly of the Institute for Cattle Breeding Research IVO), Lambert van Es (Groningen Institute for Archaeology), Harm Piek (Vereniging Natuurmonumenten (Association for Nature Reserves)), Cis (T.) van Vuure (researcher) and Marijke Dohmen (Science Shop of Wageningen UR). We would like to thank them for their enthusiasm and valuable involvement. We hope that this book will contribute to a more objective idea about the aurochs, and a clearer image of its near-magical proportions, both physical and with regard to its impact. In addition to this, we hope it

Black Cyan Page 13

PREFACE

will contribute to the discussion about the role of large herbivores in the management of nature reserves. Frank Berendse (Nature Conservation and Plant Ecology Group of Wageningen UR) Herbert Prins (Tropical Nature Conservation and Vertebrate Ecology Group of Wageningen UR) Ruud Lardinois (Stichting Kritisch Bosbeheer) Koen De Smet (Ministry of the Flemish Community, Nature Division) Marijke Dohmen (Science Shop of Wageningen University)

13

Black Cyan Page 14

14


WORD OF THANKS Because my research of the aurochs, its habitat and its offspring took such a long time, and owing to the extent of the material and the time-consuming publication of this book, I asked, and was given, the help of many without whose advice and support this book would not have its present size and shape, if it had appeared at all. I would like to take this opportunity to express my appreciation to all those who contributed to this project. First of all, I would like to thank the Stichting Kritisch Bosbeheer for their support during the 1997 recruitment campaign, which led to the recognition and execution of this study and, eventually, to this English publication. I want to thank the Science Shop of Wageningen University for its role as mediator in the development, funding and supervision of the Aurochs Project. The Science Shop constituted an indispensable link between the Stichting Kritisch Bosbeheer and the University. After Ans Hobbelink had led the project through its initial stages, Marijke Dohmen took over her work. Marijke, thank you for all your help and advice, and for your dynamic leadership of this research project in a field so peculiar to you. I thank Frank Berendse and Herbert Prins for allowing this somewhat divergent type of research to take place within their respective Ecology Groups. I would like to thank Han Olff and Sip van Wieren, who provided the practical supervision of the project for those Ecology Groups, for their gracious cooperation. The unexpected addition of this project to their often-hectic schedules does not seem to have caused too much upheaval. To the supervisory committee (see the Preface), I owe gratitude for their regular and critical assessment of my research, as well as for supplying me with information. I will not soon forget those meetings, which so often proved enervating to me. Without the funding, I would have had to carry out all the research in my spare time; now it could take place at an accelerated pace. Of the sponsors who supported the research, first of all I would like to thank

Black Cyan Page 15

WORD OF THANKS

Wageningen University, both for picking up a call for help and for creating a study framework and part of the funding. The Ministry of the Flemish Community (Nature Division) was the second sponsor, no less important both because of the funding it provided and because it eventually acted as publisher of this book. I also extend my sincere thanks to the third sponsor, the World Wildlife Fund. Aurochs research extends across several languages. I thank all those who helped me translate where my command of languages was deficient, in particular Maria Verhallen for translating the (Latin) letter of Anton Schneeberger, Mr L. Okken for translating the Middle High German in the 14th-century ‘Litauische Wegeberichte’ and Tracie Harris for the correction of my homepage into good English. Karin van den Berg carried out the creation of the English version of this book. I thank her for our pleasant, yet thorough cooperation. To all who contributed, made or had made photographs and drawings, done electronically, digitally or by hand, I am very grateful. As is true of many books, the illustrations are the most tempting part. I also thank all those who provided (scientific) information and tips that added to the reliability and depth of my work. In particular, I would like to thank Frits Laarman (ROB) and John de Vos (Museum Naturalis) for showing their aurochs material, and Martin Kockmeyer for showing material in two Munster museums. To Margret Bunzel-Drüke, I owe thanks not only for the breeding data and the photo she provided, but also for showing and explaining to me her own breeding-back experiment at the Lippe, for which she uses various Mediterranean cattle breeds. I hope this experiment will succeed. Poland has a special position with regard to aurochs research. Not only did the last aurochs live there; its close relative, the European bison, also lived there for a long time, and again now. To Prof. Z. Pucek and his staff at the Mammal Research Institute at Bia³owie¿a, I extend my sincere thanks for their help with the research and the gathering of information in 1972 and 1998. My stay and research in the Bia³owie¿a Forest are unforgettable, and have proved decisive for my vision on the

15

Black Cyan Page 16

16


ecology of large herbivores that live in the forest; Bia³owie¿a na zawsze! I am also grateful to Prof. F. Kobryñczuk in Warsaw, in whose veterinary institute I was allowed to spend a day among the aurochs- and European bison bones. I thank Wiktor Kotowski for his help at Warsaw in obtaining data about the former Jaktorów Forest. I am likewise grateful to Mr K. Heymanowski, who analysed the management of the last population of aurochs at Jaktorów, and with whom I was able to conduct an enlightening correspondence. Spanish fighting cattle had my special interest during this research. For the information I was able to gather about them, I owe a great deal of thanks to Mr Agustín García Sánchez, who obligingly took me round Salamanca province, in the world of the Spanish fighting cattle. I hope he will manage to retain the original Spanish fighting cattle. I thank Francis van Son for our pleasant cooperation when she interpreted for me there, and for the experience of Salamanca, ‘la ciudad del toro ibérico’. In the Basque Country, I saw and experienced the Betizu (wild cattle) in 1985 under the guidance of Jean-Pierre Seiliez, manager of a subpopulation. I thank him for this special experience, which gave me a glimpse of the original aurochs. My appreciation also goes out to the Lending Dept. of the Jan Kopshuis of Wageningen University, which dealt expertly with my neverending flood of literature requests. Literary data constituted the basis for this study; the department’s regular supply of correct information has clearly been indispensable. Roel Cosijn I thank for his ‘book tip’ and his stimulation over the years. Unfortunately, it had to take a little longer, Roel, but it was worth it. Without the help of those at home this would all have taken longer still, which is why I owe a lot of thanks to Wiebe van Vuure, who composed the homepage (which yielded many contacts and information), showed me the ropes on the computer and solved many related problems. I also thank Bonnie van Vuure for her many computer tips, and because she let me use it every now and then. And last but not least, I would like to thank Marijke Neijzing, without whom I would not have been able to do any aurochs research whatsoever.

Black Cyan Page 17

INTRODUCTION

1. INTRODUCTION The initiators Stichting Kritisch Bosbeheer This foundation (SKB) has originated from the Landelijke Werkgroep Kritisch Bosbeheer (LWKB), which was founded in 1977. The immediate reason for the foundation of said LWKB were the ideas about forest management inside Dutch nature conservation circles, which were outdated in its opinion. The principal objective of the SKB is: ‘To drastically push back the high rate of unnaturalness in a large part of the Dutch forests, and to promote their indigenous character, eventually omitting human interference as much as possible’. Particularly in the starting period of the LWKB and SKB, such ideas really flew in the face of the prevailing nature management, which is based on human interference, such as the production of timber and the maintenance of unnatural types of landscape. The SKB tries to propagate its ideas by giving lectures and organising excursions, by issuing publications and by conferring with the managers of nature reserves. In this way, the SKB aims to let nature in the Netherlands become more self-regulating, and to diminish man’s influence on these ecosystems. Eventually, man will no longer be manager, but spectator of natural processes. Interference through timber harvesting, water management and hunting, which currently disturb the natural processes, will no longer take place, and nature will determine which plant- and animal species occur in what densities (Van der Lans en Poortinga 1986). Already since the 70s, a discussion has been going on around this issue of more self-regulating ecosystems (LWKB 1980). First, as a result of a number of storms, the main topic was a more natural way of forest management; later, the introduction of large herbivores such as horses and cattle was added. The distribution of Scottish Highland cattle, and

17

Black Cyan Page 18

18


especially the arrival of Heck cattle in the Netherlands, caused great commotion in the press. Nature management institutions carefully began to focus on a different kind of nature management, and experiments were started. Although many things have improved in the meantime, innovation still proceeds. Too slowly, however, in the opinion of the SKB, partly because traditionally developed management insights are not easily replaced by a fundamentally different vision. Another reason for the slowness of this process is that in fact, too little is as yet known about the appearance of the natural landscape of the Netherlands and Europe, and about the parts played by herbivores and carnivores. To gain greater clarity about this natural landscape, and the role of large herbivores in particular, the SKB wanted to instigate research into the life of the aurochs in Europe’s primeval nature. Only with the help of information gained by such a study may decisive answers be found about several aspects of the current functioning of feral cattle in Dutch nature reserves, including their influence on the vegetation, choices of food, densities and manageability. The correlation between the aurochs and its habitat also affected the appearance of the animal, since this is determined in part by such factors as climate, food and predators. The SKB wanted to find out which existing domestic cattle breeds were still like the original aurochs as regards their physical characteristics. Additional research may reveal what can be done to regain an animal that is adapted to the circumstances in a self-regulating ecosystem, both physiologically and physically. Such research may yield information that can lead to further or renewed breeding back at a later stage.

Ministry of the Flemish Community, Nature Division The Nature Division gives advice with regard to the conservation and development of nature. It manages the nature reserves of the Flemish District. In addition to this, the Nature Division ensures financial support for the purchases and management of nature reserves by accredited associations.

Black Cyan Page 19

INTRODUCTION

The Nature Division takes care of the preparation and execution of the Flemish policy on nature; in this way, it contributes to the development of the Vlaams Ecologisch Netwerk (VEN, the Flemish Ecological Network), Ecologische Impulsgebieden (Ecological Impulse Areas), Regionale Landschappen (Regional Landscapes) and Natuureducatieve Centra (Nature Education Centres). The Nature Division plays a coordinating part in the application of international guidelines regarding the protection of species and habitats, and with respect to biological diversity. The slogan of the Nature Division is: ‘Nature, we take care of it’.

Background of and reason for the research Ever since the early 70s of the 20th century, cattle and horses have been used in the Dutch management of nature reserves. Originally, they were intended as means to limit the increased cost of management; gradually, however, and partly because of the ideas of the SKB, these animals were considered increasingly as ecological replacements for the wild cattle and horses that had disappeared. Other large mammals, such as elk and wolves, have also disappeared from our country. Research in reference areas such as North America and Eastern Europe has shown that large mammals may fulfil important functions in nature. The absence of such animals may lead to a reduction of natural values, which can only partially be replaced through management regulations instigated by man. From the early 80s onwards, cattle that were distributed as derivatives of the extinct aurochs have increasingly often been breeds that can withstand bad weather conditions. If possible, year-round grazing (without supplementary feeding) is aimed for, as well as life in natural herds. The aurochs that was indigenous to Europe used to be hunted intensively by man. In the Netherlands, it probably disappeared in the beginning of the Middle Ages, and the species definitely became extinct in 1627, when the last animal died in Poland (£ukaszewicz 1952). What remains of this animal species are bones, horns, pictures and descriptions. In addition to this, the genetic inheritance has survived in cattle breeds closely related to the aurochs (French 1966).

19

Black Cyan Page 20

20


From 1983, the Dutch Forestry Commission has used Heck cattle in a few large nature reserves. These animals are the result of an attempt by the German Heck brothers, carried out in the 20s and 30s of the 20th century, to crossbreed certain cattle breeds in order to ‘breed back’ the aurochs (Heck, H/L 1934). In some aspects, Heck cattle are similar to the original aurochs. The ideas of the SKB caused quite a lot of turmoil at the time, and the introduction of Heck cattle evoked similar reactions. A social discussion ensued, and is still going on, about the use and necessity of deploying this specific cattle breed. The idea behind this introduction, i.e. the creation of circumstances comparable to those in primeval nature by year-round grazing, was not readily accepted by the more traditional conservationists and breeders. The latter emphasised the impossibility of reintroducing domestic cattle in the wild and the danger of transferring cattle diseases; as it happens, the Heck cattle in the Oostvaardersplassen are not inoculated. The discussion about the use of cattle for nature management was recently given a new impulse through research carried out by Vera (Vera 1997, 2000), who studied the possible shaping influence of large herbivores on the vegetation, and the natural landscape that may result from such an influence. The deployment of Heck cattle in the 80s was accompanied by all kinds of scientific research into the social behaviour and choice of food (Cornelissen and Vulink 1995) of these animals, but was never preceded or followed by any investigation of the life and functioning of the wild ancestor of our domestic cattle, the aurochs. It might have been logical to check the life of the cattle in the large Dutch nature reserves against the conditions in which the aurochs lived in its time. The problem surrounding the ecology of cattle is more alive in the Netherlands than in other countries of Europe. It so happens that in the Netherlands, cattle have been distributed in a feral state, also in large nature reserves. In other countries, Heck cattle may be found only in zoos and small wildlife reserves. In this regard, the Netherlands acts as a pioneer. Together with the growing publicity regarding the Heck cattle in Europe, and partly as a result of the changing nature management, criticism on this cattle breed has also increased. From the moment of their devel-

Black Cyan Page 21

INTRODUCTION

opment, there has always been criticism on the alleged similarity of Heck cattle to the original aurochs (Herre 1953, among others). At the Aurochs Conference at Mettmann (1997), this criticism was once again extensively brought forward. Clearly, very different ideas exist about the characteristics with regard to which Heck cattle differ from aurochs. As a result of this, in some nature reserves culling takes place on the basis of selections made without proper know-how, in favour of a animal that, with regard to colour and horn shape, does not resemble the aurochs. On a limited scale, by crossbreeding Heck cattle with other breeds, the further development of this cattle breed has already been initiated in Germany. The great public interest for the offspring of the aurochs and their use in nature reserves outlined in the above, as well as the need for information and fragmented availability of that information, which prevents a clear overview, have provoked this research.

Definition of the problem Aim The distribution of cattle (Scottish Highland- and Heck cattle) in Dutch nature reserves has been, and is still being, monitored through investigations regarding the behaviour and choice of food of these animals. All the areas in which such cattle are distributed and in which such research takes place are environments interfered with and influenced by man. Important natural aspects like predation and a food supply determined by nature are completely or partly missing. In spite of the seemingly natural character of these areas, man is still dominantly present in the supplementary feeding and (limited) culling, and determines the size of the living area. As this unnatural situation continues, issues that come increasingly to the foreground are: What would happen if man really stopped interfering in every way? How would the landscape develop, what would the density of cattle be, how would their appearance and their choice of food change, and to what extent do landscape and density determine the animals’ behaviour and manageability?

21

Black Cyan Page 22

22


The aim of this research is to clarify, on the basis of information about the aurochs (appearance, habitat, way of living), combined with data obtained through modern research (food- and behavioural research of cattle, pollen research), what kind of animal the aurochs was and how it lived, so as to answer as adequately as possible the questions that arise in the current management of cattle in large nature reserves, such as: 1. Ecologically speaking, do the present cattle function in the same way as the earlier aurochs, for example with regard to their choice of food, behaviour, and effect on the vegetation? 2. What will the natural landscape that is aimed for look like, eventually? 3. What will the density of the cattle in such a landscape be? 4. Which cattle breed is best suited to year-round grazing in extreme circumstances? 5. Which selection criteria should apply if a Heck cattle population needs to be interfered with (physical (aurochs) characteristics, health and the like)? Answering these questions will enable area managers and breeders of Heck- and other cattle, in the Netherlands and elsewhere in Europe, to conduct a better-founded management if they aim to realise a certain type of landscape with a specific breed. Information about the aurochs is very fragmented, divided over various disciplines. Combining it to a coherent whole may provide a good overview of two centuries of research.

Questions An answer will be sought for the following questions: Main questions: 1. What did the aurochs look like? 2. What was the aurochs’ way of living? Related questions: 1. What did the natural landscape of Europe look like?

Black Cyan Page 23

INTRODUCTION

2. What was the influence of large herbivores (including the aurochs) on the vegetation? 3. In what way, with the help of information about the aurochs, can more clarity be obtained with regard to issues related to the management of cattle in current Dutch nature reserves? These issues may include the population density, choice of food, behaviour, landscape to be aimed for, and breeds to be used. 4. To what extent do the current cattle used for nature management resemble the aurochs, physiologically as well as physically, what are their relations, what is the genetic composition of Heck cattle and how does this express itself; which information can be supplied to facilitate further or renewed breeding-back at a later stage?

Method of research The method of research, analogous to the questions, will focus on both the appearance and the way of living of the aurochs. The information that will be brought together derives from various disciplines, such as pollen research, archaeology, animal ecology and linguistics. For the appearance of the aurochs, 16th-century written sources (in Wrzeœniowski 1878, among others) as well as archaeo-zoological research of the mortal remains of the animal (La Baume 1944, among others) have been consulted, added to by data obtained in modern genetic research (including Minkema and De Rooy 1982), from the study of primitive cattle breeds and related bovine species (e.g. the banteng) and from caves research (e.g. at Lascaux). To gain an insight in the way of living of the aurochs, both 16thcentury written sources and data from ecological research of cattle (and other mammals) in the Netherlands and abroad (Poland and North America, among others) (including Cornelissen and Vulink 1995). Since the original natural landscape of Europe played a crucial role in the life of the aurochs, and because the appearance of this landscape is the topic of a lot of discussion in the Netherlands at the moment, a great deal of prominence will be given to it here. The data for the description

23

Black Cyan Page 24

24


of this landscape stem from pollen research, insect research, texts by Roman writers (including Caesar circa 55 BC) and descriptions of the ‘last wilderness of Central Europe’, in the former East Prussia. Wherever possible, the original versions of such sources have been used. Thus, this study is based mainly on a literature search. In addition to this, it is based on information obtained during visits of Heck cattle breeders and the populations of cattle managed by them in the Netherlands, Belgium, Germany and Austria, and breeders of Spanish fighting cattle in the Spanish province of Salamanca. Beside this, data were gathered during visits of museums and research institutes in the Netherlands, Belgium, Germany and Poland. Additional visits were paid to nature reserves, to observe wild and feral cattle, among other places in the Oostvaardersplassen, the Bia³owie¿a Forest (Poland) and the western Pyrenees of France. In Poland, moreover, the Forest of Jaktorów was visited: the living area of the last aurochs.

Black Cyan Page 25

SOURCES FOR AUROCHS RESEARCH

2. SOURCES FOR AUROCHS RESEARCH Research into an extinct animal species like the aurochs is not restricted to the bounds of biology, but extends into other fields of science. Besides biology, which includes important aspects like the ecology of plants and mammals, mammal morphology, entomology and palynology (pollen research), sciences like archaeozoology, palaeontology and historical ecology are very useful to such research. Of these three disciplines, the first two are needed when it comes to judging old bone material, while the latter is indispensable for tracing the how and why of the changes that have taken place in the occurrence of animal species and landscapes in the course of history. In addition to these, the related sciences of archaeology and history are also important: they generally focus on the origin of and changes in human cultures, which may have had their influence on animal ecology. Apart from these directly relevant fields of science, philology is important, especially where the ‘old’ languages are concerned that were used to describe the aurochs when it still existed; specifically to explain old names and texts, mainly from Latin, but also from the mediaeval Germanic and Slav languages. None of the fields of science mentioned here is able separately to provide decisive information regarding the physical appearance and ecology of the aurochs, since the evidence is scarce. In addition to this, the information is scattered over the above-mentioned fields of science; to obtain the maximum number of jigsaw puzzle pieces, therefore, research has to be undertaken in a variety of sciences. What is generally known in one science is often unknown in another, and combining the various facts may lead to unexpected insights. Data concerning the physical appearance of the aurochs may be obtained from the study of body parts that have survived (bones, hairs, horns), as well as from pictures and descriptions of the animal when it was still living. An understanding of its ecology may be obtained through historical ecological research (descriptions of the animal’s way of life

25

Black Cyan Page 26

26


and of its environment) and through modern ecological research into wild cattle species that still exist and into vegetative processes. Modern research has the advantage of usually being verifiable, which is not the case with ancient descriptions and pictures. Of the latter, it is uncertain whether their authors or artists actually saw the animal, relied on data supplied by others, or invented or copied information. Of the Roman writers Seneca, Vergilius and Plinius, for example, who summarily described the aurochs, probably only Plinius actually saw the animal, since he served as an army officer in Germania. Even this last fact does not really constitute proof, however. Julius Caesar, who gave the most extensive description of the period, probably did not see the animal himself, although he certainly saw its horns. Mention and descriptions of wild cattle, both aurochs and European bison, were also made during the Middle Ages (400 – 1500). As the aurochs declined in number, however, it became increasingly difficult to differentiate between these two bovine species, as is evident from the descriptions of the period. In the late Middle Ages and the 16th century there was a disconcerting confusion between these animals (see Chapter 6.3), both as regards their appellations and the inadequate descriptions given of them, which has made it increasingly to assess the reliability of these data. Little reliable information survives from the Middle Ages; more valuable data stem from the 16th century. The latter include, first of all, Inspection Reports1 concerning the organization of the management of the last aurochs population in the Forest of Jaktorów (Poland). These reports were rediscovered in the course of the 19th century, when people started to realize their importance. Moreover, data from the 16th century have been handed down by three informants: Bonar, Von Herberstein and Schneeberger, in order of increasing importance. The letters of Schneeberger and Bonar were published in Gesner (1602 and subsequent editions) (see Appendix). Wrzeœniowski (1878) offers the following information about the backgrounds of these three authors: a. Johann Bonar. Information is limited. He was involved in hunting and wrote about the elk, among other things. It is not quite certain 1 These reports record the management of the forests near Jaktorów (Central Poland) and the aurochs population living there (see Chapter 5.2.1). They were published in K¸edzierska (1959 and 1965), and are currently being preserved in the Main Archive of Old Documents (Archiwum G§lówne Akt Dawnych) in Warsaw.

Black Cyan Page 27


whether he saw the last aurochs population at Jaktorów. If he did not, he must have had a very good informant. b. Sigismund von Herberstein. This German diplomat travelled to Poland and Russia between 1516 and 1553. He is unlikely to have seen any live aurochs, but he definitely saw the skin and the horns of an aurochs bull, which he received as presents. These gifts date from beyond 1548. He did see European bison alive. c. Anton Schneeberger. A student of Conrad Gesner, with whom he kept in regular contact. Studied medicine and travelled in many forest and mountain areas of Poland. He worked at recording folk medicine and was known for his conscientious observations. His description of the aurochs population at Jaktorów reveals that he was an eyewitness, or at least spoke extensively with the local managers of the population. The date of his letter to Gesner, which contains this description, is unknown. He may have visited the area in 1559, during a trip from Cracow to Königsberg (Kaliningrad) and Wilno (Vilnius), which started in April of that year (Hryniewiecki 1938). The description of the last aurochs population in his letter to Gesner is inserted as an Appendix to this book. Inquiries at the Zentralbibliothek Zürich and the Staatsarchiv of the Kanton Zürich, to find out whether the letter still exists and contains more data, were unsuccessful. For a long time after the death of the last aurochs in 1627 in Poland, rumours circulated in Europe concerning aurochs still living in Eastern Europe. In 1634, a book by Œwiêcicki was published posthumously by his son, which contains some information about the aurochs. Later editions of Conrad Gesner’s book, published in the course of the 17th century, still mention the descriptions of Schneeberger and Bonar without rectification. The curiously mixed aurochs/bison description given by Von Fleming in 1719 may well be the result of this confusing situation. During his military and hunting service, in Poland among other places, the latter author studied the plant and animal life of Europe. Linnaeus, in the 18th century, must also have read or heard rumours that the aurochs was still alive in Poland (see Chapter 3.1). Oral tradition, gratuitous copying, ignorance and poor communications caused lot of obscurity concerning the aurochs. Eventually, partly by means of the In-

27

Black Cyan Page 28

28


spection Reports, the earlier existence of the aurochs during the Holocene2 was definitively determined and generally accepted. Research into bone finds of the aurochs started in the late 18th/early 19th century. From the turn of the century onwards, these studies were characterised by three milestones: 1827 – the description of an aurochs skeleton led to the conclusion that the aurochs had existed as a separate species (Bojanus 1827). 1878 – analyses of old written accounts showed that the aurochs had lived in Europe during the Holocene together with the European bison (Wrzeœniowski 1878). 1927 – analyses of extensive bone material proved that the aurochs was the sole wild ancestor of present-day cattle and that there was a considerable difference in size between the aurochs bull and the aurochs cow (Von Leithner 1927). During the 19th century, the literature concerning the aurochs was mainly in German, even when the research was executed by non-Germans like Pusch and Wrzesniowski. Also in the 20th century, German was the language in which most items concerning this subject were published. In the late 19th/early 20th century and again, mainly in Germany, researchers such as Nehring, Hilzheimer, Adametz and Duerst tried to present an image of the aurochs based on bone material and historical sources. Szalay’s name should be mentioned in this connection, since his extensive research in the field of nomenclature of animals in old written accounts has contributed much to the acquisition of a reliable picture of the aurochs, and to clarification with regard to the confusion concerning the names of the aurochs and European bison. In his time, Szalay must have had extensive literary sources at his disposal, from which he summarily cited his references. His promise to one day publish these 12,000 literature references under the title ‘Die Weltliteratur des Wisents und Ures’ was unfortunately never redeemed (Vörös 1985). It is not known whether the file still exists, or if it does, where it is; nonetheless, Szalay’s works are of great value. After Von Leithner’s research in 1927, gradually more detailed bone research was accomplished. Through the research of Bohlken (Germa2 The Holocene is the period after the last ice age, and started about 10,000 years ago (see Fig. 30). The Pleistocene began about two million years ago, and lasted until the Holocene

Black Cyan Page 29


ny), Degerbøl (Denmark), Lasota-Moskalewska (Poland), Grigson (England), Bökönyi (Hungary) and many others, more became known about the form and size of aurochs bones. These studies continue today, not only because new bone material is still being found but also because new techniques are being developed to research them. In present-day Japan, for instance, Polish aurochs bone material is the subject of DNA research. Compared to the research of bone material, which has been going for two centuries, studies into the ecology of the aurochs are of relatively recent date and only few have as yet been done. This is partly because relevant data are scarce, partly because interest regarding the functioning of bovine animals in a natural setting and their possible impact on the vegetation has only developed in Europe in the past 20 years. Since historical data about the ecology of the aurochs are almost completely unavailable, it is worth looking at the ecology of wild cattle species still in existence. From their manner of life and choice of food, conclusions may probably be drawn regarding those of the aurochs. Such data are available for a number of related bovine species, although they are far from complete for many others. In Germany during the 1920s and 1930s, the Heck Brothers carried out experiments to recreate the aurochs (see Chapter 10.2). At the time, knowledge about the aurochs was less extensive than it is today, but sufficed to form a fair image of certain details of its physical appearance. The reasoning and arguments of the brothers are nevertheless characterized by an almost complete lack of references to the sources of their knowledge. Heinz Heck did not even provide a single reference to his sources, while Lutz Heck made only very brief mention of them. Once or twice he gives a nearly complete literature reference; generally, however, he is very vague, making the reader guess the basis of his statements. The Heck brothers were able to acquire their knowledge of cattle types and bovine species in the Berlin Zoo, where their father had built up a large collection of these animals. As to aurochs remains, Lutz Heck mentioned that ‘one’ could see many bones, a number of horns and even a skeleton of this animal. Whether and, if so, how extensively he himself saw and researched these has never been clearly shown. He knew the aurochs descriptions of Von

29

Black Cyan Page 30

30


Herberstein and Schneeberger and gives a good translation of that of the former, but an incomplete and, above all, faulty rendering of Schneeberger’s description. In Chapters 10.2 and 10.3, the inaccuracies in the arguments of the Heck brothers concerning the physical appearance of the aurochs will be discussed in more detail. For any religious and cultural-historical aspects of the relationship between man and bovine animals, few of which will be mentioned in this book, the reader is referred to the work of Von Lengerken (1955). He collected much source- and picture material concerning these aspects.

Black Cyan Page 31

SYSTEMATICS AND EVOLUTION

3. SYSTEMATICS AND EVOLUTION 3.1. The scientific name When Linnaeus classified the animal kingdom in the 18th century, he gave the European domestic cow the scientific name ‘Bos taurus’ (Linnaeus 1758, page 71). That he realized that the wild ancestor of this cow had once lived in Europe, and was possibly still alive, is evident from the fact that he ranked the ‘urus’ 3, which Julius Caesar (58 – 52 B.C.) described in Germania, under this same species name. Moreover, Linnaeus added: ‘The wild aurochs is living in the grassy lowlands of Poland’. He had obviously read, or learned in some other way, that the animal was still living there. In the course of the 17th and into the 18th century, confusion about this alleged continuing existence of the aurochs prevailed. In fact, the last aurochs had died in 1627. Linnaeus considered the domestic cow and the urus to be related to such an extent that he regarded them as one species. The memory of this wild bovine animal had become almost completely blurred by Linnaeus’ time. The discussion about the possible former existence of more than one wild cattle species in Europe began around that time (see Chapter 6.3), and even those who regarded the aurochs as a separate species did not rule out the possibility that the animal still lived somewhere in 18thcentury Europe (Daszkiewicz 1997). At the end of the 18th and the beginning of the 19th century there were two parties to the discussion. One group thought that by nature, there had only ever been a single wild bovine species in Europe, the European bison, while the other opined that there had been two: the European bison and the aurochs (Wrzeœniowski 1878). Bojanus belonged to the first. Around the beginning of the 19th century, many aurochs bones had already been dug up and there was also a nearly complete skeleton, mounted in a museum at Jena (Germany). On the basis of this skeleton, Bojanus described a new animal species, which he named ‘Bos primigenius’ (Bojanus 1827). He did not realize that 3

urus - aurochs (see Chapter 6.1).

31

Black Cyan Page 32

32


he had described the skeleton of the historic aurochs, which had lived beside the European bison in Europe in historical times. Since he thought that this bovine species had already become extinct long before, in prehistoric times, he added the word ‘antediluvialis’, or ‘antediluvian’ to the species name. In addition to this, Bojanus did not know that Linnaeus had already ranked the historical aurochs under the species name Bos taurus, and he thought that Linnaeus indicated the European bison with the word ‘urus’. The name ‘Bos primigenius’ remained in use after Bojanus’ description, to indicate both the Pleistocene and the Holocene aurochs. In recent times, however, this was temporarily changed, since the rule of priority, in accordance with which species names given later lapse in favour of names that already exist, prevailed when a list of mammals was published in 1993. In this list, based on the ‘Code of the International Commission on Zoological Nomenclature’ (Wilson & Reeder 1993), the name Bos primigenius, given by Bojanus, was changed into the name Bos taurus, given by Linnaeus, since the aurochs and the domestic cow (which originated from it) may be considered one and the same species. This decision to change the name has led to criticism, however (Gentry et al. 1998). The authors in question wanted to make an exception to the rule of priority for domesticated animals. Because of the criticism, this changing of the name Bos primigenius into Bos taurus was reversed and the original name Bos primigenius reestablished in 2003 (International Commission on Zoological Nomenclature 2003, Gentry et al. 2004). Nowadays the name Bos taurus is reserved for the domestic derivative.

3.2. Evolution The conditions that caused the extinction of the dinosaur about 65 million years ago also brought about extensive changes in the vegetable kingdom. The gymnosperms (mostly conifers and ferns), which had been predominant up to that point, were replaced more and more by angiosperms, which include grasses, herbs and broad-leaved trees (see Fig.1).

Black Cyan Page 33


Dicotyledonous plants (herbs, broad-leaved trees) dominated at first, and during the Palaeocene, the monocotyledons appeared, but without any grasses. The formerly dominant conifers and ferns became extinct or receded to dry, acid or cold areas. During the Miocene, as a result of the cooling down and drying up of the global climate, dry areas developed beside the already existing forests. These were sparsely overgrown with trees, which offered many opportunities for the initial appearance and expansion of grasses. Along with these changes in the vegetable kingdom, those in the animal kingdom were characterised by the emergence of the mammals, which were going to play dominant roles. The dominant

Fig. 1. Overview of the origin and development of the angiosperms and the various groups of ungulates from the time at which the dinosaurs became extinct, at the end of the Cretaceous period, onwards (after Van Soest 1982, revised by D.P. Bosscha Erdbrink). A – represents the ancestral artiodactyl common to both pigs and ruminants. B – represents the common ancestor to the artiodactyla and perissodactyla.

33

Black Cyan Page 34

34


appearance of wholly new plant orders and families evoked the separation of herbivorous mammal species that were going to specialise in the digestion of specific plants and their components (Webb 1977, Van Soest 1982). As a result of this, perissodactyls such as horses, rhinos and tapirs began to appear during the Eocene. They would play dominant roles during the Oligocene and the Miocene, in as far as their numbers of species and living areas were concerned. Later on their shares would diminish, as a result of the rise of competitive animal groups like elephants, deer, antelopes, bovids and camels. The first representatives of the animal group relevant to this discussion, the ruminants (which include the aurochs), appeared in the tropical forests some 40 million years ago. The emergence and subsequent separation into species of these ruminants followed the evolution of the angiosperms, and the ruminants, on account of their competitive ability, are likely to have occupied an increasingly prominent position among the ungulates. Already during the Miocene, they would go and take over the perissodactyls’ dominant role. The reason that ruminants were going to form a dominating group, rich in species, should probably be sought in their special digestive system. The digestion of food with the aid of several stomachs, combined with rumination, enables a ruminant to break down and utilize the many appetite-reducing and hard-to-digest components of grasses, herbs and broad-leaved trees. Although the ruminants, which at first lived mainly in the forests, were exclusively browsers4, the appearance and global expansion of savannas and grasslands, about 25 million years ago, resulted in the emergence of grazers4, ruminants that are geared to eating grass. In the beginning, the first species of grass-eating ungulates were not quite adapted to grazing (Webb 1977). Their teeth were not nearly as hypsodont (see Chapter 9.1.2) as those of the later, more specialized species. The specialized grazing ruminants did not develop until the end of the Miocene, and they had the hypsodont teeth pre-eminently suited to digest grass growing in dry, open areas. Their voluminous system of stomachs allows such grazers to 4

See Ch. 9.1.2 for an explanation of the words ‘browser’ and ‘grazer’.

Black Cyan Page 35


adequately digest the cellulose components of grasses5. It is not the grazers themselves that produce chemicals to break down cellulose, but the specific micro-organisms that live in symbiosis with the grazers. In non-ruminating grazers like horses, which possess only one stomach, the main part of digestion takes place in the relatively large blind gut (caecum), also with the help of micro-organisms. About 2 million years ago, a period in which extreme cooling down and warming up of the global climate alternated began. During this period, the Pleistocene, extensive dry grasslands appeared in many areas, especially during the coldest periods, the ice ages. During the warmer periods in between, their area was much diminished. These grasslands, which could be found mainly in the northern hemisphere, were also called ‘cold steppes’ or ‘mammoth steppes’. They became known especially by the presence of mammoth, steppe bison, giant deer and woolly rhinoceros. The special conditions just before and during the Pleistocene resulted in the emergence of many new species of herbivores, including the grazers of the genus Bos. The earliest representative of the genus Bos is Bos acutifrons Lydekker 1898. Remains of this species, which were more than 2 million years old, were first dug up in the Siwalik hills in Northern India (see Fig. 2). All later species of the genus Bos allegedly stem from this early species, which inhabited Europe, Asia and North Africa during the Pleistocene (Bohlken 1958, Thenius 1980). The horns of Bos acutifrons were

Fig. 2. Remainder of a skull with horn cores, of over 2 million years old, of Bos acutifrons from the Siwalik hills (North India) (after Lydekker 1898). Bos acutifrons is considered the direct ancestor of the aurochs. The missing parts of the horn cores have been drawn in.

5 This system of stomachs consists of the real stomach (rennet stomach) and the three fore stomachs rumen, reticulum and omasum. The rumen is the largest and harbours an extensive flora and fauna of micro-organisms.

35

Black Cyan Page 36

36


very long; the horn core could measure over 2 m (Grigson 1973). The section of the horn core was oval. Due to a lack of bone remains, little else is known about the size and the anatomy of this animal. Bos acutifrons still lived in India until the middle of the Pleistocene. The aurochs is likely to have developed from this species between 1.5 and 2 million years ago. In the course of the Pleistocene, aurochs spread from India to, for instance, Europe, where they were first identified in the Tiber delta (Cerilli & Petronio 1991). By its southern route, the aurochs arrived in Southern Europe much earlier than in Central Europe, which it probably reached by a route through Russia. It appeared in Spain 700,000 years ago (Estévez & Saña 1999). Its earliest presence in Germany was demonstrated on the basis of a skull from Steinheim an der Murr (Württemberg) (Lehmann 1949), which dates to about 275,000 years ago, an exceedingly warm period between the Elster and Saale ice ages6.

3.3. Systematics Bos primigenius belongs to the order of the even-toed ungulates (Artiodactyla) and within this order to the family Bovidae, the subfamily Bovinae and the Bovini tribe. The (originally wild) bovine species, mentioned in Table 1, are ranked under the Bovini. Their nomenclature follows the classification of Wilson & Reeder (1993), which is based on the names prescribed by the International Commission on Zoological Nomenclature. Roughly three subspecies of Bos primigenius are distinguished: ‘primigenius Bojanus 1827’ (Europe and the Middle East), ‘namadicus Falconer 1859’ (India) and ‘mauretanicus Thomas 1881’ (North Africa). As to the specifications ‘primigenius’ and ‘mauretanicus’, these indications roughly refer more to distribution areas, rather than any great differences between these subspecies. Such differences may have been present, but this has not been unambiguously shown on the basis of bone material as yet (Gautier 1988). The deviant namadicus subspecies 6 Today, the classical division of the Pleistocene into four cold ice ages (glacials) and three (relatively) warmer interglacials has been replaced by a division into a larger number of large ice ages, alternated by smaller ice ages and warmer periods (Kahlke 1994, Von Koenigswald 1999).

Black Cyan Page 37


Table 1. List of all the bovine species of the Bovini tribe and their distribution areas. Scientific names according to Wilson & Reeder (1993) and the International Commission on Zoological Nomenclature (2003).

Scientific name

English name

Distribution area

Bison bison Bison bonasus Bos primigenius †7 Bos gaurus Bos javanicus Bos sauveli Bos arnee Poephagus mutus Syncerus caffer Bubalus depressicornis Bubalus quarlesi Bubalus mephistopheles †7 Bubalus mindorensis

Bison European bison Aurochs Gaur Banteng Kouprey Water buffalo Yak African buffalo Lowland anoa Mountain anoa Chinese water buffalo Tamarau

United States, Canada Europe, West Russia Europe, Asia, North Africa India, IndoChina, South China IndoChina, Indonesia East IndoChina India, IndoChina Central Asia Central and South Africa Sulawesi Sulawesi Central China Mindoro

was described by Grigson (1973). The size of its horn cores and the nature of some of its skull features clearly distinguish namadicus from the other subspecies. Up to this point, however, only Pleistocene material of namadicus has been found. Conclusive remains of the Holocene namadicus, probable ancestor of the zebu, is still lacking. With the help of mitochondrial DNA research, scientists have been able to show that namadicus and primigenius probably separated in India between 610,000 and 850,000 years ago (Mac Hugh et al. 1997). In spite of many studies, the Latin nomenclature for, and the relationships within the Bovini continue to be subjects for discussion that still evoke many questions. It is still debatable, for instance, whether Bos and Bison should be regarded as separate genera or in fact belong to a single genus. The yak, for example, could either be classified under a separate genus (Poephagus), or remain under Bos. Neither is it clear yet whether the kouprey is a separate original species or a cultural type.

7

† - extinct.

37

Black Cyan Page 38

38


These questions are closely related to the evolutionary developments that took place within the Bovini tribe during the past 3 million years. Which species emerged when and from which ancestors, and who is most related to whom are subjects for research. Despite morphological studies of skulls (Bohlken 1958, Groves 1981), haemoglobin research (Schreiber & Göltenboth 1990) and research into the composition of a certain enzyme produced in the mitochondria (Janecek et. al. 1996) no clear conclusions, consonant in every aspect, have been reached about the phylogeny of bovine species. The various studies contradict each other; the placing of Syncerus and Bubalus is unambiguous, but the places of the other members of the Bovini tribe vary. The cause of this lack of clarity may well lie in the fact that the origins of the Bovini, and the specialisation within the tribe, are of

Photo 1. European bison (Bison bonasus). At one time, this bovine species was extinct in the wild; nowadays, however, it lives in a wild state once again in several places (Photo: Z.A. Krasiński).

Black Cyan Page 39


rather recent date (the past 2 – 3 million years) (Buntjer 1997). As evolution becomes a longer process, species will grow further apart and the genetic differences between them will become more obvious. In terms of evolution, 2 –3 million years are a relatively short period, for which not every genetic research method is able to clarify differences at the species and genus level. However, with the aid of nuclear DNA research such as that carried out by Buntjer (1997) and Buntjer et al. (2002), in which the genetical distances between the various species could be established, such clarification may be established (simplified rendered in Fig. 3).

Fig. 3. Most likely relationships between the species and genera inside the Bovini tribe (after Buntjer 1997 and Buntjer et al. 2002).

The results of his research roughly agree with the morphological research that Groves (1981) carried out on skulls of bovine species. Figure 3 shows that the Bubalus/Syncerus group separated itself from the other bovine species at an early stage. The remaining animals fall into two groups: bison/European bison/yak and banteng/gaur/aurochs. Besides a genetic separation, a division in climate areas seems to have taken place. The animals of the first group inhabit temperate and cold zones and live most northerly, those of the second group inhabit temperate and tropical zones and live farthest to the south. As regards its adaptation to climate, the aurochs inhabited the overlap area of these

39

Black Cyan Page 40

40


two groups; it mainly lived in temperate and subtropical zones (southerly till South India) and was less adapted to extreme cold than the yak and Bison species.

Summary In 1758, Linnaeus gave the scientific name Bos taurus to the domestic derivative of the aurochs; the latter one got the name Bos primigenius from Bojanus in 1827. The aurochs probably emerged as a species between 1.5 and 2 million years ago in India, in a period in which the global grass area expanded greatly. From India, the aurochs spread to other parts of Asia, Europe and Africa. It inhabited mainly temperate and subtropical zones. Though the relationships between the different species of the Bovini tribe have not been explained completely as yet, the aurochs is probably most related to the gaur and the banteng (see Fig. 3). Table 1 provides an overview of the species that belong to the Bovini.

Black Cyan Page 41

DISTRIBUTION AREA

4. DISTRIBUTION AREA The total distribution area inhabited by the aurochs at one time is very extensive, including nearly all of Europe and large parts of Asia and North Africa. The period of the animal’s existence occupied one and a half million years. As its existence is further in the past, and less digging for an animal’s remains has taken place in a specific country, the data concerning its occurrence, and the period in which it lived, are bound to be less exact. A larger number of more accurate data are known from Europe than from other continents, therefore, and there is more information from the Holocene than from the Pleistocene. Relevant to the existence of the aurochs, the Pleistocene period occupied roughly three large ice ages and the in between periods, with diverging climates and vegetations.

Fig. 4. The former distribution area of the aurochs, as based on data of many authors and the author’s own interpretations (see text). – – – – – The outer most border of the distribution area during the Pleistocene. For Europe, North Africa and West Asia this border roughly holds for the Holocene period as well. – · – · – · Probable outer most East- and South East border during the Holocene.

41

Black Cyan Page 42

42


Although finds from the Pleistocene are less numerous and detailed than those from the Holocene, it is safe to conclude that during the Pleistocene, the distribution area of the aurochs was larger than in the subsequent Holocene. During the ice ages, when the climate in Europe and Asia was colder and drier, this area was greatly diminished particularly in the north; during the warmer interglacial periods it extended again. In the periods most favourable to the animal, the boundaries of its Pleistocene distribution area may have been even less restrictive than is known today. During the ice ages, the European aurochs stayed in the Mediterranean area. The precise northern border of its presence there in those periods is not known. Fig. 4 gives the total distribution area of the aurochs for both the Pleistocene and the Holocene. The data were taken from many authors, as will be shown below.

North Africa The present-day landscape of North Africa, mainly determined by the Sahara desert, looked totally different during the last ice age and the early Holocene. At the time, a more humid climate prevailed, and the vegetation was that of a savannah landscape with some scattered tree growth ((Ritchie & Haynes 1987), apart from a wooded zone along the northern rim of Africa. The fauna of this landscape showed a strong resemblance to that of the present-day East African savannah landscape (e.g. Serengeti), and included zebras, giraffes, lions and African buffaloes (Gautier 1993). The aurochs had no place here. The aurochs bones allegedly found here, like those from the Ahaggar mountains in South Algeria (Mc Burney 1960), definitely stem from domesticated cattle (A. Gautier, pers. comm.). As it happens, this savannah landscape was inhabited by cattle-raising tribes at the time. Their domesticated cattle may still be found in Saharan rock paintings (Muzzolini 1983).

Black Cyan Page 43

DISTRIBUTION AREA

In the course of the Holocene period, around 4000 BC, precipitation in this area began to diminish (Ritchie & Haynes 1987) and the savannah gradually changed into the desert of our times. Around 2500 BC, the situation more or less stabilised. The fact that no aurochs remains have ever been found in the Sahara seems to indicate that the animal never lived here during the Pleistocene. An area along the northern rim of Africa, from Morocco to Egypt, was certainly fit for the aurochs. Bone finds and rock paintings (Pomel 1894, Gautier 1993) confirm the animal’s occurrence here (see Figs. 5 and 6). Its presence in Egypt was not confined to the coastal area but extended far more south than anywhere else in North Africa (Churcher 1972). The most southerly finds are from Wadi Halfa (North Sudan). The reason for this southerly presence is the River Nile: the aurochs inhabited the marshy floodplains on both sides of the river. Together with the hartebeest (Alcephalus busephalus), it was one of the most characteristic large mammals there (Gautier 1988).

Fig. 5. Drawing of the remainder of a Holocene aurochs skull (with horn cores) from North Algeria (after Pomel 1894).

43

Black Cyan Page 44

44


Fig. 6. Picture from ancient Egypt (2800 BC). The pharaoh, rendered as an aurochs, knocks down his enemy (after Von Lengerken 1955).

Asia The regular occurrence of the aurochs in the Middle East was revealed by many bone finds and man-made pictures (Von Lengerken 1955, Uerpmann 1987). There is likely to have been a nearly continuous distribution of this animal throughout the area, from the Mediterranean Sea to India. Whether there was a clear distinction, or rather a gradual transition between the two subspecies primigenius and namadicus cannot be determined with certainty due to the current lack of comparable skull material. Quite a large number of finds are known from the eastern Mediterranean coast (Israel and West Jordania), but there are none from the rest of Jordania, and further east finds do not occur until the land of Euphrates and Tigris (Mesopotamia).

Black Cyan Page 45

DISTRIBUTION AREA

No aurochs finds are known from the Arabian Peninsula, which suggests that circumstances here were comparable to those in the former Sahara. During a number of Pleistocene periods, the Arabian Peninsula had a more humid climate than today; also here, there may have been a savannah landscape (Harrison 1964). Notwithstanding this more humid climate, other factors seem to have hindered the occurrence of aurochs in this area, since the animals probably never lived here. Comparatively few finds are known from India, where the subspecies namadicus lived; these all stem, with any certainty at least, from the Pleistocene (Grigson 1973). The animal must have lived here during the Holocene as well, however, since the (domesticated) zebu is likely to have evolved from this subspecies. Excavations at Mehrgarh (Pakistan, 6000 BC) have yielded remains of bovine animals that probably belonged to the namadicus form; if so, these are the only Holocene finds of namadicus (Badam 1984). Benecke (1994b) considers Mehrgarh as the centre of origin of the zebu, as small clay statues representing bovine animals with shoulder humps were made there as early as 4000 BC. Although as far as the Holocene period is concerned, there seems to have been a somewhat arbitrary boundary through India, the distribution during the Pleistocene period seems fairly clear cut. Namadicus was not only found in northern India, on the Kathiawar Peninsula and along the Ganges, for example, but also in the south, in the Deccan area along the Khrishna River. Nevertheless, most finds stem from the north, from the area of the Namada River, from which the name of this subspecies derives. Future bone finds may well help to shed more light particularly on the Holocene distribution of the aurochs. No aurochs remains were found in Southeast Asia (D.A. Hooijer pers. comm.). Of other members of the Bovini tribe, like gaur and water buffalo, fossils were found here, which strongly suggests that this was the southeastern-most border of the aurochs distribution area. To reach Central China (where its presence has been shown) the animal would have had to cross or round the high southeastern embranchments of the Himalayan mountains. Since high mountains probably did not belong to its habitat, and in view of the fact that no remains of

45

Black Cyan Page 46

46


the animal were ever found in Indo-China, this must be considered a highly unlikely route. China was probably reached by a much more northerly route, which would also explain why the aurochs in China is more similar to the primigenius than to the namadicus form (Boule et al. 1928, Gromova 1931). In Central Asia the aurochs seems to have avoided the high mountains of the Himalaya and the Tibetan Tableland, since circumstances there were probably too cold and too dry. Up to now, no bone finds are known from these areas. In the area of Minusinsk (South Siberia) rock paintings were found from the transition period from the Neolithic period to the Bronze Age, which show aurochs, bears and elk (Francfort et al. 1993) (see Fig. 7). How far east the aurochs occurred in Holocene Central Asia cannot be determined with cer-

Fig. 7. Rock-drawing on the site Ust’-Tuba II 6, in the area of Minusinsk (Central Asia); Neolithic/ Bronze Age transition period. The drawing shows aurochs, bears and elk (after Francfort 1993).

Black Cyan Page 47

DISTRIBUTION AREA

tainty. Since its presence in China during the Holocene has not been ascertained (see below), the eastern boundary on the map is rather arbitrary. Distribution in China is likely to have been limited to the Pleistocene, since very many finds are known from this period, but none from the Holocene (see Photo 2).

Photo 2. Aurochs skull with horn cores from the Baikal area, represented both from the front (top) and the side (bottom) (after Gromova 1931). Such skulls from East Asia bear a striking resemblance to those from Europe.

Based on extensive bone material, Boule et al. (1928) described the animal as generally present. Bohlin (1938) mentions aurochs remains from Northeast China. According to Zong Guanfu (1984), all Chinese aurochs remains were in North and Central China, in the area between 1050 EL and 330 NL, apart from a single find from Taiwan. Since it was found very far from the main distribution area and this would not be the first time that aurochs bones had become mixed up with those of domestic cattle, that find must be considered suspect.

47

Black Cyan Page 48

48


No bones of this animal are known from the Holocene period in China. It is true that Jiu Lampo and Wei Qi (1980) claim to have found unique remains of ‘Bos primigenius’, but their assessment is very unreliable. As it happens, a large mass of tree trunks and animal bones were found together in the bed of the Sangganhe River (North China). The trees were dated to the period 2500-1700 BC. The bones included those of the Przewalski horse, the woolly rhinoceros, the Asian elephant and red deer. This is likely to be material from both the Pleistocene and the Holocene periods, which was dislodged by the river and lumped together elsewhere. No Chinese aurochs remains from the Neolithic period have ever been shown (Zhang Minghua 1984, Nelson 1990, Kwang-Chih Chang 1986), nor is there any evidence of local domestication. Watson (1961) describes how domesticated cattle appeared rather suddenly in the Yellow River area, around 2000 BC. Transition phases from wild cattle to domesticated cattle were not found in this area. Domesticated cattle, like many other culture elements, may well have been introduced from more westerly regions of Central- and West Asia. The Chinese name for aurochs consists of four characters, which signify ‘West-European-wildox’ ( ; Zheng Yi Li & Cao Cheng Xiu 1984). Also this is no indication in favour of an early, Holocene presence of the aurochs in China, unless it disappeared very early. The Chinese script began to develop in the course of the second millennium BC. If the animal had lived there at the time, a specific separate character would probably have been devised, instead of the four later ones known to us today, which were constructed together to form the animal’s name.

Europe The northern boundary of the distribution area went through Russia, from the Baltic to the Japanese Sea. Along this entire border, bone material was found (Gromova 1930). The most northerly find was discovered at 600 NL, at the south border of Lake Ladoga. The northern boundary roughly coincides with the present boundary between

Black Cyan Page 49

DISTRIBUTION AREA

the northern boreal forests and the more southern mixed coniferous/ deciduous forests. The Central Asian corridor probably only existed during interglacials, since the area was too cold for habitation during the ice ages due to the presence of tundras and glaciers. The entire aurochs population in the north-easterly part of the distribution area (China included) may have disappeared during these ice ages because of such unsuitable living circumstances, to be re-established from the west during interglacials, when the climate got warmer. Naturally this is merely speculation, since evidence in the shape of bones has not yet been found. The Holocene northeastern boundary of the distribution area also went through Russia (Gromova 1931). Fig. 4 gives an approximation of it; sufficient enough exact data are lacking. No finds are known from Finland, and the boundary of the distribution area probably lay south of this country. From Estonia, to the south of Finland, many bones were found both from the Pleistocene and the Holocene periods (Paaver 1965). Rather more data are known from Sweden, but only from the southern part of this country (Ekström 1993). It is likely that the northern border of the aurochs’ distribution area used to run along the plantgeographical border limes norrlandicus, through Värmland and Darlana (Aaris-Sørensen 1999). Very few data are known from Norway: not a single aurochs bone was found there. As to place-names, Szalay (1915) allegedly found four but does not indicate clearly how they are connected to the aurochs. In view of the distribution in Sweden and the fact that in the south of Norway, mixed coniferous-deciduous forests occur, it is still possible that the animal lived in the southernmost part of Norway. The western boundary of the distribution area probably lay between Ireland and England. Neither bone finds nor place-names are known from Ireland; many bones were found in England (CluttonBrock 1986).

49

Black Cyan Page 50

50


4.1. Occurrence inside the outermost boundaries Within the outermost boundaries of the distribution area outlined in the above, the occurrence of the aurochs naturally varied during the ice ages and interglacials. The northern Eurasian boundary was not only the northern boundary during the Holocene period, but probably also the approximate boundary during certain interglacials. During ice ages, the European northern boundary ran a much more southerly course, possibly through Southern France, Northern Italy and the Balkans (Von Koenigswald 1998). The northern tree limit at the time was in roughly the same position (Kahlke 1994). Only very few data exist about the animal’s occurrence in the high mountains during the Holocene. According to Söffner (1982), the European occurrence of the aurochs can be demonstrated up to the tree limit, but evidence for this is weak. In China it allegedly occurred no higher than 1000 m (Zong Guanfu 1984). For the Alps, Chaix (1994) mentions that most of the finds were discovered below 1200 m; only three of the 23 Holocene finds were between 1400 and 1500 m. As to its occurrence in steppe areas, the animal has already been shown not to have lived in the warm North African steppe, nor did it occur in the cold northern steppe during the ice age, which was inhabited by animals like the mammoth, the horse and the rhinoceros. Heptner & Naumow (1966) suggest that in historic times, the aurochs did live in the steppes, for instance in the Ukraine, but evidence for this is obscure. Whether this was a presence in the middle of the steppes or in the border zones or river areas is not clear. Rather extreme conditions prevailed in these steppe areas, such as severe winter cold, a thick layer of snow, extreme drought and poor-quality food for extended periods. As will be shown in Chapter 9.1.3., such factors probably kept the aurochs from using such areas as their habitats. The forest steppes, where somewhat milder living conditions usually prevailed, may have just qualified as aurochs living areas. Whether the aurochs occurred on any of the islands inside this distribution area probably depended on their accessibility from the main-

Black Cyan Page 51

DISTRIBUTION AREA

land and their size. In view of the fact that related wild cattle species, as well as domestic cattle, are able to swim, it is likely that the aurochs was also able to do this. However, bridging very large distances in this way must have been problematic, since no bone material has ever been found on the remoter islands. Only if an island was big enough, and connected to the mainland for a certain period, would there have been possibilities for the aurochs. The only large mammals able to reach the very remote islands in the Mediterranean Sea were good swimmers such as elephants, deer and hippopotamus (Sondaar 1977). Islands such as Cyprus, Crete and Malta were never reached by aurochs, although the said good swimmers did go to these islands (see Fig. 8). Because of its remote position Japan never had an aurochs population either. During ice ages, the northern part of Japan was connected to the mainland by a land bridge (Kahlke 1994). During such cold periods, the Chinese aurochs would withdraw to the warmer regions in the south. An island like Sicily was definitely connected to the mainland, Italy, during certain periods; in this way, the aurochs was able to reach this future island. When the land bridge was lost in the Middle Pleistocene period, however, the species remained there but was diminished by about

Fig. 8. Bull play as found in a wall painting in the former palace of Minos on the Isle of Crete (ca. 1600 BC; after Von Lengerken 1955). The bulls that were commonly used in such plays were domesticated descendants of the aurochs, who did not occur naturally on this island.

51

Black Cyan Page 52

52


20 % (Brugal 1987). Similarly, an island was formed in Denmark: the Island of Sjaelland, which was formed around 6000 BC when the sea level started rising after the last ice age. The aurochs population that lived on this island became extinct, however, together with other species like the bear, the lynx, the badger and the elk (Aaris-Sørensen 1980). Apparently, the aurochs could not survive here as an island population. The combination of island formation and hunting may well have been the cause for its disappearance. The Gobi desert, which lay inside the northeastern distribution area, was obviously not a suitable living area for the aurochs. As yet, very little is known about the vegetations that may have been present in this area in the early days.

Summary On the basis of bone finds, descriptions and place-names, the occurrence of the aurochs has been traced. Fig. 4 shows the distribution area during the Pleistocene and Holocene. The distribution area of the aurochs stretched across nearly all of Europe and large parts of Asia and North Africa. During the existence of this animal as a species, cold periods (ice ages) and warmer periods (interglacials) alternated; both had consequences for the size of the animal’s distribution area. The area diminished in cold periods and grew again in warmer ones. Because of their extreme living conditions, the high mountains, deserts and steppes within the distribution area did not belong to the living area of the aurochs. Remote islands were not suitable because of their inaccessibility.

Black Cyan Page 53

DECLINE AND DISAPPEARANCE

5. DECLINE AND DISAPPEARANCE 5.1. Outside Europe As is evident from the description of the distribution area, data about the occurrence of the aurochs become increasingly scarce as the animal lived further away from Europe. The Middle- and East Asian population did not survive the Pleistocene period; the aurochs became extinct in those areas. Of the population in India after the Pleistocene only the alleged namadicus finds from Mehrgarh are known. It is fairly certain that the aurochs lived here during the Early Holocene, in view of the fact that the domesticated zebu stems from that approximate area. Already in pictures from the Mohenjo-Daro culture along the Indus (4000-2000 BC), zebus are depicted in the way we still know them nowadays, with a large hump at the withers, a large hanging dewlap and typical, upward-pointing horns (Von Lengerken 1955). Beside this zebu type, people in this area already knew other cattle breeds, which looked a lot like the domestic cattle breeds in areas more to the west. About the aurochs in these regions, however, nothing from that period is known. There are data from the Middle East and North Africa, but they are insufficiently exact to provide a clear picture of the period in which the aurochs occurred and of the course of its decline. Pictures of the aurochs are not necessarily indicative of its presence; artists of the period often used motives and pictures from earlier cultures. The course of the animal’s disappearance can only be charted on the basis of bone material, sufficient quantities of which are lacking. Many depictions of hunting scenes are known from Mesopotamia, however, and they are relatively easy to date, since the periods of office of the kings in these pictures are known. The Assyrian king Assurnassirpal II (883-859 BC), for example, managed to catch 50 aurochs during a hunt along the upper course of the Euphrates (see Fig. 9). Hunting was done on chariots pulled by horses (Von Leng-

53

Black Cyan Page 54

54


Fig. 9. Picture of an aurochs hunt in Mesopotamia (9th century BC). King Assurnassirpal II hunts this animal from his chariot (after Von Lengerken 1955).

erken 1955). Senacherib (704-681 BC) likewise still hunted ‘wild cattle’ in Northern Mesopotamia. There is no information about later hunts in these regions. With regard to Egypt, the last occurrence of the animal is often related to a hunting party by pharaoh Ramses II (1197-1165 BC) in the marshes of the Nile delta. Further investigations have revealed that the hunt in question took place in the marshes along the Euphrates or the Tigris in North Mesopotamia (Von Lengerken 1955). The Greek historian Herodotus (± 485- ± 425 BC) mentioned the presence of bovine animals that were doomed to graze walking backwards, because of their long forward-curving horns, in neighbouring country Libia (‘Histories’, Book IV, Chapter 183). Whether these were really originally wild cattle, or indeed aurochs, is very uncertain. The lack of data clearly makes it difficult to determine how the process of decline took place outside Europe. The aurochs may be considered to have disappeared from the individual countries of the Middle East and North Africa roughly in the course of the first millennium BC.

Black Cyan Page 55


5.2. Inside Europe The decline and disappearance of the aurochs in Europe proceeded from the south and west of Europe to the northeast, and eventually ended in Poland. At present there are no bone finds, place names or descriptions from southern parts of Europe like Spain, Middle and South Italy and the South Balkans that indicate up to which period the animal still occurred there in the wild. Italy. A stone is known from Etruria, from around 350 BC, which shows a hunting scene (probably with aurochs) (Von Lengerken 1955). Mentions are few and far between also from later periods. In his description of the animal world of Germania in 53 BC, Julius Caesar records the aurochs as if it was an animal that had (almost) ceased to exist in his homeland. He himself apparently never saw the animal, since his description is not entirely realistic. From the first century BC, Vergil (circa 30 BC) mentions the presence of ‘wild aurochs’ in the north of Italy (Po River area), which were caught there and tamed to be used as draught animals, for lack of domesticated draught oxen (Georgica II, 374; III, 532). At the time of the Roman Empire (up to circa 400 AD), very many wild animals, including aurochs (Szalay 1917, page 46-53), were caught on a regular basis from around the empire and transported to Rome or other towns to be used in arena fights. In view of the huge effort that was made to catch and transport wild animals, and of the great need for them, so as to entertain the people, the aurochs is likely to have become extinct in Italy in the beginning of our era. England. The aurochs probably became extinct much earlier in England, if the age of certain bone finds is any measure. No aurochs bone finds later than circa 1300 BC (Early Bronze Age) are known from this country (Clutton-Brock 1986). Extinction must have occurred rather early, therefore, compared to other countries in West Europe. Several aurochs drinking horns found in kings’ graves do date from the period 620-640 AD (Stone 1961). One of these was found at Taplow, seven others at Sutton Hoo. The latter include the largest keratin aurochs

55

Black Cyan Page 56

56


horn known. These richly decorated horns are likely to have been brought from the mainland during the Migration Period, so they do not stem from local animals. Netherlands. The earliest written data concerning the animal date from the Roman period. Tacitus (117, Book VI, Chapter 72) reports administrative problems in the land of the Frisians, in 28 AD. At the time, these people inhabited the northwest and north of the country, which were occupied by the Romans. The officer Olennius had imposed a tribute in the form of cattle hides the size of aurochs hides. When the Frisians were no longer able to comply a revolt broke out, which eventually forced the Romans to retreat. Apparently the aurochs still occurred there to a certain extent. Further reliable mentions from later periods are lacking. There is a report from 1344 of ‘two wild oxen and a deer from the Forest of Haarlem’ (Acker Stratingh 1844), but this most likely refers to cattle that had run wild there. The research of bone finds gradually started as early as the 19th century, when mention was made of aurochs remains that had been found. Aurochs skull remains and horn cores were found in the soil that was turned up by the water when several dikes broke, for example at Weurt (Van Hasselt 1805), Eembrugge (De Fremery 1831) and Genemuiden (Van der Vlerk 1942). Systematic archaeological research, which gradually took shape in the late19th/ early 20th century, yielded an increasing quantity of data from the beginning of the 20th century onwards. Aurochs bones appeared from mounds (Van Giffen 1913), bogs (Clason 1961, 1965, Kruytzer 1949) and clay sediments (Creemers 1927, Van Es 1990). Not only bones but also horns were dredged up from the North Sea (Stolzenbach 1984 (see Photo 3), Bosscha Erdbrink 1986, Nederlof 1995). As more remains are known and conserved, comparative studies become more feasible. As early as 1909, Rutten described a limited number of aurochs remains that were kept in the then Rijksmuseum van Geologie at Leiden. Van der Vlerk studied the collection of this same museum, which had already been extended considerably, in 1942. Van Es (1990) analysed the occurrence

Black Cyan Page 57


of the aurochs in the north and centre of the Netherlands on the basis of a large amount of material, both from museums and from private collections. De Jong (1998) traced the distribution of bone finds in the province of Noord-Brabant.

Photo 3. The aurochs horn of Stolzenbach, dredged up from the North Sea. This is a part of the right horn sheath and a part of the horn core that belongs to it (Photo D.P. Bosscha Erdbrink). The piece of horn core is rendered in approximately the matching position. The holes in the horn sheath were made by stone borers.

Notwithstanding the many finds, up till now only a rough picture of the occurrence of the aurochs in this country has emerged. It is true that finds are known from the northern coastal area, for instance from mounds, but the animal does not seem to have occurred there generally (Clason 1965). Relatively many finds stem from river valleys, like that of the River IJssel (Van Es 1990) and the river area in the east of the Netherlands (Lauwerier 1988), where the animal was not rare. Lauwerier still records aurochs from the 4th century. Finds from (raised) bogs date from a period when fen vegetation was present locally, in which the animals died. What the distribution on the higher, drier soils was like cannot be determined as yet. Bones are preserved rather more poorly in dry, acid, sandy soils than in wet clay- or peat layers. In itself, the fact that far more finds stem from wet than from dry areas does not demonstrate that the animal occurred more frequently in wet areas, but rather

57

Black Cyan Page 58

58


that its bones were better preserved there. The most that can be presumed is that the animal did occurred regularly in such wet areas. A map that shows the places in the Netherlands where aurochs bones were found does not necessarily represent the actual original distribution of the animal. If anything, it shows where the digging has been done, and where bones have been preserved well, which is why bones from lowland areas and from wet peat and clay layers dominate. Moreover, not all finds in the Netherlands have been properly mapped yet. The pressure on the aurochs population is likely to have risen with the increase of the human population. Aurochs finds from the postRoman period (after 400 AD) are not known. The aurochs bone that was found some years ago at Wijk bij Duurstede, which was dated to circa 900 AD on the basis of the accompanying pottery (De Vries 1996), turned out to be from the period 124-236 AD when a 14C- dating was done a few years later (Clason et al. 2000). In view of its length (33.5 cm), the silver-mounted horn that is part of the church treasure of the Onze Lieve Vrouwe Kerk at Maastricht, and which dates from the Viking period (circa 900 AD) (Roes 1940), almost certainly does not originate from an aurochs. Belgium. Aurochs finds of both the Pleistocene and the Holocene periods are relatively scarce in Belgium. As in the Netherlands, however, many remains of other Pleistocene mammals (mammoth, woolly rhinoceros, horse, steppe bison) have been found (Gautier 1996). A national systematic review of aurochs finds is still lacking. It is known of some areas that the aurochs lived there. In the Escoural cave (area of Liege), both Pleistocene and Holocene remains of the animal have been found, of which no more specific dates are known, however (Gautier 1996). In the northwest of Belgium (the Flemish Valley), particularly when the Channel from Gent to Terneuzen was dug, many bone remains were found, which included some of the aurochs (Gautier 1993b). The systematic digging and preservation of these left much to be desired, however. The Pleistocene aurochs finds from Overmere and Zemst (Germonpré 1993) and Oostwinkel (Germonpré & Ervynck 1988) also stem from the Flemish Valley. These remains were found in sandy lay-

Black Cyan Page 59


ers. The most recent aurochs finds, in Tongeren and Oudenburg, date from the 1st down to the 4th century AD (A. Ervynck pers. comm.). France. The Pyrenees and the northeast of this country constituted the last living areas of the aurochs (Guintard 1994). This was found out on the basis of descriptions that were handed down; for the Pyrenees, for example, the aurochs is still mentioned from 400 AD (Szalay 1917, page 149). That the animal still occurred in the same area around 1600 (Gesner 1620) is likely to be a misunderstanding, and probably refers to the feral cattle that used to live in the Provence, possibly the Camargue area, where such bovine animals lived of old (Szalay 1917, page 150). Middle-Aged hunting descriptions from the northeast of the country show that the animals had already become so rare in the 6th century that hunting them had become the privilege of kings. Thus, a description of a conflict from 590, under king Gontran, concerns the killing of an aurochs by a non-royal (Szalay 1917, page 148). Reports from this area are known down to the 9th century; Charles the Great, for instance, still hunted an aurochs ‘with huge horns’ here. There is no written evidence after the 9th century, though for a long time after that, aurochs may have moved from Germany to France, or from Switzerland, where it was still mentioned around the year 1000 (Szalay 1918a, page 71). The 14th century nobleman Gaston Phoebus, in his extensive work on French game animals, does not mention the aurochs at all (Phoebus et al. 1994). Denmark. The oldest Holocene aurochs finds in Denmark are dated as early as 8500 BC. This was the Young Dryas Period, in which a cold dry climate prevailed, with semi open wooded landscape (Aaris-Sørensen 1980). Aurochs research in this country has become known especially through the work of Degerbøl & Fredskild en Aaris-Sørensen. As the ice retreated, the animal moved further north. Because of the initially low level of the sea, Denmark was still connected to Sweden by a land bridge for a considerable time. In this way, many animals including the aurochs could reach Sweden. When the sea level rose the land bridge disappeared, which created the Danish

59

Black Cyan Page 60

60


islands. As a consequence, the aurochs disappeared on these islands earlier than in Jutland, i.e. around 5900 BC. Exactly why it became extinct on Sjaelland has not yet been fully explained. The main cause is likely to have been the isolation from the mainland (Aaris-Sørensen 1980). The bear and the elk disappeared from this island at about the same time as the aurochs, but animals like red deer, roe deer, wild boar and wolf were able to survive much longer, as well as the beaver, which held its own for a very long time after the island was formed (Aaris-Sørensen, in lit.). The conditions for survival on the mainland (Jutland) were more favourable, although the existence of the aurochs ended here, as well, around the beginning of our era. Sweden. Data for Sweden are similarly scarce. According to a description by Adam von Bremen the animal still lived here in the 11th century (Isberg 1962), but it is not clear whether this author refers to his own observations or to folk tales, in which the animal is described as ‘living under water like polar bears’. Research of place names does not provide a clear picture of its last occurrence either. It is true that some parts of place names may stem from certain periods (such as ‘-röd’, from 800-1000 AD, and ‘-hult’, from the 12th and 13th centuries), but this does not prove that the prefix ‘urs-’ (‘aurochs’ or a person’s name?) indicates the occurrence of this animal in the same periods. Olaus Magnus, who had seen aurochs for himself in Poland, mentions in 1555 that the animal no longer occurred in Sweden at that time (Isberg 1962). Bone finds show a different picture, however, and seem to point to much earlier extinction. Mathiasson (1988) mentions a bone find from Stora. Förö (at the coast of Southwest Sweden), which was 14C-dated to about 2450 BC. According to the same author, rock drawings from a more recent period, the Bronze Age, depict aurochs. Aaris-Sørensen (1999) on the other hand dates the extinction of the aurochs in Sweden to a much earlier period, i.e. around 5400 BC. Probably the Stora Förö aurochs was a drowned animal that had drifted in from Denmark (Ekström 1993).

Black Cyan Page 61


Photo 4. Nearly complete skeleton of an aurochs bull from the peat of Hammarlöv, which can now be found in the Zoological Museum of Lund (Sweden) (after Von Leithner 1927).

Hungary. After the year 1000, the aurochs has not been determined with any certainty beyond Central Europe, with Germany and Poland as the most important distribution areas. The youngest bone find from Hungary dates from the 12th century (Bökönyi 1962, Vörös 1985). Szalay (1938) considers the animal to have become extinct there before 1250. The aurochs is not mentioned in a single native Hungarian source, probably because of the late (10th century) appearance of the Hungarian people in this area and the absence in the Hungarian language of an original word for ‘aurochs’. It is true that some Hungarian museums possess aurochs horns from after 1250 (Bökönyi 1956); these have turned out to be of foreign origin (East Prussia), however, and were given to the Hungarian king Sigismund in the beginning of the 15th century.

61

Black Cyan Page 62

62


Germany. The research of written sources for aurochs occurrences in the German language area is greatly impeded by the confusing appellations given to the animal and to its closest relative, the European bison (see also Chapter 6.3). At first, up to the 11th century, the old Germanic word ‘ur’ was still used for the aurochs. After this the word gradually changed to ‘Auer’ and ‘Auerochs’, a change that took place at the same time as the final disappearance of the aurochs from Germany. The aurochs survived for quite a while longer only in the remoter area of East Prussia, which was to belong to Germany from 1230, but the nomenclature did not become any clearer there, either. Since the namechange took place between the 11th and the 13th century (Szalay 1915), and the word ‘Auerochs’ was used to indicate the European bison during this period, it is risky to try and distinguish the aurochs and the European bison on the basis of their name during that period. Many authors have been misled by this. Besides name research, bone finds may be studied. Vörös (1985) has reviewed the finds from Central Europe, including Germany. The latter finds are dated roughly to the period from the 9th to the 13th century, and stem from Lower Saxony, Saxony and Berlin. Since this is rather a long period, it is difficult to determine how the disappearance of the aurochs proceeded and when the animal eventually vanished from these areas. In former East Prussia, the aurochs was able to hold out for a long time in the Great Wilderness. That it indeed occurred there has been proven by many bone finds, which include a complete skeleton (La Baume 1944, Frevert 1957). From the period in which the German Order dominated the area (after 1230) bone finds have also become known, which have been dug up and related to that period of inhabitation. The presence of the aurochs can be deduced not only from bones, but also from horns found there. The books that were kept by the German Order (the so-called ‘Treßlerbuche’), which have been preserved for the period from 1399 to 1409, reveal that as many as 15 aurochs horns were gilded or silvered and given as gifts between 1406 and 1408 (Nehring 1898b). It is true that they were recorded as ‘Wisenthörner’, but they actually turned out to be aurochs horns. Some of them were rediscov-

Black Cyan Page 63


ered in Hungary (Bökönyi 1956). After 1408 a mere two mentions from Stella (1518) and Maciej z Miechowa (1521) have survived. Stella reports and distinguishes both aurochs and European bison, which he presumably never saw himself, since his descriptions are based mainly on those by Caesar and Plinius. The Polish historian and geographer Maciej z Miechowa, who lived on the Polish side of the German border, also mentions both aurochs and European bison in his description of the Great Wilderness8: ‘Because those forests are so large, many large wild animals may be seen and caught in abundance: aurochs and forest bovine animals, which in their language are called ‘thuri’ [aurochs] and ‘zumbrones’ [European bison]’. As far as East Prussia is concerned this is the last mention. Russia. Only a few data are known with regard to the west of Russia. According to Vörös (1985), aurochs bones like the ones found in the Central Dnjepr area are not quite reliable, also because many European bison bones were found there as well. If they are really aurochs bones, their occurrence could indicate the animal’s presence here up to the 12th century. This might well be correct, since descriptions have survived of aurochs hunts in the same area, at the time of grand duke Vladimir Monomakh (early 13th century) (Dementiev 1958). Dementiev presumes that the animal became extinct here in the 14th or 15th century, but does not indicate what his supposition is based on. Poland. The very last living area of the aurochs was in Poland. Several parts of the country have yielded bone finds from the 9th to the 13th centuries (Vörös 1985), including Silesia (Wroc³aw), Masovia (Pu³tusk) and Pomerania (Szczecin). The animal also occurred in the northeast, along the Narew River (Górski & Gru¿ewski 1994). The finds from the last wild population at Jaktorów include a horn from 1620, which is now in the Livrustkammaren Museum at Stockholm (see Photo 7). In addition to this there are several eyewitness accounts, of people who either visited the last population at said village, such as Schneeberger and Œwiêcicki, or reported dead or liv8

The Great Wilderness was an extensive area of forests and marshes in what used to be East Prussia (see Chapter 9.1.1).

63

Black Cyan Page 64

64


ing specimens of the animal elsewhere in Poland, such as Von Herberstein and Gratiani a Burgo. Table 2. Approximate moments of extinction of the aurochs in several areas. The exact year is known only for Poland.

Area

Moment of extinction

England Middle East Italy Denmark Netherlands France Hungary Germany Russia East Prussia Poland

Circa 1300 BC. 1st millennium BC 1st century BC 1st century Circa 400 10th century 13th century 13th century 13th century Circa 1500 1627

5.2.1. The last living area9 Around the 14th century the kingdom of Poland consisted of several duchies, of which the duchy of Masovia (with Warsaw as its centre) was one of the most important. Apart from the earlier East Prussia, the southern half of which now belongs to Poland, the territory of Masovia during that period was the only area in Poland where the aurochs still occurred in the wild. No reliable bone finds or written sources that indicate its occurrence in the wild are known from any other part of Poland, for that or later periods. There were not only wild aurochs still living in Poland; people also held them in captivity. As such they were present in a zoo at Zamoœæ (Southeast Poland) for many years, possibly even after 1627, the year in which, as far as is commonly known, the last aurochs died. 9 The data for this chapter derive from Wrze´sniowski (1878), §Lukaszewicz (1952), Heymanowski (1972) and K¸edrzierska (1959, 1965).

Black Cyan Page 65


Also in Masovia, however, their numbers diminished more and more. From an animal that could initially be hunted freely, it became more rare and the hunt for it increasingly became the privilege of the nobility. In written records of such privileges the aurochs holds a special place among the various types of game. Differently than in the German-speaking areas, Polish and Latin writings from Poland clearly distinguish between the two related bovine species, aurochs and European bison. These were called ‘tur’ and ‘¿ubr’, respectively, in different manners of writing, and never confused with each other. A noble privilege of this kind has been passed down from as early as 1298, when Boles³aw, duke of Masovia, granted several villages with grounds in the district of P³ock to one knight Paulecz, reserving, however, the hunt for the aurochs in those areas for himself. In 1359, Siemowit IV, duke of Masovia, allows the duchess of Wyszogród to hunt on his entire territory, excluded the hunt for aurochs. Another document of the same tenor is known from 1383. The animal may still have occurred in the east of Masovia around 1436, but its occurrence was gradually restricted more and more to an area southwest of Warsaw, where there were large forests and where an aurochs population could still survive due to protective measures. The area in question was a forest area of roughly 25,000 ha, divided into several smaller forests, administratively speaking (see Fig. 10). The complex included the Forest of Bolemów (circa 6000 ha), the Forest of Jaktorów (circa 5900 ha) and the Forest of Wiskitki (circa 13.600 ha). In the course of the period in question, the latter was divided into smaller leased territorities, the Forests of Miedniewice and Korabiewice. To the north, this whole forest area connected to the Forest of Kampinos, which is one of Poland’s present-day national parks. In the early 15th century, the Polish king Jagie³³o was still invited by the duke of Masovia, who owned the forests at the time, to hunt for aurochs there. In 1476, however, management of the forests reverted to the Crown; at the same time, the Royal Family came in possession of one of the two last wild aurochs populations. This population was to remain and exist there for another 150 years. If the aurochs is supposed to have become extinct in the Great Wilderness around 1500, the very

65

Black Cyan Page 66

66


Fig. 10. The various forest parts that together formed the living area of the last aurochs population (after Heymanowski 1972). The map of Poland that shows where this area was situated in the 16th and 17th centuries has been added. –––– probable border of the Royal Forests ........ probable subdivision - - - - supposed subdivision g property of the king n property of the nobility t property of the church

last wild aurochs population must have occurred in the Polish Royal Forests in the early 16th century. At that time the animals still roamed the entire forest area, but in the second half of the 16th century they occurred exclusively in the Forests of Wiskitki and Jaktorów. Documents concerning these animals, from 1602, 1620 and 1630, only refer to the latter forest. The organisation of the management of the aurochs population in these areas goes back to the 13th century, at which time there was already a well-organized gamekeeping service, in those days under ducal authority. In 1476 this reverted to the Crown, together with the Royal Forests. The gamekeeping service employed several so-called ‘hunters’, who played an important role in the aurochs management. Most of the

Black Cyan Page 67


hunters lived in the village of Koz³owice; after circa 1500 they lived there exclusively. Their number had originally been fixed to 16, and they belonged to a privileged group that had been exempted from paying taxes and together tilled some 8 w³óka (= 134 ha) of farmland around Koz³owice around the year 1510. Their most important task was to ‘…..keep an eye on the aurochs, let them graze, collect the hay from Jaktorów from the subjects, to supply the aurochs with this in winter, to know the number of animals and give a report to the district manager or the tenant every three months’. In addition to this, they sometimes had to catch the animals with nets. In accordance with their important status they carried out their tasks ‘provided with good weapons and cloths and sitting on good horses’. Thus equipped, they kept the animals under daily supervision to guard them and to keep them inside the boundaries of the forest area. If the aurochs would cross these to graze the meadows and fields beyond, they would be driven back to the forest with the help of dogs. Beside these hunters, the farmers from Jaktorów village formed an important, similarly privileged group of people. Jaktorów was mentioned as early as 1425, and a document from 1597 reports the village to have been established at the time exclusively ‘for the aurochs and their welfare’. This is emphasised when problems emerge later on. The hay that was brought to the feeding places by the hunters in winter was made in the meadows at Jaktorów in summer. A third group of people concerned with the forests consisted of foresters, who lived in the villages around the forests. In principle, wood was cut on a limited scale in these forests; as the 16th century proceeded, however, this happened more extensively, and the foresters became increasingly involved in the growing corruption. Administratively speaking, the whole forest area fell under the district manager (‘starost’) at Sochaczew. Some of the Inspection Reports that were made in the course of the 16th century have been preserved. They constitute one of the most important sources of information with regard to the changes in the numbers of animals and the causes for these changes. This picture may be complemented with other data, to show how an initially well-organised protection service eventually de-

67

Black Cyan Page 68

68


generated into a hollow, ineffective organisation as a result of negligence and corruption (see Fig. 11). The first Inspection Report that records the number of aurochs dates to 1564. There is one earlier mention of their number, however; Schneeberger (in: Gesner 1602) writes about this: ‘It is said that in the 57th year [i.e.1557] several animals perished from the fierce cold, but now it is suspected that there are once again over 50, since the accurate number is being concealed; they do this led by some kind of superstition’. For which year this number of ‘over 50’ holds is not exactly known. Nor is it known whether this is a good estimate, not intentionally overdrawn to obscure the real number. As it happens, the ‘official’ 1564 report mentions 38 animals, more than twelve less. This is no small difference, so either the number of ‘over fifty’ does not tally, or Schneeberger’s letter was written not long after 1557, possibly in 1559, after which the number diminished drastically in the subsequent few years. In the sixties of the 16th century, the Inspection Reports began to show clearly the problems concerning the management. The 1564 report, for example, mentions that: ‘much of these forests has already been destroyed by the people who have the right to cut wood, but many people, coming from neighbouring noble-, as well as church villages, cause a lot of damage; they ride violently and do not let themselves be caught; the officials and the foresters fell trees, so that the animals that come there cannot live in peace, the way they were used to, since in former times nobody was admitted to cut even a single piece of wood, unless at the edge. In addition to this, herds of cattle of the nobility and the farmers are found in these forests, which graze there, as a result of which, according to the hunters, the aurochs do not breed well; since they do not have a stable where they can get hay, the calves die and few survive the winter. The right to cut wood freely in these forests belongs to lords Sandomirski, Maciejowski and Okuñ from Grodzisk and the wife of the starost of B³onie’. The 1564 report also states: ‘Among those cows there is one very old, thin one, which will survive the winter only with difficulty. When we asked them why the au-

Black Cyan Page 69


rochs were so thin and why they did not breed well, as a cause for this they told us that many horses and herds of cattle come to the places where the aurochs graze as well, as a result of which the aurochs cannot thrive, though there are quite a few calves’. ‘…in this forest there are numerous large trees, suited to build with, but also splendid for bee nests, but they [the forests] have been damaged in part. When we inquired who had damaged them, some foresters said that during the time of the late lord Borek the oaks had been cut for the coopers; others also said that they had been cut and taken to Oryszew for the present lord of Sandomierz’. Of old, people apparently were allowed to cut wood in this forest on a limited scale, and to graze cattle here, but this took ever-increasing proportions. In 1565, more drastic measures were already being considered: ‘The aurochs, which are in the forests of Wiskitki, need better living conditions; a shed for the winter needs to be built for them, but also, not so many herds of cattle should be permitted as is the case at present’. This is elaborated on again in the following year (1566): ‘As to the aurochs that are in the district of Sochaczew, the supervisors report that these need better living conditions and care, that is to say, a shed for the winter, where hay can be taken for them as well. That the people who are allowed to fell trees there do not pass through their habitats to get timber, also that there less cattle will come to these forests, brought there by the royal, archiepiscopal and country-noble herdsmen. In view of the fact that the aurochs no longer occur anywhere in the kingdom, and that much care ought to be bestowed upon them, His Royal Highness should order the district head to take care of everything, and to do what should justly be done to enable the aurochs to live well, and by rights to be defended by the district head against the harmful cutting of trees as well as the grazing of so many herds of cattle’. The situation did not improve, however; if anything it became worse. In 1570, the only mention of aurochs was that these had been found. In addition to this, mention was made of pigs in the forest: ‘When there are many acorns in these forests, they come from many royal and country-noble villages and drive their pigs in there; they give

69

Black Cyan Page 70

70


two bushel of oats for seven pigs and for what they gather in addition they also give geese, capons and chickens’. For over 20 years, no report was passed down. The situation had not improved, however, since the 1597 report reads: ‘We demand that the inhabitants of said village do not herd cattle and do not mow grass for their own use there, where the aurochs live and where they have their meadows and habitat, because at the time, this village was not established and given liberties so much for their cattle, as for the aurochs and their welfare. The starost has to ensure that our forests, where the aurochs live, are not destroyed by said subjects, so that the aurochs may retain their former habitat’. In 1599, the total number of aurochs had shrunk further: ‘Of these aurochs, now, not more than 24 in all may be found, including the young animals. When we asked the hunters of Koz³owice what the cause could be, that there are so few of them left, they said that partly, they killed each other, partly, that no small number of them had perished during the past winter’. At the same time, an inventory of the same year shows that the shed, which had been written about as early as 1566, had been built in the meantime. As the population started to approach its end the warnings and admonitions became more severe. Of the hunters of Koz³owice, for example, the report states: ‘According to the old revised statute there should be 16 of them, each on half a w³óka. Now these hunters have multiplied to such an extent that they have divided those 8 w³óka into small parcels of land, though they do not honour their obligations. This is why it is necessary that they are called to order and have their obligations pointed out to them’. Even in 1604, when the situation was already hopeless, a decree from the King arrived: ‘For the increase of the aurochs in our forests the said starost in his capacity as manager should provide protection carefully and with all possible means, so that neither their maintenance, nor their urgent protection ever diminish, since this region and these animals belong to the glory of the Empire’. The 1602 Inspection Report mentions only four animals left:

Black Cyan Page 71


‘In these forests at Jaktorów, aurochs are hiding, of which at this moment there are only four, which we have seen for ourselves, three bulls, one cow. The hunters let us know that before, there were more of them, but many of them have died through other cattle contaminated with the pest10. We asked about their hides and horns, and an official of the starost told us that they have been sent to the king. However, these aurochs died while the district was being managed by managers given to them by the king. Under the present starost only one was shot, which is the subject of the legal inquiry’. In 1620, the last aurochs bull died and only one cow was left. It so happens that the 1620 Inspection report mentions that there was only one cow left in that year. In addition to this, it is written on an aurochs horn from Poland, which is currently being preserved in a museum in Stockholm, that it is the horn of the last aurochs bull, which died in 1620 (see Photo 7). The last cow eventually died in 1627, because the 1630 Inspection Report states: ‘In the last report [from 1620] it was written that there was one aurochs cow, but now the inhabitants of this village [Jaktorów] said that she died three years ago’. It is not known of what this animal died. If it was the same cow as the one that was mentioned already in 1602, she had become more than 26 years old and probably died of ‘old age’. Perhaps it was a cow that had been born between 1602 and 1620. Presumably the records were not kept too meticulously, also in view of the fact that the death of the last aurochs in 1627 was not mentioned until 1930. That this cow was killed by a poacher, as is sometimes claimed, is based on fantasy. Beside this wild population, a number of animals lived in captivity for a long time, in a zoo at Zamoœæ in Southeast Poland that was owned by the Zamoyski family. Both aurochs and European bison were represented here, as a letter is known from the year 1610 of someone who wanted to visit this zoo and wrote beforehand: 10 · Which disease was meant here cannot be concluded from the Polish text (‘powietrze zarazeni’). Possibly it really was rinderpest, which was imported from Russia. It also could have been a different disease, since the diagnosis was not always clear.

71

Black Cyan Page 72

72


‘How many aurochs and European bison do you have, will you please let me know, and how many males and females’. Earlier, this person had already written about the aurochs: ‘They are nowhere to be seen any more, except in the Zamoyski zoo’. It is not known how long the animals survived there, and whether they outlived the wild population.

5.2.2. Causes of extinction That the aurochs was actually exterminated by man over a period of thousands of years, may be evident from the preceding chapters. Thus, the animal has gained the dubious honour of being the first documented case of extinction11. This process of extinction consisted of two main elements, hunting and ousting, both initiated by man. The local climate in those thousands of years may well have played a role, too. As a result of the climate getting dryer, for example, forest steppes could change into steppes, where the aurochs could not keep its ground. Marshes dried up, which eliminated another living area. The main factor was man’s presence, however, and the hunting aspect was the most striking. The aspect of ousting was probably just as important. This entailed the expulsion of the animal from its preferred habitat by man and his cattle. Aurochs and man probably preferred similar living areas. Especially in summer, the aurochs lived in wet habitats, such as river valleys, lake fringes and marshy forests. Man also had a strong preference for these areas. As soon as man started to live in permanent settlements, these were always in the vicinity of water. To provide food for the cattle, meadows were created in these wet habitats if they were not there naturally. Such a situation may still be seen in the forests at Bia³owie¿a (Poland). The meadows here are situated along small rivers (Narewka, Leœna, Hwo¿na, £utownia) and have partly taken the place of alder-ash (Circaeo-Alnetum) and ash-elm forest communities (Fraxino-Ulmetun) (Faliñski & Faliñska 1986). The villages are situated some11 The second documented case is that of the dodo (Raphus cucullatus Linnaeus 1758). This large cursorial bird lived mainly on the Island of Mauritius and became extinct about 1665 through man (Wissen, B. van (ed.) (1995): ‘Dodo – Raphus cucullatus [Didus ineptus]’. Catalogue of the exhibition ‘Het lot van de dodo’ (‘The fate of the dodo’). Zoological Museum, University of Amsterdam.

Black Cyan Page 73


what higher up, not far from the meadows. A similar situation occurred at the time of the reclamation of the Great Wilderness in East Prussia (Mager 1960, part I, page 274). The areas that were of vital importance for the aurochs were increasingly occupied by domestic cattle. The domesticated form took the place of the wild one. The situation at Jaktorów may serve as an illustration of this process. The meadows that were situated mainly along rivers (Heymanowski 1972), where for the greater part the aurochs could find the food they needed, were increasingly taken up by the cattle of the local farmers. As a result of this, sufficient reserves could no longer be built up for the winter, and many extra victims fell in this difficult period. Any temporary and local favouring of the aurochs by man, caused by the opening of the forest through woodcutting and shifting cultiva-

Photo 5. In 1972, in the village of Jaktorów (Central Poland), a memorial stone was unveiled in memory of the last aurochs population that lived in the local forests. The last aurochs, a cow, died there in 1627 (Photo K. Jaworski)

73

Black Cyan Page 74

74


tion, eventually detracted little from the process of extinction. People were unlikely to cut and burn significantly more forest than was necessary to provide their cattle with food, since this was a very time-consuming activity. Moreover, he will undoubtedly have tried continuously to keep the aurochs away from these laboriously acquired meadows. Apart from its exclusion from its most important source of food, hunting must have constituted a continuous threat to the animal’s existence. In the beginning, the hunting pressure will not have been very strong, because of the low density of the human population. As this population grew, the pressure became stronger and stronger. A large animal like the aurochs, which, moreover, probably lived in specific habitats, must have been easily traceable. The hunt for it was nevertheless regarded as dangerous, and whether the aurochs was the most preferred kind of game is very doubtful. Bones of red deer and wild boar have been found in much larger quantities in former human settlements than aurochs bones. Such animals, as well as smaller ones, could probably be caught more easily and with less risk. During periods of crisis in human cultures, the pressure on the aurochs may have mounted (HüsterPlogmann et al. 1999). Hunting took place with the use of pitfalls, nets, spears, and arrows and, in the 16th century, even with fire weapons. As the animal became rarer during the Middle Ages, the hunt for it was increasingly claimed by the nobility. This was the case in Poland, for example, where both the hunting and ousting in the Jaktorów forests levied their toll on the animals. Which factor actually turned the scales in the animal’s extinction is not clear, however. Of the Polish kings, Jagie³³o († 1434) definitely hunted here at least three times; less is known about later kings, though both Zygmunt I († 1548) and his son Zygmunt August († 1572) are known to have shown no special interest in hunting aurochs (Heymanoski 1972). The officially appointed hunters sometimes killed animals; these were bulls that had covered domestic cows or killed other bulls in fights they had amongst each other. The meat, hides and horns of such animals were given as presents to important relations of the king. Thus, the diplomat Von Herberstein (1557) received a nearly complete hide, which he exhibited in his house in Vienna after he had had it stuffed. Live animals

Black Cyan Page 75


were also given as presents. In 1563, Gratiani a Burgo saw aurochs calves in captivity at Königsberg (East Prussia), which probably stemmed from the forests at Jaktorów (Szalay 1938, page 58). The same happened with European bison. The hunting may also have entailed poaching; the 1602 Inspection Report clearly shows that there was a legal investigation into an alleged case of poaching. Poaching may well have taken place on a larger scale than was known, and it may have been done not only by poachers, but also by the appointed hunters. In view of the growing corruption among them and the great power they wielded, this is not unthinkable. As far as can be ascertained, however, the hunting pressure exercised by the Polish kings must have been fairly limited, comparable to the hunting pressure on the European bison in the Forest of Bia³owie¿a in the 19th century. Also here, the Russian Czars, who had appropriated the exclusive right to hunt European bison exercised this right only sparingly, making the availability of food the determining factor for the size of the population of these animals. The aspect of ‘ousting’ shows from the competition with cattle and horses, which grazed in the same places as the aurochs. This fact is mentioned regularly in the Inspection Reports, which shows that people did realize this at the time. In autumn, the pigs were chased into the forest to eat acorns. Acorns formed part of the food of the aurochs (Schneeberger in: Gesner 1602), and their consumption by pigs will have contributed to a certain degree to the poor state of nutrition of the aurochs population. It is true that supplementary hay was provided in winter, but it is not clear whether this was always sufficient to get and to keep the animals in good condition during that period (5 to 6 months) (see Chapter 9.1.2). As a result, during severe winters, such as those of 1556/ 57 and 1598/ 99, more animals perished than usual. Apart from this, there were years in which the hay harvest failed completely or in part due to bad weather or some negligence of the farmers, like in the summer of 1596. The royal decree from the year after this warned the negligent officials involved and threatened to punish them should this happen again. An attendant aspect is the susceptibility to disease, transferred by domestic cattle grazing in the same area. The 1602 Inspection Report records many animals dying from this, although it does not give details about the

75

Black Cyan Page 76

76


disease in question. There is no indication that it was the well-known rinderpest that would demand so many victims in later centuries. It may have been a way to disguise poaching by officials (£ukaszewicz 1952). The Inspection Reports indicated that negligence and corruption in the lower official ranks increased. The hunters did not honour their duties and their numbers grew steadily, while those of the aurochs diminished. The number of foresters grew too, since timber was being harvested on an increasing scale. The forest was divided into ever-diminishing lease units; a reflection of how intensively it was being used. A third group that saw its chance was that of the farmers. From the moment they were allowed to herd their cattle on the meadows in the forest, they took advantage of this. They paid (bribes?) to be allowed to bring more cattle into the forest, which was detrimental to the aurochs that needed protection. A factor which may have played a role especially near the end is inbreeding. There is no doubt that this must have increased in the everdiminishing aurochs population, though no evidence for such a fatal development has been recorded; no mention is made about clearly deviant or sickly animals as a result of inbreeding. In addition to all this, the supreme authorities, i.e. the Polish kings that ruled in succession, were not free from blame. A king who cares about, and will do anything to preserve, the object of his hunt may achieve a great deal. Basically competent kings like Zygmunt and his successor Zygmunt August did not take any special interest in hunting, as a result of which the royal grasp on the protection of the aurochs may have slackened more and more, allowing free scope to the various profiteers. After 1572, a very turbulent period started in Poland, in which various kings reigned in quick succession and there were a lot of interior struggles. The organisation of the protection of the aurochs was increasingly undermined, and the influence of the kings had on it was fading. Shortly after 1600, when there were only a few aurochs left, and it was ordered (in 1604) by royal decree that everything should be done to protect the aurochs and its living area, the village of Baranów was founded in the northern part of the Jaktorów Forest, where there were important pastures.

Black Cyan Page 77


Two places in the graph show a marked decline of the number of animals: the periods 1559-1564 and 1599-1602. After the first decline the number still seems to have stabilised between 1564 and 1599, with possible (but unknown) ups and downs. The strong decline between 1599 and 1602 nevertheless turned out to be fatal. Apparently something happened which eventually made the downfall of the population inevitable. In the 1602 Inspection Report, a disease is suggested as the main cause for this. The fact that the foundation of the village of Baranów in the middle of the living area of the aurochs was made possible shortly after 1600 suggests that around 1600, the corruption in the management organisation was already quite extensive.

Fig. 11. Changes in the numbers of the last aurochs population in the forests at Jaktorów (Central Poland). Data used for this graph: 1559 – In this year, their number is given as ‘once again over 50’. For making the graph, the number of 53 was chosen. Between 1557 and 1559 there was a rising trend. 1564 – In this year, 38 animals are mentioned: ‘8 old bulls, 22 old cows, 3 young bulls, 5 young calves’. 1599 – In this year, 24 animals are mentioned. 1602 – In this year, the fact that there are 3 bulls and one cow left is mentioned. 1620 – The last bull dies and one cow is left. 1627 – The last cow dies. (drawing: E.S. Bakker)

77

Black Cyan Page 78

78


After the extinction of the aurochs, the deforestation of its last living area progressed steadily. From a late 18th century map (see Fig. 46), it is evident that the Forests of Bolemów and Wiskitki were still largely intact forested areas at the time, but the larger part of the renowned Jaktorów Forest had been deforested, especially the northern half, and changed into farmland. Nowadays nearly the whole former forest area has been changed into pastures and cornfields, with poplar-lined lanes and straightened brooks. Only here and there have small pieces of forest survived.

Summary On the basis of bone finds and descriptions, the decline and the eventual disappearance of the aurochs were traced. Hardly any data about this are known from North Africa and Asia, since the animal already became extinct there rather early. However, as regards the situation in Europe, where the last aurochs lived, many data have survived. In Europe, its decline proceeded from the south to the north and from the west to the east. Table 2 shows roughly the moments of its disappearance from the various countries. The decline and eventual disappearance of the aurochs were caused by man, who both hunted and ousted the aurochs from its feeding grounds by introducing domestic cattle. Poland was the last country in which the aurochs continued to live for rather a long time. In Central Poland, in the forests near the village of Jaktorów, a well-organised management was conducted in which the aurochs held a central position. In the beginning the animals were ducal, later royal possessions. They enjoyed protection there, and received supplementary food in winter. Since their management was reported to the king on a regular basis, the situation at the time is well documented. Fig. 11 offers a graphic representation of the changes in the last population of aurochs. Eventually the aurochs population perished also in this area, through a combination of disinterestedness, corruption, cattle diseases, competition for food (from domestic cattle) and, to a lesser degree, hunting. The last aurochs died in 1627.

Black Cyan Magenta Yellow Page 79

LANGUAGE AND SYMBOLISM

6. LANGUAGE AND SYMBOLISM 6.1. Etymology Like many other words in the English language, the word ‘aurochs’ was adopted directly from German, in which the term ‘Auerochs’ is used. No local name has survived in the British Isles, since the animal became extinct there at a very early date (see Ch. 5.2). The word ‘aurochs’ has a long history. The Germanic tribes that inhabited Central and Northern Europe called the animal ‘ur’ (pronounced: ‘oohr’). We know this from various written sources including the Nibelungenlied (ca.1200), in which the animal is mentioned in connection with a hunting party held by the hero Siegfried (Szalay 1915, Franz 1967): ‘dar nach sluoc er sciere einen wisent und einen elch, starker ure viere, und einen grimmen scelch’ ‘after this he quickly killed a European bison and an elk, four strong aurochs, and a fierce stallion’. The word was passed down in a document from around 800, in which the Saxons make a pledge to their supreme deity, should they be victorious in the battle against Charles the Great (Szalay 1915): ‘ik tif ti in ur tu scapa un tat rof’ ‘I will give you an aurochs, two sheep and all the loot’ In addition, it is known from an Old-English runic poem (see Ch. 9.1.3) and from the North Germanic heroic songs of the Edda (Beck 1973), as well as for its continued use in many toponyms. The Romans, including Julius Caesar, turned the Germanic ‘ur’ into the Latin word ‘urus’ – in any case, that was the word used by official writers and scientists. In Italy at the time, Vulgar Latin had the name ‘bubalus’ for

79


80


the aurochs. The use of this name in other places in Europe, later in the Middle Ages, has contributed to the confusion about the difference between the aurochs and the European bison. Since the animal was called ‘ur’ in Old German, Old Swedish and Old English, it may be assumed that it was also labelled thus in Dutch. The word underwent a change in the course of time: the ‘u’-sound was lengthened to a diphthong, which changed into ‘aue’ in the German regions around the 11th to the 13th century. Thus ‘ur’ became ‘auer’, to which ‘- ochs’ was then added (Szalay 1915). The German ‘Ochs’ and Dutch ‘os’ are both old names for ‘bovine animal’. The sound developed similarly in other words: German ‘mur’ turned into ‘Mauer’, ‘sur’ became ‘sauer’. Words that look alike may have very different origins. German ‘Aue’ and ‘Auer’, for example, are completely unrelated. As was explained in the above, ‘Auer’ derives from ‘ur’. ‘Aue’, however, belongs to the group of words that also includes Dutch ‘ouw’ (as in ‘landouw’- pasture) and ‘ooi’ (‘soil along the river’). ‘Aue’ and ‘ooi’ actually mean ‘land situated by or in the water’. A transformation also took place in Swedish, where ‘ur’ changed into the present ’uroxe’. In Danish, this became ‘urokse’. In English and French, the German word was adopted almost without change: ‘aurochs’. During the transition of the Old Germanic ‘ur’ into ‘auerochs’, words like ‘urrind’ en ‘urochs’ existed for brief periods, one after the other, in the 10th and 11th centuries. In the 19th and 20th centuries the word ‘Urrind’ was used sporadically in Germany. There is no simple theory about the origin of the word ‘ur’ (De Vries 1987, Mayrhofer 1986). There may be a link with the red morning sky and the colour red, or a possible connection to water and masculinity. Thus, ‘ur’ may either mean ‘the red animal’ or ‘the impregnating, male animal’. Neither theory can be called convincing, since the genesis of the word ‘ur’ still has not been explained. The word ‘ur’ for ‘aurochs’ may also be present in the name of a related bovine species, the gaur. This word is sometimes explained as ‘gai-ur’, in which the Hindustani term ‘gai’ means ‘bovine animal’ (Ly-



dekker 1912). In that case, the two bovine names may have had common Indo-European origins.

6.2. Toponyms Toponyms are names of places, rivers or other landscape features from which the presence of animals or human activity may be derived. Thus, a toponym with a name for ‘aurochs’ could mean that the animal lived there; a possibility only, however, since one has to be careful with explaining toponyms: changes in the ways of writing and pronouncing words can easily create confusion. Like many other animals, the aurochs has left its mark in toponyms. In spite of its initially large distribution area, toponymic research with reference to this animal is only known from Europe. This is probably related to the fact that Europe formed the last territory of the aurochs. If its occurrence in a certain area is further in the past, toponyms are more likely to have changed their shapes or even to have been replaced completely. This means that aurochs toponyms in Poland and Lithuania are fresher and, therefore, clearer, than elsewhere. Researchers of aurochs toponyms in the Netherlands and Flanders are faced with two problems. First of all, it is very long ago that the aurochs lived in these areas; it probably disappeared well before the year 1000. As a result, any toponyms that existed at the time will either have disappeared, or been greatly changed. The fact that the syllables ‘ur(s)’ or ‘oer’ of a toponym need not always be indicative of the word ‘oeros’ (aurochs) is another problem. To illustrate, ‘ur(s)’ cannot only refer to the aurochs, but also to ‘hors’ or ‘hros’, earlier names for horse. ‘Oer’ can also mean ‘ferriferous soil’, ‘over’ (over) or ‘oever’ (riverbank), and ‘Ur’ may indicate the Christian name ‘Uri’ (Gysseling 1960). This often makes it difficult to be sure of any aurochs-related meanings toponyms may have. In spite of this, Gysseling considers toponyms such as ‘Ursel’, ‘Oorbeek’ and ‘Oorschot’ to have derived from ‘oeros’12. The Dutch aurochs toponym ‘Voerendaal’, which Szalay (1915) came up 12 ‘Ursel’ from: ‘ur’ and ‘lauha’ (coppice on elevated sandy soil). ‘Oorbeek’ from: ‘ur’ and ‘baki’ (brook). ‘Oorschot’ from: ‘ur’ and ‘skauta’ (wooded piece of sandy soil in marshy territory).

81


82


with, has no value. The ‘voer’ in Voerendaal cannot be explained in a satisfactory manner. It probably has to do with water, but seems unrelated to the animal name ‘ur’ (Moerman 1956). Other alleged aurochs toponyms, such as ‘Ureterp’ (South-east Friesland) actually have other meanings, too. The name of this village, which, incidentally, is not situated on a terp (mound, knoll), means ‘elevated village’. Many aurochs toponyms survived further to the East. Szalay (1915) mentions quite a few for the German regions, such as ‘Urbach’, ‘Uraha’, ‘Ursberg’, ‘Urdorf’, ‘Urheim’ and ‘Urthal’. There are also a number of Swedish aurochs toponyms, such as ‘Urshult’, ‘Uranäs’ and ‘Ursberg’ (Isberg 1962). In Poland and Lithuania, where the aurochs survived longest, the animal was called ‘tur’ and ‘tauras’, respectively. Especially in Poland, these words may be found in many toponyms, including ‘Turady’, ‘Tureczka’, ‘Turza gni³a’ and ‘Turynka’ (Wrzeszniowski 1878). Another three aurochs toponyms are known from the Bia³owie¿a Forest: ‘Turycz b³oto’, ‘Turowisko’ and ‘Turzycowisko’13 (Rokosz 1995, Faliñski 1968). Lithuanian examples include ‘Taurage’, ‘Tauragnai’ and ‘Tauralaukis’ (in lit. W.J. Morrison). From Latvia, where the animal is called ‘taurs’, names likes ‘Taurupe’, Taurene’ and ‘Nitaure’ are known (in lit. Z. Andersone). Toponyms, especially if they are part of a series, may constitute definite proof that an animal lived in a certain area. Whether they can provide any information about the relation of the animal in question with its environment is far less certain. In the case of the aurochs, the names of villages, mountains, valleys and forests are basically vague indications, which often fail to lay a link with any type of vegetation. A toponym can only offer a clue if it is related to a very specific, as yet unchanged feature of the landscape. Thus, a single Polish toponym refers to a marsh (‘Turycz b³oto’ – aurochs marsh), and the ten Lithuanian and Latvian toponyms that have survived include four that represent names of rivers or lakes. These could be indicative of wet aurochs habitats. This is not entirely certain, however, since important rivers may simply have been named after important animals. Not a toponym, but a word in which a name for the aurochs may be found, is the term used for a species of grass, i.e. sweet grass (Hierochloe 13 However, ‘Turzycowisko’ is not a definite aurochs toponym, since the ‘turzyca’ part can either mean ‘aurochs cow’ or ‘sedge’.



odorata). In Poland, this grass is called ‘¿ubrówka’ (‘¿ubr’ - European bison) and it is used to add flavour to the vodka of the same name. The earlier name for this grass would have been ‘turówka’ or ‘turza trawa’, in which the word ‘tur’ (aurochs) may be recognized (Wrzeœniowski 1878). In Lithuania, it is known as ‘taurele’ (‘tauras’ - aurochs). Whether the aurochs used to enjoy eating this grass, which grows in marshy grassland, is not known. The European bison eats it occasionally, although it does not prefer it to other things. What is true for toponyms may also apply here: a striking plant (with a pleasant smell) may have been named for a striking animal.

6.3. The aurochs-European bison confusion After the aurochs became extinct in 1627, the only surviving large bovine species in Europe was the European bison, which held its ground for quite a while, especially in Eastern Europe and the Caucasus. In the late 18th and early 19th centuries, as the memory of the aurochs dimmed and more and more finds of bovine bones became known, a discussion started around the question whether the European bison was the only original wild bovine species in Europe, or whether there had been another, or even several others, at the same time (Wrzeœniowski 1878, Szalay 1918b). Adherents of the ‘single bovine species’ theory stipulated that only the European bison had inhabited Europe in historic times. The bovine bones that had been found were supposed to be from long extinct species; at best, they belonged to domesticated cattle that had gone wild again later on. Allegedly, the names ‘oeros’ and ‘wisent’, ‘Ur’ and ‘Auer(ochs)’, ‘tur’ and ‘¿ubr’ were all names for one and the same bovine species, the European bison. The main representative of this theory, Pusch (1840), thought he could substantiate it on the basis of philological and historical arguments. Initially, the increased number of bone finds in the 19th century did not result in a more accurate idea about the number of wild cattle species and their kinship. In 1878, after a more extensive study of historical documents, such as Inspection Reports,

83


84


correspondence and eyewitness accounts, as well as investigations into the reliability of the various authors, Wrzeœniowski was able to definitely conclude that besides the European bison, there had been at least one other wild bovine species in historical Poland, the aurochs. In retrospect, his research showed that in their writings, the Polish people had always distinguished clearly between ‘tur’ and ‘¿ubr’, aurochs and European bison, in contrast to the Germans, whose naming of the animal during and after its disappearance led to great confusion about the difference between these two animals. As was described in the above, the German language confusion was caused by the philological development of the Old German word ‘ur’. From the 11th to the 13th century, the ‘u’-sound was lengthened to ‘aue’, and ‘Ur’ became ‘Auer’ and ‘Auerochs’. This process coincided with the final disappearance of the aurochs from Germany (Szalay 1915). Strangely, the new word ‘Auer(ochs)’was transferred to the surviving European

Fig. 12. This is a picture that contributed to the confusion about the physical appearance of aurochs and European bison. The drawing was intended to represent an aurochs, but shows more of a crossbreed of the two species (beard and horns of the European bison). This picture was published in 1669 but is found in several publications by Conrad Gesner during the 16th and 17th centuries (Source: Library Wageningen UR).



bison. According to Szalay the aurochs was never called ‘Auer’; the word referred to the European bison from the start. In later times, the term ‘Wisent’ became obsolete in Germany. Whenever Von Herberstein (1557) wanted to demonstrate to his German readers that there was a difference between the aurochs and the European bison he would call the aurochs ‘aurox’ and the European bison ‘bisont’ (see Fig. 15). The latter word derives from the Latin ‘bison’, which was used to indicate the European bison in Latin writings. Halfway through the 16th century, when Von Herberstein displayed aurochs and European bison hides prepared as stuffed animals, he no longer knew the term ‘wisent’. Although this resulted in depictions that were far from perfect (the horns of both animals, among other things, do not present a true picture), the difference became clear in broad outlines. A comparable case of name confusion with reference to the aurochs took place in the Middle East. The Old-Hebrew word ‘reem’, related to the Mesopotamian ‘rimu’, means ‘aurochs’ (Unger & Ramey 1966, Duerst 1899, Gispen et al. 1975). During or after the extinction of the aurochs in these regions, this name was transferred to other animals. In present-day Arabic and Hebrew the word ‘reem’ is used to indicate the oryx (Oryx beisa), and in North Africa the word has been passed on to the rhim gazelle (Gazella leptoceros). Like the European bison, and in spite of the fact that these two animals will probably had names of their own, they were given the name of a formerly known, extinct animal that obviously appealed to the imagination. A comparable case is that of the modern Caucasian ibexes (Capra caucasica and Capra cylindricornis); local people call them ‘tur’, which in fact was the former Russian name of the aurochs. The words ‘Ur’/ ‘Auerochs’ and ‘Wisent’ (European bison) were not reinstated in Germany until the end of the 19th century (Nehring 1898b). In East Prussia, where the aurochs survived until ca. 1500, this confusion over names was especially awkward and unfortunate, since the aurochs and the European bison continued to exist side by side for a long time. There was no effort to clearly distinguish between such words as ‘Euwir’, ‘Uwer’, ‘Auer’, ‘Auerochs’ and ‘Wisent’. Well-known examples of this are the ‘Wisenthörner’ (horns of a European bison) already mentioned in Ch. 5.2, which turned out to be aurochs horns.

85


86


The confusion over names has often caused the many ‘Auerochs’references in the literature to be applied to the aurochs, although the European bison was concerned in almost all those cases. Claims of authors who assume that the aurochs still lived in a certain area, on the basis of the word ‘Auerochs’, should be regarded with suspicion. Descriptions by authors who did not themselves see the animal also lead to confusion. Descriptions of the animals would sometimes get mixed up, or be applied to the wrong animal. The description Von Fleming (1719) gives of the ‘Auer’ is an example. He adds a picture to his description, of an animal that is unmistakably a European bison. However, the description itself contains all kinds of details that have no bearing whatsoever on the European bison, but seem to refer more to the aurochs. It seems likely that the author did not know the European bison well, but still had written or verbal sources at his disposal at the end of the 17th century which provided information about both animals. Because of all the confusion about the identities of these species he unsuspectingly combined the information, writing, among other things, that the ‘Auer’ usually lives in marshes, and that its skeleton looks like that of a domestic cow. Both details definitely do not apply to the European bison, but may be descriptions of the aurochs. Although Wrzeœniowski showed that the aurochs and the European bison were two different species, this did not end the discussion about the total number of wild cattle species that had lived in Europe after the Pleistocene. Which domesticated cattle breed was supposed to have descended from which wild species was subject for debate. There were a series of theories about this, which used a corresponding multitude of Latin names (Wrzeœniowski 1878, Von Leithner 1927). The three most important theories were those of Nehring, Keller and Adametz (Von Leithner 1927). Nehring assumed that all domesticated cattle had descended from the aurochs. Keller and Adametz thought that another wild species besides the aurochs had been involved in the domestication. According to Keller, our long-horned cattle breeds had descended from the aurochs, and the short-horned ones from the banteng (Bos javanicus). Adametz (1925) was under the impression that besides the aurochs, Bos brachyceros had lived in Europe, a short-horned wild bo-



Fig. 13. Picture of an ‘aurochs’ (after Jonstonus 1657), probably drawn after the picture that Von Herberstein had commissioned (see Fig. 15).

vine species that was ancestor to our short-horned domestic cattle. Many of the bovine species presented in the various theories were described on the basis of a single, or a few, skulls. Initially, there was little material for comparison. Not until 1927 was Von Leithner, in an extensive study based on a much-increased quantity of bone material, able to test the various theories. He reached the definite, later generally accepted conclusion that there had only been one other wild bovine species in Holocene Europe besides the European bison: the aurochs. Afterwards, Adametz’ Bos brachyceros turned out to be a domestic cow. The many obscurities that had arisen at the time of the earlier, inadequate bone research turned out to have resulted from mistaken dates that had been attributed to the finds, and from faulty interpretations of the characteristics perceived. Initially, there was no knowledge of the obvious difference in size between the aurochs cow and the aurochs bull. Moreover, the bones of young animals could not be clearly distinguished from those of grown animals. In addi-

87


88


tion, as also sometimes happened with other domestic animals, the domestication process and the formation of new breeds had brought about all kinds of changes in the skeleton, particularly the skull, for example brachycephalism. Von Leithner still named the Pleistocene aurochs Bos trochoceros, to distinguish it from the Holocene Bos primigenius. Research by Lehmann (1949) showed that, apart from their horns, there was no demonstrable difference between the two types. Comparison of the lengths of the bones (see Ch. 7.1) indeed shows that there was no, or hardly any, difference in size between these two types. This research consigned the various theories about the origin of our domestic cattle based on more than a single wild ancestor to the wastepaper basket. Especially on account of their unfamiliarity with the longextinct aurochs, however, many people, including biologists, remain uncertain about the difference between the aurochs and the European bison. They think that these are two names for a single species, the European bison, and that this is the wild ancestor of our domestic cattle. Thus, the aurochs-European bison confusion has not yet been completely dispelled.

6.4. Nature and symbolism With reference to people, the word ‘character’ indicates the personality and nature characteristic of a certain person. Every human being has his own, individual character, shaped by heredity and education. Individual differences in character may also be seen in animals, albeit only after long-term behavioural research. In this context, ‘character’ does not refer to an individual aurochs feature, but to the complex of behavioural characteristics of the animals, especially as it was displayed in contacts with human beings. Such contacts took place during the hunt, or when people tried to chase the animals from their fields and pastures. The behaviour displayed by the animal on such occasions induced hunters to attribute certain characteristics, a certain nature to the animal, obviously always comparing its behaviour to that of people



in a similar situation, since traits such as ‘brave’, ‘fast’ and ‘proud’ are naturally always relative or subjective, and speak eloquently of the admiration of the hunter for the animal. It is remarkable that so many expressions and references regarding this ‘character’ have been saved about, specifically, the aurochs; this is not the case with, for example, the European bison. Roman writers already referred to the special character of the aurochs, the animal was commended in the Bible and mentioned by mediaeval man, and the Polish and Russian languages retain many expressions about the animal. Of the European bison, reports of mainly physical characteristics, such as its size, weight and hair, have survived. Through the ages, meetings with the aurochs and, more specifically, the aurochs hunt have obviously made a profound impression on people, more so than seems to have been the case with other animals. The Gilgamesj-epic (Mesopotamia, 3rd millennium B.C.) already mentions the heroic battle of the hero, Gilgamesj, with a bull sent from the heavens. The impression the aurochs made during hunting parties and its way of fighting, with its big horns, were partly responsible for the wide-spread bullfights, such as were and are still being held in Turkey, Egypt, on Crete and on the Iberian Peninsula. The aurochs hunt was practised in various ways. In prehistory, weapons definitely included the bow and arrow and the spear, since stone arrowheads were found among some of the aurochs remains. Sometimes the animal had survived an earlier injury, as demonstrated by an arrowhead that had concrescended with the bone. Caesar mentions the use of pitfalls for the aurochs hunt by the Teutons. In the Jaktorów Forest in the16th century, firearms were used besides spears, and probably also occasionally nets. The many references usually show that characteristics that are also greatly appreciated among humans were attributed to the aurochs. Thus, the Bible contains several passages (including Numeri 23, 22) in which the power of God is compared to that of the aurochs: ‘God has led them out of Egypt; his powers are like those of an aurochs14 ’. As early as 53 B.C., Julius Caesar said about the aurochs: ‘They have great power and speed; they will neither spare a human being, nor a wild animal they have seen. They [the Teutons] kill these 14 In original Hebrew texts, the word ‘reem’ is used to indicate the aurochs (Unger & Ramey 1966, Duerst 1899, Gispen et al. 1975).

89


90


after they have caught them in pitfalls. The young men season themselves to do this work and they practise this kind of hunt and those that have killed the most are greatly praised, after the horns have been displayed in public as evidence’. This passage shows that Caesar not only records the special physical capacities of the animal, but also describes how a lot of praise could be gained with the hunt of the animal, apparently because it was seen as a dangerous and fascinating hunt. Young men in particular could show their courage in this way. In addition to the maned European bison, Plinius (ca. 70) mentions the ‘exceptionally strong and fast aurochs’. In an Old English runic poem from the Early Middle Ages, the animal is called ‘fearless’, ‘fierce’ and ‘brave’ (see Ch. 9.1.3). The Russian language still contains several words and expressions that refer to the speed, the power and the courage of the aurochs (Dementiev 1958). The Russian word for aurochs can still be traced in verbs like ‘to hurry’ and ‘to be fast’. In Russian epic poems, the heroes are compared with the ‘powerful aurochs’, and they can change into aurochs to be able to run faster. The voice of the aurochs is also frequently mentioned. Its volume must have been impressive, certainly compared to that

Fig. 14. Two pictures that show how people used to imagine the ‘fierce’ character of the aurochs. The right one was first depicted in the ‘Hortus Sanitatis’ in 1491 (after Von Lengerken 1955), the left one is depicted on the ‘Carta Marina’ by Olaus Magnus from 1539. The location on this old European map is the area of the present East Poland/Byelorussia (Source: Library of the Amsterdam University).



of the rather taciturn European bison. In Russia, a person who behaves rowdily when drunk is said to be behaving like an aurochs, and a burly, stalwart woman is said to walk ‘like an aurochs cow’ (Pusch 1840). The Polish language also still has many words and expressions that recall this legendary animal. There are expressions like ‘strong as an aurochs’ (‘mocny/ silny jak tur’), ‘he walks like an aurochs’ (when a man or a horse walks proudly or fervently; Bonar in: Gesner 1602) and ‘a bloke like an aurochs’ (said of a person who is big and strong; Skorupka 1968). That language also contains the expression ‘tura skakaæ’, which means something like ‘to be untameable’ or ‘to act recklessly’ (Kar³owicz et al. 1919). All these expressions emphasize the alleged great power, speed and ferocity that the aurochs was able of displaying in certain circumstances. Naturally, these kinds of comparisons would sometimes give rise to exaggeration. Thus, Œwiêcicki (1634) reported about the aurochs: ‘In addition, they are so dazzlingly fast that after they have defecated, they even manage to playfully catch their manure with their horns before it reaches the ground’. This kind of tall story, in which almost supernatural characteristics were attributed to the animal, probably held a lot of appeal. From the exaggeration of character traits to the world of magic is only a small step, and the aurochs, like so many other animals, also played a part in the latter. As a natural result of the image of power and courage projected by the animal, magical powers were attributed to various body parts of its body. Accordingly, Plinius (cited by Szalay 1930) reports: ‘When the blood that a tick has sucked from the body of the black forest ox is rubbed onto the loins of a woman, she will become averse to the joys of lovemaking’. Apparently, this necessitated a powerful remedy in the shape of blood from a live aurochs. A passage from Schneeberger (in: Gesner 1602) shows that there was probably even greater interest in two other parts of its body. When the animal had been captured during the hunt, and was still alive and on the ground, the following scene could take place: ‘Farmers will continuously split pieces of wood, which they will throw against, slantwise and straight on top of it, thus surrounding the animal

91


92


while standing firmly on it, so that it will no longer move to resist, and while it is held thus and is still alive, they will pull off the skin which covers its forehead between the horns, and which by its curly hair makes the animal horrible to behold; when this has been ripped off they will finally kill it: immediately after it has been killed they will take out the heart, when they have cut this in half they will find a small bone15 in the shape of a cross, which they will send to the king together with the skin that has been torn off the head, and they greatly value both. Some say that this bone is advantageous to pregnant women, and to those who suffer greatly when giving birth’. Thus, parts of the heart (heart bones) and the head (forehead skin) were symbolic for the power of the animal. Possession of these objects gave the possessor strength and confidence. Schneeberger is the only author to mention the use of the heart bone. Several authors mention use of the skin of the forehead. Bonar (in: Gesner 1602) reports, without further detail, that ‘the skin’ is torn into belts, girded by which ‘pregnant women can give birth with miraculous ease’. Other authors, such as Schneeberger, Mucante, Von Herberstein and Œwiêcicki stress that the ‘belts for women’ in question were made exclusively from the skin that is situated on the forehead and between the horns of the aurochs. These belts cut from the skin of the forehead were very valuable and were sometimes presented by the Polish king as gifts to important persons. Thus, the diplomat Von Herberstein received two such belts from queen Bona, for services rendered. As the aurochs became more rare, these belts probably became more precious. The skin on the forehead was covered in curly hair, which must have given the aurochs a fierce look. Schneeberger already mentioned this before in his letter: ‘The forehead, because of the curly, frizzy hair, makes them terrible to behold’. Like Schneeberger and Von Herberstein (1557b), Œwiêcicki also reports that the skin of the forehead was torn off the aurochs while it was still alive: ‘Nothing is coveted more by the hunters than the middle part of the forehead with its twisted, shaggy hair………torn off dying aurochs after the skin has been cut loose in a circle’. 15 The heart of a bovine animal contains two small, so-called ‘heart bones’ (the ‘ossa cordis’; Sisson & Grossman 1953). In the calf, these still consist of cartilage, in the grown animal they have hardened to bone.



According to Œwiêcicki there was even a special Polish word for this piece of skin with hair: ‘turzywicher’ (‘aurochs-windstorm’), which was a clear reference to the fierce, curly appearance of this hair. It is possible that the exclusiveness of the belts was safeguarded by the fact that they could be recognized by these hairs. Regardless of all the magic it is doubtful whether the aurochs was really such a fierce and unpredictable animal as the above would seem to suggest. Both Œwiêcicki (1634) and Schneeberger (in: Gesner 1602) were able to report about its nature. Œwiêcicki wrote: ‘When not provoked, they will allow a human being or even a wild animal easy passage, when provoked they can be exceedingly fierce’. Schneeberger, who probably visited the last aurochs population at Jaktorów, reports more extensively about the animals: ‘An aurochs is not afraid of humans and will not flee when a human being comes near, it will hardly avoid him when he approaches it slowly. And if someone tries to scare it by screaming or throwing something, this will not scare it in the least, but while it stays in its place it will actually open its mouth, widen it and close it again quickly, as if it is making fun at the human for his attempt. When it is standing in the road or somewhere else, one must go around it, even if one is driving a carriage, since it will not move off the road by itself. When challenged they become very hot-tempered, but if the person who has provoked it stretches out on the ground, nothing bad will happen to him, since they spare those who are stretched out, just like lions, with remarkable kindness, but if he does not stop challenging them, they will attack with their horns and throw him up into the air with them’. Hunters appointed by the Crown, whose task it was to supervise them, visited the last aurochs population every day, so the animals were confronted with people on a daily basis. Moreover, although their frequency is not known, the king sometimes held hunting parties in which animals were killed. Between these hunts, occasionally bulls were shot that had either covered domestic cows, or too often displayed aggressive behaviour towards other bulls. Animals who had left the wooded areas to graze the adjoining fields and pastures were also sometimes herded back by the hunters with the help of dogs. Their food was sup-

93


94


plemented with hay in winter. On the one hand, therefore, the animals were used to people, since the latter were present on a daily basis in a protective capacity. On the other, however, man showed his aggressive character by killing the aurochs, driving them away or catching them. There was probably also some poaching, be it on a limited scale. The animals may have had reason to distrust humans, but this does not seem to have led to timidity. If we are to believe Schneeberger and Œwiêcicki, they were easily approachable and certainly not shy. Only if one provoked and tormented them could they become very bad-tempered and fierce. In that state they could throw humans, and even a horse and its rider, up into the air with their horns (Œwiêcicki 1634). In 1472 a Lord Chamberlain was given a sum of money by the Polish king in compensation for a wound that had been inflicted on him by an aurochs a year before (Heymanowski 1972). Such scenes are strongly reminiscent of the bullfights in Spain. Spanish fighting bulls have been held for centuries for the sole purpose of the final fight in the arena. For this purpose, bulls as well as cows are tested and selected at the age of two for their fighting spirit and courage (Fullton 1971). The animals that possess these characteristics to a sufficient extent go on to fight in the arena. During such tests, cows can reveal as much fighting spirit as bulls. In spite of this

Photo 6. One of the heart bones from the heart of a domestic cow. In the old days in Poland, people attached considerable value to the heart bones of the aurochs. They attributed great magical power to them (Photo: Beeldgroep Wageningen UR). In the background a division in centimetres is given.



continuous selection, during which the aptitude for aggressive behaviour seems to be fostered, these cattle have not become hyperagressive. The extent to which fighting bulls in breeding farms are approachable for people differs from farm to farm (author’s observation). They may be easy to approach and not aggressive on the one farm, and quite the opposite on the other. This may be related to the management of the farms and the social structure of the groups of bulls. As long as the bulls are in a familiar environment with companions they will not feel threatened, and will usually not be aggressive. If they are isolated, however, and taken into a different, unfamiliar environment, they will become stressed by the uncertainty of their situation. If they are hurt and provoked in addition to this, their mood can change into a blind fury. In that mood they are apt to attack anything that moves, not just a moving red cloth but also people, cars and even trains (Fullton 1971). Their fury may subside when the object that evoked the aggression no longer moves. Schneeberger already described how the aurochs would leave a person who was on the ground and who no longer moved, undisturbed. This is a known fact also among bullfighters. Although it does not apply in every case, the best chance to survive for a bullfighter who is injured and on the ground is to lie still (Fullton 1971). Running away will actually make the bull aggressive; for the animal, movement in itself is clearly a characteristic of the danger it feels it has to combat. The behaviour of the Spanish fighting bull, therefore, is quite similar to the behaviour Schneeberger described for the aurochs. If we look at the behaviour of domestic cattle gone wild, such as the Betizu in the South of France, we see that they are completely wild and timid (Van Vuure 1998), in the way we know about of the red deer on the Veluwe (Holland). This may have something to do with frequent disturbances in the form of hunting (on the Betizu), tourism and sheepfarming. These wild bovine animals are difficult to approach. Occasional confrontations with humans have taken place over there, when a cow has felt cornered by hunters. The Heck cattle, which are held in a feral state in the Dutch Oostvaardersplassen, inside gates and with occasional supplementary feed-

95


96


ing in winter, generally do not present any problems where encounters with people are concerned. They are completely used to human beings and easily approachable. The first Heck cattle, when it was released in 1938 at Rominten, caused a lot of problems in the beginning (Frevert 1957). The animals brought about devastation and attacked people. Later on, they became progressively wilder and eventually timid. In 1942, a number of these animals was transferred to the Bia³owie¿a Forest. There was an instance of a man allegedly attacked by an injured bull there (Klaus 1969). It looks as if the measure of disturbance determines the approachability of the animals. The more they are hunted and chased away, the more timid they become. Seen in that connection, the aurochs at Jaktorów had apparently become used to people and had had too few bad experiences with them to become timid. Œwiêcicki already suggested this at the time: ‘Now that the aurochs are kept in a limited wooded area, they unavoidably become tame’. A comparison can also be made with European bison. Of the European bison that lived near Bia³owie¿a in the beginning of the 19th century, in circumstances similar to those of the aurochs at Jaktorów (little hunting pressure, supplementary feeding in winter), Brincken (1826) writes: ‘The old European bison does not run from humans; on the contrary, it stops when it is approached, and does not budge; it only attacks man when it is irritated, in which case it is fierce and dangerous. In winter, one can approach the animal to twenty paces, and it frequently happens in this season that passers-by are forced to wait until it pleases the European bison to leave the road it is blocking before they can continue on their way. It is much more timid in the summer, since it can find food almost anywhere’. This description is strongly reminiscent of the behaviour of the European bison in the 20th century, in the same area, with similar stories of solitary animals that would block the road, and the same difference between summer- and winter behaviour (Stechow 1929).



Summary From the Old Germanic word ‘ur’ for aurochs, which was used everywhere in the Germanic regions, by way of the word ‘Auerochs’, evolved in Germany out of ‘ur’, the Dutch word ‘oeros’ (aurochs) finally developed in the 19th century. If the animal had survived in The Netherlands, it would now be called ‘uuros’. In many European countries, including Sweden, Germany and Poland, aurochs toponyms have survived: names of towns, rivers and mountains in which the local term for ‘aurochs’ may be recognized. In the Dutch regions, there probably are also still a few of such toponyms. Toponyms have proved to be of little value where the determination of the natural habitat of the aurochs is concerned. At best, it may be concluded that a toponym is a global indication of an animal’s presence. For centuries on end, there has been confusion about the possible coexistence of two wild bovine species in post-Ice Age Europe. Because of the inadequate descriptions and portrayals of the aurochs and the European bison, and because the word that was originally used for the aurochs, ‘Auer(ochs)’ had been transferred to the European bison, it was hard to determine the species of animal in question. Proof that the Holocene aurochs and the European bison were two separate species was not offered until 1878. From the descriptions and expressions that have survived it may be concluded that the aurochs, more specifically the aurochs bull, must have made a great impression on man in the early days. As a result, it was called fierce, courageous, brave and fast, and certain of its body parts played magic roles. However, descriptions of the last aurochs population at Jaktorów (Central-Poland) show that the animals there displayed little aggressive behaviour. Only when they were hunted or otherwise tormented could their calm behaviour suddenly turn into blind fury. The behaviour of Spanish fighting bulls may still be comparable to this.

97


98


Photo 7. The right horn of the last aurochs bull, who died in 1620. The horn was wrought with a mounting of gilded silver and was used as a hunting horn. The (Polish) text on the horn says: ‘aurochs horn – from the last aurochs bull – from the Forest of Sochaczew – from the wojewode of Rawa Stanis law Radrzieiowski – at that time starost of Sochaczew – in the year 1620’. In the 17th century, the horn was robbed from Poland by the Swedish; it is currently displayed in the museum Livrustkammaren in Stockholm (Photo Göran Schmidt – Livrustkammaren). §

Fig. 15. Pictures of woodcuts of aurochs and European bison in Von Herberstein (1557a). In his house in Vienna, this German diplomat displayed stuffed hides of aurochs and European bison to show the differences between the two species. At that time, the aurochs was hardly known anymore. Above the aurochs (left) it is written: ’People call it ‘bison’/ I, however, ‘aurochs’. Above the European bison (right) it is written: ‘People call it ‘aurochs’/ I, however, ‘bison’ (Source: Universitätsbibliothek Wien).



Photo 8. Banteng bull, cow and calf (Bos javanicus). Not only in the aurochs did a marked difference in coat colour between male and female animals exist (sexual dimorphism); in the banteng such a difference in colour occurs as well (Photo: T. van Vuure).

Fig. 16. Comparison of bulls and cows of the aurochs and modern cattle (after Boessneck 1957, modified and coloured). The differences in body size, udder size, coat colour, body proportions and horns are clear. See the text for coat colour nuances.

99

Black Cyan Page 100

100


7. CHARACTERSTICS OF THE AUROCHS A description of a new, recently discovered animal species will usually include a detailed summary of its dimensions, nuances in colour, sex differences and other more or less significant morphological characteristics. Researchers who try to reconstruct a long extinct animal species like the aurochs cannot fall back on any detailed descriptions that offer a reliable picture of its appearance. The people that were this animal’s contemporaries either did not think a description was important or, if they did, would commonly adopt descriptions given by others, not from their own observations. As the aurochs became more rare, an increasingly number of reports was made. In particular several eyewitness accounts of the last population near Jaktorów have been preserved, the most valuable of which are those of Von Herberstein and Schneeberger. Only when it was too late did people become aware that something special was disappearing; once the aurochs had become extinct, the collective remembrance of it also disappeared quite rapidly. Hardly any records are known after 1627. The material that forms the basis of aurochs descriptions consists mainly of a very large number of bones from the Pleistocene en Holocene periods. In addition to this, a number of horns and a tuft of hair constitute tangible remains. A number of pictures, both in black and white and in colour, which date from the last Ice Age to the 16th century, have also survived. It is no sinecure to form an unambiguous description of the aurochs on the basis of this material, as will become apparent from what follows. Descriptions, even from eyewitnesses, may be open to more than one interpretation and pictures may be based on subjective interpretations. Any hereditary material from the aurochs still present in domestic cattle must be regarded critically, since it is not always clear what truly stems from the aurochs and what has developed as a mutation after domestication.

Black Cyan Page 101

CHARACTERSTICS OF THE AUROCHS

Despite these difficulties it is still feasible to sketch a reasonably reliable picture of the aurochs, unclouded by the characteristics of its descendants today.

7.1. Size As the area that forms the habitat of an animal species is larger, and the period in which it lives there more protracted, the variations in the size of this animal, caused by the place and time of its existence, will become more marked. This is partly due to the climate, the nature of the habitat and the presence of predators and competitors, and is true, for example, of the African buffalo (Sinclair 1977) and the bison in North America (Geist 1996), as it was true for the aurochs. The periods in which the animal existed (Pleistocene and Holocene) together lasted about 1.5 million years, and the area of its distribution stretched from the Atlantic to the Pacific Ocean and from North Eurasia almost to the Equator. From the Pleistocene, bone remains and pictures (including those from Lascaux) have been preserved; from the Holocene there are also descriptions by contemporaries.

Comparison of the Pleistocene and the Holocene aurochs Depictions of the aurochs, such as those in the caves at Lascaux and Altamira, can provide valuable information about the appearance of the animal but are not particularly useful as a basis for determining its physical size. At best, these pictures will warrant the conclusion that the aurochs belonged to the larger category of animals. Depending on the space available for the drawing or on its importance for the artist in question, the animal was shown smaller or larger. Due to the absence of sufficient bone remains from the various periods of the Pleistocene in Europe, it is not yet really possible to present a clear picture of the developments in size undergone by the aurochs in these periods. From its appearance in Europe, during the first intergla-

101

Black Cyan Page 102

102


cial period, its distribution area shrank to Southern Europe during the ice ages, later to be expanded again to the north. Cerilli and Petronio (1991) examined the length and thickness of the metapodes (metacarpus16 and metatarsus16) of the aurochs to determine whether any changes in size had taken place during the various periods of the Pleistocene. Based on these measurements they thought it safe to conclude that the aurochs, from the time of its appearance in Europe, initially increased in size until it reached its maximum dimensions in the Riss Ice Age, subsequently to get smaller again. They quote a maximum metacarpus length of 275 mm during the Riss Ice Age. Degerbøl & Fredskild (1970), however, give 278 mm as the length of the metacarpus of the Holocene bull of Grejs Mølle, which would then be the larger. The average metacarpus and metatarsus lengths found by Cerilli & Petronio do give a slight difference in size in favour of the animals from the Riss Ice Age. However, this difference in size might also have been caused by the fact that a relatively larger number of metapodes were preserved from the Holocene than from the Pleistocene, which results in a larger proportion of smaller-sized bones in the Holocene period and produces a smaller average. Whether the metacarpus and the metatarsus are the bones best suited to determine the size of the animal by is uncertain. It might be better to include other bones of the skeleton in order to get a good picture of the possible changes in size. However, there is a desperate shortage of Pleistocene material for comparison, both in quantity and in quality, from areas in the north as well as the south. Only skulls have been collected in relatively large quantities. From these skulls, the obvious differences between the Pleistocene and the Holocene aurochs can be seen quite clearly. Von Leithner (1927) examined the characteristics of six Pleistocene skull fragments of bulls from Central Germany and compared them with Holocene skulls of aurochs bulls and cows from Sweden, Germany, Poland and Austria. He discovered that the smallest forehead width of these Pleistocene bulls was about 10 % larger than that of the Holocene ones. The greatest difference was found in the horn cores, however. The Pleistocene horn cores were noticeably longer and thicker than the Holocene ones and more oval in cross-section. According to Von Leithner, the girth of 16

Metatarsals of foreleg and hind leg, respectively.

Black Cyan Page 103


the horn core at its base was on average 20 % larger and the length of the horn core, measured on the outside, as much as 50 %. Based on their studies of Pleistocene aurochs from Italy (see Table 3), La Baume (1947) and Portis, quoted by the former, confirm that particularly the horn cores of the Pleistocene aurochs were noticeably bigger than those of the Holocene. If the data of Von Leithner, La Baume en Portis for the Pleistocene, as set out in Table 3, are compared with those of Degerbøl & Fredskild (1970) and Ruprecht (1976) for the Holocene, a noticeable difference is indeed revealed regarding the size of the horn cores, both for bulls and cows. With reference to the length of the horn core, it may be added that the Holocene horn from Sutton Hoo (England) measured 107 cm (Stone 1961). This horn, the largest found up to now, probably had a horn core of about 80 cm. From the little information available, which is particularly scarce for the Pleistocene aurochs, it may be concluded that the body measurements of the Pleistocene aurochs probably exceeded those of its Holocene counterpart, though not by as much as is often maintained. Von Lengerken (1955), for example, spoke of ‘Riesenure’ (giant aurochs) when he heard about the larger Pleistocene horn cores. If the body size had been in proportion to the horn size, they would have been extremely large animals indeed. This proportionality need not always be a fact, however. From the evolution of the Pleistocene bison in North America forms are known with very large horns, and others with small horns, though the size of their skulls and bodies were about the same (Guthrie 1990, p.188). As far as the smallest width of the forehead is concerned, there seems to be little difference in size between the Pleistocene and the Holocene forms. Since Von Leithner’s material is rather too limited (6 skulls) and La Baume (and Portis, whom he quotes) do not give any averages, it is difficult to make a comparison between the Pleistocene and the Holocene skulls. Apart from the measurements of the horn cores, the other dimensions of the skull do not as yet warrant the conclusion that the Pleistocene aurochs was very much larger than the Holocene. The Pleistocene skulls are usually too incomplete to take accurate measurements

103

Black Cyan Page 104

104


of their basal lengths. Moreover, a somewhat larger subspecies may have been alive during a certain period in the Pleistocene in Italy (the origin of La Baume and Portis’ skulls). Comparison of the Pleistocene material with Holocene material from Central Europe is not yet sufficiently possible, since the quantity of Pleistocene material is still too limited for such a comparison. The possible height of the withers of the Pleistocene aurochs will be discussed in more detail at the end of this chapter. Table 3. Comparison of the skull- and horn core characteristics of the Pleistocene and the Holocene aurochs, for the European distribution area.

Characteristic

Pleistocene (data: Von Leithner (1927), La Baume (1947) and Jánossy & Vörös (1981)) (material: 38 skulls and 20 separate horn cores)

Holocene (data: Degerbøl & Fredskild (1970) and Ruprecht (1976)) (material: 98 skulls)

335 - 485 250 - 275

275 – 445 185 – 275

horn core length † (mm) ‡

630 - 1200 490 - 590

450 – 820 330 – 530

basal length of the skull (mm)17

580 - 600 (3 skulls) —-

525 – 612 455 – 535

238 - 294 195 - 214

207 – 289 170 – 205

horn core girth (mm)

† ‡

† ‡

smallest forehead † width(mm)18 ‡

Up to this point, the subject of discussion in this chapter has been the primigenius-subspecies of the aurochs in Europe. Much less is known about the namadicus-subspecies from India, which lived there during the Pleistocene. Most of those finds have been skulls. On the basis of similar skulls, Grigson (1973) has been able to indicate differences with 17 Length measured from the tip of the incisive bone (Os incisivum), across the roof of the palate, to the lower edge of the occipital gap (Foramen magnum). 18 Measured above the eye sockets at right angles to the longitudinal axis of the skull.

Black Cyan Page 105


the Holocene primigenius. The most noticeable differences were in the dimensions of the skulls and the horn cores. The maximum horn length of the bulls was 1150 mm; comparable, therefore, with what was found in Europe during the Pleistocene. On the basis of the skull, Grigson discovered a great variation in size. The small namadicus-specimens were significantly smaller than even the relatively small female Holocene primigenius. It is true that none of the larger namadicus-specimens had dimensions outside the limits for the Holocene male primigenius, but on average they were smaller. Apparently the Indian Pleistocene aurochs were just as varied in size as the European ones, but they were smaller on average. It may be expected for the southern species of aurochs to have been smaller than the northern ones. Any animal species in the Northern Hemisphere is usually larger as it lives more to the north than its counterparts that live more to the south. For a long time, Bergmann’s law was quoted as an explanation of this; it states that larger animals are better resistant to cold than smaller animals on account of their relatively smaller body surface. At present, however, this rule is very much under discussion (Partridge & French 1996). It is definitely not true for all animals (e.g. weasels; Ralls & Harvey 1985) and whether temperature is a predominant factor is also highly debatable. Thus, although the northern aurochs was larger than the southern species, this was not true of the horns of these animals, which were actually larger in the Indian species than in the populations living more to the north. On average, therefore, the Pleistocene Indian species was smaller than both the Pleistocene and Holocene species in Europe, but had horns as large as those of the European Pleistocene aurochs. The social function of the horns is also likely to have influenced their size, just as it influenced other external features (Guthrie 1990).

The Holocene aurochs To determine the size of the Holocene aurochs we have pictures, descriptions and bone remains. Pictures cannot be used for this purpose because there are no sizes to compare against, and descriptions only

105

Black Cyan Page 106

106


give a very global indication. The most reliable, therefore, are conclusions based on the size of the bones and on the skeletons constructed on the basis of the latter. Our earliest description of the aurochs, from the Roman general Julius Caesar, dates from 53 B.C. His accounts of the battle against the Gauls and the Teutons also include descriptions of several animal species that inhabited the Hercynian Forest19. One of them was the aurochs, of which Caesar says (Book VI, Ch. 28): ‘These are little smaller than elephants in size….’ It is highly questionable whether he ever actually saw the animals in the flesh. He had also been told about characteristics of other animals there that were far from the truth. Elk, for example, were allegedly unable to bend their legs, so they had to sleep standing up. His description does seem to indicate that the aurochs must have been an impressive animal, which was of particular interest to the Germanic tribes. The aurochs emerges just as impressively, though hardly realistically, from a description by the Spanish Arab Abu Hamid al Andaluzi al Garnati (Bartosiewicz 1999), who visited Hungary in the mid-12th century and describes the aurochs there as follows: ‘A large wild bovine animal is living in Bašgird; it is as big as an elephant ……. Its horns are big and long, like the tusks of an elephant’. Another report dates back to the 14th century and was written by Thomas Cantipratensis, who describes a number of European animal species (Hilzheimer 1910). After his description of the European bison he goes on to mention the aurochs: ‘There is another species of European bison, which the Poles call ‘thuroni’ [aurochs], smaller in size but faster’. Whether this author had seen the animal himself or heard about it from others is not certain, either. A similar case of quoting dubious sources is presented by Cramer (Daniel Cramerus) (Hilzheimer 1910). Around 1600, this author reported a hunting trip in Pomerania in 1364 during which a European bison had been killed that had allegedly been said to have been larger than an aurochs. Cramer’s report turned out to be based on reports from writers who had adapted the original story according to their own imagination (Szalay 1938, p. 50). The original version made no mention of either the year or the comparison between the European bison and the aurochs. There are still more ‘descriptions’ that are too unreliable to men19

This large forest stretched from the Black Forest (Germany) to Romania.

Black Cyan Page 107


tion. Szalay (1915, 1938) analysed many of these texts and checked their truthfulness. Von Herberstein (1557), who actually saw hides from the aurochs and European bison, does not mention any differences in size. Two comparisons of the size of aurochs and domestic cattle have survived. The first is Schneeberger’s (in: Gesner 1602), who may well have visited the last aurochs population himself. He reports: ‘They look a lot like domestic cattle, but they are much larger….’ and: ‘The cows are smaller than the bulls and not as long’. Œwiêcicki (1634), who was probably also an eyewitness, wrote: ‘They are much larger than our own domestic cattle, but their build is very similar’. All in all, these descriptions offer little to go on if one wants to determine the actual size of the aurochs. The most we can conclude from them, if Cantipratensis, Schneeberger and Œwiêcicki are to be believed, is that the aurochs was much bigger than domestic cattle but smaller than the European bison. This is obviously a rather vague indication since there was a significant margin between the size of 16th century domestic cattle (which were smaller than present-day cattle) and that of the largest European bison.

Fig. 17. Diagram in which the lengths of horn cores of the Holocene aurochs (‘B. prim.’) and domestic cattle (from the Neolithic) (‘B. t. dom.’) are compared. The horn cores originate from Denmark, England and Germany (the different-colour squares) (after Degerbøl & Fredskild 1970).

107

Black Cyan Page 108

108


Schneeberger observed what Von Leithner later confirmed for the first time: on the basis of their bones, a difference in size between the aurochs bull and the aurochs cow may be ascertained. This phenomenon, which is called ‘sexual dimorphism’, was confirmed by subsequent authors (including Degerbøl & Fredskild 1970, Boessneck 1957). It did not just occur in the aurochs but may also be seen in bovine species that still exist, such as banteng, gaur, yak and bison. Sexual dimorphism can be seen, not just in size but also in a difference in colour between the sexes, as was the case for the aurochs (see Ch. 7.2). This difference in size is not just confined to certain parts of the body, but may be found in all bones, not just in the horns or in the size of the head but also in the bones of the limbs, the shoulder blades and the vertebrae (see Fig. 17 (horn core lengths) and Table 4 (bone lengths)). Table 4. Comparison of the lengths of bones (in mm) from the limbs of the Pleistocene and the Holocene aurochs. The sex of the Pleistocene finds is not known.

Bone

humerus radius ulna metacarpus femur tibia metatarsus

Pleistocene (Germany and England). According to data from Jánossy and Vörös (1981), Vörös (1987), Lehmann (1949) and Döhle (1990) (mm) †† and ‡‡ 395 – 47120 370 – 434 247 – 285 485 – 510 465 – 543 278 – 328

Holocene (Central and Western Europe). According to data from Degerbøl & Fredskild (1970), Ekman (1972), Martin (1990) and Van Es (1990) (mm) †† ‡‡ 386 – 440 345 – 392 461 – 500 239 – 278 470 – 524 452 – 488 270 – 315

355 - 367 315 - 345 407 - 465 230 - 253 427 - 445 400 - 420 268 - 283

Sexual dimorphism is least apparent in the length of the row of molars. That of the aurochs cow is entirely inside the range of that of the aurochs bull (Degerbøl & Fredskild (1970). Grigson (1978), too, observes that there is hardly any difference between the two sexes with 20 This size (471 mm) is the probable greatest length (GL) of the humerus in the Neumark-Nord aurochs. It was determined by dividing the height of its withers (195.1 cm; Döhle 1990) by 4.14 (the humerus conversion factor in Matolcsi’s method; see Ch. ‘The height of the withers’).

Black Cyan Page 109


regard to this characteristic. In the bull the length varies from 146 to 183 mm (167 average), in the cow from 148 to 169 mm (159 average). As regards the width of the molars there is hardly any difference, either. In modern domestic cattle, sexual dimorphism between bulls and cows still exists; the difference in size has decreased, however. Apart from the length of the row of molars, little or no overlap can be seen in any of the bone characteristics of the aurochs bull and the aurochs cow; in domestic cattle, however, there is always an overlap, and a much bigger one than in the case of the aurochs (Grigson 1978; see Ch. 8). After domestication, not only did body size decrease but also the difference in size between the sexes. Sexual dimorphism is found particularly in animals living in herds, whose adult males only spend part of the year with their cows and young. This social structure is familiar to us from red deer, wild boars and elephants. The males only join the other animals during the mating season. They do not play a leading role in the herd, unlike horses, whose males and females stay together and where the difference in size (between stallions and mares) is small.

The height of the withers The body size of the original aurochs is an aspect that has long appealed to the imagination, and has been a subject of discussion for just as long. The Heck brothers, who started their breeding-back experiments in the 1930s, often describe the wild species in meticulous detail as to colour, body weight, horn shape and choice of food. These descriptions often come unaccompanied by any mention of their source, and may well based on personal interpretation. They are noticeably vague, however, about the size of the animals and the differences in size between the sexes (H. Heck 1934, 1949; L. Heck 1934, 1952a, 1952b). Scientific research of bone material up to around 1930 had revealed that the wild aurochs was significantly larger than the domestic cattle of the same period. The Heck brothers suggested that there was a geographic diversity in the height of the withers, but made no further definite state-

109

Black Cyan Page 110

110


ments about this. A possible explanation for this may be that it did not suit them to be pinned down on any definite size, since they may well have suspected that there were no animals of aurochs proportions among the species of domestic cattle they used in their experiments (see Ch. 10.3). Not until 1980 did Heinz Heck announce that the larger aurochs subspecies must have had withers of almost 2 m high (Heck 1980). Herre (1953) was the first to give a reasonably precise description of the animal’s size in a critical review of the breeding-back experiment. He portrayed an aurochs bull with a withers height of 2 m and an aurochs cow of 1.80 m, but failed to mention how he had calculated these results. Von Lengerken (1955) adopted Herre’s suggestion, but commissioned a painting to be made on the basis of bone material. This shows an aurochs bull with a shoulder height of almost 2 m. A few years later Boessneck (1957) depicted an aurochs bull and an aurochs cow with withers heights of 1.75 m and 1.50 m, respectively (see Fig. 16). These pictures were made on the basis of extensive comparative bone research, in which the author was one of the first people to use the length of the metacarpus and metatarsus in order to determine the height of the withers. In an attempt to bring the continual uncertainty about the height of the withers of the aurochs and prehistoric domesticated cattle to an end, Matolcsi (1970) undertook an experiment in Hungary to show a possible connection between the shoulder height of cattle and the length of certain bones of the limbs. Of the 186 live animals that made up the original material for this experiment, about half were Hungarian Steppe cattle; the others included Ukrainian Steppe cattle, as well as Braunvieh and Charolais and Jersey cattle, both cows and bulls. First, the shoulder height, body length and body weight of every animal were ascertained. After the animals had been slaughtered, the bones from the limbs and vertebrae were boiled and cleaned, after which they were used to determine their relation to the height of the withers, body weight and body length. Only the determination of the height of the withers will be discussed here. The percentages of all the bones in the shoulder height of the animal were determined. On the basis of these data, Matolcsi found a connection between the length of the bones from the

Black Cyan Page 111


limbs and the height of the withers. The conversion factors for the bones in question are listed in Table 5 below. Table 5. Conversion factors for the various bones from cows and bulls to determine the height of the withers (according to Matolcsi 1970).

Bone

humerus radius metacarpus femur tibia metatarsus

Conversion factors †† ‡‡ 4.14 4.30 6.33 3.23 3.45 5.62

4.14 4.30 6.03 3.23 3.45 5.33

The length of a certain bone, multiplied by the conversion factor for that bone, should give the height of the withers. This table shows that the humerus, radius, femur and tibia form the same percentages of the shoulder height in both sexes. A difference according to sex can only be seen with the metacarpus and metatarsus. Matolcsi considers the practical value of the metapodes greater than that of the other four bones. At the same time, he is aware of the limitations of the use of conversion factors for calculating the height of the withers. In actual fact, they only reflect the commonest proportions in the breeds investigated and cannot simply be applied to their wild ancestors. Other authors have attempted a similar approach to this problem. In their calculations, Bergström & Van Wijngaarden-Bakker (1983) and Bartosiewicz (1985) all came to the conclusion, however, that of all bones, the metapodes are not the most suitable for determining the shoulder height, as Matolcsi thought, but rather the socalled long bones of the limbs (humerus, radius, femur and tibia). In the early growth of cattle, the bones that are furthest from the body, such as the metacarpus and metatarsus, are the first to reach their full length, unlike the long bones near the trunk, which only reach their full length at a later stage. The bones in the trunk part of the

111

Black Cyan Page 112

112


skeleton are the last to stop growing. The metapodes stop growing well before sexual maturity sets in, while the other bones only stop growing when this phase has been reached or completed. The most marked sexual dimorphism where absolute length is concerned, therefore, is found in the bones closest to the trunk; the least obvious sexual dimorphism in the metapodes, although clear sexual dimorphism may be seen in the thickness of these bones. During the entire period of growth, the contribution of the long bones to the height of the withers is very consistent. Matolcsi already discovered that the percentage share of these bones in the withers height was the same in both sexes. As far as the influence of breed is concerned, it should be stressed that these data refer to the usual medium-sized breeds. It is still not known whether these conversion factors also apply to extremely large or small breeds and specimens. Whether these conversion factors can simply be used for the wild ancestor of domestic cattle is as yet unknown. The sexual dimorphism in aurochs is more marked than in domestic cattle where size is concerned, in addition to which the wild version reaches maturity later than domesticated cattle (see Ch. 8). It so happens that growth stops at an earlier stage of life in domestic cattle, which makes them sexually mature at an earlier age (Steehouwer 1987). Such growth processes may bring about other proportions and conversion factors for the wild species. To be able to ascertain whether the conversion factors found by Matolcsi are applicable to individual aurochs specimens, complete skeletons as well as bones that belong to one and the same foreleg are available for research. By applying the conversion factors to the individual bones from a single animal, it can be shown whether identical results for the original shoulder height may be obtained for each individual animal. Table 6 shows the results of these calculations. The shoulder heights of the animals were calculated according to Matolcsi, on the basis of the foreleg bones humerus, radius and metacarpus, which are directly related to the withers height. The aurochs used for these calculations included three mounted skeletons; moreover, measurements of two domesticated Simmentaler bulls, one small and one large (Koch 1927), were recorded.

Black Cyan Page 113


First of all, the table shows that conversion of the length of the three bones in question to the original height of the withers hardly ever produces the same shoulder heights, although according to Matolcsi, this should be the case. In a few cases two bones give the same height; apart from that, results vary. It also appears, as Degerbøl had already discovered, that the proportions of the various bones of the limbs are not constant; there may be differences between animals. A large humerus need not necessarily be combined with a large radius and metacarpus. The large aurochs bull from Braunschweig, for example, has a very large humerus and radius, but only a medium-sized metacarpus. A large humerus may also be combined with a medium-sized radius. Even the left and right forelegs of a single animal can have humeri and radii of different lengths, as may be seen in the Düsseldorf aurochs bull. Determination of the height of the withers on the basis of a single bone seems inadvisable, therefore, if these variations in the lengths of the bones are taken into account. Such a bone should always be seen in combination with other bones from the same animal. Table 6. Calculation of the height of the withers on the basis of various bones, based on Matolcsi’s conversion factors (1970).

Place found and bone

Length (mm)

Britsum † humerus radius metacarpus Terp † humerus radius metacarpus Taastrup ‡ humerus radius metacarpus

(Van Es 1990) 386 372 239 (Degerbøl 1970) 430 363 251 (Degerbøl 1970) 367 345 245

Withers height according Actual to Matolcsi (1970) (cm) height (cm)

160 160 151 178 156 159 152 148 148

113

Black Cyan Page 114

114


Table 6. Continued

Place found and bone

Length (mm)

Sørø † humerus radius metacarpus Store Damme † humerus radius metacarpus Ullerslev ‡ humerus radius metacarpus Düsseldorf † humerus l. r. radius l. r. metacarpus Braunschweig † humerus radius metacarpus ‘Aktionär’ † humerus radius metacarpus ‘Hartmann’ † humerus radius metacarpus

(Degerbøl 1970) 410 362 258 (Degerbøl 1970) 410 363 258 (Degerbøl 1970) 359 320 246 (Martin 1990) 405 390 350 360 245 (Martin 1990) 440 392 259 (Koch 1927) 413 347 239 (Koch 1927) 373 313 233

Withers height according Actual to Matolcsi (1970) (cm) height (cm) 172 (skeleton) 170 156 163 175 (skeleton) 170 156 163 150 (skeleton) 149 138 148 168 162 151 155 155 182 169 164 167 (alive) 171 149 151 150 (alive) 154 135 148

Black Cyan Page 115


Matolcsi’s conversion factors are based on averages and are therefore not intended for individual specimens. The table also shows that the radius and metacarpus alternately give the lowest withers heights and that the humerus always produces the greatest21. Moreover, it turns out that the humerus gives the shoulder height closest to those of the two domesticated bulls, as well as those calculated by Degerbøl with regard to the mounted skeletons. It seems reasonable to conclude, therefore, that application of the length of the humerus gives the closest approximation to the height of the withers in cattle. In itself, the fact that one specific bone correlates well with the withers height is not an unknown phenomenon. In the same way, the humerus was discovered to be a useful tool for calculating the height of the withers in domesticated sheep (Von den Driesch & Boessneck 1974). The radius in horses, however, came closest to the actual results. Table 4 may be used to calculate the limits of the shoulder heights of aurochs. The data suggest that the height of withers in bulls may vary from 160 to 182 cm and that of the cows between 147 and 152 cm. We can compare these data with the results Degerbøl obtained on the basis of the mounted skeletons and the bone combinations from the foreleg he used for comparison. He discovered an average of 175 cm for the bulls and a variation of 148 to 157 cm for the cows. Degerbøl’s calculations result in a shoulder height of a few centimetres higher than the one calculated according to Matolcsi’s method. In both authors, the difference in the height of the withers between cows and bulls is about 20 cm. The variation in shoulder height appears to be more marked in bulls (20 cm) than in cows (10 cm). If we were to calculate the withers height of the Pleistocene aurochs, by means of the conversion factor for the humerus, the maximum humerus length of 471 mm (see Table 4) would engender a shoulder height for bulls about 13 cm higher than that of the Holocene aurochs22. This cannot be calculated for the cows because of a lack of Pleistocene data. As a matter of fact, both methods are open to criticism. Matolcsi’s method was originally derived from and intended for domesticated cat21 Van Wijngaarden-Bakker (1997) found this phenomenon in Heck cattle, too. Döhle (1990) got roughly the same result when he compared the various bones of the Pleistocene Neumark-Nord aurochs to those of 17 other Pleistocene and Holocene aurochs. Apart from a few cases, humerus lengths yielded the greatest shoulder height; only in three cases did metatarsus lengths result in one that was ca. 1 cm greater. 22 From the length (GLC) of the (partial) humerus in the research by Döhle (1990; see footnote above) a shoulder height of 195.1 cm was ascertained using Matolcsi’s method.

115

Black Cyan Page 116

116


tle; it may not take into account cattle with a more marked sexual dimorphism, which reach maturity later. As to Degerbøl’s method, it is not always clear whether a skeleton has been mounted correctly, in addition to which there is no agreement on how many centimetres have to be added for hooves, muscles and skin to determine the actual height of the withers from the skeleton height. If one looks at close relatives of the aurochs among the Bovini, their measurements are found to be comparable. The shoulder height of the European bison and the length of its bones are well known, for example (see Table 7). Table 7. Length of bones of the European bison (Bison bonasus). According to data from Koch (1927) and Lehmann (1949).

Bone

humerus radius metacarpus femur tibia metatarsus

Lenght (mm) ††

‡‡

311 - 368 280 - 339 196 - 218 357 - 416 374 - 439 240 - 268

313 – 430 302 – 375 204 – 224 370 – 469 392 – 491 250 – 276

If it is compared to Table 4, this table shows that the bones of the European bison are either the same length or shorter than those of the aurochs, both for cows and for bulls. The height of the withers of the European bison bulls varies between 158 and 188 cm (Krasiñska & Krasiñski 2004) and is on average 172 cm 23. In the cows it varies from 134 to 167 cm, with an average of 152 cm; the rump is much lower, however. The big difference between the rump height and the shoulder height of the European bison can be largely explained by the large neural spines of the thoracic vertebrae. This may also explain the large variation in withers heights. In the banteng bull, whose shoulder height · are likely to be Measurements of the height of the withers and the body weight of the European bison in Bia §lowieza fairly precise, since the local people have years of experience in catching and transporting these animals, which gave them the opportunity to measure them It is not clear how the measurements of the banteng were obtained; they were probably taken from dead animals lying on the ground.

23

Black Cyan Page 117


barely exceeds that of its rump, the height at the withers varies from 150 cm to a maximum of 191 cm (Hoogerwerf 1970). The fact that its shoulders are about the same height as its rump makes the banteng comparable to the aurochs, the shoulder height of which is also known to have barely exceeded the height of the rump. This can be seen in the rock paintings at Lascaux, in the picture of the so-called Augsburg aurochs (see Fig. 23), and in the way Von Herberstein had an aurochs hide set up in Vienna (see Fig. 15). Little is known about the size of the neural spines of the thoracic vertebrae in aurochs, since these have never been the subjects of thorough research. One of the reasons for this is that few have been preserved in good condition. From these few, however, it appears that said spines were clearly shorter, and broader (seen on the front), than those of the European bison (Ekström 1993). Since the aurochs occupied a large habitat, size differences within the species will have occurred on the basis of their geographical position, as is common in other animal species. Only from Europe, sufficient bone finds are known to be able to make statements about this aspect, though conclusions may only be drawn very carefully even here. In the first place there was probably a difference in size in Europe between animals in the northern parts and those from areas more to the south, in that the latter were smaller. Von den Driesch & Boessneck (1976) examined bone material from the Iberian Peninsula and compared it with bone material from other parts of Europe. They came to the conclusion that the size of the aurochs there was comparable to that of aurochs in areas like Hungary and Switzerland. The animals from more northerly areas like Denmark were bigger. The actual difference in the height at the withers cannot be deduced from their comparisons, however, and whether this difference in size was caused by a different climate, different food or competition from other kinds of herbivores has not as yet been explained. In the larger part of Europe, the Holocene aurochs tended to be smaller in build than the Pleistocene one, but this does not seem to hold true for the Iberian Peninsula. From research by Estévez and Saña (1999) in that region, the average size of the aurochs there would appear to have increased gradually after the Pleistocene period, all during the Holocene period up into the Bronze Age. An explanation for this development has not yet been found.

117

Black Cyan Page 118

118


Just as there were probably differences in size between northern and southern populations, allegedly there were also such differences between the populations in Western Europe and Eastern Europe (Lasota-Moskalewska & Kobryñ 1990): the latter were somewhat larger. These authors compared bone material from Northwest-, North- and East European countries. The difference seems to apply to female animals in particular. A similar tendency is also known for red deer and wild boars. It is not improbable, therefore, that the same went for the aurochs, though to what extent the western and eastern populations differed from each other is not yet entirely clear. This kind of research requires a great deal of bone material from all the countries involved and requires that measurements are obtained in the same way in all of them. As yet, very little is certain about the size of the aurochs in the Middle East, due to lack of sufficient bone material for comparison. As for North Africa, based on only few long bones Linseele (2004) cautiously concluded that aurochs there were somewhat smaller than the Portuguese aurochs researched by Von den Driesch & Boessneck (1976). As part of his research into aurochs body size, Matolcsi also attempted to determine the length of the body on the basis of bone lengths. His method proved too uncertain a basis for conclusions. Skeletons would provide a better one; Matolcsi quotes research by Duerst. The latter examined a number of skeletons in England, Germany and Sweden and came to the conclusion that the length of the trunk was slightly more than the height of the withers. He expressed this in the equation: shoulder height . 100%, trunk length which gives a result between 91.2 and 98 %. This shows that the aurochs was almost ‘squarely’ built24. Body proportions have changed in contemporary cattle breeds, whose shoulder heights have become smaller in the absolute sense while their trunk lengths have become relatively larger. The ratio of the two varies from 73 to 94 % (Matolcsi 1970), which makes domesticated cattle longer (‘longer in the body’) than their wild ancestors. This change is also shown by the aurochs tracks found on the coast near Liverpool (see Ch. 7.4, ‘Hooves’). From this print it appears that the hind legs 24 The reason for this square build in comparison to that of domestic cattle probably lay in the greater able-bodiedness, manoeuvrability and speed that would be required in a wild state.

Black Cyan Page 119


were always placed in the prints of the forelegs, as is common with roe deer and red deer. With domestic cattle, the prints of the forelegs are always in front of those of the hind legs (Lawrence & Brown 1974), because the legs are relatively shorter than those of aurochs. Mounted aurochs skeletons and the picture of the Augsburg Aurochs (see Fig. 23), for example, also show that the front part of the body (the forehand) is clearly more strongly developed and heavier than the back part (hindquarters). This difference has become much smaller in contemporary domestic cattle (see Fig. 16). With regard to the weight of the animals, Matolcsi (1970) and Bergström & Van Wijngaarden-Bakker (1983) all tried to make a connection between the weight of certain bones in domesticated cattle and their total body weight. However, this connection appeared to be strongly influenced by age, breed and level of nourishment, so it was difficult to determine. Moreover, the bones in question were fresh, which made it difficult to establish any possible relationship that would apply to the (sub)fossil aurochs bones that had been found. If we look at the weight of full-grown European bison and banteng, it appears that the weight of free-living European bison cows varies from 340 to 540 kg (from 320 to 640 kg in zoo animals) and that of free-living European bison bulls between 436 and 840 kg (from 580 to 920 kg in zoo animals) (Krasiñska & Krasiñski 2004). Hoogerwerf (1970) records a weight of 500 to 920 kg for banteng bulls. Weights obviously varied considerably, and although it seems probably, on account of its comparable size, that the weight of the aurochs was similar to these, this is still conjecture.

Summary One aspect of the aurochs that has long been a focus of interest and a subject of discussion is its size. Early descriptions give only vague indications about this. Numerous aurochs bones have survived, even a number of almost complete skeletons. Since the dimensions of these bones are well known it has proved possible to reconstruct the original size of

119

Black Cyan Page 120

120


the animal from them. For a long time the metapodes (the metacarpal and metatarsal) were used for this, since a large quantity of these had been found. From the above analysis, however, it would appear that the length of the humerus is probably a more appropriate tool, since it provides a more reliable measurement for the height of the withers, which has been proved to have varied between 160 and 180 cm for the (Holocene) aurochs bull, and to have been around 150 cm for the (Holocene) aurochs cow. For the Pleistocene aurochs the height of the withers was probably well over 10 cm greater.

7.2. Horns Although a great number of aurochs horn cores have been found, only a few dozen of the horn sheaths that originally covered the horn cores have survived. Generally speaking, bone material is more successfully conserved than the soft keratin of the horns. Of the horns that still remain, some were dredged up from the North Sea (Stolzenbach 1984, Bosscha Erdbrink 1986, Nederlof 1995), while others stem from peat sediments (Nehring 1900, Ewald & Laurer 1911). As it happens, certain acidic peat layers do conserve the keratin, but not the bones (which mainly consist of lime). However, the vast majority of the horns were saved because they were used for specific purposes, given as gifts or inherited. In this way they were kept dry. They were sometimes used as signal-horns, like in the Roman army (Hilzheimer 1910, p. 74) or during the hunt (Pöschl & Hasenhüttl 1995), or to contain oil or other liquids during church ceremonies. Mostly, however, they were used as drinking cups for beer or mead. For this purpose, they were often beautifully tooled with gold or silver and given as presents to important persons. Caesar already mentions this as a Germanic custom. In this way, the attractively wrought aurochs horns of Sutton Hoo (Stone 1961) were provided as burial gifts for the king. In later periods, like the Middle Ages, this custom was retained (Bökönyi 1956). As the aurochs became more rare, the value of the horns probably increased, and it is likely that only the nobility still used them as interment gifts, in the same way that the aurochs hunt became their privilege.

Black Cyan Page 121


Almost exclusively aurochs horns were beautifully tooled; in a few cases, horns of European bison were used (Grüß 1931). The latter may be distinguished from aurochs horns by their lesser length, different shape and different colour, i.e. solid black (Pucek 1986). On account of its limited length (33.5 cm; see Table 8), the drinking cup from the Viking period, which is among the church treasures of the Onze Lieve Vrouwe Kerk at Maastricht (Roes 1940), cannot be regarded as an aurochs horn. The European bison, or a domesticated cattle breed, would be more likely candidate for its origin. Four aspects are important for the description of aurochs horns: the size, the colour, the shape and the position. On the basis of these features, an accurate idea about the appearance of the horns may be obtained.

Size As was already indicated in Ch. 7.1, the dimensions of the horn cores of the aurochs are well known. Many of these survived, from many different countries, and they can be distinguished by sex. The keratin horn sheath, the actual horn, surrounded the horn core. The horns that survived are hollow, therefore, and by subtracting the length of this hollow from the

Photo 9. A right aurochs horn from the peat of Treten (Germany) (after Nehring 1900). In the horn sheath a part of the horn core had been preserved. The length of the horn is 76 cm; its girth at the base (probably) 38 cm.

121

Black Cyan Page 122

122


total length of the horn, the length of the solid part in the tips of the horn may be determined, which is not inconsiderable. Unfortunately, however, this is an aspect that has rarely been considered, and has been reported even less infrequently. Of two horns that belonged to the same animal, with probable reconstructed lengths of 72 cm, Ewald & Laurer (1911) measured solid parts of around 20 cm. On the basis of horns of Hungarian Steppe cattle, Bartosiewicz (1997) calculated the solid horn part that belonged to an aurochs horn core of 50 cm as 17 to 20 cm. The aurochs horn remains (with horn core remains) that were dredged up from the North Sea (the Stolzenbach horn; Bosscha Erdbrink 1986) contain a solid part of about 25 cm (see Photo 3). The solid parts of two other right horn remains, which were also recovered from the sea (the horns of Van Noort (Bosscha Erdbrink 1986) and Nederlof (1995)) were 33 and 19 cm long, respectively (author’s observation). It would seem obvious for the solid part to have been longer as the horn was longer and thicker. A little incongruous, in comparison with the dimensions mentioned here, is the length of the solid part of a 75 cm-long horn from the Baderup peat (Germany) (Requate 1957). This solid part measures 5 to 6 cm. If this is the original length of the solid part, this seems to have been an extremely variable aurochs feature. In the African buffalo, there seems to be no connection between the length of the horn core and the length of the solid part of the horn itself, either (H.H.T. Prins pers. comm.) In order to determine the maximum horn length for bulls, therefore, one cannot simply add the largest horn core length (82 cm) to the largest known solid keratin horn part (33 cm). The ratio between these two lengths is likely to have been different for each individual animal, which makes it impossible to determine the maximum length in this manner. The largest surviving horn is that of Sutton Hoo (England). This reconstructed horn had a length of 107 cm (Stone 1961). To determine the girth of a horn at its base, the diameter of the horn corn should be increased with 0.5 to 1 cm, depending on the girth of the horn core itself; 0.5 cm for small horn cores, 1 cm for larger ones. Table 8 lists estimates of the girth and length of aurochs horns, made on the basis of these data. For this table, diameters were calculated as if the bases of the horns were round. In reality, the horns are not quite round, but somewhat

Black Cyan Page 123


Table 8. Approximate largest and smallest diameters (at the base) and lengths of the horns of aurochs bulls and cows.

Characteristic

Horn diameter Horn length

Dimension (cm) ††

‡‡

10 - 18 60 - 107

7 - 10 40 - 70

oblong at the base. The dimensions given here for the diameters, therefore, should be seen as averages of the largest and smallest diameters per horn. The largest diameter (18 cm) refers to the horn of Sutton Hoo. This horn has a volume of 6,8 litres. The table seems to reveal that the horns of both bulls and cows are equally slim. The ratio of their length and diameter would be about 6:1. However, the thickest horn need not automatically be the longest, or the thinnest the shortest. The ratios found here are averages, therefore.

Colour Determination of the original colour of the aurochs horns by the horns themselves is best done on the basis of horns that have not been conserved in wet soil layers, since these may have been quite darkly discoloured under the influence of the underground water, which makes it impossible to recognize their original colour. Untreated horns that have been kept dry would be best suited for this, but their occurrence is unlikely. The horns have almost always held beer, wine or oil, which have given the horns a different colour, or they have been covered with a coat of dark varnish (Hilzheimer 1937), whose residue has soaked into the horn. Hilzheimer describes such a horn as having a reddish-brown colour with yellowish stains and a greyish tip. Bökönyi (1956) describes three horns from Esztergom in Hungary, which were sent from Poland and East Prussia as presents to the Hungarian kings during the 14th and 15th centuries. They were never varnished, but may have contained liquids. The colour of the three horns is

123

Black Cyan Page 124

124


yellowish white to light brown, and black towards the tip. According to Mertens (1906, p. 84), the so-called ‘horn of Treten’ from Hinterpommern (see Photo 9) has a comparable colour. He describes it as yellowish brown, blackish (‘schwärzlich’) at the tip, colours which remind the author of the horns of the Augsburg aurochs. In spite of the fact that this horn stems from a peat bog, its contrasting colours are still clearly recognizable. Another horn that is suitable for colour description belongs to one of the last aurochs (Photo 7), which died in the Jaktorów Forest (Central-Poland) in 1620. The horn, which was beautifully wrought and used as a hunting horn, is greyish yellow, turning into black towards the tip (Catalogue of Livrustkammaren Museum, Stockholm). Another way of obtaining information about the colour of the horns is by studying pictures of the aurochs. To this purpose, we can use the Augsburg aurochs (Smith 1827), and rock paintings like those in the Lascaux caves. Smith, who saw the original painting (the copy of which is represented in Fig. 23), describes the horns as ‘pale in colour with black tips’. Such colours may also be seen in the paintings at Lascaux; here as well, the general impression is of light-coloured horns with black tips (Guintard 1988). On the basis of these data, it is safe to conclude that the aurochs had horns with contrasting colours, light (probably greyish white to pale yellow) with dark tips. As was mentioned in the above, this makes them clearly different from those of the European bison, not just in length but also in colour. The European bison has relatively small, dark, inconspicuous horns, while those of the aurochs are large and striking – probably the more so because of their contrasting colours.

Shape The shape of the aurochs horn has long been a subject for debate, since many authors were not sure which sources to use to determine it. Really the only way to discover the right shape is by studying horn cores that are still attached to the skull. For the sake of completeness, however, an idea of the way the shape of the horns was interpreted and described in the past, and the mistakes that were made in the process, is offered below.

Black Cyan Page 125


Descriptions of aurochs contemporaries indicate that its horns were different, both from those of the European bison and from those of domestic cattle. As Caesar records: ‘the size of the horns and their shape and appearance are very different from the horns of our cattle’. Caesar is unlikely to have seen a live aurochs, but he undoubtedly saw the horns, and held them in his hands during his meetings with Gaulish and Germanic kings, when he drank from such horns or was given them as gifts. Von Herberstein (1557a) calls the horns of the European bison ‘shorter and thicker’ than those of the aurochs. He probably did not see the aurochs alive either, though he did see its hide and horns. More mention of the horns may be found in the novel ‘Banialuka’ by the Polish author Morsztyn from around 1600. He writes about the ‘broad-horned’ aurochs. The Roman writer Seneca (1st century A.D.), in his poem ‘Phaedra’ (line 65) also mentions ‘wild broad-horned aurochs’. In an Old English runic poem, the aurochs is called ‘large-horned’ (see Ch. 9.1.3). In the most detailed aurochs description available, that of Schneeberger (in: Gesner

Photo 10. The head of a Spanish fighting bull, in the neighbourhood of Salamanca (Spain) (Photo: T. van Vuure). The horns of this bull have roughly the same form and position as those of aurochs. In Spain, this horn shape is called ‘brocho’. It still regularly occurs in fighting cattle.

125

Black Cyan Page 126

126


1602), the horns are also mentioned. Schneeberger reports: ‘…, they have two horns, which are curved forward and slim’. Generally speaking, all these written sources are rather vague and ill suited to provide an accurate picture of the original shape of the horns. The pictures that represent the aurochs are somewhat clearer. These are ancient; those at Lascaux, for example, are at least 15,000 to 20,000 years old. There are colour paintings as well as engraved pictures. Other depictions, of a much later period, are known from the Middle East (Egypt, Mesopotamia), and still later ones from the end of the Middle Ages. The picture from the latter period that was provided by Von Herberstein (1557a) cannot be taken seriously (see Fig. 15), as the aurochs horns depicted in it are identical to those of the European bison in the same picture. As it happens, the hide which Von Herberstein received as a gift from the Polish king came without the horns, and the artist seems to have portrayed a pair of horns fashioned after his own fantasy, which Von Herberstein, who did not know their correct position either, approved. A far more accurate representation may be seen in the picture of the Augsburg aurochs (see Fig. 23). It also reveals the problem artists of the period must have had in depicting such horns. As will be shown later on, an aurochs horn has various curves. Depiction of such three-dimensional curves on a flat surface is only possible by using shading, as was done in the case of the Augsburg aurochs. The artist probably used a trick even here, however, since he put the left horn a little higher than its actual position so as to make its curves more apparent. The problem of depicting the horns can also be seen in the prehistoric cave paintings (see Fig. 48 and Photo 11). It is clear from these Pleistocene representations that the horns are relatively large, that they are curved and that they are clearly pointed forward. However, the actual position of the horns, and exactly how they are curved, is not clear. In almost all of these pictures the heads, therefore also the horns, are seen from the side. A frontal view is only offered in very few pictures, such as the one from the Dordogne, which was engraved in limestone (Lorblanchet & Welté 1994; see Fig. 18). It shows an aurochs head, including the horns, which clearly point inward, i.e. towards each other, as well as forward. Artists from the Middle East had the same problems. The horns were depicted in the oddest positions, just to make clear that they

Black Cyan Page 127


Fig. 18. Engraving of an aurochs head in stone (site: Limeuil, Dordogne, France). One of the few pictures in which an aurochs’ head can be seen from the front, with the horns curving towards each other (after Lorblanchet & Welté 1994).

were curved inward, as well as pointed forward (see Figs. 6 and 9). In Mesopotamia, when the animal was shown from the side, the back horn that was supposed to be behind the front one was left out completely, so that the animal seemed to have only one horn. The first domestic cattle breeds that were used in Egypt had horns that were clearly different from those of later breeds, and looked more like those of their wild ancestors. Petrie (1920) describes engravings of domesticated cattle from the Prehistory of the Egyptian Empire (10,000 – 3,000 B.C.). He reports: ‘In all these engravings, the horns usually curve forward, sometimes downward, once only upward, but they never stand out sideways. The early species seems to have inward-curving horns in the same plane, and with upward as well as downward variations. The later species with horns that stand out to the sides …..is not known from any engravings’. Schäfer (1896) shows a few statuettes from the same period that has such a shape and position of the horns (see Fig. 19). Pictures of cattle from the period of the pharaohs (after 3000 B.C.) usually depict horns that protrude and are more upward than forward pointing.

127

Black Cyan Page 128

128


The horns of domesticated cattle, which were still quite close to their wild ancestors at the time, still had the same shape and position. In one of the centres of domestication, Çatal Hüyük in Turkey, the horns of the first domesticated cattle breeds in the lower habitation layers were the same size and shape as those of the local wild aurochs (Mellaart 1975) (see Fig. 20). As the domestication process went on, the size and shape of the horns changed. It is not always clear if the artists actually saw the wild aurochs they depicted, especially from the pictures of the last period the aurochs existed in the Middle East, which do not always seem realistic. The picture of the aurochs hunt of Ramses III in Mesopotamia, for example, shows an animal with horns that do not point forward. Nor are they curved in the way that must have been detectable in real aurochs from this period (Von Lengerken 1955). The artist apparently depicted the horns of domesticated cattle, without suspecting any difference. References to the Crete aurochs cannot be used for the determination of the shape of the horns, either, since the aurochs originally did not exist on far-away Crete (Sondaar 1977). It was brought there in its domesticated form by men, for the bullfights, among other things (Nobis 1993). Both the colour of the fur and the shape of the horns of the bulls depicted there show characteristics of domestication. The only reliable method to find out how the horns looked, therefore, is to study the horn cores that were preserved, preferably attached

Fig. 19. Statues of domestic cattle from predynastic Egypt (before 3000 BC), in which the horn shape of the aurochs can still be recognized (after Schäfer 1896). As domestication progressed the horn shape changed increasingly

Black Cyan Page 129


to the skull, and keratin horns. Very many horn cores were found, but only a few dozen horns. Horn cores are known mainly from Europe; much smaller quantities survived in other parts of the aurochs distribution area, including China, the Middle East and North Africa. Horns are known only from Europe, not from any other continent. Von Leithner (1927) gave the first exact description of the shape of the horn cores. He did this on the basis of a study of extensive bone material, which showed that the horn cores of both bulls and cows have characteristic features. Bohlken (1958) and Grigson (1976) confirmed this in later years. Seen from the basis, where it was attached to the head, the direction of the horn core was first sideways and upward. After about a quarter of its length it would then curve forward and inward, in the last quarter further inward and upward. To Von Leithner, the characteristic, constant feature of the horn cores was their consistent inward curve. This means that the distance between the tips of the horn cores was always smaller than the distance between the outside curves. This was true for bulls as well as cows, although the difference in distance was smaller (i.e. 49 - 180 mm) for cows than for bulls (65 - 280 mm). In other words, the horns of a bull were curved more distinctly inward than those of a cow, in which their position was more ‘open’. Von Leithner saw the shape of the horn corn as something of a spiral; Grigson confirms this image and calls it the ‘primigenius spiral’. This shape is easy to recognize in the large horns of the Franqueiro-oxen in Brazil (see Fig. 27) and those of the Black Pied cow in Photo 12. The horn core was surrounded by the horn sheath. As was discussed in the above, this was easily 20 cm longer than the core, which made the spiral shape of the horns even more distinctive, since the tips would curve inward, forward and upward even more, making the distance between the tips of the left and the right horn even smaller. The development of the curves, which determined the shape of the horns, would vary, so that the spiral shape could be very conspicuous in some, and less so in other horns. The horns from Hungary that Bökönyi (1956) described, for example, were curved, but on a practically flat plane, from which only the tips protruded. Ewald & Laurer

129

Black Cyan Page 130

130


Photo 11. Wall painting of aurochs bulls in the Chauvet Cave (after Chauvet et al. 1998). Also here, the difficulty in representing the curves of aurochs horns in a flat surface is obvious.

(1911) described similar horns. Other horns, such as those depicted by Nehring (1900) (see Photo 9) and Tratz (1958), have exuberant, threedimensional curves. A cross-section of the horn core, at the base of the horn itself, was somewhat oval rather than completely round (Grigson 1973), at least in adult animals. It was still round in younger animals, but would grow more oval with age (Von Leithner 1927). The longer axis (the largest diameter) was practically parallel to the surface of the head. Its size would be 125 to 140% of that of the shorter axis. At its base, the horn core of the horn of Treten (Nehring 1900), for example, had a large diameter of 10, and a small of 7 cm. This oval shape may also be used to distinguish aurochs horns from the horns of European bison, whose sections at the base are round (Pucek 1986). A characteristic already noticeable from the grooves in the surface of the horn core, which are occasionally quite deep, is the fact that these are spirally twisted, a feature that is also recognizable in the keratin horn (Bosscha Erdbrink 1986) (Photo 3). The grooves and lines form a

Black Cyan Page 131


spiralling pattern on the surface of the horn(core). This feature is not found in, for example, the European bison and its ancestor, the European Steppe bison, the grooves on whose horn cores go straight from the basis to the tip. In fact, aurochs horns have two types of curves: those that involve the entire horn (the most distinctive), and the spiral twisting, which only involves the surface of the horns.

Fig. 20. Excavated former temple at Çatal Hüyük (Turkey) (7000 BC). This site was one of the first domestication centres of the aurochs. In this temple, clay bull heads had been placed onto which real aurochs horns were attached. Of these horns only the horn cores are left (Mellaart 1975).

Position Another feature of aurochs horns is their characteristic position, which determines their ‘forwardness’. On the basis of five skulls, Ewald & Laurer (1911) determined the angle of the horn cores in relation to the surface of the head. This angle varied between 50 and 60 degrees. For

131

Black Cyan Page 132

132


the Russian distribution area, Gromova (1931) determined an average angle of 63 degrees. Mensuration of six aurochs skulls at the Natuurhis-

Fig. 21. Aurochs skull at which the most prevailing angle between horn cores and forehead is pointed out. This is the characteristic ‘forwardness’ of aurochs’ horns (see also Fig. 16).

torisch Museum Naturalis in Leiden (author’s observation) revealed that this angle varied between 50 and 70 degrees, with an average of 59 degrees. It is represented in Fig. 21. Although the majority of aurochs remains were found in Europe, other areas, such as North Africa and East Asia, have also yielded sufficient remains by which to determine the shape and position of the horns. Only a single pair of Pleistocene horn cores is known from North Africa (Fayum, Egypt; Hilzheimer 1917), but there are many more Holocene ones (Pomel 1894, among others). From East Asia there are only Pleistocene skulls with horn cores from which to draw conclusions (Gromova 1931, Bohlin 1938, Boule et al. 1928, Zong Guanfu 1984), since the aurochs no longer existed there during the Holocene. For reasons already mentioned in Ch. 4, the East-Asian aurochs is probably genetically closely related to populations in West Asia. The horn cores from both East Asia and North Africa are remarkably similar to those from Europe. They do not only

Black Cyan Page 133


have the characteristic, inward-curving horn cores in common, but their position on the skull is also essentially the same as in the European aurochs. Between the various local populations inside Europe, there is no real difference with regard to these characteristics, either. Since this type of shape of the horns could be found throughout the distribution area, and can be called remarkably constant, it was obviously very efficient and functional. Of course, the basic shape of the horns, with its inward, forward curves, did allow for some variation, but strong deviations from these basis horn shape features seem to have been consistently removed from the aurochs population. For at least 1,5 million years, this type of horn has managed to hold its own in a very large area. The type can be found even in the Indian namadicus type, which lived there during the Pleistocene (Grigson 1973). Of all subspecies, the namadicus sub-species is probably removed furthest from the European sub-species. As yet, there is no satisfactory explanation for the stability of this type of horn. The defence against predators, such as wolves, or the way

Photo 12. This 14-year-old piebald cow possesses horns in the form and position like those of aurochs. Thus, even in modern cattle breeds features of the aurochs can still be found. (Photo: Elsevier, Doetinchem).

133

Black Cyan Page 134

134


in which bulls as well as cows fought with each other and the position in which they held their heads during these fight, or a combination of these two factors, may have had something to do with it. Bison bulls, which have relatively small horns, butt their heads against each other and then push their adversary away (Guthrie 1990). Aurochs probably linked their horns, after which the pushing and pulling would begin. Their horns were not suited to butt against each other, but could be linked together (Guthrie 1990). For fighting bulls in Spain, who still often have horns that are pointed forward and curved inward, this is the usual way of fighting (Domecq y Diez 1986). Bulls with such type of horns know exactly where the horn tips are and feel very insecure when these have been shortened by even a few centimetres. Spanish fighting

Photo 13. A photograph of two aurochs skulls with horn cores from Denmark, seen (top to bottom) from the front, from the side and from the back (after Degerbøl & Fredskild 1970). In the left one, the horn cores are curved inward only a little; in the right one strongly. In this way, a good impression of the three-dimensional curves of aurochs horns may be obtained.

Black Cyan Page 135


bulls have the tendency to make an upward movement with their horns. They are sometimes able to throw a horse with its rider up into the air. The aurochs was also wont to do this (Œwiêcicki 1634). For banteng, which have curved horns that are pointed more upward and backward, fighting consists of pushing and butting their heads against each other, and occasional sideways linking of the horns (Halder 1976).

Summary The numerous aurochs skulls with horn cores, and a limited number of keratin horns, show that the horns of this animal had a characteristic shape and position. This typical shape and position may be found throughout the distribution area, which shows it to have been very stable. The horns were pointed forward and curved inward. In this chapter, the shape, position, size and colour of the horns are discussed. Although the shape of the horns was very characteristic, there were some variations in length, girth, extent of curves and position in relation to the forehead. These variations occurred within specific margins.

7.3. The colour of the fur After the description of the colour of the horns in the previous section, this part of chapter 7 will deal with the colour and the markings of the fur of the aurochs; a discussion based on early descriptions and pictures of the aurochs, with the addition of a feasible genetic explanation. The correct and objective naming of the fur colours of wild animals is no easy task, if only because the colour of a species may vary with its place of occurrence inside the distribution area, the animals’ age, the season and their nutritive state. Seasons may show a great diversity in the temperature, the state of the vegetation and the intensity of the light, which may have their influence on the colour and nature of the fur. The nutritive state may also have an effect on the intensity of the colour and the shine of the fur. Moreover, colour may vary with sex; the

135

Black Cyan Page 136

136


colours and markings of male animals may be different from those of their female counterparts (sexual dimorphism). Another aspect which may interfere with the objective naming of colours is the personal interpretation of the observer, and way he communicates his observations to his audience and readers. In the early days, descriptions of rare species were usually hearsay; as a result, descriptions based on personal observations are very rare, particularly in the case of the aurochs. The original colour of an animal was wont to change completely in the course of stories repeatedly quoted. In 1460, for example, an author described the NorthPolish European bison as ‘lemon-coloured’ (Szalay 1930). Although this is obviously an extreme example, it does show to what extent colours may change when species are described that hardly anybody knows anymore. For the sake of convenience, people often have the tendency not to describe colour in detailed nuances, but by using the closest basic colour. Naturally, black people are not really black, red-haired people do not have red, but reddish-brown hair, and blue sheep do not have blue, but greyish fur. Red deer are not really red, but have reddish-brown fur in a specific season. This manner of describing colour has contributed to the numerous 19th-century discussions about the ambiguous existence of two wild bovine species in historic Europe, the European bison and the aurochs. Colour descriptions like black, brown and greyish do not evoke in the minds of all readers the nuances of colour actually perceived by the author, the more so since one and the same author sometimes uses several names for the same colours. It is the more confusing because there is really only one colour for the European bison, while for the aurochs there are two. In this way, distinguishing the aurochs and the European bison on the basis of earlier, unintentionally vague colour descriptions has become increasingly difficult. However, since the most chaotic, unreliable reports were ignored in favour of the more reliable ones, the following discussion may yet evoke a reasonably sound idea of the true original colours of the aurochs.

Black Cyan Page 137


Pictures and descriptions The earliest data regarding the colour of the aurochs may be found in rock paintings at various places in the South of France and in Spain (including Lascaux, Altamira, Chauvet). In the many caves, which served Pleistocene man as dwellings and, possibly, for magic rituals, countless animals have been depicted either in monotint or in colour. Together with animals like horses, mammoth, bison and deer, also the aurochs has been portrayed many times. In ‘monotint’ actually means that a black dye was used to indicate the outline of the animal, which was not filled in with colour. Sometimes, the outline has been partly filled in with small or larger spots. In some other drawings, the back of an animal’s body is missing: only the front part has been depicted. An explanation for these ‘artistic styles’ could be that there was a lack of dye, or that one was content with just the contours. Quite often, animals were drawn as in a character sketch, with a few simple lines, almost abstractly. Lascaux in particular has become famous for its beautiful, sometimes unnaturally large pictures of aurochs (see Photo 40). Grigson’s (1978) analyses of rock paintings from nine different SouthFrench locations have led to a number of conclusions: It is quite possible to distinguish two sizes of aurochs among those depicted. The larger are black, sometimes indicated only by contours, some of which have been filled in with stains. The smaller are mostly reddish brown (Grigson describes their colour as ‘red’), occasionally partly black. The larger type is not just different in colour, but also made to look sturdier, with bigger, thicker horns and clearly recognizable male genitals. The smaller, reddish-brown animals are slimmer and more elegantly built than the black ones, with thinner, smaller horns. They lack male genitals and a very few seem to have udders, although these are barely visible. Sometimes, though this is not certain, such a reddish-brown bovine animal seems to be followed by a calf. It seems reasonable to conclude that in the caves that were studied, male (large black) as well as female (smaller reddish-brown) aurochs have been portrayed. A very few reddish-brown bovine animals also have black body parts, for example at Lascaux, where a reddish-brown cow is de-

137

Black Cyan Page 138

138


picted with a black head and neck. Moreover, the cow in question has a broad light zone on its back. This could be the eelstripe, although other interpretations are possible. The snout is very rarely coloured, and some male animals have light rings around their eyes (Guintard 1988). Generally, however, the heads are solid black or reddish brown. As this subject will also be discussed in relation to the breeding-back experiment of the Heck brothers (see Ch.10), what is known about the colours of Egyptian wild cattle will be briefly mentioned here. As it happens, particularly Lutz Heck referred to the colour of the aurochs in Egypt in order to describe the colour of the animal’s fur. Besides all kinds of other wild animals, the many wall paintings in the royal graves in Egypt from the period of the Pharaohs (3000 - 1000 B.C.) also often portray bovine animals, either in hunting scenes, in which people hunt them with bow and arrow and lassos, or being attacked by lions. At first sight, therefore, there seems to have been an originally wild bovine animal that belonged there, and that looked a lot like the aurochs (see Fig. 22). However, if the various paintings are compared it becomes clear that the bovine animals in the various graves were not all depicted in the same way. Various ‘types’ emerge, with different colours and markings. The following is a survey: - Grave of king Sahu-Re (Hilzheimer in: Borchardt 1913): Paintings of a bull, a cow and a very young calf with uniform colours and markings. They are mainly reddish brown, with light yellow ‘saddles’ on their backs; their bellies, the insides of their legs and their tails are also light yellow, their snout, hooves and horns are black, and their polls are white. - The graves of Beni Hassan (graves 2, 5, 15, 17 and 29) (see Fig. 22) (Duerst 1899, Hilzheimer 1917, Newberry 1894): These show bulls of a reddish-brown colour, with white saddles on their back and yellow horns and hooves. Their bellies are white, which gradually turns into the reddish brown of their flanks. Some are reddish brown with only white bellies. There are also solid dark- or light-coloured bovine animals. Such single-colour wild cattle are also known from other graves (Hilzheimer 1917). Duerst describes a bovine animal hit by an arrow from grave 2. It has yellow horns and hooves, the back of its head is black, its face and

Black Cyan Page 139


neck are yellowish brown, the underside of the chest and belly white, the back, flanks and legs are all yellowish brown with black spots. - Grave of Ptahhotep II at Saqqara (Leclant 1979): The east wall of the grave shows a bull that is being attacked by a lion. It is mainly a dark reddish-brown colour, but the face, the chest, the inside of the forelegs, the genital area and the saddle on its back are a lighter reddish brown. The wild cattle portrayed in the various paintings are obviously not uniformly coloured and marked, but show all sorts of variations and colours. Of an originally wild species, in a limited area and over not too long a period, a lot more uniformity might reasonably be expected. The non-consistent colours and markings do not really call to mind a wild bovine animal; moreover, the shape of the horns of the bovine animals depicted and discussed here, which point upward, are lyre-shaped, and turned outward, brings to mind that of the horns of Egyptian domestic cattle rather than that of real aurochs. As was discussed before, the horns of the earliest, prehistoric Egyptian domestic cattle were more similar to aurochs horns than the horns of the later cattle breeds. In the period of the Pharaohs, there were already many kinds, sizes and colours of domestic cattle breeds in Egypt. On the basis of the colour and the shape of the horns, it seems logical to conclude that the wild cattle in question were actually domesticated animals, which either went wild entirely (for which theory there are many indications) (Duerst 1899)) or which may have been kept inside cattle grids for religious purposes, to be caught or killed. Some of the bovine animals depicted were caught alive. In accordance with an ancient Egyptian custom, bulls, which had an important role in religion, were caught in the wild and subsequently sacrificed (Von Lengerken 1955). It seems likely that initially, these were wild aurochs, from which the custom was later transferred to animals gone wild or kept in a feral state. There are many pictures that show how such animals were caught with lassos and lots of manpower. The bovine animals may also have been used for bullfights, which were popular for a period in Egypt as well as on Crete. During such bullfights the bulls would fight each other. There is a picture of such a fight between two bulls of similar colour and markings (Fraser

139

Black Cyan Page 140

140


Fig. 22. Pictures of wild cattle from ancient Egypt in the graves of Beni Hassan (after Newberry 1894). From top to bottom: Tomb nr. 5, tomb nr. 5 and tomb nr. 29. Given the various colours of these cattle and their lyre-shaped horns, these were domestic cattle gone wild that were hunted by people.

Black Cyan Page 141


1972, p.17). The bulls portrayed are black or very dark, with light reddishbrown heads, necks, saddles, bellies and behinds. On Crete, domesticated cattle were imported at the time, to be used in bullfights, among other things. These animals were generally black- or red pied. The colour of the fur of the original aurochs in Egypt cannot be determined on the basis of the bovine animals depicted in the old graves. By that time, the real aurochs had probably become so rare in the Middle East that it was barely known any more. Its place had been taken by domestic cattle gone wild, which showed all kinds of characteristics of domestication, such as changed colour, markings and horn shape. While there are few reliable pictures of the aurochs from the Greek and Roman periods, European pictures do not reappear until the end of the Middle Ages. Although the real aim of such pictures was to represent the animal in question, they were often copied from other sources, or sketched on the basis of verbal communications with more or less reliable eyewitnesses. An example is the 13th-century so-called ‘World Map of Ebstorf’, made by a monk from that village. In an area west of Kiev on this map, an animal has been depicted with the name ‘urus’ (aurochs) (Hilzheimer 1910). It has a reddish-yellow colour. The reliability of the representation is doubtful for several reasons. First of all, it is unlikely that the colours on such an early map are original; they may well have altered with age. Secondly, the monk probably never saw the animal in the flesh; it was drawn from different sources, which was often the case in those days. Moreover, according to Szalay (1930) the animal is supposed to represent a European bison instead of an aurochs. Pictures that offer a little more to go on did not appear until the 16th century. In the first place there is the ‘Carta Marina’, first made by Olaus Magnus 1539, which shows the northern half of Europe. Approximately at the height of the Polish-White Russian border, the map shows a rider on a horse that is being attacked by an aurochs (see Fig. 14). This aurochs is dark brown on the original coloured map, but it is not clear what sex it is. Olaus Magnus claimed to have seen the aurochs population at Jaktorów (Isberg 1962).

141

Black Cyan Page 142

142


In addition to this, there are the pictures commissioned by Von Herberstein. This diplomat had been given hides of an aurochs and a European bison by the Polish queen Bona for the great services he had rendered (Hilzheimer 1910). They lacked the horns and the forehead skins, however; the heads, therefore, seem likely to have been missing completely. Von Herberstein obviously did not know what an aurochs horn looked like, though he did know how the forehead fur looked, since he had also been given two belts, which allegedly had been cut ‘from skin which covers the forehead’. He had both skins stuffed to look like the original animals. Two artists made pictures (woodcuts) of the stuffed animal that was supposed to represent an aurochs, one of which is shown in Fig. 15. In consultation with Von Herberstein, the forehead skin and the horns in both pictures were drawn (cut) in. One artist made the woodcut in such a manner that it would provide a black print of the animal (see Fig. 15); the other confined himself to a woodcut that made a ‘black-and-white’ print. Although Von Herberstein approved and published the prints, the looks of the animal portrayed should not be attributed too much value to, since he never saw it in the flesh. The pictures from Von Herberstein’s publications were adopted by many authors, some of who adapted them to their own views. They can be found in Gesner (1602, 1620; see Fig. 12), Jonstonus (1657) (see Fig. 13) and various others (Pyle 1994). The repeated copying probably made the pictures increasingly unreliable. A picture that was very probably made of a real aurochs is that of the so-called ‘Augsburg aurochs’. In the early 19th century, the English biologist and art connoisseur Charles Hamilton Smith found an oil painting on wood at an art dealer’s in Augsburg (Germany). According to his appraisal the painting dated to the first quarter of the 16th century (Smith 1827). Smith had a copy made of the original (see Fig. 23). He describes the colour of the animal in the original as ‘entirely sooty black’; only the chin was ‘white’. Evidently, these were the only recognizable colours. Whether the original 16th-century artist painted in colour, or only in ‘black-and-white’, is not revealed by Smith’ description. A corner of the original painting still showed the remains of coats of arms and the word ‘Thur’ (aurochs) in German gold lettering. Judging from these latter

Black Cyan Page 143


details, the whole painting may probably be considered to have been rendered in colour. Smith had the original and the copy in his possession until his death, after which they were sold (Nehring 1898a). Whether they still exist, and where, is unknown. This concludes the discussion of the pictures. References to the colour of the animal may also be found in written descriptions. Roman authors such as Caesar, Plinius and Seneca do not report anything about the colour from their own observations (Szalay 1930). Plinius does quote an earlier source about the ‘black forest ox’ (‘bos silvestris niger’), but it is unclear how reliable this information is. From the post-Roman period, the first thing that stands out is that in the Old-Germanic hunting laws, recorded in the ‘Lex Baiuvariorum’ (ca. A.D. 800), for example, the aurochs was ranked among the ‘suarzwild’, which included the very darkly-coloured game, such as the wild boar (Szalay 1930).

Fig. 23. The Augsburg aurochs (after Smith 1827). This painting is a copy of the original that was present at a dealer’s at Augsburg (Germany) in the early 19th century. The original probably dates to the beginning of the 16th century. It is not known whether the original or the copy still exist.

143

Black Cyan Page 144

144


The early popular traditions also include some mention of colour. Romanian folk tales, for example, tell of the ‘black aurochs’ (‘bohor negro’) (Szalay 1930), Russian stories from the Central Dniepr area (near Kiev) mention the ‘reddish-brown aurochs with the golden horns’ (Dementiev 1958), and the Polish language has the expression ‘red like an aurochs’ (‘czerwony jak tur’). In the latter language, a passage from a 17th century wedding song reports: ‘and a red aurochs calf [‘czerwone turz¹tko’]will be the roast’ (Pusch 1840, p. 121). Of the five eyewitnesses known to have recorded their experiences with the aurochs, two are rather vague about its colour. One of them is Gratiani a Burgo, who saw aurochs calves in captivity in East Prussia in 1568 (Szalay 1917, pp. 113 and 114). He uses the Latin term ‘subniger’ to describe the colour of the fur of these animals. The word means ‘somewhat black, dark’, but is really a vague term that can be interpreted in more than one way. It is not known what age the calves were at the time. As will be explained later on in this chapter, the colour of aurochs bulls changed when they were about six months old. The colour of the cows remained almost the same. A second eyewitness is Mucante, who travelled in cardinal Gaetano’s entourage and was sent an aurochs by the Polish king on 30 September 1596, during a stopover in Warsaw. He describes the colour of the animal as ‘grau’ (Pusch 1840, p. 75); in any case, this is the word used in the German text, which evolved from a Polish translation of the original Latin text. It is impossible to find out which Latin term led to the translation ‘grau’; to get even a rough idea of the colour Mucante was trying to describe is even less feasible. In English, ‘grau’ means ‘grey’ or ‘greyish’. Further on in the text, the same author describes a number of European bison he observed as ‘like black oxen’. He either did not take accurate description of the true colour very seriously, or the original colour may have been altered in the course of several translations. The other three eyewitnesses are Von Herberstein (1557a, 1557b), and Bonar and Schneeberger (in: Gesner 1602; see Appendix). In addition to this, Gesner himself described a belt cut from aurochs skin (1620, p.146). Von Herberstein probably did not visit the last aurochs at Jaktorów, though he did hear them mentioned by people who had seen them. Both

Black Cyan Page 145


a German (1557a) and a Latin (1557b) version appeared of his report, which makes it possible to compare the colour descriptions. Von Herberstein is really the only person who is known definitely to have held an aurochs skin in his hands. If we add the fact that he took the trouble to inform the public about the aurochs (and the European bison) by writing about them in Latin and German and by exhibiting a ‘stuffed’ skin, he should be considered a serious observer. In the Latin version, he reports that aurochs do not differ from domestic cattle at all: ‘…except that they are all black [‘nigri’]; on their backs they have a marking in the shape of a line mixed with white [‘ex albo mixtum’]’. In the German version, he also mentions the colour: ‘seind ganz guetschwarz, allein am ruckhgrad ainen grahlaten strich nach derleng’ ‘they are entirely black, with only a lengthwise grey/greyish stripe over the spine’ The Old-German word ‘grahlat’ is best translated into modern German as ‘grau’, which means ‘grey’ or ‘greyish’ in English. In the Latin version, he clarifies the colour of the stripe by mentioning that the black fur is mixed with white hair in that place. Obviously, to judge from his description, black (in any case, very dark) hairs and white hairs are mixed in the eelstripe, to form what may be called a grey or greyish line. Further on in the German text he adds, in his description of the European bison, that this animal is ‘not as beautifully black as the aurochs’ (‘nit so schön schwarz als der Thur’). In the picture Von Herberstein commissioned of the stuffed aurochs skin in his house (see Fig. 15), the back of the animal shows a thin, light, lengthwise stripe. It seems reasonable to assume that the artist intended to depict the eelstripe in this manner, as any colouringin with smaller lines was intentionally omitted here. Von Herberstein does not mention any other, divergent aurochs colour. As was mentioned before he possessed two belts, which had been given to him by the Polish queen. Such belts were traditionally cut from the forehead skin (see Ch. 6.4). Since Von Herberstein makes no remarks about an aberrant, for example lighter colour of these belts we have to assume that the curled hairs on the forehead and between the horns were the same colour as the rest of the ‘black’ fur.

145

Black Cyan Page 146

146


It could be concluded from Bonar’s report that he saw the animals himself or, at least, had a reliable informant, since he philosophises about the animals’ origin and reveals a good grasp of the differences between domestic cattle, aurochs and European bison. He wonders whether aurochs stem from ‘outstanding (domestic) cattle’ or from crossbreeding between European bison and domestic cattle. Although he does not take the trouble to give a detailed description of the aurochs, he does say about its colour: ‘When the skin of this animal has been cleaned it is covered with very fine black [‘nigris’] hairs’. The most extensive description we have of the colour of the aurochs is that of Schneeberger. Whether he visited the aurochs at Jaktorów is not entirely certain either, but his often very detailed description seems to show that he either saw the animals himself, or that he recorded verbal communications from those who were in charge of the animals on a daily basis for years on end. Even if he did visit the animals himself, he must have spoken to the managers, since they were the only people who had information and specific knowledge. Schneeberger uses three languages for the description of the colour of the animals: Latin, German and Polish. This is a translation of the passage in question, with the original text about the colour in ‘square brackets’. ‘When a bull is born, its hair is a chestnut colour [‘castanei coloris’] (blackish brown, dark brown or the colour of black ash [‘schwarzbraun, dunckelbraun oder schwarzaschenfarb’]; ‘plowy’ as the Polish call it); within half a year they become completely black [‘nigrescunt’], however, while the line over the spine, some two fingers wide, remains a lighter colour [‘subnigra’]. The cows always keep the colour described in the above, and black [‘nigrae’] cows are found very rarely’. That is all Schneeberger had to say about the colour. He really only mentions the general colour of the fur, the changes it would go through, and the eelstripe. His use of three different languages for the description of the colour shows that he seems to have been well aware of the difficulty of describing colours objectively. In spite of the trouble he took, his explanations still lack clarity in places. The Latin term ‘castanei coloris’ and

Black Cyan Page 147


the Polish word ‘plowy’ (there is no doubt that the Polish term ‘p³owy’25 is meant here) are well matched, but the German description ‘schwarzbraun, dunckelbraun oder schwarzaschenfarb’ seems to indicate a very dark colour, dissimilar to the other two. He may have wanted to indicate that darker colour nuances of reddish brown occurred besides light ones. As was explained before, the Latin word ‘subnigra’, which is used to describe the colour of the eelstripe, is a vague term that one can interpret a little lighter or darker, at one’s own discretion. Gesner (1620, p. 146) himself describes a belt cut from aurochs skin, which was sent to him from Poland by Schneeberger. The hairs on this piece of skin were ‘…of a black colour, but if you look closer they turn out to be mixed with a few reddish-brown hairs’. Obviously, at first sight the colour could be called black, but if one looked more closely some brown hairs would show. This is what is known about the colour from the descriptions of real or alleged eyewitnesses. In addition to this information, data that have resulted from dozens of years of genetic research of domestic cattle may be used.

Genetic aspects of the fur colour Current cattle show a rich variation in colours and markings. The nearly 1,000 breeds that developed in the course of the past centuries all have their own characteristically coloured appearance. All these colours stem from the original set of genes that made up the wild aurochs. As the domestication process went on, and a cattle breed was subjected to line breeding, the ‘wild-type’ colour would disappear more and more. However, nowadays remains of this colour are found not only in socalled ‘primitive’ cattle breeds; some of it still shows also in the more ‘modern’ breeds. To gain an insight in the various colour-determining genes in cattle and the ways they affect each other, long-term observation is necessary to record both the of the animals and their colour 25 The accepted meaning of this word is ‘flaxen, buff’. Used for wild animals, however, it means ‘fox-coloured, reddish brown’, as in roe deer and red deer (Bulas et al. 1961, Doroszewski 1958 – 1969); in Polish language, ‘zwierzyna p§lowa’ is a collective term for roe-, red- and fallow deer and elk.

147

Black Cyan Page 148

148


patterns. The results of such observations were analysed by Olson & Wilham (1982). The data used were collected at the Wichita Mountain Wildlife Refuge, where a herd of Texas Longhorn cattle (mainly of Spanish-Portuguese descent) was followed descent during a period of 47 years in order to study their colour and descent. In addition to this, data from a research centre, where long-term crossbreeding experiments were conducted with fifteen modern cattle breeds, were included. Observation revealed that various genes are instrumental in the development of the definitive colours and markings. Although the effect of some of the genes is not yet clear, there is a general idea about how the colour of the fur comes about. The genes that determine the colours of cattle include one gene that plays a central part. It is indicated by the letter E+ and is also sometimes called the wild-type colouration gene, since it must have been present in the aurochs. The E+-gene and the genes that developed from it determine the basic colours black and red, which appear in many cattle. Although it is not always very apparent, the gene may still be found in many cattle breeds, including the zebu, the Texas Longhorn, Jersey cattle, Braunvieh and Heck cattle. Animals that have this gene in duplicate (written E+E+), since they received one from each parent, are born reddish brown. Within six months, the bull calves will have changed their colour to a very dark, blackish-brown. The female animals do not change their colour; it remains more or less the same. The gene, therefore, contributed to the difference between bulls and cows, since the double colour is partly due to the male hormone testosterone. The bull’s production of this makes the reddish-brown colour turn into blackish brown. If the testosterone is absent the reddish-brown colour remains. This effect was shown in particular by a number of experiments in which the process was inverted. Jersey bulls, whose colour had changed to black and who had subsequently been castrated, so that the production of testosterone had stopped, largely faded to a lighter colour. The black that was originally perceptible on their heads, ears, polls, trunks, legs and tails, only remained on their legs and tails. Evidently, another gene for black was effective in these areas (Gilmore et al. 1961). Colour changes following

Black Cyan Page 149


castration are also known from various zebu breeds from India and Pakistan, including the Bhagwari (Joshi & Phillips 1953). A probably comparable principle may be found in banteng. In Bali cattle, which descend from the banteng, it was also apparent that the black colour disappeared after castration and that the colour of the animal faded (Rouse 1970). A similar phenomenon was observed in wild banteng. Hoogerwerf (1970) reports a case of natural castration of a banteng bull in the nature reserve Udjung Kulon on West Java. After an injury to its scrotum the bull, whose production of testosterone had ceased as a consequence, had taken the colour of the cows. In the course of the domestication process, changes (‘mutations’) to this E+-gene developed on an ever-increasing scale, and remained thus in various breeds. Mutations have been recorded both towards the ‘black’ and the ‘red’ colouring. As to the black, the Ed-gene evolved, which causes a deep black colour in Black Pied, Aberdeen Angus and Galloway breeds. On the red side the e-gene developed, which determines the deep reddish-brown colour of the Maas-Rijn-IJssel cows and the Simmenthal and Hereford breeds. In view of the fact that the deep black colour that is caused by the Ed-gene occurs in cows as well as bulls, expression of this colour appears to be independent of testosterone levels. In addition to the E+-gene for the basic colour there are also genes that affect the intensity of the colour, the more or less prominent presence of red and black pigments in the hairs (Olson & Wilham 1982). In domesticated cattle, mutations in these genes cause the original colours to be ‘diluted’, as is the case in Jersey cattle, Braunvieh (Brown Swiss), Limousin and Scottish Highland cattle breeds. Sometimes the pigments have all but disappeared, as in the Chianina (see Photo 49), zebu and English Park cattle (see Photo 44). There is yet another gene that determines whether the cattle have solid colours or not. This wild-type gene S+ used to cause the colour to be distributed evenly across the body. Mutations of this gene in domesticated cattle cause all kinds of spotty patterns, as seen in Black Pied and Red Pied cattle, Lakenvelder, Witrik and Belgian Blue cattle. Which gene or genes determine(s) the wild-type characteristics of eelstripe and pale snout is not quite clear. As was said in the above, the

149

Black Cyan Page 150

150


eelstripe is a characteristic of the bull. It may be found almost anywhere in the world, not just in Europe, but also in North Africa, East Asia, India and North and South America. A feature that is probably related to this occurs in domesticated cattle and is called ‘patterned blackish’ (Minkema & De Rooy 1982, Olson & Wilham 1982). Calves that possess the gene (or genes) for this colour reversal are born reddish brown; the bull calves as well as the cow calves change to black between 3 and 6 months, however. This black colour has a reddish tinge and both sexes may show paler eelstripes, snouts and inner ears. For this colour reversal to take place, the e-gene must be present in duplicate (ee), yet the E+-gene should be missing. Thus, cows may start to show the colours and markings of bulls. The development of this type has not been fully explained yet; in the future, however, such unusual mutations may shed more light on the effect of the genes on the colour of the eelstripe and the pale snout.

Comparison with related bovine species If the colours of the aurochs are compared to those of the related bovine species inside the Bovini tribe, a number of similarities are revealed. In the first place, calves of many bovine species have a reddish-brown colour. This is true for European bison, bison, gaur and banteng. The calves of the African buffalo are reddish brown to dark brown. Schaller (1998) reports that wild yak calves have a dark brown colour, different from the deep blackish-brown tinge of the adult animals. The function of such a reddish-brown colour may be partly explained by the social status of a calf. Its conspicuousness among the adult animals serves to emphasize the vulnerability of calves. On the other hand, the reddish-brown colour may be useful for a calf that is laid up in the forest for a short period, since the colour may help to camouflage it. Instinctive concealment of a new-born calf was probably done by aurochs (Schneeberger; see Appendix)26, like also by gaur (Hubback 1938) and banteng (Halder 1976). 26 Schneeberger describes how the aurochs cow would isolate herself during the period just before the birth of the calf until a few days afterwards. It is known that the Camargue cattle (Schloeth 1961) and the Spanish fighting cattle (Domecq y Diez 1986) hide their young calves, which will then remain on the ground without moving.

Black Cyan Page 151


As the calves grow bigger, the reddish-brown colour usually changes. In European bison and American bison it turns dark brown; in gaur bulls black to blackish-brown (sepia) and in gaur cows dark (chocolate) brown. Banteng go through a colour change comparable to that of the aurochs: the bulls turn a deep blackish brown, while the cows keep the colour of calves. The time at which the colour change takes place varies with the bovine species. As is true of the aurochs, gaur calves change to their adult colour before they are six months (Hubback 1938). For banteng this takes a little longer; the bull calves change colour around age 1 (Hoogerwerf 1970). In the cow calves, however, the reddish-brown colour from the heels to the toes changes into a light, almost white colour as early as 1 to 2 months after birth. The adult animals, both cows and bulls, also have such white ‘socks’, as well as white behinds (‘mirrors’). The white socks also occur in both sexes of the gaur; occasionally, they may be a khaki (very light brown) colour. Eelstripes occur sporadically in the other bovine species; the reddish-brown banteng calves, for example, have a dark, almost black stripe on their backs from birth (Hoogerwerf 1970). Wild yak also have eelstripes when they are young. Prschewalski (1877), who studied the latter in Tibet, reports that young animals have a narrow, silvery grey stripe along their backs. It is not clear whether this stripe is limited to only one sex. Schaller (1998) makes no mention of the eelstripe, but limits himself to the general colour of the fur. Divergent colours may also occur on the head. The snout of the yak often bears a pale grey to white zone, and the black hairs from its forehead to its snout have silvery-grey tips. Both banteng and gaur often have white zones on their snouts and chins. The forehead of the gaur bears a light grey zone. Like the domestic cattle that descend from the aurochs, other domesticated cattle show characteristics of domestication in the colours of their fur. This process is particularly advanced in the yak, whose colours not only include black and brown but also light brown, white, pale yellow and all kinds of colourful spotted patterns. Colour mutants that would have disappeared through natural selection as unfavourable may continue to exist in domesticated cattle. Man will make selections

151

Black Cyan Page 152

152


with reference to milk production, pulling power and such, but colour plays no really significant part. In such a situation, divergent colours are tolerated and sometimes even preferred, and they will remain as a consequence.

Reconstruction of the fur colour of the aurochs The above discussion about pictures and descriptions of the aurochs, the genetic study of domestic cattle and the comparison with the fur colour of related, still existing cattle breeds, may result in a description of the colour of the fur of the aurochs. Basic body colour The aurochs calf was born with reddish-brown fur, which in bull calves would turn a deep blackish brown, not entirely black, within half a year. For convenience’s sake this colour would be called ‘black’’. The cows more or less retained the reddish brown of the calves, a colour that was called ‘red’. In cows as well as bulls there are likely to have been colour variegations, in that the reddish brown and the blackish brown may have been a little lighter or darker on certain zones of the body. Naturally, colour derivations may have occurred from time to time; there may have been very pale animals, such as also sometimes happened among the North American bison. Moreover, like Schneeberger records, there may have been black cows in rare cases, most likely due to the presence of the Ed -mutant mentioned before. There is no reason to assume that these basic colours were clearly different outside Europe, e.g. paler, or reddish brown for both sexes (also see the section about the eelstripe below) Eelstripe On the basis of the descriptions from ancient Poland and in view of the fact that the eelstripe in domestic cattle occurred almost exclusively in bulls, this should be seen as a male characteristic. It must have been a narrow stripe with a colour that was greyish or drab, at least in Europe.

Black Cyan Page 153


The early descriptions and pictures do not reveal what the situation was like outside Europe. In view of the fact that the E+-gene also occurs in zebus and that various zebu breeds have pale-coloured eelstripes as well (Joshi & Phillips 1953), which is also the case in North African breeds (Joshi et al. 1957), it seems likely that the colours of the European aurochs could also be found in the animal outside Europe. At most, there may have been a difference in nuance. Besides the main body colours brown, reddish brown and blackish brown, Bovini always have paler markings, i.e. white, grey or khaki. On the basis of this fact, very dark eelstripes should be considered domestication features. Dark reddish-brown eelstripes sometimes occur, for example, and even black ones in light-coloured breeds like the Jerseybrown Criollo in South America (in lit. J.E. Rouse). Among the grey colour variations of Spanish fighting bulls, such as the Cárdeno Claro, black eelstripes also sometimes occur (Rodríguez Montesinos 1994). Snout The pictures of the Augsburg aurochs and the fact that domesticated cattle often have a very light zone at the snout make it likely that something similar may have been the case in aurochs. The white chin of the Augsburg aurochs is the only indication that this is true. In gaur and banteng, a light zone at the snout may also be seen. In his report on the aurochs, Schneeberger does not mention this; artists from Lascaux, among other places, did not take the trouble to depict such a feature. In view of what was said in the above section about the eelstripe and comparable characteristics in Bovini, the colour of the area around the snout is likely to have been a dirty white. Other zones The poll, the rims of the eyes, the insides of the ears, the belly area and the insides of the legs are zones that are often a different colour in the current breeds. In domestic cattle, the forehead and the area between the horns sometimes has fur of a lighter colour, for example white, yellowish, reddish brown or pale brown. Neither the Augsburg aurochs (whose

153

Black Cyan Page 154

154


poll and forehead show dark curls), nor the aurochs depicted by Von Herberstein, nor pictures from caves in South Europe show any lighter poll colours. Various sources (Œwiêcicki, Von Herberstein, Schneeberger and Mucante) reveal that magic belts were cut, quite particularly from the skin on the aurochs’ forehead (which bears curled hair). The curls were left on the belts intact and must have been dark. Gesner (1620, p.144) reports having heard from a ‘respectable’ Pole that this hair was ‘very dark’ (‘nigerrimus’). Whether the ‘magic’ was only in the skin of the bull, or if the cow was also used for this purpose, is not clear from the reports, which are rather general. The fact that the belts were black could be an indication that only bull skin was used. This would also mean that the polls, of bulls in any case, were very dark, not pale. In the experiment carried out by Gilmore et al. (1961), the black polls turned out to have been caused by the E+-gene together with testosterone. Banteng do not have a light area on the forehead, though gaur do. Domestic cattle have a number of genes that remove either the red or the black pigment (Olson & Wilham 1982). The ‘chinchilla gene’, for example, removes red pigment, while the ‘zebu-tipping gene’, removes the black pigment from the tips of the underbelly hairs. There is also a ‘Brown Swiss-tipping gene’, which removes the black from the poll, the ears, the chest, the belly and the back. In the Augsburg aurochs, the rims of the eyes and ears have a lighter colour. Whether this means that they were really this pale, or that the artist found it easier to make the eyes and ears stand out against the darker background in this way, has yet to be resolved. The Augsburg aurochs, the cave drawings at Lascaux or at any other cave do not give any indication of light ears, pale underbellies or pale insides of the legs. Light rims around the eyes sometimes show. Horses portrayed in the same caves often do have pale bellies and insides of the legs, features that are still visible in the Przewalski horse. The fur of the aurochs is unlikely to have been a perfectly solid colour. Other wild bovine- or deer species have lighter and darker areas in their fur, and this may well have been true for the aurochs. Rock paintings sometimes show cows with somewhat darker legs or heads than the rest of their bodies, which may indicate that such aberrations occurred from time to time (see Photo 38). That this is not to be found in the

Black Cyan Page 155


descriptions probably indicates that, if there were such areas at all, they were probably not conspicuous enough to warrant mention. A representation of the most remarkable general nuances would then have sufficed. The genes that used to be responsible for the lighter areas on the fur may have mutated in domestic cattle, to genes that pale the areas in question quite considerably, to form clearly lighter or white areas.

Summary On the basis of paintings from caves (those of Lascaux, for example) and pictures from the 16th century (including that of the ‘Augsburg aurochs’), combined with early descriptions of the aurochs, contemporary genetic studies and comparisons with wild cattle species that still exist, it has proved possible to draw a relatively clear image of the fur colour of the aurochs. There was a clear difference between the colour of the cow and that of the bull. The bull was blackish brown to black, with a narrow, lighter stripe (eelstripe) on the back. The cow was reddish brown, like the calf. Both sexes probably had lighter areas around their snouts.

7.4. Other physical characteristics

Hooves The characteristics of the hooves of even-toed ungulates are directly related to the nature of the soil they live on. On hard soil, the hooves will have a different structure and shape than on soft soil; on bare rocks in mountainous areas, for example, ungulates such as mountain sheep and goats will have hard, narrow, fairly inflexible hooves. Soft, wide, spreading hooves are better suited to soft soil types, such as swamps, and may be found in elk (Alces alces), reindeer (Rangifer tarandus), and in African swamp animals such as the sitatunga (Tragelaphus spekii) and the lechwe (Kobus leche).

155

Black Cyan Page 156

156


Little is known about the hooves of aurochs. According to Wrzeœniowski (1878, p. 525), a poem by the Polish poet Morsztyn (1st half of the 17th century) mentions ‘broad-footed aurochs’. The original text by this poet, however, records ‘broad-horned aurochs’ (‘tury szerokorogie’) (Kar³owicz et al. 1919), and contains no reference to the hooves. Schneeberger (in: Gesner 1602), who may be considered a reliable source, wrote the following description: ‘Ungula fissa, magis cava quam domesticarum’. ‘Their hoof is cloven, more hollow than in domestic cattle’. There is no further explanation of what is actually meant by this. The author, accompanied by caretakers of the last aurochs population at Jaktorów, may have been able to compare hoof prints in the area with those of domestic cattle, which also lived there. A somewhat comparable, though more detailed description may be found in the curiously ‘mixed’ aurochs/ European bison description given by Von Fleming (1719). He reports: ‘Finally I will say of its footprints……… Since, according to the description of its characteristics, it is almost the same in all aspects as domestic cattle, apart from the fact that it is wild, that its hooves are much longer, more crooked and harder, as well as more arched, more closed than in domestic cattle……….., moreover, the footprints of the wild bovine animal are straighter and more even’. This description does not really give a clear impression of the hooves, either. Von Fleming was very well accustomed to the hunting rituals and game in Europe at the end of the 17th century and may have had information from oral or written sources with regard to the aurochs, for example from the Great Wilderness. Whether the aurochs hooves were shaped differently than those of domestic cattle can no longer be ascertained, since no keratin hooves of the former have ever been found. Some claw bones (to which the keratin hooves were attached) were preserved, but they have the same shape as those in domestic cattle (author’s observation). A possible explanation for Schneeberger’s somewhat vague description could be that the word ‘cava’ should not be translated as ‘hollow’, but as ‘deep, sunk’ (in lit. G. Roberts). In that case, ‘ungula’ should be translated as ‘hoof print’ instead of as ‘hoof’. In this manner, the caretaker of the area

Black Cyan Page 157


may have wanted to emphasize the difference between the hoof print of the aurochs and that of the Polish domestic cow of the period in question. This explanation came about as a consequence of the find of a series of aurochs hoof prints, with a total length of over 11 metres, on the coast of Liverpool (England), which was dated to the transition period from the Mesolithic to the Neolithic period (circa 3200 B.C.) (see Fig. 24 and Pho-

Fig. 24. Aurochs track at Formby Point (on the coast at Liverpool, England). This over 11 m long track was deposited in the tidal area, some 5000 years ago (drawing: G. Roberts).

Photo 14. Prints of the right hind leg and right foreleg from the track in Fig. 24 (at the ‘7 m’ mark). The prints of the smaller dew claws can be recognized as well (Photo: G. Roberts).

157

Black Cyan Page 158

158


to 14) (Huddard et al. 1999; in lit. G. Roberts). Deeper (Mesolithic) layers in the same area have yielded comparable aurochs hoof prints. Comparison of the latter with hoof prints of domestic cattle from the Iron Age (in soil layers deposited later) shows that those of the aurochs were deeper (‘more hollow’), probably as a result of their greater body weight. The trail of hoof prints shows that the hind legs stepped into the prints of the forelegs; the prints of the smaller dew claws are also visible. The animal seems to have walked calmly. The average width of the spoor, measured between the centres of the left and right hoof prints, is 40 cm. (measurements taken by G. Roberts). The length of the hoof prints (without the dew claws) is around 16 cm on average; the width is about 14 cm. The pace length, measured between the front edges of two subsequent prints of the same hoof, is 220 cm on average, and varied between 190 and 265 cm. Comparisons of these data with the pace length of present-day domestic cattle are scarce. M.F. Wallis de Vries (in lit.) found a pace length of 160 cm for MRIJ-oxen. Herlin (1994) found an average pace length of 170 cm for Frisian dairy cattle walking with a speed of 1,4 m per second. If more extensive studies were to be done of the pace length and other cattle trail characteristics, and the resulting data were to be related to the shoulder height of the animals in question, a more accurate idea about the height of the withers of aurochs might be obtained.

Udder Consideration of the numerous current cattle breeds reveals a large variation in udder sizes, from small udders, which mainly consist of four teats, to very large ones. Particularly in many European breeds, the size of the udder can be quite considerable. As it happens, stringent selection criteria were applied with regard to milk production on this continent. In areas such as the Middle East, India and Africa, cattle have been used mainly as draught animals and meat providers. The udders are generally small in these regions. Selection for milk production has certainly increased udder size. Any idea of what aurochs udders looked like is almost entirely dependent on paintings from caves, such as those at Lascaux (see Fig. 25),

Black Cyan Page 159


since these do not only depict bulls, but cows as well. If the udders had been distinctly visible, like the male genitals, they would also have been portrayed clearly. Apart from a few uncertain cases, however, they are not visible (Grigson 1978, Guintard 1988). Even a painting from Lascaux, which depicts a cow followed by a calf, does not show the udder, although it might well have been somewhat enlarged in a nursing cow. It seems reasonable to conclude, therefore, that the udder of the aurochs cow was apparently too inconspicuous to be portrayed.

Fig. 25. Drawing in the La Mairie Cave (Dordogne, France), depicting an aurochs bull and two cows. The udders of the aurochs cows are small and not visible in this way.

In domesticated cattle breeds that have not undergone selective procedures with regard to their milk production, udders appear to be very small and not visible from the side, since they remain hidden between the hind legs. Particularly Spanish fighting cattle, which for centuries have been selected for their fighting spirit and efficiency, but certainly not for their milk production, have such small udders. Their calves are relatively small, some 20 to 25 kg at birth, and may be adequately fed with the milk from such ‘small’ udders (French et al. 1966) (see Photo 15). The extent of the milk production in these cattle is unknown. In a different breed, Hungarian Steppe cattle, which were originally used as draught animals, milk production varies between 1400 and 2400 kg. per lactation period (French et al. 1966). In spite of the fact that some limited selection in favour of milk production does take place, as it used to in the past, their udders may still be called small as compared to those of, for example, Black Pied dairy cattle, whose average milk production is 7000 kg. per lactation period (Vink & Wolbers 1997).

159

Black Cyan Page 160

160


Existing related bovine species show the same tendency as Spanish fighting cattle. European bison and banteng also have small udders that are invisible from the side (see the banteng cow in Photo 8). European bison calves weigh between 15 and 35 kg. at birth (Krasiñska & Krasiñski 2004), which is also the range Spanish fighting cattle calves would fall in. The milk production of such a small udder is sufficient to feed one calf, and sometimes even two. The ‘hidden’ position of the small, yet sufficiently productive udder may be essential as protection against damage and frost to this vulnerable body part.

Photo 15. Cow and calf of Spanish fighting cattle (Province of Salamanca, Spain). The cows of this cattle type have small, barely visible udders (Photo: T. van Vuure).

Fur Not very many descriptions of the fur of the aurochs have survived, either. Schneeberger (in: Gesner 1602) reports that the aurochs, compared to domestic cattle, was covered with ‘longer hairs’. Bonar (in: Gesner 1602) writes that the skin is ‘covered with very fine black hairs’. His record should also probably be considered as a comparison with 16th century domestic cattle. Gesner (1620, p. 146) himself reports hav-

Black Cyan Page 161


ing been sent a belt from aurochs skin from Poland by Schneeberger. He writes about the fur: ‘…the hairs are really very soft (surprisingly so), like the wool of sheep, close together…’. Such remarks only give a rough indication of the fur, however. If the aurochs is mentioned together with the European bison (by Plinius (circa 70) and Von Herberstein (1557a), for example), the latter is often called hairy and bearded, unlike the aurochs. Apparently, the longhaired and bearded appearance of the European bison (on the forehand, in any case) was worth mentioning, which was not the case for aurochs. Judging from the paintings found in caves, as well as the Augsburg aurochs, the fur was really reasonable evenly divided over the body; spots with markedly longer hair are nowhere to be found. The hairs were only really different between the horns and on the forehead. Both Schneeberger (in: Gesner 1602) and Œwiêcicki (1634) report that the hairs in these places were curled and twisted, which must have made the animal ‘terrible to behold’ (see Ch. 6.4). These curled hairs on the forehead are clearly visible in the picture of the Augsburg aurochs. They were definitely a feature of aurochs bulls; whether cows had them as well, and to what extent, is unknown. They may have been an aspect of sexual dimorphism. More concrete information is known about the structure of the fur, since a tuft of hair from an aurochs that lived in 13th-century Poland, near Pu³tusk (40 km north of Warsaw), has been preserved. It is still connected to the end of a shinbone; its greatest width measures 77 mm (Ryder 1984), and since this varies from 48 to 68 mm in domesticated cattle, the shinbone must have belonged to an aurochs. A sample of one hundred hairs was used to determine the diameters of the hairs and their distribution. They have been listed in Table 9, where they are also compared to those of prehistoric and present-day cattle. The table shows that the hair diameter increased in the course of several centuries, while the occurrence of double fur became more rare (Ryder 1980, 1984). In wild animals, such as the European bison, deer and sheep, this fur consists of two layers, the outer and the under coat. Under coat hair is thin, woolly and relatively short, while outer coat

161

Black Cyan Page 162

162


Table 9. Comparison of hair diameters of a number of cattle breeds and bovine species of various origins and periods (compiled from Ryder 1980 and 1984).

Origin

Aurochs (Pu ltusk) European bison Heck cattle Domestic cattle: Skara Brae (England) Rochford (England) Aberdeen (Scotland) Sebay (England) Walcheren (Netherlands) Chillingham (England) Shorthorn (England) Highland (Scotland) §

Period

Hair diameter (µm)

Average Double fur diameter (µm) (yes/no)

13th century Present Present

18-64, 70, 72 12-66 10-60

36,6 39 30

Yes Yes Yes

Neolithic Middle Ages Middle Ages Middle Ages A.D. 1000 Present Present Present

8-34, 60,80 20-74, 82 12-70, 78, 86, 90 10-124 28-118 22-160 14-100 14-72

15 38,1 33 35 75 56 44 41

Yes Yes Yes No No No No No

hair is thicker, stiffer and longer. They can be clearly divided into two diameter groups. While during the domestication process of sheep, a conscious selection in favour of thinner, woolly hairs (the ‘wool’) took place, there was an unconscious selection for thicker hair in domestic cattle. The aurochs and cattle from the Neolithic period and the Bronze Age still had double fur. This clearly began to change during the Middle Ages, and the division between the two types of fur was almost entirely lost in later cattle breeds, because all kinds of intermediate thicknesses started to occur. The proportion of thick hairs increased at the expense of the proportion of thinner hairs. The opposite took place in (wool)sheep, where selection took place in favour of the woolly hairs. In the older sheep breeds, which underwent little wool selection, some of the original double fur may still be found. Apparently, it seems to have been less important for domestic cattle to retain their double fur. The extreme demands made by the northerly climate could be met by stabling the animals in winter. In a warm, humid stable, double fur is probably a disadvantage for the regulation of

Black Cyan Page 163


the body temperature, so a different type of fur from that of their ancestors may have been more suitable to domestic cattle. As a result, the double fur gradually disappeared. If it is reasonable to assume that aurochs had double fur, like so many animals in cold areas, there was probably also the occurrence of summer fur and winter fur. The winter fur is thicker and longer, and is shed in spring to be replaced by the shorter summer fur (Ryder 1976). In European bison, this summer fur is fully developed around the mating season (August) and later grows into the longer winter fur (Pucek 1986). In view of the close relationship between aurochs and European bison, these fur changes probably also took place in aurochs. The moulting process is likely to have been a little more detailed than is often recorded, and the hairs may not simply have been shed only in spring. Studies of the hair growth and moult in present-day cattle (Black Pied dairy cattle) show that besides the spring moult, moulting also takes place in autumn, though on a smaller scale than in spring (Udo 1978). The fur of the cattle in question underwent the following changes in the course of a year: - In spring, the great majority of hairs are renewed. The long and short winter hairs are shed and the shorter summer fur appears, from which the thin woolly hairs are almost entirely lacking. - In autumn, approximately half of the (summer)fur is replaced. Woolly hairs appear once again, and the hairs that were already present in the summer grow into longer winter hairs. The autumn moult is far less marked than the spring one. - Apart from the spring- and autumn moult, some hairs are shed regularly in the course of the year. The general image presented by many kinds of animals is that their hair diameters, as a result of the many woolly hairs, are smaller in winter than in summer (Ryder 1976). In summer, the fur is less long and dense than in winter. In short, the summer fur is thinner and lighter than the winter fur. A double fur of outer and under coat would occur particularly in winter, therefore.

163

Black Cyan Page 164

164


As the nuances of the process of autumn moulting are difficult to ascertain, it may well take place in a far larger number of animals than is generally assumed, including the European bison and its relation, the aurochs.

Hide Bonar (in: Gesner 1602) remarks the following about a specific part of the skin of the aurochs: ‘The ‘corrigium’, however, is very thick, so that on account of its hardness it is not fit for anything but a horse’s bridle and -saddle, since it does not tear at all’. The author does not explain which part of the skin is meant with ‘corrigium’. Nowadays, horse’s saddles are preferably made from the part of the cow’s hide that used to cover its back, since this is considered the heaviest and strongest part. For the sake of comparison: European bison skin can generally be used quite well for making belts. The back part in particular is seen as the best, and therefore the most valuable (Heptner & Naumov 1966); in the early days, European bison poachers would often only take home this part. It seems reasonable to assume, therefore, that Bonar used the word ‘corrigium’ to indicate the part of the aurochs hide that used to cover its back.

Summary On the basis of often-scarce data, descriptions are given of a number of physical characteristics of the aurochs, such as its hooves, udder, hair and hide. With regard to the hooves, no definite difference can as yet be indicated with those of current domestic cattle, excepting their size. The udder of the aurochs cow was small and barely visible. The structure of the fur was studied on the basis of a tuft of aurochs hair. Only very small details could be noted about the hide.

Black Cyan Page 165


7.5. Summarising table Characteristic

The appearance of the aurochs on the basis of the results found in Chapter 7

Fur colour

†† Deep blackish brown to black. In winter, the fur is likely to have been longer and denser than in summer.Along the length of the back, a narrow stripe of a lighter colour, probably greyish.Black curled hairs between the horns and on the forehead. The area of the snout was light-coloured. ‡‡ Reddish brown. Black in rare cases.The winter coat is likely to have been longer and denser than the summer fur. Calves Reddish brown. Within half a year (after birth), the male calves would turn the colour of adult bulls. The female calves kept their reddish brown colour. Outside Europe There is no reason to assume that the colour of the aurochs outside Europe was clearly different than inside.

Horns

The horns of the aurochs always showed a characteristic shape and position. The spirally curved horns were pointed forward and curved in. The horns were light-coloured with dark tips.

Size

The bulls had a shoulder height between 160 and 180 cm. The withers height of cows was around 150 cm. Body weight possibly like that of European bison and banteng.

Udder

Small, hardly visible.

Body shape

Since the height of the withers was approximately equal to the length of the trunk, the animals were squarely built. The withers were barely or not at all raised.

165

Black Cyan Page 166

166


8. CHANGES RESULTING FROM DOMESTICATION Apart from the wolf, which had already been domesticated to a dog during the Pleistocene, the aurochs was one of the first animals to be domesticated in the Early Holocene period, along with the sheep and the goat. The domestication process was very gradual, and can only be dated approximately. At its start, the newly created animal bore a strong resemblance to its wild counterpart, since its body size had only just started to decrease. Even when the domestication process has already proceeded significantly, this is not necessarily visible in the size of the bones. In addition to this, sexual differentiation and castration may interfere with the clarity of any assessment of the bones found. People probably started to domesticate wild animals to have them, and their produce, more readily at their disposal. Hunting is full of dangers and uncertainties; keeping the animals nearby made it easier to attune supply and demand, and decreased the risk of shortages. The domestication of the aurochs is assumed to have taken place, not in a single area, but in several centres. Reports seem to indicate that domestic cattle may originate from various places in the Near East and Southwest Asia (Payne 1991). The domestication process probably started around 7000 B.C. (Brentjes 1967, Bradley et al. 1996). Judging from the pictures of that period, it took at least several thousand years before specific cattle breeds could be distinguished. Besides the development of different breeds, a division into two main groups evolved: domestic cattle with humps at the withers (zebu cattle), and domestic cattle without (taurine cattle). The earliest domestic cattle are all long-horned but hump-less; their horns still resemble those of their wild ancestors. It originates from countries like Turkey, Iraq and Iran (Payne 1991, Troy et al. 2001, Edwards et al. 2004). Not until around 3000 B.C. did short-horned domestic cattle, as mutants resulting from the domestication process, appear in Mesopotamia. Around 4000 B.C., the first zebus appeared in Mehr-

Black Cyan Page 167

CHANGES RESULTING FROM DOMESTICATION

garh (Baluchistan, Pakistan), where they developed in a drier and warmer climate. From their areas of origin, the newly bred cattle were spread around the earth to Europe, Africa, India and Southeast Asia. Two routes may have led to China, one from West Asia to North China (short-horned cattle) another from Southeast Asia to the South (zebus) (see Ch. 4 and Li et al. 1991). Zebus were not only distributed to the East, but also to the West, to Africa. In addition to the domestication centres mentioned here, the aurochs is thought to have been domesticated in other places, including Northeast Africa (Bradley 1996, Troy et al. 2001) and Hungary (Bökönyi 1974). The fact that Neolithic bone material allegedly contained socalled ‘transitional bones’ is an argument in favour of the Hungary theory. These are bones whose size is between that of aurochs- and domestic cattle bones, which could indicate a smooth transition between the wild and the domesticated species. However, research in South and Central Germany shows that domestic cattle were already clearly distinguishable from aurochs in Early-Neolithic cultures (Linear Pottery Culture), and that the two species had very few features in common (Benecke 1994a). The only similarity seems to have been that a small number of domestic bulls and castrates were the same size as aurochs cows (see Fig. 17). Neolithic cattle not only distinguished themselves by their significantly smaller size, but also by the extent of their sexual dimorphism, which was much more limited than in aurochs (40% less; Benecke 1994a). Limited sexual dimorphism is a well-known characteristic of domestication in cattle (Bohlken 1962). Naturally, crossbreeding between domestic cattle and aurochs in Europe cannot be completely ruled out. Domestic cows may either have escaped the farmers’ attention or been kept in a feral state, where they may have been covered by aurochs bulls. Schneeberger (in: Gesner 1602), Commendon (in Flechier 1695; Commendon saw aurochs in Königsberg, in 1563) as well as Œwiêcicki (1634) report that male aurochs would visit and cover domestic cows. Schneeberger says: ‘But they hunt the aurochs that have been spotted covering domestic cows, since the latter, although they will conceive, will either miscarry, or give birth to non-viable young.’

167

Black Cyan Page 168

168


Moreover, aurochs calves were sometimes caught and used. Schneeberger already wrote: ‘More than once, they separated the calves from the aurochs and brought them to domestic cows to be raised by them; this attempt failed, however, because they all died.” Reading these quotations, one cannot escape the thought that aurochs and domestic cattle may have become incompatible due to their long time apart. Genetically they would have been too separate to crossbreed well. However, since taurine cattle can easily crossbreed with zebu cattle, in spite of the fact that their wild ancestors separated some 700,000 years ago (MacHugh et al. 1997), genetic incompatibility cannot be the cause. The reported failure of aurochs-cattle cross-breeding rather seems to indicate poor husbandry (illnesses, infections), to point at the difference in size between (aurochs) bulls and (domestic) cows, or to result from the unwillingness of the farmers. Neolithic farmers will generally have been little inclined to crossbreed their cattle with their wild ancestors (Benecke 1994a). In fact, they are more likely to have tried to prevent this: relatively speaking, their cattle were a high-quality product with respect to their adjusted size, manageability (docile), productivity (larger fat- and milk yield), and fertility (more frequent and at an earlier age). Crossbreeding with wild counterparts will inevitably degenerate the breed in such a case. Besides, mitochondrialDNA research on European aurochs- and Neolithic cattle bones has shown that there is a clear genetic separation between the two groups, which points to a small imported founder population of cattle, and no hybridization with aurochs afterwards (Bollongino et al. 2003). As the domestication process was more prolonged and man became more selective, domestic cattle became increasingly different from their wild ancestors. The changes that took place not only had to do with the ‘tameness’ of the animal, but also with all kinds of bodily characteristics, such as colour and size. Wildlife colours disappeared more and more and were replaced by an almost unlimited range of colours, which had actually lost their function. The first thing that stands out when one considers the dimensions of domestic cattle bones is the fact that they have become smaller than the bones of the original aurochs (see Table10). The horn core and the skull, as well as the bones of the limbs are an illustration of this fact.

Black Cyan Page 169


Table 10. Survey of the dimensions (in mm) of the bones of domestic cattle, compared to the dimensions of Holocene aurochs bones.

Bone27

horn core: girtha horn core: lengtha skull: basal lengthb skull: smallest forehead widtha humerus: lengthc radius: lengthc femur: lengthc tibia: lengthc metacarpus: lengthc metatarsus: lengthc

Domestic Cattle ††

‡‡

Aurochs (Holocene) (from Tables 3 and 4) †† ‡‡

160 – 281 190 – 480 355 – 510 159 – 221 325 – 381 291 – 348 409 – 480 357 – 433 207 – 244 239 – 276

92 – 202 174 – 305 354 – 510 133 – 210 302 – 355 293 – 336 399 – 440 349 – 413 204 – 248 229 – 274

275 – 415 450 – 820 525 – 612 207 – 289 386 – 440 345 – 392 470 – 524 452 – 488 239 – 278 270 – 315

185 – 275 330 – 530 455 – 535 170 – 205 355 – 367 315 – 345 427 – 445 400 – 420 230 – 253 268 – 283

This survey does not list the more extreme dimensions of the bones of dwarf and giant breeds, nor of the bones of hornless and extremely large-horned cattle breeds. The dimensions for domestic cattle derive from: a – Grigson 1974 b – Grigson 1974 and Bohlken 1962 c – Matolcsi 1970 The most important domestication characteristics in cattle breeds are as follows: Decreased height of the withers. The height of the withers has decreased as a consequence of the smaller limb bones (Matolcsi 1970). This feature was already very apparent during the Neolithic period. In the Iron Age and in Roman times the average bone size increased again, until it reached a lower level than in the Neolithic period during the Middle Ages. After that, the average bone size increased to that of contemporary cattle breeds. 27

See Table 4 for an explanation of the Latin terms.

169

Black Cyan Page 170

170


Together with the shoulder height, the trunk length, compared to that of the aurochs, decreased initially (Matolcsi 1970). It followed the same temporary variations as the height of the withers. The trunk length of contemporary cattle breeds is now more or less equal to that of the aurochs (Matolcsi 1970). As a consequence of the marked decrease in the shoulder height, combined with more or less equal trunk length (compared to the aurochs), the legs have become shorter, also in relation to the length of the animal. As a result, contemporary cattle breeds seem to be longer than the aurochs, which were rather more ‘squarely’ built (see Ch.7.1). Decreased skull length. Domestication also resulted in a shortening and narrowing (relatively speaking: a broadening) of the skull (Bohlken 1962, Grigson 1974). This is a common feature in domestic animals (e.g. dog, pig). Conscious selection resulted in extreme shortening of the skull in canine breeds like the bulldog and the Pekinese. The South American Niata presents a bovine example of a shortened skull (Benecke 1994b; see Fig. 26).

Fig. 26. Comparison of the skull of an aurochs (left) with that of a South American Niata (right). In domestic cattle the domestication process led to shortening of the skull, with the Niata as an extreme example of that process.

Black Cyan Page 171


The decreasing skull size also resulted in smaller brain content. The brain content of the aurochs bull was 760 - 830 cm3, of the cow 650 780 cm3 (Von Leithner 1927). Studies of domesticated cattle have shown that their brain content has decreased by approximately one third compared to that of aurochs28. Quantitatively speaking, the brain content decreased in proportion to the decrease in skull size. However, there was also a decrease in quality. Domesticated animals such as cats, dogs, and pigs are known to have changed their ways of finding food, their ways of mating and their social behaviour, compared to their wild ancestors (Herre & Röhrs 1990). Moreover, their senses have deteriorated: domesticated animals generally have a poorer sense of sight, smell and hearing than their wild ancestors. Animals in the wild probably need more brain content and a longer skull to fully make use of their senses. The differences in this respect between domestic cattle and aurochs are not known, but will undoubtedly exist. Changes in the shape of the horns. During the domestication process, all kinds of new horn shapes developed. Increasingly, the original aurochs horn shape receded into the background. Instead of their original forward and inward position, the horns started to point upward, outward and backward. Hornless-ness was no longer prey to natural selection and occurred more and more frequently. Decreased sexual dimorphism. The difference in size between the sexes is much smaller in domestic cattle than in the aurochs. The dimensions of comparable bones of domestic cows and bulls overlap to a much greater extent than they did in the case of the aurochs. Sometimes the difference has almost completely disappeared, for example in the metacarpus. Man eliminated the mechanism of natural selection, which creates much bigger bulls than cows, in domestic cattle. In a cultured state, man adapts the size of his animals to his wishes; marked sexual dimorphism is no longer functional in that situation. On the basis of the length of the humerus (conversion factor 4.14), the difference in the height of the withers between the domesticated bull 28 Nickel et al. (1975) report that the weight of the brain of domesticated cattle (with a body weight of 500 to 600 kg) varies around 410 to 480 grams. Klatt’s research (1912) shows that the brain weight in grammes is more or less equal to the brain volume in cubic centimetres. Thus, the corresponding brain size would be around 410 to 480 cm3. Brain size may vary considerably with body size and nutrition.

171

Black Cyan Page 172

172


Fig. 27. In domestic cattle domestication led to more diversity in the size of the horns. Both polledness and extremely large horns occur, as in the Franqueiro in Brasil (after Nehring 1888). These horns have a length of ca.150 cm and a girth of 48 cm at the base.

and cow turns out to be much smaller than in the aurochs: 10-11 cm (8 %) compared to 13-30 cm (9 to 20 %). The minimum and maximum values represent the differences in shoulder height between the smallest cow and bull on the one hand, and between the largest cow and bull on the other (derived from Table 10). Increased variation in size. Domestication generated nearly a thousand different cattle breeds and -varieties, which show great differences in size (Mason 1996). There are dwarf breeds like the Dexter (shoulder height 80-110 cm; Felius 1995) and giant breeds like the Chianina (bulls of over 180 cm; Frahm 1982). Extremes also exist with respect to the size of the horns: there are hornless cattle (Galloway cattle, Fjäll cattle), but also large-horned breeds, like many African cattle (Watusi, N’Dama) and the Franqueiro in South America. The latter’s horns may measure up to 150 cm (Nehring 1888) (see Fig. 27). Obviously, the variation in bone size in domestic cattle is larger than in the aurochs (Bohlken 1962, Kobryñ & Lasota-Moskalewska 1989). The variation in the overall dimensions of the aurochs was also smaller than is the case with domestic cattle.

Black Cyan Page 173


Increased variation in colour. The colour of the fur of the aurochs bull differed from that of the cow. For the separate sexes, colour varied within specific restricted limits, which were established by the predators, the nature of the habitat, and the social relationships within the species. These forces cease to exist the moment domestication begins, after which only man may still want to select by colour. Colours being no longer limited, and all sorts of colour mutations allowed to continue to exist, a whole range of colours of domestic animal evolved, such as white, grey, yellow, brown, red and black (see Photo 16). In addition to and in combination with these colours, all kinds of markings develop, consisting of small and large white spots and zones. The same process may be seen in the domestic yak in Central Asia. Early maturity. Lowering the age at which the animals would reach maturity and would be able to mate was one of the most important selection targets with regard to domestic cows, in particular in the earliest farming communities. Early maturity presented various advantages (Steehouwer 1987). The cows were able to calve more often and from an earli-

Photo 16. A group of Hungarian steppe cattle in the former East Berlin zoo (Photo: T. van Vuure). The coat colour of these cattle varies from light grey to dark grey. This cattle type has left a clear mark on the breeding of Heck cattle

173

Black Cyan Page 174

174


er age, both the bulls and the cows would remain smaller (smaller shoulder height) and they would amass fat at an earlier stage (relatively small muscle mass). Early-mature cattle breeds developed particularly in Northwest Europe, and included both dairy breeds (Jersey) and meat cattle (Aberdeen Angus). In Southern Europe, late-mature cattle breeds may be found (Chianina, Charolais, Steppe cattle), since these are better suited to fulfil their roles (as draught animals), and to eat the food available (agrarian waste). The aurochs probably mated at a relatively late age, like other related bovine species (Mason 1984)29. The cows of early-mature cattle breeds often calve at 24 months or even earlier. The cows of late-mature breeds generally calve at age three, sometimes four (Frahm 1982). According to Frahm, the bulls of some very late-mature breeds, such as the Maremmana and the Pugliese, only reach maturity when they are between the ages of five and seven. Besides genetic factors, good nutrition may influence growth and advance the time an animal reaches maturity. Increased udder size. The udder of the aurochs was very small, and could feed one calf. Nowadays, udders of that size are still found in the cows of Spanish fighting cattle. During the past millennia, selection for greater milk production has resulted in domestic cows with much larger udders.

Summary Probably around 7000 B.C., the aurochs was turned into a domestic animal (‘domesticated’) in the Middle East. Domestication could secure a more stable source of food than hunting. As a result of domestication, as well as intentional and unintentional selection, the external and internal characteristics of the aurochs changed into those of the hundreds of cattle breeds that were produced by these processes. The variations in body size, fur colour, horn shape and udder size increased. The differences between cows and bulls (sexual dimorphism) decreased. Physiological processes, such as early or late maturity, reproduction, and milk production changed as well, and were adapted to the needs of man. 29 In European bison, cows calve for the first time at 48 months on average (Krasiński & Raczyński 1967). Physical maturity of cows is actually reached at the age of five, in bulls at the age of seven (Krasińska & Krasiński 2004). However, skull size and horn size may continue to increase slightly until a very old age (Empel 1962). Between the ages of six and nine, the length of the humerus and the femur may still increase slightly as well (Kobryńczuk & Kobryń 1975).


ECOLOGY OF THE AUROCHS AND OTHER WILD CATTLE

9. ECOLOGY OF THE AUROCHS AND OTHER WILD CATTLE 9.1. Habitat 9.1.1. The natural landscape of Europe

Introduction The ecology of an animal species is directly connected with the landscape surrounding it and the inherent possibilities for finding food, water, hiding places and the like throughout the year. At the time, the aurochs in its large distribution area encountered various climates, types of vegetation and soil types. Within this distribution area, the animal will have sought suitable stands to fill its needs. Because most is known about the European aurochs, and the circumstances in the Low Countries during this early periods are of particular interest here, the natural landscape of Central and West Europe, which played a crucial role in the life of the aurochs, will be focused on. Elucidation of the European aurochs habitat may be illustrative and clarifying, moreover, of its habitat in other continents, about which there is very little information. Twenty or so years ago, the appearance of the European landscape during the various climatological periods of the Holocene was not a topic for discussion. Generally speaking, the prevailing opinion was that closed forests formed the main component of the natural vegetation; as a result, these largely determined the outlook of the landscape. At most, these forests were allegedly alternated by large peat bogs. This view was mainly based on pollen research undertaken on soil samples (Bottema 1987; see Fig. 28). Around 1980, a different view started to take root: the earlier European forests had not been closed and immense at all, but very open. According to this view, large grasslands had been alternated by brushwood, patches of forests and solitary trees (Van de Veen 1975, Poortinga 1981). This type of landscape would have been created

175


176


and maintained by grazing and browsing large herbivores, such as aurochs, European bison, wild horse, red deer and others. Vera (2000) tried to give scientific substance to these ideas about an open, park-like landscape, which took shape in the early 80s, on the basis of palynological and forest-ecological data, replenished with information from historical sources from the Roman period and the Middle Ages. His re-

Fig. 28. Brief overview of the climate-, vegetation- and habitation history of the Low Countries during the Holocene. To the left: time scale in years BC (Source: Groninger Instituut voor Archeologie GIA/ former BAI).



search evoked lot of criticism. To add clarity to the discussion, and because additional information continues to be found, also here an explanation will be offered of any evidence that may throw more light on the type of natural vegetation in Europe at the time. The fundamental question, whether large (and small) herbivores are able to open up forest vegetations and to keep them open, will be dealt with in more detail in Chapter 9.4. The data used here stem from scientific fields like entomology (insect research), palynology (pollen research) and the study of written sources from the Roman period and the Middle Ages.

Insects An aspect of the former natural vegetation that has not yet been the subject of much research, compared to pollen research, is the occurrence of insects in earlier times. In spite of this, Dutch, French (Andrieu et al. 1997, Ponel 1995, Ponel et al. 2000, 2001) and, more particularly, English research have produced a relatively clear picture of the insect species that occurred in the course of the Holocene and of their relation to the vegetation in that period. The occurrence of certain insect species and their relative numbers may warrant conclusions about the vegetation at the time. In the Netherlands, Brussaard (1985) researched the occurrence of the minotaur beetle (Typhaeus typhoeus). This beetle is found on sandy soils, preferably in heather fields and in pine forests, more particularly along the paths and in the open spaces there. It lives from the dung of larger mammals (rabbit-sized and larger), without which it cannot exist. The research of fossil remains and digging traces of this insect shows that it still occurred in this country up to 6800 BC. No traces have been found from the period between 6800 and 3000 BC, but they were once again clearly present after this date. An explanation for this may be that from 6800 BC, forests became increasingly dense (based on pollen research), which may have been unfavourable to this beetle, which occurs in open vegetations, preferably on bare soil. A different explanation could be that

177


178


the decline of the mammal populations, caused by the more intense afforestation, resulted in less dung, so that the dung beetles also decreased in number. Both explanations are directly or indirectly connected to the increasing afforestation. The decline of the forest area after 3000 BC is ascribed to the fact that cattle-raising man opened the forest at a large scale. He brought many cattle and sheep along, which resulted in a lot of dung for the dung beetles. That no traces of these beetles have been found for the period 6800-3000 BC does not mean that they did not occur. The difference between the periods before and after this period is very striking, however, and seems to indicate, if any, a very rare occurrence of the animal in the intervening period. English research into the early occurrence of insects has been published in the works of Osborne (1978) and Girling (1989), among others. Osborne’s research of several places reveals a clear distinction between the various phases of the Holocene, as regards the composition of the accompanying insect fauna. The material found shows that, from the Early Holocene up to circa 3000 BC, the insect fauna was completely dominated by species specifically tied to trees. Especially the species associated to dead trees, in various phases of decomposition, were prominently present. In the beginning, during the Early Holocene, these were species that lived on willow and birch. As the climate became warmer the number of species grew. After England became divided from the mainland by the Channel, around 5000 BC, the number of new species dropped to a minimum. During the Atlantic period, characteristic species such as Ernoporus caucasicus (on linden), Dryophthorus corticalis (on oak), Melasoma aenea (on alder) and Hylesinus species (on ash) occurred. Insects typical for coniferous trees were also present, such as Hylastes brunneus and Eremotus species. Of the species living in dead wood, particularly, many no longer occur or are extremely rare in England today. Osborne draws the conclusion that in the period in question, England must have been covered predominantly with dense forests, and a habitat like dead wood would then have been present in abundance. From around 3000 BC, the range of species started to change. Occasionally, the changes took place so quickly that certain soil layers reveal a marked contrast between two successive periods, with regard to the insect



species involved. During the period in question, the occurrence of species that are characteristic for grasslands with large grazing animals is conspicuous. Insect species that live off trees were not or scarcely present. In this period, dung beetles of the genera Aphodius and Ontophagus and

Fig. 29. Overview of the relative occurrence of several types of landscape during the successive periods of the Holocene in England, based on the number and the species of insects found at the various sites (after Dinnin & Sadler 1999). As man occupied more ground surface, the number of tree-tied insects diminished.

179


180


others first appeared in the dung of large grazing herbivores; they were lacking before this time. Other new species, such as Adelocera murina and Phylloperta horticola, lived on grass roots and are characteristic of high grassy vegetations. From a site of 1000 BC, an insect fauna is also known that is typical for an open meadow landscape. Apart from very occasional finds of insects that lived on dead wood not a single insect species was found that was associated with trees. Not until the Roman period did a composition of insect species appear, similar to the ones we still know nowadays, and which is attached to a mosaic landscape of meadows, fields, hedges and scattered thickets. During that period, species that are characteristic for largescale cereal production and large cereal stores, such as the grain weevil (Sitophilus granarius), appeared for the first time. Such cereal-eating species are often of subtropical origin, brought along by the Romans. Girling (1989), at an excavation at Hampstead, was able to establish the succession of insect faunas from the Mesolithic situation up to this point. Parallel to this, the development of the vegetation could be charted on the basis of pollen found at that site (Greig 1989). The most striking change in the vegetation found there took place at the transition from the Mesolithic, in which man lived as a hunter-gatherer, to the Neolithic, in which agriculture and the raising of cattle began to take place. Archaeological finds showed that this transition took place there around 3000 BC, up to which point the landscape must have been densely forested with oak, linden, elm and hazel. The proportion of grass in the pollen was about 5% at the time (see ‘Pollen research’). After 3000 BC this proportion increased markedly, while that of forest declined strongly. The situation is also clearly reflected in the insect species. Before 3000 BC, species attached to forest dominated, including a large proportion of dead wood consumers. After 3000 BC, the latter category of insects barely occurred any longer. The proportion of insects that occur on living trees clearly continued to decrease. In contrast, species that live on graminoids and forbs increased in number and diversity. After 3000 BC, dung beetles appeared as well, though they were not found before this time. Man and the cattle that accompanied him apparently created the right living conditions for this group of insects.



To sum up, it may be said that the insects found show that up to around 3000 BC, forests, which contained a lot of dead wood, dominated England and part of Continental Europe. After this date, man started to systematically fell these forests and change them into grasslands. In the Roman period, the large-scale man-made landscape that we still know today appeared. The composition of insect species in England has hardly changed during the past 2000 years. A survey of the research undertaken by Osborne and Girling, and of many other English entomological studies, has been provided by Dinnin & Sadler (1999). The results of this survey are shown in Fig. 29, which charts the relative occurrence of the various habitats indicated by the insect species found for the periods of the Holocene.

Pollen research Plant pollen, provided it is well preserved in soil layers, may remain unchanged and recognizable to species for a long time. As a result of this, pollen of, sometimes, tens of thousands of years old (up to, even, millions of years; Janssen 1974) may still be found and provide data about the plant species that were present during a certain period (Guthrie 1990, Kahlke 1994). In addition to this, the scale on which they occurred at the time may also be gathered to a certain extent from pollen research. Conclusions about the prevailing climate (temperature, precipitation) may thus be drawn. Pollen research may provide an insight into certain aspects of the early natural landscape of Europe. Since the aurochs must be considered a specialised grazer (see Chapter 9.1.2), its habitat must have contained graminoids (grasses, sedges and such) in sufficient quantity to supply its needs. As such plants occur especially in open, more or less treeless areas, the first issue is to establish the degree of openness of the forest vegetation. The more open (more treeless) vegetation is, the more graminoids will grow there; because of their wind pollination these are easy to detect in pollen analyses. It needs to be determined, therefore, whether it is possible on the basis of pollen research to conclude if, and

181


182


if so, to what extent, graminoids occurred in a specific vegetation, and how open the landscape in question was30. Several researchers have asked themselves whether the rate of openness of certain vegetation may be derived from a pollen diagram (Janssen 1974, Aaby 1994, Zoller & Haas 1995). In such a diagram, distinctions can be made between the percentage of arboreal (AP) and non-arboreal pollen (NAP): the pollen of trees and of grasses and forbs, respectively. On the basis of European and Russian research, Zoller & Haas have established that NAP percentages of between 40 and 50 or over indicate an absence of closed forest. The Russian study shows NAP percentages in steppe areas of between 70 and 75 and in tundra vegetations and steppe forests of between 45 and 50. NAP values below 40% already indicate the presence of forest. Research in North China by Liu et al. (1999) and research by Mitchell (2005) corroborate such a relation. Aaby (1994), also based on research by Jonassen, has found comparable results. He compares four stages of openness to the current pollen precipitation produced in the landscaped in question, i.e. forests, (small) open spots in forests, (large) fields and meadows in forested areas, and totally open areas. This shows that the percentage of NAP in closed forests is generally lower than 20. In forests with a very open structure, or in open spots in forests, this percentage lies between 20 and 40, depending on openness and tree species. As the latter produce more shadow, the forest floor will be more scarcely overgrown. In Pinus forests, the NAP percentages were higher than in Picea forests. Fields and meadows in forested areas, with a distance of 0.2–1 km to the forest edge, already yielded more than 50% NAP. Very open ground areas showed values of between 45% and 100% NAP. The latter value is not always 100% because certain species of tree pollen from very remote areas may be blown in by the wind. It became evident that, with the help of the 30 Important support of pollenanalytical research is formed by research on plant macrofossils (fossil seeds, fruits, branches etc.). It was found that flanking research on macrofossils (but also on insects, see preceding Chapter, or molluscs (Gedda et al. 1999)) not only confirmed the results from pollenanalytical research, but in some cases could provide more precise and accurate interpretations as well, or fill gaps in the pollen research (Birks & Birks 2000, Svenning 2002). Moreover, Svenning found a strong statistical (Spearman) correlation, showing that both methods correlate well. Such research on both pollen and plant macrofossils may be found in: Andrieu et al. 1997, Hannon 1999, Zazula et al. 2003, Jackson et al. 1997, Watts & Winter 1966, Bos et al. 2004, Goetcheus & Birks 2001, Regnell et al. 1995, Garry et al. 1990, Hölzer & Hölzer 1988, Lynch et al. 2004, Gobet et al. 2003, Ammann et al. 2000, Kneller & Peteet 1999, Barber, K.E. & Clarke, M.J. 1987, Baker & Drake 1994, Hu & Davis 1995, Lagerås & Bartholin 2003, Brombacher & Jacomet 1997, Barnekow & Sandgren 2001, Litt 2000 and others.



composition of the pollen, the rate of openness of the vegetation could be ascertained. From pollen diagrams of the period just before European man started with agriculture and the raising of cattle, the percentage NAP is revealed as 15 at most, and often even less than 5 (Zoller & Haas 1995). This would indicate a densely forested landscape. As man increasingly opened up this landscape during the Neolithic period (4500 – 2000 BC), by cutting and burning the forest, the percentages of NAP in the pollen diagrams rose accordingly. The NAP mainly contained species that are characteristic for fields and meadows (Janssen 1974). The inversed process, i.e. the closing up again of fields and meadows with forest, also took place, as is reflected in pollen diagrams for many places in the South of the Netherlands in the period AD 400 – 500 (Teunissen 1990). These diagrams clearly show that the original open landscape closed up again with forest after the departure of the Romans from these areas, which became depopulated as a result, after which the fields closed up with trees. In later periods, the forest once again had to yield to the increasing population, after which the final deforestation took place. This information found in the pollen diagrams is supported by archaeological data that indicate a decline and subsequent increase in population (Teunissen 1990, Kooistra 1996). Major changes in the proportions of AP and NAP also took place as early as the transition from the last ice age to the Holocene, about 10,000 years ago. This is shown by data about Europe (Kahlke 1994) and North America (Guthrie 1990) as well as Siberia (Hahne & Melles 1997, see Fig. 30). During the last ice age, an extensive zone of cold, dry steppes was present in Europe, Asia and North America, which was inhabited by a very diverse mammal fauna, including four large specialised grazers (woolly mammoth, woolly rhinoceros, steppe bison, horse) and two large browsers (giant deer and reindeer). Only locally bushes and trees were present along lakes and rivers (Bradshaw et al. 2003). Pollen diagrams of that steppe vegetation show an NAP percentage of 90. This NAP derives from grasses, sedges and forbs (especially Artemisia species). Alder, willow, birch and pine form the remaining 10%. On the basis of these proportions, it is reasonable to assume that this was a nearly treeless area, comparable to still existing Russian and North

183


184


American steppes as far as openness is concerned (Guthrie 1990). In such an ecosystem hardly influenced by man, the grass and forbs vegetation was able to produce a lot of pollen, in spite of the many species of herbivores that fed upon it. Vera (2000), as an explanation for the low pollen proportion of grasses in many pollen analyses from areas grazed by large herbivores, proposes that this was due to the many large herbivores, which kept the grassy vegetations so short that grasses could not flower and produce pollen. The example of the grassy plains in the northern ice age mentioned here, as well as the pollen diagrams of African savannah areas at Victoria Lake (Kendall 1969), clearly show, however, that populations of large herbivores regulated by nature are not able to prevent the massive flowering of grasses. In natural circumstances, grass vegetations are able to produce large quantities of pollen. Also in cultivated circumstances, grass may be identified in proportion to its occurrence by pollen analyses, as was demonstrated by McAndrews (cited by Janssen 1874) and Aaby (1994). Only where human manipulation resulted in overgrazing by cattle (as for instance in the Imbos-experiment by Van Wieren (1988)), little grass pollen could be identified in proportion to the grass area (Groenman-Van Waateringe 1993). There the (very) high density of cattle prevented massive flowering of grasses and thus the formation of pollen. About 10,000 years ago, the steppe vegetation changed in a relatively short period, under the influence of the warmer climate. The proportions of grass and forbs decreased markedly, while that of forests increased. In Europe, initially these were birch and pine forests, later followed by elm, oak, hazel, linden and alder, among others (Bottema 1987) (see Fig. 28). Beech and hornbeam forests followed at a much later date. As has been shown, the NAP percentage during the Atlantic period in Europe was usually 15 at most, often even less than 5 (Zoller & Haas 1995), which indicates a great degree of forestation. The NAP in such a case derives from plants that grow under trees or occur in small open spots in the forest. When such open spots grow, this is reflected in the pollen diagrams. Tamboer-Van den Heuvel & Janssen (1976) were thus able to recognize clearly the local mountain meadows in the pollen of



Fig. 30. Pollen diagram from Lake Lama (Central Siberia), which shows the transition from the grassand herb steppe during the ice age (zone L1) to the warmer period of the Holocene (from zone L7). The L5/L6 boundary lies at ca.10200 BC. (after Hahne & Melles 1997). The occurrence of the poplar (Populus) can be recognized in this diagram as well.

185


186


the beech forests in the Vosges. Meadow pollen dominated in the meadows, forest and fern pollen in the forests. In North America, based on present-day pollen, Anderson & Brubaker (1986), Oswald et al. (2003) and McAndrews (quoted by Janssen 1974) could interpret the character and openness of the forest vegetation. Large quantities of grass and forb pollen were indicative of prairie or tundra, limited amounts of such pollen indicated coniferous or broad-leaved trees. In the former habitat of the aurochs in Europe, certain vegetation types in particular were of special interest to the animal. These were vegetations in river valleys and vegetations subject to the influence of beavers (Castor fiber). In river valleys, the establishment of trees like willow and alder is often difficult because of the extreme wetness of the soil and the possible effects of ice. In beaver areas, settlement is influenced both by the wetness of the soil and by the feeding of beavers, who use trees for food and as building material. An example that shows the open vegetation of a river valley is the pollen diagram of the River Rhine at Wijk bij Duurstede (Steenbeek 1990) (see Fig. 31). The pollen dates from the Atlantic and Subboreal periods, and shows vegetation that indicates open marshy circumstances in which sedges (Carex sp.) were particularly dominant, especially in the PW-3/4 zone. Beside these, grasses played an important role. The water level in such marshes varied, and the soil that produced this pollen is mainly composed of sedge peat. This early, dynamic situation along the Rhine shows a lot of resemblance to the flood plain vegetation along the Missouri in the beginning of the last century, described by Weaver (1960), where sedges also played an important part. As will be shown later, such sedge vegetations probably were an important habitat for the aurochs. The influence of beavers may be found in a pollen diagram from North America (Ohio) (see Fig. 32) (Garrison 1967). This shows that around 13,000 14C years ago (ca. 14,000-13,000 BC), beavers (Castor canadensis) settled a Picea/ Abies forest as a consequence of the warmer climate. Their presence was established from the soil there, which yielded branches of willow (Salix), spruce (Picea) and juniper (Juniperus) that had been gnawed by them. As a result of the beavers feeding,



Fig. 31. Pollen diagram for the banks of the Rhine at Wijk bij Duurstede (Netherlands) (Section WbD 271-1; after Steenbeek 1990) from the Atlantic and Subboreal periods. Here the distinct presence of sedge vegetations (Cyperaceae) can be recognized in the PW-3/4 zone (515 – 370 cm depth).

the proportion of forest decreased and those of grasses, forbs and bushes increased. Grasses in particular showed a marked growth. Comparable beaver activity was also ascertained in other places in Ohio (Garrison 1967). The pollen diagram shown here clearly reveals the influence beavers may have on a forested area, and the link that may be laid to grazers like aurochs. A tree that is hardly, or nor at all found in European pollen diagrams, since its pollen does not keep all that well, is the poplar (black poplar (Populus nigra) and aspen (P. tremula)) (Janssen 1974). The fact that the presence of poplar is thus difficult to prove is used by some authors (Vera 2000) to deem pollen diagrams inadequate for determining the rate of forestation and the species composition of the forest. Poplar pollen, however, has turned out to be well preservable under oxygen-free conditions, such as found on the bottoms of deep lakes, for example (Mott 1978). North-American pollen research has shown that P. tremuloides (related to aspen) played a rather important role during the Late Pleistocene in North America (Mott 1987). Apparently, very advantageous circumstances for this poplar prevailed at the time, either climatologically or for reasons of competition. In North America, the presence of this poplar to this day

187


188


Fig. 32. Pollen diagram for a site at Columbus (Ohio, United States), showing the influence of beavers on the riverside vegetations. Higher up in this profile the proportion of trees (Picea/Pinus/Quercus) decreases in favour of the proportion of grasses (Gramineae) and bushes (after Garrison 1967).

has been charted in pollen diagrams (Mott 1978, Vance 1986). The aspen occurring in Europe and North Asia also played a larger role than is the case today, in large parts of those areas during the Early Holocene (Godwin 1975, Bottema 1987, Hahne & Melles 1997) (see Fig. 30). Seen against this background, the aspen turns out to be a tree that thrives in the more continental colder areas also today, for instance in the northeast and north of Europe, where it plays an important part as a forest tree and attains larger dimensions than elsewhere in Europe. Occurrence of the black poplar is limited to the edges of large rivers, so it plays a subordinate role in the total forest picture. It is naturally present in the most dynamic parts of the flood plain in white willow-poplar forests (SalicetoPopuletum), where it favours the somewhat higher, drier areas on account of its relatively great drought tolerance (Splunder et al.1996, Ellenberg 1986). Depending on the character of a river as regards water drainage, flooding intensity and deposited soil material, the numbers of this poplar may differ strongly from river to river.



An important aspect in the discrimination between open, park-like forests and closed forests is constituted by the plant communities of the mantle and fringe vegetations (Prunetalia spinosae), which are characteristic for forest edges, where closed forests border on open ground, as well as for hedges (Groenman-Van Waateringe 1978). The plants in question love light and warmth, and are tied to this border zone. The forest is too dark for them and the feeding (of animals) or clipping (by man) prevents them from colonising open areas. Characteristic bushes and trees in such vegetation include the blackthorn (Prunus spinosa), hawthorn (Crataegus spec.), dogwood (Cornus sanguinea), common spindle tree (Euonymus europaeus), roses (Rosa spec.), holly (Ilex aquifolium), Guelder rose (Viburnum opulus), traveller’s joy (Clematis vitalba), honeysuckle (Lonicera caprifolium) and brambles (Rubus spec.) (Groenman-Van Waateringe 1978). Many of these are thorny or prickly plants; a characteristic that offers a certain measure of resistance against feeding animals. Beside these bushes and trees, there are a number of forbs that are characteristic for the mantle communities that border on the said communities, such as St. John’s wort (Hypericum perforatum), wild garlic (Allium ursinum), wild strawberry (Fragaria vesca) and hogweed (Heracleum sphondylium) (Troels-Smith, quoted by Groenman-Van Waateringe 1978). The combined presence of these forbs has turned out to be strongly correlated to the combined occurrence of a number of bushes in the mantle vegetations. On the basis of this, it has proved to be possible to trace such communities in pollen diagrams (Groenman-Van Waateringe 1978). It was also revealed that these communities did not appear for the first time before the Neolithic period, when their occurrence was influenced by the activities of man, who opened up the European forest for agriculture and to raise cattle. Only then did the right conditions apparently develop to prompt the appearance of these vegetation types; before this time, the forest was still too dense and too dark to allow the plant species already mentioned to appear together on a rather large scale. Other authors also indicate the existence of such mantle and fringe vegetations. Godwin (1975) states, based on many studies of charcoal finds, that a number of bushes and small trees that are characteristic for the said communities either did not occur until the Neolithic period in

189


190


England, or showed a marked increase right around that time (see Table 11). Before the Neolithic, these species were hardly or not at all present there. Table 11. Trees and bushes that were ascertained for the first time in the Neolithic period in England (with asterisk), or were already present before that period and expanding at the time of the opening-up of the forest (Godwin 1975).

* hedge maple (Acer campestre) * common spindle tree (Euonymus europaeus) * purging buckthorn (Rhamnus catharticus) * southern boxwood (Buxus sempervirens) * common gorse (Ulex europaeus) * rose (Rosa spec.) blackthorn (Prunus spinosa) * wild cherry (Prunus avium)

* whitebeam (Sorbus aria) * apple (Malus sylvestris) hawthorn (Crataegus spec.) * hornbeam (Carpinus betulus) * beech (Fagus sylvatica) * brittle willow (Salix fragilis) ash (Fraxinus excelsior) black elder (Sambucus nigra)

In Switzerland, agriculture was practiced on a small scale already during the Late Mesolithic, and parts of the forest were burnt down for this, as has been proven by Erny-Rodmann et al. (1997). In the course of the Neolithic period a development could be shown here during which open locations increasingly emerged. Pollen and macrofossils of certain plant species indicate that during the Early Neolithic period the arable fields were still small, kept open by man and surrounded by woods (Brombacher & Jacomet 1997). In the Late Neolithic, the arable fields increased significantly (as shown by the increase of (mainly) winter annual weeds). An increase of sloe (Prunus spinosa) and Rosa species indicates the increase of forest edges and hedgerows. A growing number of treading-resistant plants like Prunella vulgaris, Plantago major and Potentilla reptans points to the grazing (treading) of fallow fields. Indicators of pastoral woodland, like heather (Calluna vulgaris), juniper (Juniperus communis) and holly (Ilex aquifolium) are encountered more frequently in pollen spectra. In Germany (Kückhoven, Westphalia), research was done on a well from the time of the Linear Pottery Culture, the earliest agricultural phase



in Central Europe (Kalis & Meurers-Balke 1998). The first building phase of this well took place in 5090 BC, the second one around 5072 BC. In this well, many remains (pollen and plant parts) of plants growing there at the time were found. The interesting aspect is that also many remains of insect-pollinated plants were found. Usually, a relatively small amount of such pollen is found, since it is not dispersed by wind. Vera (2000) uses this argument to question the reliability of pollen diagrams. In the case of Kückhoven, however, it turned out to be possible to ascertain the proportion of plant species pollinated by insects (beside that of wind-pollinated plants), which is not always the case. The landscape in which this early agricultural settlement was established, was characterised not only by openness (with a large proportion of grasses and forbs), but also by its great richness in plant species characteristic for forest edge- and hedgerow communities, many of which are insect-pollinated. These species must have been present in the village or in its surroundings. Comparison with a pollen diagram of a contemporary neighbouring forest area shows that the latter yielded hardly any grasses and forbs (but many ferns) and only few plant species that are characteristic for mantle and fringe vegetations. Clearly, this forest had not been opened up by man (yet). It seems evident, therefore, that the communities at the edge of a forest, in the form in which they are present nowadays in many nature reserves and primitive agricultural landscapes, are a product of the agriculture and cattle raising carried out by man (Groenman-Van Waateringe 1978, Kalis & Meurers-Balke 1998). These communities could not only be found at forest edges, but also in the hedges people used to plant around their villages or fields to keep wild animals or the enemy at a distance (see ‘Frontier forests’). Plant species that naturally occur in limited quantities, at specific stands in the forest, together may find new stands at the forest edges and hedges in the landscape of meadows, fields and bushes opened up by man. Man created a type of landscape that had scarcely or not been present before the arrival of agriculture and cattle raising. The survival of the vegetation types in question, which have a history of over 7,000 years in Europe, have remained dependent on man; if man withdraws, as he did from the South of the Netherlands

191


192


after the Roman period as well as from the Great Wilderness (East Prussia), the forest will take renewed possession of the cultivated land, and the forest edge- and hedgerow communities will disappear.

Roman writers In order to know what the European landscape looked like some 2,000 years ago, the writings of Roman writers are often reverted to. During the period in question, the Romans tried to subject areas like Gaul (France and Belgium) and Germania (Germany and the Netherlands) to their authority. Caesar conquered Gaul around 55 BC; attempts were made to conquer Germania during the reign of emperor Augustus, around the beginning of our era. Roman writers are often reviled, since their descriptions allegedly fail to represent the truth and the writers themselves mostly referred to eyewitness accounts provided by others. It is true that historians such as Tacitus and Dio Cassius and the geographer Pomponius Mela naturally obtained their information from numerous eyewitnesses, but they had to be careful not to tell their Roman readers any nonsense, since many of the latter knew the areas described. Other writers are known definitely to have visited Gaul or Germania themselves. Plinius (circa 70) stayed in Germania as a cavalry officer, and Julius Caesar conquered Gaul when he was a general. Caesar in particular must have been well informed about the nature of the terrain in Gaul. As general of a large, well organised army, he knew the exact sizes of the forests and the marshes and how to march through them, the positions of the fordable places in rivers, where to find cereals, and how the various tribes related to each other politically. He received all this information from scouts, traders, deserters and such. His descriptions of landscape elements possess a large degree of reliability, therefore. The accounts clearly sketch the difficulties the Romans faced during their warfare (Timpe 1989). There were large problems, especially in Germania, so much so that eventually, lasting occupation of this country was abandoned and the Rhine was held as a border from then on. A



large, ‘modern’ army like the Roman one needed a good infrastructure (roads, bridges), surveyable battlefields and safe supply lines for food. These conditions were met in Gaul. The Gauls were good cartwrights and had established a good road network (the present French word ‘chemin’ (road), is of Gaulish origin). In addition to this, they had a lot of agriculture (cereal cultivation) and often concentrated in the cities. Such cities often formed targets for the Romans, whose siege techniques were organisationally and technically superior to those of the Gauls. Germania was different. There was no thorough road network, nor any indications of the formation of towns. The Teutons practiced little agriculture; they were more occupied with cattle raising and hunting (Caesar, Book VI, 22 and 29; Tacitus 98). Corn for the Roman army continually had to be ordered from Gaul and was either supplied by land or by the Weser and Elbe Rivers. Since the Romans wanted to extend their territory to the Elbe River, the supply lines were long. If Germania had had an open, park-like landscape, the problem would have been less. All sources, however, as will be explained below, seem to indicate that the Germanic landscape was very difficult to pass through, especially because of its dense forests. A term that is used again and again to describe the natural landscape of Europe is ‘forests and marshes’. Both Roman and Medieval writers used these words (‘silvae et paludes’; Caesar, Tacitus, Mela, Strabo (in: Goetz & Welwei 1995), Paravicini 1989/ 1995). According to Forcellini (1940), the Latin word ‘silva’, as the Romans used it, has the meaning of ‘forest’ or ‘wood’ as we still know it nowadays. Words like ‘shadowy’ (umbrosa, opaca), ‘dark’ (nigra), ‘deep’ (profunda), ‘dense’ (densa) and ‘rugged’ (aspera, horrida) were used to describe it. Beside this, the word was also used metaphorically, as in ‘a wood of spears’. That a ‘silva’ consisted entirely of trees is demonstrated by the saying ‘in silvam ligna ferre’ (Forcellini 1940). This literally means ‘[it is like] taking timber to the forest’, i.e., ‘it is like carrying candles to the sun’. That the meaning of the word ‘silva’ is clearly different from the one given by Vera (2000, page 120 ) (‘mosaic landscape with groves and grasslands’) may be additionally demonstrated by a practical example mentioned by Caesar (Book III, 28). When Caesar had conquered all of Gaul

193


194


at a certain point, only the territories of the Morini and Menapii remained to be occupied. Their territories were the present-day province of West Flanders (Belgium). The tribes in question did not wait for a confrontation and withdrew into ‘uninterrupted forests and marshes’, as Caesar describes it. When Caesar arrived at the edge of these woods, he writes: ‘In the next days, Caesar decided to cut the forests [silvas]. To prevent a flank attack on his unarmed and unsuspecting soldiers, he ordered that the felled timber be placed in such a way that it was turned toward the enemy, like a wall piled up on both sides. In a few days time, a large part of the forest was felled at an incredible rate’.31 The cutting of the forest had to be stopped, however, since a tremendous storm broke out (this was autumn) and the soldiers had to stay in their tents because of the continuous rain. Caesar eventually contented himself with destroying the villages of the tribes. He was unable to chase these tribes from their hiding places in the forest. In the land of the Eburones, the Ardennes, Caesar also had trouble finding the enemy in the dense forests (Book VI, 34), as: ‘the forests, because of their unreliable and hidden paths, hindered the entrance in closed [army] formations’. The same problem arose in the fights with Ariovistus in East Gaul. At a certain point, a great fear had developed among the Roman soldiers, not so much because they feared the enemy, but rather the narrow roads and the large forests that stretched out between them and Ariovistus (Book I, 39). There was clearly no question of an open, clear landscape here. One of the most renowned forest areas at the time was the Hercynian Forest (Silva Hercynia). Caesar describes its size as follows (Book VI, 25): ‘In width, this Hercynian Forest extends to a nine day’s journey for a good pedestrian….. And there is no one in this part of Germania who will say that he reached the beginning of the forest, even when he has covered a 60 day’s journey, or who has heard reported where it begins’. This forest stretched from Southwest Germany to Romania, a distance of some 1,500 km. If a 60 day’s journey was needed to cover its length on foot, day’s journey must have stood for about 25 km, a dis-

31 The felling of the forest here means the cutting of broad paths in order to allow the army to pass through. Later on, in Germania as well, this was frequently the way to get control over a forest area (‘limites’ (forest roads); Timpe 1989). In this way, the Roman emperor Domitianus had constructed over 120 miles of roads in the forest to chase the enemy from its hiding places and to gain control over the battle (Bendz 1963).



Fig. 33. Detail of Trajan’s Column in Rome. On this column the conquest of Dacia (Romania) is portrayed (ca. 100 AD). Several pictures on the column, like the detail rendered here, show how Roman soldiers clear (cut) forests to accommodate the construction of roads. Originally the hands of the sculpted soldiers held small metal axes.

tance similar to one covered in an average day’s journey through the Great Wilderness, a large primeval forest area in what used to be East Prussia, which was notably difficult to pass through, and about the appearance of which a lot is known (see further along in this Chapter). This same apparently impressive Hercynian Forest, through which the Roman general Germanicus had still managed to cut a road at the beginning of our era (Timpe 1989), Plinius described as ‘a large extensiveness, untouched by the centuries and as old as the world’. About the Germanic forests he continued to write: ‘These cover all the remainder of Germania and augment the [natural] cold by their shadow’. Looking at these descriptions, it becomes clear why the Romans did not hold with the Germanic ‘forests and marshes’, as these strongly hampered their warfare. Indeed, nourished by this aversion Tacitus (98) wrote:

195


196


‘Though the land(scape) rather varies in appearance, still in general it is either rough42 because of its forests or ugly on account of its marshes’. In the same century, the geographer Mela (± 40) reaches a consonant conclusion about Germania: ‘The land itself is difficult to pass through because of numerous rivers, rugged on account of many mountains and for the largely impassable because of forests and marshes’. The attempt to subject Germania eventually led to the defeat of the Romans in the so-called ‘Battle in the Teutoburg Forest’ in AD 9. At some point, when the Romans unsuspectingly walked through the forests in a column of several kilometres, they were suddenly attacked. Dio Cassius (± 200; Book LVI, 19,20) describes the battle: ‘…..they came upon Varus in the midst of forests by this time almost impenetrable……..The mountains had an uneven surface broken by ravines, and the trees grew close together and very high. Hence the Romans, even before the enemy assailed them, were having a hard time of it felling trees, building roads, and bridging places that required it. They had with them many wagons and many beasts of burden as in time of peace; moreover, not a few women and children and a large retinue of servants were following them – one more reason for their advancing in scattered groups. Meanwhile a violent rain and wind came up that separated them still further, while the ground, that had become slippery around the roots and logs, made walking very treacherous for them, and the tops of the trees kept breaking off and falling down, causing much confusion. While the Romans were in such difficulties, the barbarians suddenly surrounded them on all sides at once, coming through the densest thickets, as they were acquainted with the paths’. Eventually, after four days of fighting, three Roman legions under the leadership of Varus were defeated there by the Teutons under Arminius (Wiegels & Woesler 1995), in a narrow passage north of the Teutoburg Forest, between the forested Kalkrieserberg and a large peat bog (‘Grosses Moor’). All this happened on a terrain unsuitable for the Ro32 The original text is: ’Terra etsi aliquanto specie differt, in universum tamen aut silvis horrida aut paludibus foeda, …..’. Vera (2000) translates ‘horrida’ in this context with ‘thorny’. The literature source he cites (Muller, F. & Renkema, E. H. (1995):’Beknopt LatijnsNederlands Woordenboek, 12th edition, page 408), however, does not give the meaning ‘thorny’, but ‘rugged, rough’. Other Latin dictionaries, such as Forcellini (1940) and the Thesaurus Linguae Latinae, agree that the latter is the correct meaning. Gramatically speaking, in the text used by Tacitus ‘horrida’ goes with ‘terra’. The word ‘horrida’ thus refers to the (in view of the forests and marshes) rough, uncultivated appearance of the landscape (Thesaurus Linguae Latinae).



man type of warfare, though eminently useful for an enemy that knew the terrain and applied guerrilla tactics (Clavel 1967, Timpe 1989). With some respect, Julius Caesar at the time already said about the Teutons: ‘Marshes and forests do not form a hindrance to these warriors grown up in wars and predatory expeditions’ (Book VI, 35). The Roman writer Frontinus (1st century; Bendz 1963) also already stated that: ‘….the Teutons, as is their custom, attacked our troops from mountain forests and dark hiding places, and again and again had a safe retreat into the depths of the wood….’. It is worthwhile in this respect to mention the presence of sacred forests in the landscape of that period. In such forests all human influences were forbidden, except the worship of the gods. No hunting, wood cutting, agriculture or cattle pasturing took place; in fact, therefore, such forests were left to nature. Their thickly wooded character may be deduced from an account by the Roman writer Lucan, who describes how in 49 BC Julius Caesar, in view of the siege of Marseilles, commissioned the cutting of a sacred forest (Lucanus & Duff 1928, Book III, 399-452): ‘A grove there was, untouched by men’s hands from ancient times, whose interlacing boughs enclosed a space of darkness and cold shade, and banished the sunlight far above……This grove was sentenced by Caesar to fall before the stroke of the axe……Spared in earlier warfare, it stood there covered with trees among hills already cleared…….All [trees] lost their leaves for the first time; robbed off their foliage, they let in the daylight; and the toppling wood, when smitten, supported itself by the close growth of its timber’. If the landscape in Germania had been open and surveyable, the Romans could have carried out their military manoeuvres and troop movements in the best possible way. The Teutons knew that they were no match for them and chose a tactic that disrupted the military machinery of the Romans. They continued doing this also after the battle mentioned above, until eventually, the Germania adventure was whistled off from Rome. The Roman roads, frequently found west of the Rhine, are completely lacking to the east of this river. The Romans

197


198


never managed to construct a lasting network of metalled roads in order to open up and colonize the area. No pictures of the campaigns in Germania have survived. Pictures do exist of the Roman conquest of Romania (Dacia) in the period AD 101-106. These may be found on Trajan’s column in Rome; the many pictures on this column include several that show Roman soldiers who are clearing forests by cutting trees, in connection with road construction (see Fig. 33). As Julius Caesar mentioned that the Hercynian Forest extended to Dacia, this also offers an opportunity to gain an impression of the hindrances the Germanic forests posed.

Frontier forests That ‘forests and marshes’ were not always without obstacles even for the Teutons is shown by the fact that there were so-called frontier forests. Caesar mentions such a frontier forest, somewhere in Germania (Book VI, 10): ‘There is an immensely large forest, called Bacenis, which stretches out very far and, like a natural wall, keeps the Cherusci and Suebi [Germanic tribes] from wronging and attacking each other. The Suebi have retreated to this forest to await the arrival of the Romans’. Caesar decides no to look for the Suebi in this forest. Tacitus (98) also mentions frontier forests, since he states that some tribes in Denmark ‘are protected by rivers or forests’ (Chapter 40). Such frontier forests formerly occurred generally, where the living areas of several native tribes bordered on one another. Not only in Germania, but also among West Slavic tribes, among others in Silesia, a defence used to be based largely on broad belts of forest, whose cutting was forbidden (Kuhn 1977). On the forest inside, such a defence was fortified by a man’s high hedge of bent or cut trees, between which thorny bushes were planted. These hedges were usually of some considerable depth (width), so as to hinder and slow down any attackers as much as possible. They had to be kept up so as to keep them as impenetrable as possible. The fact that this was a general custom indicates



that they were very effective. Also in Gaul, this type of border defence was known. Caesar (Book II, 17) already describes this technique with reference to the Nervii, who lived between the Scheldt and the Sambre: ‘…they cut young trees so that the branches grow sideways, and brambles and thorn bushes stand between these, so that like a wall, these hedges form defence works that are not only impenetrable, but impossible to look through as well’. Until the late Middle Ages, such frontier forests still existed in Hungary and Serbia (Tagányi 1921). Down to the 12th century, people in the latter country kept the so-called Bulgarian Forest untouched and in its original state of ‘wildness and inaccessibility’, as a buffer zone. Also in North America, many buffer zones between Indian tribes are known, sometimes with a width of up to 200 km (Kay 1995). Here, also, their function was to ‘keep the tribes apart’ and to limit confrontations to a minimum. The interesting aspect of these buffer zones was that to prevent conflicts, these areas were visited as infrequently as possible, which left them almost completely to nature. There was no timber cutting or hunting, since these were dangerous areas. As a consequence, such frontier forests were relatively rich in game, which found a refuge there. This was the case at times of war, which were not rare. Hunting occurred during protracted periods of armistice (Kay 1995). Analogous to this, Mech (in: Kay 1995) mentions that wolves utilised the edges of their territories less intensively than the central part of their living areas, so as to avoid confrontations with wolves from neighbouring areas. The very slight predation pressure in the border areas resulted in a relatively high density of whitetail deer (Odocoileus virginianus). From experience, these knew that they were relatively safe in those zones. In view of the fact that the local tribes barely infringed on the situation that prevailed in their frontier forests, if at all, the natural situation in such large areas scarcely frequented and exploited by people is likely to have been very closely approximated. This supposition is important for an assessment of the largest frontier forest known in history, the Great Wilderness in the former East Prussia (Mortensen 1938) (see Fig. 34). For over 200 years, this extensive area constituted a buffer zone between the Germans on the one side, and the Poles and Lithuanians

199


200


Fig. 34. The Great Wilderness in former East Prussia (NE Poland/ Russia) during the 14th century (after Mager 1960). The aurochs lived here till ca. 1500; until that time this area with its complete original fauna made up the last Central European wilderness, a quite impenetrable forest- and marsh area.

on the other. This border zone was relatively little tread upon by people and a study of this area makes it possible to us to have a look into the last Central European wilderness.

The last Central European wilderness The last Central European wilderness, also called ‘Great Wilderness’ (‘Grosse Wildnis’, as the Germans named it), was partly situated in the east and southeast of the former German province of East Prussia (see Fig. 34). The remainder was in Poland and Lithuania. The total area took up between 50,000 and 60,000 square kilometres (Mager 1941). The interesting aspect of this area lies in the fact that until quite recently (up to circa 1500), it had remained nearly untouched by people and largely left to nature. For some large herbivores such as aurochs, wild horse and European bison, it constituted one of the last refuges. Nowhere else in Europe did an area of such size remain inviolate, and in



possession of its original range of animal and plant species, for such a long time. Its continued existence as such was not a result of the medieval people’s notion of its beauty; rather, the reason was political and pragmatic. The status of the area had resulted from the political complications between the different nations that lived there. Since the circumstances in the Great Wilderness still closely approximate the natural situation, and many eyewitness accounts in which the area is described have survived, it is possible to get a fairly accurate idea of the former natural landscape of Central Europe. Before looking at the descriptions, however, it is worth knowing who managed these areas and how that management was accomplished. In the beginning of the 13th century, the Southeast Baltic countries formed the place of domicile for the last heathen European tribes. Christianity had not yet gotten hold. Attempts by the Poles, in the neighbouring duchy of Masovia, to conquer the area of the four Prussian tribes that lived between the Vistula and Memel Rivers were unsuccessful. By request of the Poles, the knights of the German Order next undertook this task, with approval of the German emperor and the Pope (Anstadt 1965). These knights were an organisation of monks fighting to establish Christianity, comparable to the Templars (Paravicini 1989), for instance. In 1230, the knights of the German Order marched into the area from the southwest and were able to conquer it from the Prussians during a battle that lasted fifty years. Around 1280, they had finally gained possession of the area between the Vistula and Memel Rivers. On the other side of the Memel the heathen Lithuanians were still present, and they would stay unkindly disposed towards the Germans down to the 16th century. The relation with the Poles had not improved, either; they also regularly waged war against the Germans. Peace did not return to the area until 1525, with the signing of a truce in which the proprietary rights were settled (Szper 1913). Of old, the coastal area and the southwest of East Prussia were most densely populated and most cultivated. The interior was relatively sparsely populated, hence barely cultivated. This interior, especially the southern and eastern border areas, became partly depopulated as early as the beginning of the 13th century, as a result of Polish and Lithuanian raids. The total depopulation was brought about by the German Order, which

201


202


established its authority in this area. All villages in the area, which later was called the Great Wilderness, became depopulated in the course of the 13th century. There was no more permanent habitation of this wilderness area down to the 16th century, apart from some defence posts (Schlüter 1921, Mager 1960). The area showed the characteristics of the frontier forests mentioned above, and was going to function as a buffer zone between hostile peoples. The war situation made it a dangerous area. Moreover, its tremendous size and the fact that it was difficult to pass through made it hard to traverse. The expeditions to and from Lithuania, which often took weeks, had to be prepared thoroughly, for example with respect to the food supplies for both the men and the horses. To keep hostile neighbouring peoples at bay, but also, and especially, because the German Order lacked money and manpower, the reclamation of the Great Wilderness was delayed for a long time. In this way, the area could remain and continue to exist nearly untouched during that period. Soldiers and noblemen from many European countries marched to this area to colonise the southwest and the coast, and to fight against colonists of neighbouring peoples (Paravicini 1989). The Great Wilderness was host to a plant- and animal world that, as a whole, was no longer to be found anywhere else in Europe. Beside large ungulates such as red deer, elk, roe deer and wild boar, which could still be found in other countries, in addition the European bison (Bison bonasus), the wild horse (Equus ferus) and the aurochs (Mager 1941) still occurred here. Predators included wolf, bear and lynx, as well as the wolverine (Gulo gulo) (Maciej z Miechowa 1521). As it happens, the climate in this area had a rather continental, northern character with severe, snowy winters. The aurochs was still mentioned by Stella (1518) and Maciej z Miechowa (1521), but must have been near extinction at the time, since no more was heard of it after this. When the aurochs had disappeared, this wilderness could no longer be called ‘complete’. The wild horse presumably became extinct in the beginning of the 17th century, and the last European bison was probably shot in 1755 (Mager 1960). From that moment on, the European bison occurred in the wild only in the Caucasus and near Bia³owie¿a.



Of old, East Prussia was known for it profusion of game. In the 12th century, of the Prussians it was said (Laurent & Wattenbach 1986): ‘Gold and silver they consider of little value, they abound in strange furs’. Even in the beginning of the 16th century they still had this reputation, as the Pole Maciej z Miechowa (1521) says of the Great Wilderness: ‘Because the forests are so great, many large wild animals are seen and caught in abundance……’. During the period that the Great Wilderness existed in this way, hunting took place there. The knights of the German Order were only allowed a little hunting, on account of their vows to the Order. Others were permitted to hunt, fish and gather honey from nests of wild bees. Hunting rights were bestowed and controlled by the Order. To Mager (1941), the fact that they were was granted to ‘outsiders’ would seem to indicate that the density of the game was relatively high. Hides could be traded only through the Order, and the right foreleg of each specimen of big game had to be given up to them. Hunting in this wilderness was carried out from primitive, temporary shelters, by people who often stayed there for months. This situation is comparable to that of the trappers in the early North America (Mortensen 1938), and of the ranging hunters in the Ussuri area (East Siberia), as early as the beginning of the 18th century (Arsenjew 1924). The hunting pressure experienced by the people is likely to have been relatively low (Mortensen 1938, Mager 1941), for the following reasons: - the very stretched-out Great Wilderness had to be traversed on foot or on horseback, along narrow paths. - the area was difficult to pass through, on account of the rivers, marshes and fallen trees. - a war situation prevailed, so hunting activities had to be limited. Factors like predation, the amount of food and the usually severe winters thus had a dominant influence on the density of the game. Supplementary feeding by man was not in order. This was not done until the 18th century, to save the last European bison from extinction. Poaching probably occurred very infrequently, mainly as a result of the conscientious and severe observance of the rules by the German Order. Problems with this only arose when the political situation became safer in

203


204


the course of the 16th century, and also because the game in the neighbouring forests became scarce (Mager 1941). In the 17th century, poaching grew to an alarming extent, especially in the southern zone that bordered on Masovia. Based upon these data, a sketch may be drawn of what this Great Wilderness actually looked like, and how nature, little influenced by man, shaped it. In the Middle Ages, the words ‘wald’ and ‘wildnis’ or written variations of these were frequently used to describe the area or parts of the area. In the Old Germanic language, these two words initially had the same meaning of ‘savage, uncultivated land, as yet little influenced by man’ (Grimm & Grimm 1854-1954, Mager 1960). This could refer to different types of landscape, with or without trees. The word ‘wildnis’ always kept this meaning, but in the course of time, the word ‘wald’ took on the more specific meaning of tree stand, cultivated or otherwise. In the Great Wilderness, during the period of interest for this book (13th–16th century), the word ‘wald’, judging from the descriptions of it, was used to indicate a ‘primeval forest’, i.e. an uncultivated forest stand (Mortensen 1938, Mager 1960). These descriptions may be found in many eyewitness accounts and official documents, and in the so-called ‘Route Descriptions’ (‘Littauische Wegeberichte’; Hirsch 1863). Based on extensive research of these writings, Mager (1960, I, page 26) concludes that at the end of the 13th century, and apart from the lakes and rivers, nearly 100% of the Great Wilderness were made up by ‘forests and marshes’. The conditions that prevailed in these forests and marshes made the Great Wilderness very difficult to pass through, so all the sources from the 13th century described this area as ‘very inhospitable’ (‘höchst unwirtlich’; Mager 1960, I, page 28). The many descriptions paint a vivid picture of the often frightful expeditions, right across these primeval forests and marshes. Several names were used for the various forest areas inside the primeval forest of the Great Wilderness, which were distinguished from each other by their tree species and the moisture content of their soil. First of all, the word ‘heide’ was used. Originally, this had the same meaning as ‘wald’ and ‘wildnis’, i.e. the uncultivated wilderness (Grimm



& Grimm 1854-1954, Mager 1960). Later on, in the West Germanic areas, it received the meaning of open ‘heather field’; particularly in the East Germanic areas, like East Prussia, people started to use the word to indicate a certain type of forest, which occurred on dry sandy soils and was characterized especially by the occurrence of heather (Calluna vulgaris) and Scots pine (Pinus sylvestris). In this connection, Mager (1960, I, page 139) cites Hennenberger (early 17th century): ‘there are such places that only have spruce or thick, sweet-smelling pines and such places are called ‘Heyden’, under which the ‘Heydenkraut’ also grows readily…..’. In the Route Descriptions, these are the forest types that were easy to pass through, where only little had to be cleared33. The word ‘heide’ may still be found in the names of forest areas like Rominter Heide, Borkener Heide and Dübener Heide. In fact, heathlands could not and cannot exist in East Prussia in the way we know them in Western Europe. Heather barely occurs in the open air in East Prussia since this plant is unable to resist severe winter cold; because of this it thrives at best under a canopy of Scots pine (Steffen 1931). A second forest type is the ‘Damerau’. The name derives from the Polish word ‘d¹browa’, which means ‘oak forest’ (Mager 1960). In Poland, this was originally the name for a somewhat open grassy oak forest that was used as a Hudewald to graze cattle. Although initially, this was clearly a culturally tied name, the word was used in the writings of the German Order with reference to the Great Wilderness, to indicate woods in which the oak took an important place. Besides oak trees there could also be linden, aspen, birch and hornbeam. These forests occurred on sandy soils that were rich in loam and had rather good drainage, which is why the 16th- and 17th- century colonists preferably felled these forests, to turn them into cultivated land from them. The Damerau were also relatively easy to pass through, and they had to be cleared only a little. The ‘Grauden’ constituted a third forest type, which according to the Route Descriptions caused a lot of problems when people tried to traverse it; large-scale clearing was always necessary here. The actual meaning of the word ‘Grauden’ is not known; the explanations found 33 ‘Clearing’ (in German: ‘rumen’ - ‘räumen’) means the removal of fallen trees to make passage possible (Mortensen 1938, Mager 1960 (part I), Grimm & Grimm 1854-1954).

205


206


up to this point have all been unsatisfactory (Mager 1960). Since the forests concerned are often rather wet, and the Polish language has words like ‘gr¹d’ and ‘grud’, general names for oak-hornbeam forests, this could be a possible explanation, indicating that the Grauden may have been a wetter variant of the Damerau, in which linden and hornbeam played an important role. A fourth element that pops up regularly in the Route Descriptions is the ‘velt’ (‘field’). This was a more or less open space, where people pitched their camp in the evening and where the horses could graze (Mortensen 1938). Such fields were of crucial importance to the expeditions through the wilderness. Sometimes for weeks on end, people would travel through the forest with their pack horses and riding horses. To feed these (work) horses and the (booty) cattle that was often dragged along, sufficient grass had to be present at stated distances. To supplement the food provisions for man and beast, at certain places fixed storage buildings (‘maia’) were constructed, in which food for the people, fodder for the horses and firewood could be stored. The fact that the word ‘maia’ is of Prussian origin indicates that this system was in use already in the pre-German period (Paravicini 1995).The fields largely owed their existence to man (Mager 1960, Mortensen 1938). They were partly situated in places where the Prussian natives had had their villages; the rest are likely to have been wet river banks, if these were naturally treeless (Mortensen 1938, page 23). In many Route Descriptions, the night camp is pitched at a river, where there was often sufficient grass. Mortensen compared the situation in the Great Wilderness with the primeval forests in South Chile visited by him. There as well, people travelled the forest with horses and cattle, and there were fields in which they rested at stated distances. If necessary, the vegetation was set on fire at leaving, to keep the fields productive and open. In the Great Wilderness, people would naturally follow the best passable route, along which such fields were present. In the Route Descriptions, explicit mention is always made of the places that have sufficient grass and those that do not, which seems to indicate that it was scarce (Mager 1960, I, page 146). People often even brought fodder for the horses with them, and left it in the maia to be used on the way back.



That such fields, in places favourable for the growing of trees, had to be kept open by man is shown by the following two facts. In the first place, some shifting cultivation (‘Scheffelwirtschaft’) was practiced in the border zones of the Great Wilderness, where people would cut and burn a piece of forest and grow rye or buckwheat there for three years. The soil that was then exhausted was left uncultivated, after which the forest once again took possession of the open space. After 12-15 years, this procedure would be repeated in the same place (Mager 1960). Secondly, reports have been passed down about the early Prussian settlements in the Great Wilderness, which had been depopulated definitively by the German Order. Wherever these were not used as places for night camp during forest expeditions, they closed up again with forest. There are descriptions of this by several authors (Mager 1960, I, page 30): ‘…in Prussia there are many forests, which grow on villages devastated by wars, where farmers used to live…’. ‘…here and there one can see large forests, which used to be open fields that produced a lot of corn. Although the shape of the fields can

Fig. 35. The problematic journeys people with packhorses made through the Great Wilderness are best reflected by this picture of such a journey through Rocky Mountain forests (United States) during the 19th century (Mortensen 1963). The picture was made by one of the participants, T. Hildebrand.

207


208


still be recognized, they are covered with trees that are thick as barrels, oak, aspen, birch, beech, spruce and such…’. This is reminiscent of the period after the Romans had left the South of the Netherlands, where the forest had once again taken over of the fields and meadows, as pollen diagrams have shown (Teunissen 1990). The expeditions through the primeval forest were made on foot or on horseback. The lack of a good road network long made the use of carts impossible. At the first mention of the use of a cart, in 1390, it was explicitly added that this was the first time a cart was driven in the Great Wilderness (Mortensen 1938, page 53). In winter, sledges were used to cross frozen rivers and marshes. The roads that are mentioned in the many descriptions have to be pictured as more or less wide paths, along which some of the trees had been marked with signs that were often secret, which also used to be a custom in North America and East Siberia (Mortensen 1938, page 46 and 47). In addition to this, expert guides were needed to show the way. Nevertheless, many people lost their way in these primeval forests and never returned, or took a long time getting back (Mortensen 1938, page 16). Hirsch (1863) collected and commented on 100 of the Route Descriptions through the Great Wilderness that existed at the time. These descriptions, most of which date from the end of the 14th century, others from later periods, give an impression of how people traversed the wilderness, and what effort went into fulfilling such an expedition (Hirsch 1863, Mortensen 1938, Mager 1960). The Route Description 2 offers a good description of the various aspects of such a forest march. ‘This march will be accompanied by Eywon Spandenne and Swalgenne from Waldow. The first night camp is Rossiten, from there on to Wintburg on the other side of the Nerge [a river], the third night camp on the other side of the lagoon, the fourth night camp 3 miles34 up to the Meinie [a river]. Little needs to be cleared there, because this ‘heide’ is easy to pass through, one leaves the first food35 there; the fifth night camp is at Bareykenfelt, 5 miles further along, this distance has to be covered in one day, it is easy to pass through, and there one leaves the second food, and one has to bridge36 the Weywir [a river], and also a

34

The mile used here is 8280 m (Mager 1960, I, page 149). In the maia. 36 Bridging could mean the making of primitive bridges, but also the ‘bridging’ of marshes with tree trunks and faggots (see Fig. 17). Often, provisions had already been made and only needed repairs. Some rivers could be waded knee deep. 35



small river with bushes; from there on to the Jure [a river] it is three miles, in between there are a ‘grauden’, where some clearing has to be done, and 2 rivers that have to be bridged and improved, at the Jure one has to rest, after which there are still 2 miles to the land37. In between there are hedges38, which have to be cleared, and one has to ride those 2 miles by night….’. The length of a day’s journey could vary from 1 to 6 miles (8-50 km), but was usually between 2 and 3 miles (17–24 km). This depended on the difficulty of the march. An additional problem was the presence or absence of grass; this was mentioned regularly, for instance in Route Description 62: ‘….from there 3 miles to the Naresna [a river], in between are a forest and a river, named Geskra, there one has to bridge the width of a bowshot on both sides; from there 6 miles to a river, named Subbresla, in

Fig. 36. Cross section of a wooden trackway in former East Prussia. This road was constructed right across the valley of the Sorge River to bridge the fens located in the floodplains, and the river (after Conwentz 1897). The 1231 m long road was entirely made up of oak stems.

37 38

I.e., the land of the enemy. Constructed by the enemy.

209


210


between there is no grass, from there 5 miles to a river, named Ploska, there is sufficient grass there. From there 3 miles to the little Naresna, there is sufficient grass there, also…’. A description that very clearly represents the great problems people often had to overcome, is Route Description 42: ‘…there is over a mile to go from the hedges [of Insterburg] to Girwisken, in between there is a forest, more than half a mile has to be cleared there and 2 small rivers have to be bridged. 2 Miles from there to Boyte, in between at least half a mile needs to be cleared and there is a small river to be bridged. From there 2 miles to Helledompnen, the Pissa and Rodappe have to be bridged there, the Pissa over more than 2 rods, but the other is small, those 2 miles have to be cleared. From there to Pissenyken [is] 1 mile, that mile has to be cleared entirely, and one has to bridge 2 small rivers, also one has to bridge the Pissa once again. From there to Lüpow, 2.5 miles, in between one has to bridge 2 rivers, the Ywayde and the Lypow, also more than 1 mile has to be cleared. From there 1.5 miles to the Zelwo, those 1.5 miles have to be cleared, the Zelwo has to be bridged over 1 rod. From there 2 miles to the Serwynte, 1 mile has to be cleared there, the Serwynte has to be bridged across 2 rods. From there 3 miles to the Rawsze, half a mile has to be cleared there and over 3 rods have to be bridged as well as over 3 rods on the other side, also the Rawsze, at which one has to pass the night, has to be bridged. From there to the Suppe 3 miles, clearing is needed everywhere there and 3 bridges have to be made, each bridge a rod’s length. From there it is 9 miles to the country of Slowisso, in between one has to clear a great deal and bridge 9 times….’. The entire distance of this march was more than 223 km. These marches were often accompanied by special pioneer units with axes and saws. Of a march in 1427, Mager (1960, I, page 151) mentions 200 such ‘clearers’. In spite of this, people sometimes did not advance more than 8 km a day. If this had been an open, park-like landscape, it would not have had to be cleared and progress would have been much faster. The descriptions are reminiscent of the march of the Romans through the Teutoburg Forest as described by Dio Cassius; here, too, a lot of cutting



and bridging was needed, which also took a lot of effort too. Comparable situations in earlier centuries in the east of the United States, as well as in the beginning of the 20th century in the Ussuri area (Southeast Siberia) will be described in Chapter 9.4. In the 16th century, after 1525, when a peaceful period started in East Prussia, the definitive devastation of the Great Wilderness set in. In the 17th and 18th centuries, the reclamation went so quickly and dynamically that in 1739, the German emperor proclaimed an official prohibition on using the word ‘wilderness’ for the former Great Wilderness (Mager 1960, I, page 153). The frightful primeval forests, which were then seen as so unyielding to man, had vanished or had been brought under the control of man at that point. Part of a verse by Peter Suchenwirt that refers to a march through the Great Wilderness, to and from Lithuania, in September 1377: Straight across the wilderness the army marched. They mounted their horses, got off, crisscrossing through it, jumping, bending, crawling; Strongly the branches pulled at the collar of many a man; The wind had torn down many large trees; With great effort we had to pass over them. In the crowd there was much shouting, the Prussians hindered us so much. Horses and pack animals we pulled along, heavy-laden and in the crowd many an animal became unwell. Because they were urged on so and injured knee and leg. Pleasantry and laughing had become hushed.

211


212


Summary In this chapter, different lines of approach were used to trace the sources that can provide information about the appearance of the natural landscape of Europe during the period after the last ice age, with the help of insect research, pollen research, Roman writings and descriptions of the Great Wilderness in former East Prussia. Insect research in England and the Netherlands shows that the landscape in the areas in question, up to the arrival of the agricultural, cattle-raising man during the Neolithic (3000 BC), was covered for the greater part with continuous forests. The insect species from the period before this were all found to have been associated with trees. Especially species that live on dead trees were remarkably well represented. Not until much later did insect species appear that are characteristic for meadows and fields with cultivated plants. Pollen research has revealed the near omnipresence of forests at the time. Beside treeless peat bogs and coastal salt marshes, treeless vegetations, in which sedges (Carex spec.) dominated, were also found along rivers. The proportion of grass in the forests was relatively small. Pollen research has also shown that the communities at the edge of the forest appeared simultaneously with the expansion of human settlements in the course of the Neolithic period. This indicates a landscape with a lot of man-made forest edges and hedges. Accounts by Roman writers reveal that, at the time, in Gaul and especially in Germania, beside human settlements and fields, the landscape consisted of extensive forests. If there were no roads cleared by man in those forests, marching through them was clearly very problematic for the Roman army; this indicates the closed aspect of the forests. An area that until recently had only been described in German literature is the Great Wilderness in former East Prussia. Many descriptions of this extensive wilderness area were passed on from the Middle Ages. This area, which was hardly influenced by man, and which contained the original range of animal species, consisted of continuous forests, alternated locally with marshes. Traversing these primeval forests often presented great problems to man.



9.1.2. Food The close relationship of the aurochs with the other bovine species of the Bovini tribe (Wilson & Reeder 1993, Buntjer 1997), which all belong to the ruminants, warrants the conclusion that the aurochs was a ruminant as well. Ruminants are plant eaters, who use a variety of plant species to get nutrients from. They have four stomachs: the rennet stomach (the actual stomach) and three fore stomachs, the rumen, reticulum and omasum. Up to the age of four weeks, only the rennet stomach is active in a calf; as more solid food is eaten the other three stomachs also begin to function (Richter 1982). The rumen, into which the food enters first, is the largest of the four stomachs. On the basis of their various bodily adaptations, ruminants generally can be divided into three groups, connected to the plant species and plant parts that they eat (Hofman 1989) (see Fig. 37). These three groups are called ‘browsers’, ‘grazers’ and ‘intermediate feeders’. Browsers look selectively for forbs, leaves and branches, not grass. First and foremost, grazers are in search of grass, which forms their main food. Browsers are especially focused on the content of the plant cells, while grazers are more adapted to the digestion of cells walls (which mainly consist of cellulose). The bodily adaptations that make the three food digestion strategies possible are found especially in the rumen and in the head of the animal (Hofmann 1989). The rumen in a browser is relatively small and built to enable the fast passage of plant material and the fast intake of nutrients through large papillae in the stomach wall. The browsers include roe deer and elk, among other species. The grazer possesses a relatively large rumen, which can hold plant material for a long time. The intake of nutrients through small papillae in the rumen wall takes place slowly. The grazers include cows and sheep, among other species. With regard to their bodily adaptations, intermediate feeders fall in between these two groups as regards their adaptations; they include red deer and fallow deer, among other species. The rumen of ruminants contains many species of micro-organisms, which are able to break down the various components of the plants.

213


214


Fig. 37. Classification of European ruminants into three types. The domesticated animals among them have not been shaded. (after Hofmann 1989).

The adaptations in the head concern the lips, the incisors, the molars, the tongue and the salivary glands (Hofmann 1989; Janis 1986, 1988), all of which are designed to pick up, grind and digest a specific type of food as efficiently as possible. Compared to grazers, browsers have long lips, a soft tongue, a narrow row of incisors and large salivary glands. Grazers have comparatively large, more extensive chewing muscles than browsers, so as to be able to grind tough grass (Solounias et al. 1995). Again, these adaptations in intermediate feeders are somewhere between those in browsers and grazers. The molars of plant-eating ungulates can be recognized in the first place by the furrows (ridges of enamel more or less close to each other) on their chewing surface. Beside this, the extent of hypsodonty39 may be an indication of the type of food an animal usually eats (Janis 1986, 1988). Ungulates show various rates of hypsodonty; especially equids are very hypsodont. Bovine animals are less hypsodont than equids, but more so than most intermediate feeders and browsers. Hypsodont un39 The extent of hypsodonty is determined by dividing the height of a molar crown by its length (perpendicular to the height). Janis (1988) used M-3 for the height, and M—-2 for the length of the molar. The height M3 is measured when the molar has just appeared and is still unworn.



gulates are generally grazers, though mere hypsodonty is no proof of grass eating. Intermediate feeders like the saiga antilope (Saiga tatarica) of the Central Asian steppes and the pronghorn (Antilocapra americana) of the North American prairies are as hypsodont as bovine animals (Janis 1986, 1988). The foraging on the open dry plains plays a part here; the plants often are covered in dust and grit, and eating them requires hypsodont molars. To assess whether an animal is a grazer or not, one has to consider not just the isolated adaptations, but also a combination of them. On the basis of their choice of food and the characteristics mentioned in the above, cattle, the domesticated descendants of the aurochs, are considered grazers, like the other bovine species of the Bovini tribe (Van Wieren 1996). As to their bodily adaptations, grazers are better adjusted to digesting grass than other ruminants. This does not mean they eat grass only; they are also able to eat other plant species, like forbs or parts of trees and bushes. In the course of a year, however, grass is not always equally well digestible for all grazer species. Climatological influences (periods of drought, winter cold) may result in marked variations in the chemical composition of grass, which may affect its digestibility (Van Wieren 1987). No soft parts have been found of the aurochs, unlike the finds of frozen mammoth, woolly rhinoceros, steppe bison and horses in Siberia and Alaska (Guthrie 1990). Only certain features of the head, like the shape and surface structure of the molars and the width of the incisor bone are still assessable. The molars of the aurochs were hypsodont, as they still are in domestic cows (Degerbøl & Fredskild 1970) (see Fig. 38), and their surface structure was also the same. The decline of the length of the molar row during the domestication process kept pace with the decline of the basal length of the skull. The length of the molar row in both aurochs and cattle is always 29 to 30% of their basal skull length (measured using data of Grigson 1974 and 1978). The premaxillary width is probably directly related to the width of the teeth row in the lower jaw. In the course of the domestication process, the width of this bone in relation to the length of the skull increased, as is also

215


216


the case with other skull width measurements (Bohlken 1964). It is true that the head of the domestic cow is shorter; relatively speaking, however, is broader than that of the aurochs (see Chapter 8). Judging from the molars and the width of the mouth, there was no essential difference concerning their food strategy between the aurochs and cattle that descended from it.

Fig. 38. Drawing of a remaining part of an aurochs jaw with two molars, seen in profile (top) and from above (bottom). Site: North Algeria, Holocene (after Pomel 1894).

There is some concrete information about the food eaten by the aurochs. Their food during the summer period is reported both directly and indirectly in the various descriptions. A direct mention is known only from a Russian epic, in which the aurochs was said to eat grass that occurred in the marshes (see Chapter 9.1.3; Dementiev 1958). Indirect reports about its food are found in the Inspection Reports that were made in the 16th and 17th century about the management of the Jaktorów forests. In various places in these reports, the relation of the aurochs with the local meadows is mentioned. The farmers from Jaktorów, for example, are said to ‘mow grass, where the aurochs have their meadows’, their ‘horses and cattle graze, where the aurochs stay’ and hunters appointed by the Crown are said to ‘have to pasture the aurochs with devotion in the usual places, so that they do no harm to the agricultural fields, also in the meadows of the Crown’ (£ukaszewicz 1952). In addition to this, the fact that they judge the grazing by cattle and



horses as detrimental for the aurochs is mentioned twice. Since the domestic cows happen to come ‘where also the aurochs graze’, the latter cannot develop well because of the diminished amount of food. These reports make it reasonable to assume, first of all, that the meadows in question (mainly) contained grass, and secondly that the aurochs, like the horses and cattle of the nobility and the farmers, used this grass as their main food. The grass in the meadows at the village of Jaktorów was used to make hay, which in wintertime was taken to the forest to feed the aurochs. A certain meadow at Jaktorów was said not to be mown for the aurochs, but used to pasture cattle in exchange for geese and capons (£ukaszewicz 1952). The conclusion that the aurochs, like horses and cattle, ate mainly grass in summer, which was provided in dried form (hay) in winter, seems to be in accordance with a mention by Von Fleming (1719). In a certain passage of his, probably mixed, aurochs/ European bison description, he says: ‘What at present other cattle eat, the ‘Auer’ eats also’. Though this is likely to have referred to the aurochs, it cannot be made out unambiguously so, since the European bison is also mainly a grazer. Apart from the grass the aurochs ate in summer in the meadows in the forest, it also liked to leave the forest to graze in the meadows elsewhere. In addition to this, the mown grain that was standing in the fields of the farmers to dry was also in demand. Schneeberger (in: Gesner 1602) writes: ‘In summer they leave the forests and go into the fields and devour mown grain and when they are satisfied they throw the rest apart with their horns, unless they can be chased away by dogs’. This was probably in the month of August, when the grain would ripen and become very attractive to the aurochs; this was also the month in which the mating season started (see Chapter 9.3). During the autumn, acorns appear to have been a desirable food. As Schneeberger reports: ‘In the autumn they feed on acorns (it so happens that the forest has enormously large oak trees, they say some are bigger than 9 m around), and then they are fatter and more shiny than at other times’.

217


218


About the winter food, Schneeberger writes: ‘In winter they eat branches40 of shrubs and trees; however, they are also supplied with hay that the farmers of the above-mentioned villages have gathered for the aurochs in summer’. The fact that ‘branches of shrubs and trees’ are mentioned first, followed by hay, warrants the assumption that branches form a not unimportant type of winter food. How much hay was fed supplementarily in winter cannot be precisely ascertained. The hay was stored in stacks, of which the Inspection Reports provide no measurements. Such a haystack (indicated as ‘bróg’ in Polish; £ukaszewicz 1952) consisted of four vertical poles placed in a square, which held up a slideable roof. The type derived from the more westerly situated Germany. If it were the same type as the one that was still used in 19th-century West Poland (Tetzner 1902), which was sized 3 by 3 by 6 m, this would mean a volume of about 50 m3 of hay. For some years of the 16th century, the number of stacks of hay produced at Jaktorów is known, and also how many of these were used for the aurochs and how many were left over (£ukaszewicz 1952). Since the number of aurochs is known for the same years, the amount of hay available to each aurochs may be calculated; it varied between 0.7 and 1 haystack per winter. Based on a period of supplementary feeding of five months, this means that on average, one aurochs could dispose of between onefourth and one third m3 of hay a day. This must have been sufficient. Especially in years when the hay harvest failed, the animals were forced to look for food in the forest. For the sake of completeness, a transport of four ‘Uwer’ in April 1409 from Königsberg (nowadays Kaliningrad) to West Europe (Nehring 1898b) needs to be mentioned here. These animals, which were probably caught young, were fed with oats and hay during their journey. Whether these were really aurochs, or possibly European bison, can no longer be traced because of the confusion of the names of these animals. The food of wild cattle that are related to the aurochs also warrants some inspection. Table 12 lists a number of species of the Bovini tribe, with a survey of their food choices, to show the relationship between the various species with regard to this aspect. 40 In the original Latin text, the word ‘frondes’ is used here, which means ‘branches with foliage’. Since the winter is the period in question, and not all trees or bushes have leaves or needles then, it is probably used here in the general sense of ‘branches with or without foliage’.



Photo 17. African buffalo (Syncerus caffer caffer) in South Africa (Photo: M. Drescher).

Table 12. Survey of the research into the food choices of a number of bovine species related to the aurochs.

African buffalo (Syncerus caffer caffer) (see Photo 17).For the main part, the buffaloes in the savannas of Serengeti and Manyara eat grasses; only a small percentage of their food consists of parts of trees and bushes (Prins 1987). In general, there is no difference as to food choice between wet and dry periods. A different choice of food is found among the buffaloes that live along the open-forested Zambezi River in Zimbabwe. In the course of a year, the average proportion of grass is 45% there (Jarman 1971). Forbs, trees and bushes form the other 55% of the food; this percentage varies from 26% to 80% during the year. During the dry period, the animals frequent the riverbanks, where they can still find the most edible grass.

219


220


Table 12. Continued.

Forest buffalo (Syncerus caffer nanus) (see Photo 18).Grasses and graminoids constitute 60% of the food of these bovine species in Ivory Coast (dung analyses; Hoppe-Dominik 1988). Forbs, trees and bushes make up the remainder. Naturally, the proportion of grass is higher in areas that have a lot of grass. Yak (Poephagus mutus).During the summer, the food consists mainly of grasses, the proportion of which varies from 44% to 98% (Schaller 1998). In some cases, it consists of 30% forbs; in a single case of 56% dwarf bushes. No data are available about the winter food of the wild yak, but research by the same author of the domesticated yak, which hibernates outside, shows that this consists of 95% grasses. Banteng (Bos javanicus).Both Hoogerwerf (1970) and Halder (1976) mention grasses and graminoids as the most important foods for the banteng, though neither mentions percentages. Hoogerwerf mentions 20 species of grasses and 70 species of forbs, trees and bushes, which Halder endorses. Fruits are eaten regularly, and tree bark sometimes (Halder 1976). Gaur (Bos gaurus).In Malaysia, on the basis of observations of the food plants that were eaten, Conry (1981) concluded that the food of the gaur there consists of 41% grasses and graminoids, 23% forbs and 36% woody plants (mainly leaves). According to Conry, this would be in accordance with research by Weigum, also in Malaysia. In India, on the basis of four rumen samples taken in the dry period, Schaller (1967) ascertained that these consisted of 85% grass on average. The same author mentions the sporadic eating of tree bark and fruits. Likewise, Hubback, also in Malaysia, mentions grass as the main food for the gaur. Bison (Bison bison).The bison occurs both in dry steppes (prairies) and in treeless marshes, in predominantly forested areas (Wood Buffalo National Park). Grasses and sedges form their main food (Reynolds et al. 1978). This is supplemented with forbs and usually few trees and bushes. In the prairies, grass is both summer and winter food, while sedges fill this part in Wood Buffalo National Park. Sometimes, as in Alaska (Guthrie 1990, page 176), their summer




food consists of 50% willow (Salix), the winter proportion of which is 75% (dung analyses). In the McKenzie Bison Sanctuary bison reserve (Central Canada), willow turned out to be an important source of food during extremely dry summers (Larter & Gates 1991); its proportion from year to year during that season varies from 10 to 40%. Bison are supposed to be better able than domestic cattle to digest grasses and sedges of bad quality (low protein content, high content of crude fibre) (Hawley 1987, De Liberto & Urness1994). This is probably possible because they are able to recycle relatively large amounts of nitrogen in the shape of ureum, and to transport this to the rumen by way of their blood. In this way, any nitrogen shortage can be corrected and protein-poor (i.e., nitrogen-poor), fibre-rich food can be digested. European bison (Bison bonasus).Initially, Polish and Russian researchers concluded in 1972 that the food of the European bison during the growing season consists of 61-67% of grasses, sedges and forbs and 33-39% trees and bushes (leaves, branches and bark) (G¸ebczyńska et al. 1991). This research, which caused a lot of confusion, had been based on observations of eaten plants. The rumen research carried out later by G¸ebczyńska et al. (1991) suggests that the proportion of trees and bushes during the growing season had probably been strongly overestimated. Based on that research, these authors classified bison as grazers. The research revealed that from spring to autumn, their food consists of 68% grasses and sedges, 3,3% forbs and 8,3% · get supplementrees and bushes. Since the bison in the Forest of Bialowieza tary hay in winter, their food consists of nearly three quarters grasses also in that period. In winters without snow, none of the hay that is offered gets eaten; the animals eat what the forest has to offer. Exactly what that is is unclear. In the winter of 1916/ 1917, when there was no hay because of the war, the bison ate many branches and bark of trees felled for that purpose (Escherich 1917). In the beginning of the 19th century, when small amounts of supplementary hay were given, they would eat a lot of branches and bark of broad-leaved trees and bushes in winter (Brincken 1826, page 57). Hofmann (1989) places the European bison on the transition of the intermediate feeder to the grazer. The population of European bison in the Caucasus is not §

221


222



provided with supplementary food in winter. During the growing season, their food consists mainly of grasses and forbs, and to a lesser degree of trees and bushes (Heptner & Naumov 1966). In autumn, acorns and wild apples and pears are also eaten, as well as the fallen leaves of broad-leaved trees. According to Heptner & Naumov, in winter food is obtained mainly from the trees and bushes, especially bark. Where there is little snow, the edible parts of grasses and forbs are still used. Research in North Ossetia, carried out by Kazmin & Smirnov (1992), seems to corroborate this picture. According to this research, the proportion of bark in the food increases from 39% to 79%, and that of branches of trees and (large) bushes from 2% to 15% in the course of the winter, while those of grasses and forbs decrease from 38 to 5% and of small bushes from 21 to 1%. Bark and branches are taken almost exclusively · the eating of bark (of from linden and elm. Also in the forest of Bia lowieza, aspen, oak, hornbeam and linden) increases towards the end of the winter (Krasińska & Krasiński 1995). §

Photo 18. A group of forest buffaloes (Syncerus caffer nanus) in the Marahoué National Park (Ivory Coast) (Photo: B. Hoppe-Dominik). Wherever they occur in forests these buffaloes are strongly attached to the (already) existing open, grassy spots, such as clear cut areas, forest roads or marshes.



Now that a rough picture has been provided of the food of the aurochs and its relatives, it makes sense to look at what domesticated cattle (can) eat. This does not refer to the present-day stable cows, which are fed grass, silage and concentrated feed, but to cattle that live in a wild or feral state. It is important, moreover, to see how farmers in earlier times, especially during the Neolithic period, got their cows through the winter. Winter, especially the last part of it, was the most difficult period for matters like food, and hay harvesting and storage were not yet common practice during the Neolithic period. Various studies have shown the nature of the food of cattle that have to get through the growing season without supplementary feeding, on whatever plants are naturally available at the time. Scottish Highland cattle that lived in a very grassy Scots pine forest in the Imbos (Veluwe, Holland) from 1982 to 1987 nearly exclusively ate grasses (90100%, mainly green wavy hair-grass (Deschampsia flexuosa) (Van Wieren 1988) (see Fig. 39). The rest of the food consisted of bilberry (Vaccinium myrtillus), heather (Calluna vulgaris) and birch (Betula pendula). In

Fig. 39. Year-through food intake by Scottish Highland cattle in the Imbos Forest (Veluwe, Central Netherlands), an area of heath land and Scots pines. Food intake during the growing season is clearly greater than during the winter period. In terms of percentage, in winter more trees and bushes are consumed (after Van Wieren 1996).

223


224


another Veluvian Scots pine forest on poor sandy soil (Kootwijkerveen), the food of the Scottish Highland cattle during the growing season consisted of 60-70% grass (mainly wavy hair-grass). The remainder was made up of branches, with leaves or needles, of broad-leaved trees and conifers and little bilberry (Van Wieren et al. 1997). Information about cattle food during the growing season, in this case with regard to Heck cattle, are also known from areas that are rich in nutrients, such as the Oostvaardersplassen (Flevoland, Holland) (Cornelissen & Vulink 1996; De Jong et al. 1997). The food in these often wet, rich clay soils consists of 75-95% grasses, including a large proportion of common reed (Phragmites australis). The rest of the food consists mainly of willow (Salix spec.) and few forbs. These are data for the Heck cattle in a part of the Oostvaardersplassen that measures several thousands of hectares. In addition to this, an experiment was carried out on 20 hectares of the same reserve from 1982 till 1986, with MRIJ-, Frisian- and Scottish Highland cattle (Vulink & Drost 1991) (see Fig. 40). Here as well, grass

Fig. 40. Food intake by MRIJ-, Friesian- and Scottish Highland cattle during ten months in a part of the Oostvaardersplassen reserve (Central Netherlands) (after Vulink & Drost 1991). Until the month of March these cattle received no supplementary feed and had to make do with the natural food supply. In winter, in contrast to the rest of the year, they ate mainly willow (Salix sp.).



that included a lot of common reed turned out to constitute a large proportion of the food, i.e. 60-95%. The remainder was made up of forbs and little willow. In some summers, stinging nettles (Urtica dioica) formed an important part of the forbs. Foreign research has also shown that during the growing season, grasses are the main ingredient of the food of domestic cattle, for example in the United States (Thill 1983, Roath & Krueger 1982a, 1982b) and England (Putman 1987). As autumn progresses, it becomes increasingly difficult for cattle to supply their needs with the help of grass. Its digestibility decreases as a result of the declining percentage of proteins (from 12 to 5) and the increasing percentage of cellulose (Van Wieren 1987, Vulink & Drost 1991). The digestibility of forbs and tree leaves also decreases, though much less than is the case with grasses. In forbs, the percentage of protein declines from 17 to 15, in tree leaves from 18 to 12. During winter, moreover, above-ground grasses and forbs die totally or partly, so they become less important sources of food, while trees and bushes become more so (Wallis de Vries 1994, Roath & Krueger 1982). In autumn, acorns can still supplement the food; probably not in large quantities, however, since too many acorns and also beechnuts may cause symptoms of poisoning (Wiesner 1987). Fruits readily eaten by cows include wild apples (Malus sylvestris) (Buttenschøn & Buttenschøn 1985). In the areas where the food choice was ascertained for the vegetation period, the food choice in winter, when there was no supplementary feeding either, was also researched. The Imbos experiment revealed that the proportion of trees and bushes in the cattle food increased to about 35% and that the remainder of the food was made up of wavy hair-grass. At Kootwijkerveen, the proportion of grass declined more strongly; at the end of the winter 65% of the food consisted of conifers and broad-leaved trees. Also at the Oostvaardersplassen, the proportion of willow increased in the course of the winter; in the large reserve it increased to about 50% (De Jong et al. 1997). In the experiment described by Vulink & Drost (1991), the proportion of willow (Salix alba, S. triandra, S. cinerea) had increased to 95% by February.

225


226


The proportion of trees and bushes in the mild climate of Louisiana (United States) may also increase to 55% during the winter (Thill 1983). The intake of branches and leaves by cattle during the dry period in the tropics, when the quality of the grass declines, increases just like in the more temperate zones. In Zambia, for instance, the grass in the driest period (August) contains only 3% protein, while tree leaves contain 12.5%. During the dry period, therefore, zebus eat a lot of tree leaves and little grass (Rees 1974). A type of food not generally known with reference to cows, which may nevertheless sometimes contribute greatly to their food intake, is tree bark. This may derive from various tree species, both broad-leaved trees and conifers (Kinnaird et al. 1979, Tustin 1975). The consumption of rowan (Sorbus aucuparia), as described by Kinnaird et al. (1979), is an example of this. This took place in Scotland in the winter of 1973/1974, from December to April. The cattle there (Hereford and Hereford crossbreeds; D. Welch in lit.) ate about 1,5 kg (dry weight) of rowan bark each a day. Notwithstanding the fact that cattle there were given supplementa-

Photo 19. Cattle in nature reserves sometimes eat a lot of tree bark. Here a Galloway is pulling the bark off a willow stem (De Blauwe Kamer reserve, Central Netherlands) (Photo: J.M. Gleichman).



ry feed, and in April, a lot of grasses and forbs were newly available, a lot of bark was eaten. Since tree bark contains few nutrients and is not easily digestible, the reason for the bark stripping was not quite clear. In the nature reserve De Blauwe Kamer (Rhenen, Holland), a lot of tree bark is eaten as well, especially willow, mostly from February to early April (J.M. Gleichman, pers. comm.) (see Photo 19). The consumption of willow bark in spring may well be connected to its relatively low cell wall content during that period (S.E. van Wieren pers. comm.) Another aspect of feeding in winter is that the amount of food that is taken in declines during that period, as was already ascertained of other herbivores during that period (Van Wieren 1996, Thill 1983). In cattle, the amount of plant material can be cut by half; this is attended by slower metabolism. In spite of this slower metabolism, body weight may still decrease greatly in the course of the winter. In the Imbos experiment, Van Wieren (1988) noted a decline in body weight of about 30% compared to the situation in autumn. Especially against the end of winter the initially available vegetable food starts running short, and the most difficult period of the year sets in for many species of herbivores. As to the European bison at Bia³owie¿a in the beginning of the 19th century, Brincken (1826) noted that they were ‘very fat’ in autumn, but ‘thin’ in spring. The importance of good food and fat storage during the growing season is obvious; with the help of those, a difficult period like the winter may largely be survived well. Although cattle during the growing season have the opportunity to eat the food they are initially adjusted as grazers, with regard to their morphology and metabolism, in winter they are compelled by necessity to change to food that is more difficult to digest and contains fewer nutrients. From the moment that man started keeping wild cattle as domestic animals, this problem kept presenting itself during cold or dry periods, when no good grass was available in the fields. For centuries, in the areas of Europe that are rich in grass so that there was sufficient opportunity to make hay in summer, this was the most important winter food (Heybroek 1963, Behre 1988). In areas where climatological circumstances prevented this, people were forced to use other vegetable

227


228


resources, such as harvest waste and branches, leaves and bark of trees and bushes, to feed their animals. For a long time, especially trees and bushes formed the most important winter food in many areas (Rasmussen 1989, 1993; Brockmann-Jerosch 1918, 1936). During the Neolithic period, man was not yet able to produce and store large quantities of hay. South-Swedish research shows that this was not possible until the transition from the Bronze to the Iron Age (Gaillard et al. 1994). Before that time, hay was not an important food yet; cattle mainly had to feed on trees and bushes. This practice lasted for centuries in large areas of Europe and Asia, either because regular hay harvests were not possible in summer, or as an emergency practice after the hay harvest had failed. Especially in mountain areas, the practice of feeding tree foliage endured until way into the 20th century, and it is still done there (Alps, Behre 1988; Himalaya, Heybroek 1963). Both in the Alps and the Himalayas, nearly all broad-leaved tree species (such as oak, hazel, maple, poplar, holly) were used for harvesting foliage (branches and leaves); sometimes also conifers (yew, Heybroek 1963). The trees were lopped in summer, after which the foliage was dried and stored. Trees were usually stripped of their foliage once every two years. Of the broad-leaved trees, especially ash (Fraxinus excelsior) (see Photo 20), elm (Ulmus spec.) and linden (Tilia spec.) were used a lot, probably because these species are so nutritious. Their leaves are among the richest in protein and easiest to digest of all European tree species, comparable to good-quality hay in these respects (Nehring 1965). It is true that tree leaves that have fallen in October and November are of lesser quality, but they are still fit to be used as cattle fodder. Because the harvesting and transporting of tree foliage was probably rather more ‘labour-intensive’ than harvesting and transporting a quantity of hay of similar nutritional value, the latter was still preferred wherever possible. Another type of winter food, which was given to cattle especially in Norway and Sweden, was tree bark (Ropeid 1960). This was given raw, boiled or ground, and not stripped off living trees, but from harvested ones. The stripping started in autumn and continued through the winter. Several tree species were used, both broad-leaved trees (for instance



Photo 20. During early-day winters branches with leaves were usually fed to domestic cattle. This foliage was cut for that very reason at the end of the summer. This picture from Romania shows ashes that are cut in a 2-year cycle for the same purpose. (Photo: H.M. Heybroek)

elm, rowan, oak, apple, ash, birch) and conifers (juniper, Scots pine). Aspen and rowan were used most often, however, and most appreciated. Though tree bark was often used as winter food, it was still considered emergency fodder, to replenish shortages and survive long winters. It was mostly given only as admixture. In many areas, the bark and branches of trees formed the only available cattle food in winter. Studies of aspen and rowan bark explain why this food had to be considered emergency fodder (Nehring 1965); while the digestibility of organic matter equals that of good straw, the proteins in tree bark, which amount to 4.7% in aspen and rowan, are poorly digestible. During the Neolithic period, when hay harvesting was not yet practiced, the feeding of cattle with branches and leaves must have been

229


230


common practice in winter (Rasmussen 1989). Evidence of this was found at Early-Neolithic sites in Switzerland (4300 BC, in the starting period of agriculture in Europe), not just in the shape of some heaps of leaved twigs (of elm, birch, linden, ash and maple), but also of thick layers of animal dung, in which branch and leaf rests could be shown (Rasmussen 1989, 1993). Research showed that not only branches with leaves, but also branches without leaves, with flowering catkins (of alder and hazel) were fed to cattle. These were obviously cut both in winter and in early spring. The main tree species that were used were ash, linden, willow and alder. The not inconsiderable proportion of ivy (Hedera helix; sometimes up to 18% of the volume) in the winter food stood out in this research. Rasmussen supports the results of his research by relating them to early peasant communities in Norway, which had to survive extremely long winters (eight months). The cutting of branches in the middle of winter and in spring, when there were flowering catkins, was common practice there. Elm twigs were said to be sufficient for cows to survive on; birch catkins were also highly appreciated. Analysis of winter twigs (up to 1 cm in diameter) shows that their digestibility, compared to that of summer twigs, is a little better (Nehring 1965), due to the higher protein content (7.4% against 5.9% in summer) and the lower content of crude fibre (41.5% against 44.8% in summer). According to this analysis, there is little difference between the various tree species in winter. Feeding experiments with ground winter twigs, which were given to horses, cows and sheep, revealed that horses did not eat these, or were very reluctant to do so. Cows and sheep, however, could be fed with exclusively winter twigs for some months, although their milk yield left something to be desired. Large differences in their reactions to this food were found between individual animals, and this also showed in other experiments. Fresh twigs were received more favourably than dried ones, while some of the tree species used, such as birch, turned out to be better liked than others. The explanation offered in the above, of the morphology and food choice of both the aurochs and the domestic cow that descended, warrants the conclusion that there is no reason to suppose that any great



differences existed between the two. Unlike the external morphology of the domestic cow, man is unlikely to have influenced its internal morphology, meant for the digestion of food, and its metabolism. Through the centuries, cattle have had to digest roughly the same food as that which was available to the aurochs.

Reconstruction of the food of the aurochs in its original habitat Spring and summer: Mainly grasses and graminoids, supplemented with forbs and leaves of trees and bushes. Autumn: A declining proportion of grasses and increasing proportion of trees and bushes. Possible supplements in the shape of tree fruits and fallen tree leaves. Winter: Beside grasses, graminoids and forbs, an important part of the food was made up by branches and, to a lesser degree, bark of trees and bushes.

Summary As written sources have shown, the food of the aurochs during the growing season consisted mainly of grass. Apart form that, acorns were eaten in autumn; in winter, moreover, branches of trees and bushes were added to the menu. The last population of aurochs in the forests at Jaktorów (Poland) was provided with supplementary hay in winter. Research of the food choice of wild cattle in nature reserves shows that grass is the main food during the growing season. In winter, a large, sometimes even very large part of the menu may consist of branches of trees and bushes; tree bark is also eaten regularly during this period. Whenever early man was unable to make hay on a large scale, or could not dispose of hay for other reasons, he would give his cattle tree foliage that had been cut and dried during the summer as winter food. In some parts of Europe this practice is still in place.

231


232


It may be concluded there is probably no difference between the food choice of present-day cattle and that of their wild ancestor, the aurochs. This chapter also provides a survey of the food choices of a number of still-living relatives of the aurochs, which have been characterised as grazers.

9.1.3. The habitat of the aurochs Data about the habitat of the aurochs, such as the type or types of landscape it preferred, stem mainly from Europe and to a small extent from the Middle East and North Africa. Because of the relatively late occurrence of the aurochs in Europe, the majority of the bone and horn finds, pictures, descriptions and records that have been preserved stem from the European aurochs population. These European data, supplemented with information about its occurrence in the Middle East, are particularly useful for a portrayal of this animal’s habitat. Aurochs bones are found mostly in soil layers. The rate of preservation of those bones depends on the conditions in the soil. In wet soil layers, like wet clay and peat, where oxygen-poor or even oxygen-less conditions prevail, bones are better preserved than in, e.g., dry sandy soils, which admit oxygen so that decomposition can take place. If a bone of a specific animal is found somewhere, the chance that that animal lived there is good, especially when more of its bones are found at several sites in the area in question. However, if no bones of a certain animal are found, this does not necessarily mean that it did not occur there. It may have lived only in the drier sections of a specific area, for instance, where its bones have decomposed quickly. Whenever this is possible, in order to get a clear picture of the aurochs’ choice of habitat inside its distribution area, oral and written records that may provide some additional information have to be used as well as other sources. From North Africa and the Middle East, both bone finds and pictures have been preserved. The North African aurochs is known from bones that were found in caves as well as from rock paintings, both of which are related to the northern coastal area (Gautier 1993a). Neither bones nor rock paintings of aurochs were found in the more southerly



situated area that is now called Sahara. The most similar bovine animals depicted in rock paintings from the area are clearly domesticated and often accompanied by shepherds (Muzzolini 1983). The only area in North Africa where the aurochs occurred somewhat further to the south is the Nile valley, where the animal occurred as far as North Sudan (Gautier 1993a). Up to 3000 BC, the present-day Sahara was much wetter than today and included a savannah landscape, comparable to the present Serengeti in East Africa. The fauna was similar too, and included African buffalo, hyena, elephant, lion, rhinoceros, zebra, and baboon. Bones of these and other large mammals were found in the area (Gautier 1993a). The fact that bones of these animals were found, but no aurochs bones, quite probably indicates that the aurochs did not live there. This animal did not inhabit even the wetter parts of this area, such as riverbanks and temporary marshes. In view of the bone finds, it must have lived in the northern coastal area, which was densely forested, probably with steppe forests at the south border, where the aurochs came in contact with zebra and wildebeest. In this northern coastal zone, other animal species from more northern regions must have occurred as well, such as red deer, brown bear and wild boar. Beside this coastal zone, both the flood plain of the Nile valley and the areas directly bordering it must have hosted aurochs at one point. The aurochs, like the hartebeest (Alcephalus bucephalus), was one of the most conspicuous large mammals in this valley. The wild boar was another (Uerpmann 1987). Vegetation along the Nile, when the aurochs still lived there, was very open and low in trees. Plant growth in the flood plain of this river consisted mainly of grasses (Cynodon spec., Panicum spec.) and Cyperus species. Here and there, willow bushes (Salix spec.) occurred (Schild & Wendorf 1986). If its distribution in North Africa is considered, the habitat of the aurochs already emerges a little more clearly. The animal avoided areas that were too dry and looked for more forested, wetter areas. Its distribution in the Middle East draws a comparable picture. During the period 8000-6000 BC, when man is unlikely to have had a lot of influence on the occurrence of the aurochs, the animal lived in the valley of the Euphrates in North Syria, in the adjacent forest steppes in Central Syr-

233


234


ia, with their Pistacia trees, in the area of the springs of El Qom and Palmira and in Central Anatolia (Turkey), in the lake district of Çatal Hüyük and in the fresh and salt marshes there (Konya) (H. Buitenhuis pers. comm.). To judge by these sites, the animal occurred especially in places that could provide water and shelter; it did not occur in steppes, or at best in forest steppes. The conclusion drawn by Duerst (1899), that, in view of the Sumeric character for it41, the aurochs in the Middle East occurred mainly in mountains, needs revision. The three ^’s in this character do mean ‘mountains’, but used in this connection they indicate the ‘exotic, foreign’ character of the bovine animal (T. Krispijn pers. comm.). The animal apparently already lived well apart from human society in Mesopotamia. Such Sumeric characters were used from circa 3000 BC. During the last ice age (Würm), when the climate in the main part of Central Europe was cold and dry and this area was covered with an extensive steppe vegetation of grasses, sedges and Artemisia species (Kahlke 1994), the most southerly living areas of the aurochs were probably situated in Spain, South France, Italy and the Balkans. The exact northern border of its distribution area in these countries at the time cannot be traced as yet. The aurochs did not belong to the steppe fauna, which did include mammoth, woolly rhinoceros, steppe bison and horse. Its bones have never been found together with bones of these steppe animals; at most with those of bison and horse, dating from the Preboreal and Boreal periods. Moreover, it seems to have lived in the mountains, separate from those other species, in this period. In the Pyrenees, the aurochs did not live above 500 metres during the last ice age, although the four other species did (Clot & Duranthon 1990). The aurochs avoided the dry, cold grasslands of the mountains and limited itself to the more temperate, southerly areas. The fact that no aurochs remains from the period in question were found above the said height probably indicates that they did not visit these regions, even in the summer. With the retreat of the glaciers to the north as a result of the warmer climate, the aurochs extended its distribution area further north. In Germany, the first postglacial aurochs finds date from the Allerød period 41

In former Mesopotamia, the character M was used to indicate the aurochs.



(Baales & Street 1996, Tietz 2004). The earliest Danish and Swedish Holocene aurochs finds date from the Young Dryas period (Degerbøl & Fredskild 1970, Aaris-Sørensen 1999). The Allerød and Young Dryas periods were characterised by great climatological changes that also caused changes in the flora and fauna. There was a transition from a relatively dry, continental climate to a warmer and wetter climate. The landscape was still very open and consisted of steppe forests and floodplain forests, with birch and pine as the most important tree species and to a lesser degree also willow, aspen and juniper (Bottema 1987). In the Preboreal and Boreal periods, the conditions for tree growth improved gradually and the landscape became less open as a result. During the Atlantic period, the maximum temperature and degree of afforestation were reached. It is remarkable that most of the bone finds from both Denmark and South Sweden date from the first half of the Boreal period (Isberg 1962, Degerbøl & Fredskild 1970), while only a few were found there from the Atlantic and Subboreal periods. The conditions during the Preboreal and Boreal periods, with not entirely closed forests of tree species that demand light, were probably well suited to the aurochs (see Fig. 28). The Atlantic period that followed and the periods after that offered less favourable living conditions for this animal, possibly because the forest became more closed, with tree species that are tolerant of shade (linden, elm, beech, hornbeam). In addition to this, the hunting pressure exercised by man and the ousting by its cattle continued to increase. The vast majority of European aurochs finds stem from the lowlands; hardly anything was found in mountainous areas (Söffner 1982). Nearly all lowland finds were discovered in wet clay and peat layers, such as river- and lake banks, coastal sediments and certain peat bog layers, dating from the period in which the peat was still fen (Degerbøl & Fredskild 1970), Söffner 1982, Van Es 1990). Only a small proportion of the finds stem from human settlements, which would seem to warrant the conclusion that the aurochs occurred exclusively in those wet areas and not outside of them, since it was not found there. As was explained before, however, the superior preserving conditions of wet soil layers may lead to the wrong conclusions about the occurrence of a species, since any occurrence in dry areas may not be observed in the

235


236


same way. Nevertheless, the fact that so many finds originate from wet areas does suggest that such habitats at least constituted an important part of the local distribution area of the aurochs. That the number of bones found there is so large cannot simply be explained by the theory that these were animals that somehow died on their way through those wet habitats. Another part of the explanation for this must be sought in the assumption of their protracted stay in these areas, for instance because the aurochs were looking for food. For the sake of comparison, it so happens that the bones of a related European grazer, the European bison, were found only rarely in river sediments, for instance of the Narew River (Northeast Poland, Ruprecht 1976). Bone remains of European bison were found mainly in human settlements, for instance in Germany and Switzerland (Söffner 1982). European bison in the forests of Bia³owie¿a are known to have displayed an aversion to wet habitats. Research by Borowski & Kossak (1972) has shown that a forest type like the alder-ash forest (Circaeo-Alnetum) was not frequented by bison, in spite of the presence of suitable food. Bison did not occur in open, wet sedge vegetations, in the forest or along small rivers like the Narewka, Hwo¿na and Leœna, either (Krasiñski 1978). Clearly, it will not have stayed long in such habitats, as a result of which few remains have been found there. Some aurochs remains were in fact found in human settlements, but proportionally many more were found in places that have no relation to human habitation. Areas in Europe that were definitely frequented by aurochs are the estuary of the Severn River (Southwest England) (Allen 1997) and the more northerly coastal area at Liverpool (Formby Point, Huddard et al. 1999). Clay layers from the periods 5000-4000 BC and 4000-3000 BC, respectively, have yielded hoof prints and tracks of a very large bovine species. These were very probably left by an aurochs, since domesticated cattle did not yet occur in these areas at that point, and such wet coastal marshes are considered not to have attracted the European bison, if it occurred there at all. The earlier occurrence of the aurochs in those salty coastal marshes, which are influenced by the dynamics of the sea, should not evoke a lot of surprise. Also nowadays, domestic cattle can still manage quite well in the salty and brackish coastal vegetations of several



nature reserves in Europe (Camargue (France), Verdronken Land van Saeftinghe (Holland), isles of the Wadden shallows)(see Photo 21). In the Camargue, salt plants such as glasswort (Salicornia spec.) and seablite (Suaeda spec.) are used as food (Schloeth 1961), while the wild cattle on the Island of Swona (England) eat several species of seaweed that they find on the beach (Hall & Moore 1986).

Photo 21. Domestic cattle in the coastal salt marshes of the Isle of Schiermonnikoog (North Netherlands) (Photo: S. E. van Wieren). The aurochs likely lived in such salt vegetations as well.

Of the last two areas in Europe known from written traditions to have hosted aurochs until the late Middle Ages, i.e. the Great Wilderness in the earlier East Prussia and the forests at Jaktorów in Central Poland, the former provides the least information about a specific aurochs habitat. There are no mentions or other indications from this area that can throw any light on this. The Great Wilderness is known to have consisted of forests and marshes, it is known which tree species grew there (Mager 1960) and that the aurochs occurred there, but where exactly the animal lived cannot be traced on the basis of the data at hand today. The area was rather flat and low, and included the Mazurian lake district and the Prussian lowland plain. The climate was somewhat comparable to that of Bia³owie¿a, strongly tended towards a continental character and was characterised by severe, snowy

237


238


winters. Since the marshes and rivers were mostly frozen in wintertime, the Great Wilderness was easier to visit (for people) in that season than in the summer (Paravicini 1995). The forests of Jaktorów, Bolemów and Wiskitki together formed a much smaller forest complex (25,000 ha) than that of the Great Wilderness. The winters there are a bit milder than at Bia³owie¿a (Faliñski 1968). The height of the area varies from 90 to 130 m above sea level (Heymanowski 1972). The Inspection Reports provide information about the tree species that grew in these forests. About the forest of Jaktorów, the 1564 report notes (Kêdzierska 1959): ‘At this village there are large forests……in these forests there are numerous large trees, suitable to build with and for bee nests……there are large (Scotch) pines, very large oaks, ashes, hornbeams, (Norway) maples, elms, alders, birches, (sycamore) maples and numerous other trees’. The forest of Bolemów is mentioned in the same year: ‘At Bolemów there are large forests…..partly coniferous forest, partly broad-leaved forest; there are large Scots pines, no few bee nests, large oaks, hornbeams, lindens, alders, birches and plenty of timber of all kinds…’. The larch (Larix decidua) probably occurred here as well, since larch timber was found in neighbourhood houses from that period (Heymanowski 1972). As was already explained in Chapter 5.2.1 on the basis of the Inspection Reports, in the 16th century the forests in these areas began to be used increasingly for timber. The felling of trees was rampant. A map of 1792 by De Perthées (see Fig. 46) shows that a large part of the forest complex (especially of the Jaktorów forest) had already vanished, and that the remaining part was cut by roads and zones designed to open up and subsequently undermine the forests. Moreover, small rivers and accompanying meadows can be seen. The mapmaker distinguished between forests, bushy areas, meadows and various types of roads and rivers with the help of a legend. To this day, the forest area maintains a considerable network of brooks and small rivers (Heymanowski 1972). Judging by the situa-



tion around 1800 (Heymanowski 1972), it must have been much wetter in the old days. The present-day cultivated landscape that has derived from that is characterised by ditches and straightened brooks (author’s observations), which are clearly meant to make the area drier and easier to cultivate. Matuszkiewicz (1966) has tried to ascertain the potential natural vegetation (PNV) in a part of Masovia, on the basis of the present vegetation (scale 1: 300.000). The area of his research included the forests of Jaktorów and Wiskitki. According to this crude study, nutrient-rich wet alder-ash forests were situated along the small rivers, while the rest of the area was mainly covered with linden-hornbeam forests and, in relatively dry, nutrient-poor areas, with pine-oak forests. In the 16th century, there were meadows along the rivers and rivulets (Heymanowski 1972), though it is not clear if these were natural or man-made. The data that are available at this point merely serve to sketch a rough picture of the old situation; apart from the meadows we do not actually know where the aurochs lived at the time. In addition to the many generalising mentions of the occurrence of the aurochs ‘in forests’, as in the Forest of Jaktorów, a number of written reports have been passed down, which contain more specific remarks about this animal’s habitat. In Russian epic poems, for example, the aurochs is said to have stayed in marshy forests, feeding on grasses (Dementiev 1958). In the poem ‘Dobrynia Nikititch’, the sorceress Marina says to the knight who has changed into an aurochs: ‘Pull out the stiff grass, devour the grass with the marsh water, come to me from the black mud, from the waters of the marsh’. This suggests what was most likely considered something the audience was well aware of, i.e., that such animals not seldom occurred in marshes. A mention in which the same connection was made is found in an Anglo-Saxon runic verse, which was passed down in writing for the first time in the 9th century (Beck 1968, Rodrigues 1992). It is a poem that belongs to the rune sign that represents the ‘oo’-sound, and which was associated with the aurochs42. The verse, which was written in Old English, gives a concise description of the animal: 42 The rune G is used to represent this ‘oo’-sound. Each rune is accompanied by a verse, in which something is said about the plant, the animal or the concept associated with that rune.

239


240


‘Ur byþ anmod and oferhyrned; felafrecne deor – feohteþ mid hornum – mære morstapa; þæt is modig wuht’. ‘The aurochs is fearless and large-horned; a fierce animal – it fights with its horns – the famous marsh walker; it is a brave animal’. Here, the animal’s characteristics are described, the way many people (in the area of origin of the Anglo-Saxon language, i.e., North Germany) knew and experienced them at the time, the way the aurochs was known there. Apparently it was not unusual to nickname this animal ‘marsh walker’, presumably because the marshes formed the place where it was quite regularly seen. Plants and animals often get names that are related to their physical appearance (redbreast, weeping willow) or to their preferred habitat (reed bunting, marsh fern). The aurochs may well have received its nickname in the same manner. Another bovine species, the African forest buffalo, in the Rega language (Central Basin of the Republic of Congo), is called ‘Mbogô nyama y’ilambo’, which means ‘the buffalo is an animal of the marshes’43. This is another example of a name that refers to an animal’s preferred habitat, in this case the marshes in the rain forests situated there. In the same way, the Malaysian gaur is nicknamed ‘seladang’ (Hubback 1938); the word ‘ladang’ means ‘open place in the forest, caused by shifting cultivation’. The animal is observed regularly in such places and was named after them. The aurochs received a nickname that referred to its marshy habitat, similar to the Dutch name for the white stork (Ciconia ciconia), i.e., ‘ooievaar’. The original meaning of the word ‘ooievaar’ was probably ‘marsh walker’ (De Vries 1987); a name evoked by the bird’s preference for nutrient-rich marshes, for instance along rivers. Another clue to the animal’s habitat may be found in the mixed aurochs/ European bison description given by Von Fleming (1719). This description, in which the author unintentionally mixes up elements of both the aurochs and the European bison, includes the following passage: 43 Information provided by Malembe Mbo, Institut Congolais pour la Conservation de la Nature (Kinshasa, Republic of Congo).



‘Whatever other cattle eat, the ‘Auer’ eats as well. It is a harmful animal, very unfriendly disposed towards man, and in the wilderness, in marshes and reed lands, it usually lies deep in the water, so that only its head can still be seen’. This passage cannot refer to the European bison at all, since this animal, as was mentioned before, was not inclined to stay in such areas for long. That is why this is probably a description of an aurochs habit, i.e., the frequenting of and sojourn in marshes. The source for Von Fleming’s description may have stemmed from the Great Wilderness. The fact that this animal no longer occurred there at the end of the 17th century does not necessarily mean that there were no more stories being told about this remarkable animal. People had long been unable to distinguish between the aurochs and the European bison, which may explain how the reference to marshes came to be included in a description of the latter animal (‘Auer’). A final mention, although it was found in Old-Germanic myths and legends, originally stems from South Sweden (Isberg 1962). The legends in question, in which giants, dwarfs and witches play a role, go back to heathen, Pre-Christian times (before AD 1000). Several of these legends mention ‘giant black cattle’, also called ‘river cattle’. These terms probably refer to the aurochs, which then still occurred in North Germany (not in Sweden anymore) and of which Adam von Bremen (1072) indiscriminately wrote, that they lived ‘under water’ (‘sub aqua’), just like polar bears. Von Bremen may have been influenced by the myths that surrounded this animal. As such, even though they stem from the misty world of heathen legends, on account of their possible core of truth these data may be able to lift a corner of the veil.

Other bovine species To put the habitat choice of the aurochs in a somewhat broader perspective, it may be useful to compare it to those of the other members of the Bovini tribe. This is especially significant since it may provide an overview of the possibilities available to grazers in forest areas, and also

241


242


of the habitat choices open to the aurochs. In the first instance, like the aurochs, the other bovine species are considered grazers (Van Wieren 1996, page 175), which is not to say they would not eat anything else. Depending on the bovine species and the place and the season in question, other vegetable food is also often eaten in no small quantities. Grasses and graminoids do form their most important source of food, however (see Chapter 9.1.2 and Table 13). Table 13. Survey of research into the habitat choices of a number of bovine species related to the aurochs.

- African buffalo (Syncerus caffer caffer). This animal occurs from the lowlands (sea level) up to the high mountains (4000 m; Prins 1987); it inhabits, among other places, savannas and savannah forests, always in the vicinity of water, since it depends, more than most other African members of the family of Bovidae (antelopes and wild cattle) on water to drink and on pools to take mud baths (Prins 1987). The African buffalo has become known especially from the East African savannas, but also occurs in large numbers in the marshes and reed lands along the rivers and in the river deltas, of which the large Zambezi delta (Marromeu) is an example (Tinley 1974). - Forest buffalo (Syncerus caffer nanus). This is a small subspecies of the large African buffalo, which occurs in the West-African tropical rain forest zone, for instance in Ivory Coast (Hoppe-Dominik 1988). The habitat this animal has chosen in this zone consists mainly of riverbanks and marshes (e.g. the Azagny reserve, Ivory Coast) (Christy 1929, Hoppe-Dominik 1988, Blake 2002). It seldom occurs in primary forests, since the amount of grass is limited there. Whenever man by means of roads opens such primary forests, the forest buffalo uses the roads to go into the forests, since these are very grassy areas (Hoppe-Dominik 1988). In the tropical rain forests of Gabon (Central Africa), the animals use the elephant paths (Campbell 1991). In the Tai Forest (Ivory Coast), forest buffaloes are found only in the border zones on account of the road construction carried out there by man, and because, more so than forest elephants (Loxodonta cyclotis) and bongos (Tragelaphus euryceros), forest buffaloes are dependent on grass. In Northern Congo, forest buffaloes prefer ‘open grass and sedge dominated swampy clearings’ (Blake 2002).



Table 13. Continued

- Water buffalo (Bos arnee). Of all members of the Bovini tribe, this animal is most strongly tied to water (Klös & Wünschmann 1968). It occurs at river- and lake banks and in marshes and marshy forests, and often lies in water and slush for lengthy periods of time. - Yak (Poephagus mutus). This animal occurs in the cold, dry grass steppes of Northern Tibet, up to a height of 5200 m (Schäfer 1937); it bathes regularly in lakes and rivers. - European bison (Bison bonasus). Nowadays, European bison occur in the · and in the Caucasus, among other places. It used to occur forests of Bialowieza generally in the steppes and steppe forests of South Russia as well (Heptner & · Forest, the preferred forest types of the Naumov 1966). Inside the Bialowieza European bison are the moist and dry nutrient-rich linden-hornbeam forests and the drier nutrient-poor pine-spruce forests (Krasińska & Krasiński 1995). The least frequented forest types include the wet and nutrient-rich alder-ash forests (Circaeo-Alnetum) situated along brooks and rivulets. Bison do pass through these but seldom eat there, in spite of the fact that there is sufficient suitable food for them (Borowski & Kossak 1972). This animal’s presence in the somewhat drier forest types in the forests, and in the drier regions outside these wooded areas, seems to be reflected in the European distribution of bone finds of the European bison. In the west and south of Russia, many bones of this animal were found (Heptner & Naumov 1966); elsewhere in Europe, few were found in river sediments, but all the more in human settlements (Söffner 1982). Such settlements are often situated at the foot of mountain areas or in the vicinity of somewhat drier, higher regions. From a vertical perspective, also in Switzerland there seems to have been a distinction between the occurrence of the European bison and the aurochs. (Söffner 1982); the European bison lived higher up in the mountains, the aurochs lower. Also nowadays, the Caucasian subspecies of the European bison still lives up to great heights (3000 m) in the mountains of the Caucasus, up to the summer snow boundary (Heptner & Naumov 1966). In this way, the European bison seems clearly distinguishable from the aurochs by its life in a somewhat colder, drier habitat. This could also be an explanation for its less than general occurrence in West Europe. §

§

243


244


Table 13. Continued

Its preference may reflect the choices of its immediate ancestor, the steppe bison (Bison priscus), which inhabited the cold, dry Pleistocene steppes during the ice age.Apart from swimming across rivers, the European bison does not bathe (Heptner & Naumov 1966). It likes to roll in dry sand, not wet mud. - Bison (Bison bison). The North American bison occurs from the lowland up to the mountains (Roe 1972). The mountain bison was of smaller stature than the prairie bison. The bison became known especially in the prairies in North America. Beside this area, it used to occur also in the wooded east of the United States. Exactly where in those forests it lived has not yet been researched thoroughly (see Ch. 9.4 ‘Regions comparable with Europe’). Prairie bison lived on the prairies both in summer and in winter. Further to the north, in Canada, a number of bison occur inside the boreal coniferous zone. Wood Buffalo National Park (Northeast Alberta) is the best known. This area is entirely forested, except where it is too wet for tree growth, for example along rivers. Bison scarcely live in the forests there; they live nearly exclusively along the rivers and in the delta of the Peace and Athabasca Rivers (Carbyn et al. 1993). They stay in these places both in summer and in winter (see Photo 22). - Gaur (Bos gaurus) and banteng (Bos javanicus). The gaur lives in India and IndoChina, the banteng in Indonesia and Indo-China (Wharton 1968). Both live in Indo-China, therefore, but a distinction between their living areas may be broadly outlined. According to Wharton (1968) and Steinmetz (2004), this distinction consists of the fact that the banteng lives in drier habitats than the gaur: the savannas, savannah forests and the relatively dry deciduous forests. The gaur, on the other hand, occurs in the wetter deciduous forests and can also be found in tropical rain forests, provided that there are sufficient grasses and forbs. The open spots in the forests (‘ladangs’), caused as a result of shifting cultivation, are examples of such places. If these ladangs are not burned regularly, they close up again with forest, notwithstanding the feeding by gaur and other herbivores that takes place in them. Other grassy open vegetations in this tropical rain forest occur along rivers. Conry (1989), on the basis of his own research and that of others, was able to ascertain a strong tie of the gaur to riverbank vegetations.



Photo 22. Bison grazing in riverside sedge marshes in Wood Buffalo National Park (Central Canada) (Photo Parks Canada, Wood Buffalo National Park). Both in summer and winter these sedge vegetations make up the preferred habitat for bison.

Reconstruction of the aurochs habitat As was outlined in Chapter 9.1.1, insect- and pollen research, Roman reports and the situation in the Great Wilderness show the natural landscape of Central Europe to have consisted mainly of extensive ‘forests and marshes’. These ‘forests’ may be imagined as more or less closed complexes of primeval forests, with occasional open spots. The ‘marshes’ were either peat bog marshes, not influenced by any ground or surface water, or fen marshes along rivers and lakes. Marshes were more or less treeless. In Ch. 9.4, it will be explained that the feeding by large herbivores naturally has very little impact on the development of forests in the places that are suitable for such development. Other natural factors, such as the availability and the influence of water, have more significant effects. Another natural factor, fire, probably played a negligible role in the Central European forests, as has become evident in the case of the Great Wilderness (Mager 1960, II, page 214) and in the forests at Bia³owie¿a (Brincken 1826, pp. 38, 39). Man in particular used fire as a strategy to gain agricultural ground or to increase the production of honey; fire was very rarely caused by lightning.

245


246


Storms are able very locally to open up a forest by destroying the trees (Brincken 1826, page 30, 39; Faliñski & Faliñska 1986), but this has scarcely any impact on the forest as a whole. It remains questionable, therefore, to what extent places that had been more or less cleared by storm could really contribute to the food supply of grazers, more particularly large grazers. It is true such parts of the forest would admit an increased amount of light to fall on the forest floor, which would lead to a greater production of grasses and forbs (Reynolds 1969) compared with other, adjacent pieces of forest. The difficult accessibility of such areas for large animals will probably have resulted in their relatively limited use of them44. Brincken describes the ‘impenetrable’ chaos of just such a piece of forest overthrown by storm, where many young trees were starting to shoot up between the masses of fallen trees (see Ch. 9.4). Research by Reynolds (1959) shows that parts of a thinned-out Populus tremuloides forest, where the fallen trees were left lying, were used much less by deer and cattle than forest sections that had not been thinned out, in spite of the former’s greater biomass of grasses and forbs. The greatly increased regrowth of Populus tremuloides in the thinnedout pieces of forest does suggest that the regrowth of trees under conditions of limited feeding progresses relatively well. Both Mager (1960, II, page 208) and Brincken (1826, page 40), with regard to the Great Wilderness and the forests at Bia³owie¿a, respectively, conclude that under natural circumstances, the influence of insects was probably generally very limited. Brincken mentions that an infestation by insects in 1811, which involved pines across several dozen hectares, was considered quite exceptional. Very many trees perished at the time; 15 years later an abundant forest regrowth had shot up in their place. The forestry methods that emerged in the course of the 19th century disrupted the natural forest ecosystem in such a way that insect infestations began to occur on an ever-increasing scale (Mager 1960). Under natural conditions, however, insects at most will have defoliated small pieces of forest. The aurochs in its natural forest habitat is likely to have sought the grassiest areas. The occurrence of such places depends on the quantity 44 A comparable example is mentioned by Groenman-Van Waateringe (1993). In a certain forest area much dead wood lay for years, thereby deterring cattle from entering. Only after the removal of this dead wood, the cattle entered this forest area and started feeding on the abundant grass, not exploited before. See also Ripple & Beschta (2005b).



of light let through the forest canopy. This is affected in part by the species of trees, and also by the aging processes of the forest. If one or several trees fall down from old age, their place in the forest will temporarily become relatively grassy. In so far as such places were still accessible in spite of the fallen trees, the aurochs are likely to have taken advantage of them. The fact that the biomass of grasses and forbs in the forest was relatively slight in comparison to that in treeless areas does make one wonder where the three large grazers (wild horse, aurochs, European bison) that lived in Europe thousands of years ago found their food. As it happens, assigning the aurochs a place in the ecosystem simultaneously raises the question where the other two used to live. Although the aurochs and the European bison could still be considered rather common, the wild horse was probably much less so, and was probably characterised as quite rare (K. Aaris-Sørensen pers. comm.). Wild horse finds from the mainly forested Europe, which had scarcely been influenced by man at the time, are very rare (Uerpmann 1990). Generally, the steppes were the best-known habitat of wild horses. The amount of grass is considerable there, and there is sufficient space for escape conduct. The habitats considered most suitable for the wild horses in Central Europe were fresh- and salt-water marshes. Horses are known to be able to survive in marshy areas; the wet, salty vegetations of the Camargue (Duncan 1992), the sedge marshes by the river in Canada (Slave River Lowlands, Canada; Pringle 1987) and the marshes in Yakutia (East Siberia; Andreyev 1971, Zimov et al. 1995) are illustrations of this fact. Although they are few and far between, finds from the Netherlands that relate to wild horses appear to agree with this. These were found at two sites in Flevoland, De Hoge Vaart (Laarman 2001) and Swifterbant (Clason 1986), which are both situated in former coastal marshes. Such large marshes presumably offered sufficient food, as well as space for escape conduct. Since such conditions were present only very locally in the Low Countries, the populations of wild horses are unlikely to have been large. One of the other two large grazers, the European bison, has already been reported to have occurred not only in forests, but also generally in

247


248


steppe areas. It is questionable that the aurochs ever resorted to the cold steppes. In North Africa, it evidently did not stay in the warm dry areas, but there were probably no possibilities for it in cold dry habitats, either. To illustrate: the bison in North America is able to withstand the low temperatures, combined with wind, much more successfully than the local domestic cows (which include Hereford cattle) (Smoliak & Peters 1955, Peters & Slen 1964). In addition to this, bison (Hawley 1987, Peden et al. 1974, Schaefer et al. 1978) and horse (Andreyev 1971, Van Wieren 1996) are much better able to digest food with a high cell wall content and low protein content (e.g. dead grass) than domestic cattle. Bison and horses are able to survive winter steppe conditions without supplementary feeding, in contrast to domestic cattle (Roe 1972, page 190; Pringle 1987; Andreyev 1971). If there is a thick snow layer, bison (Geist 1996) and European bison (Heptner & Naumov 1966) can push aside the snow with their heads; horses can do the same with their feet (Smoliak & Peters 1955). Domestic cattle do neither and face a lot of problems to get their food out from under the snow. This may well be the reason why aurochs only very rarely resorted to the steppes and always stayed in the vicinity of forests, where the living conditions were somewhat milder and vegetable food other than grass was also available. That the European bison was unwilling to live in forests and could not actually survive there, is belied by this animal’s century-long presence in the Great Wilderness, where it managed to survive under the natural (forest) conditions (Mager 1941). The climate and the food there were different than on the steppes, but it was still able to hold its own, by adapting to the circumstances. The smaller biomass of grasses in the forest may naturally have resulted in a much lower density of European bison than on the steppes. Obviously, European bison and aurochs both occurred in forests. Since the amount of grass in the forest is limited and both species prefer to eat grass and are of similar size, an ecological partition45 between these two species, to avoid excessive competition, would seem logical. Roughly speaking, such a partition has already been shown to exist between gaur and banteng: the banteng frequents the somewhat drier areas, the gaur the somewhat wetter ones (Wharton 1968, Steinmetz 2004). A similar ecological 45 ‘Ecological partition’ means that two animal species live in the same area, but exploit different resources within this area. Their relation is in a dynamic balance, based on their individual competitive power through physical and physiological properties. Changes of circumstance will change the balance.



· (East Poland). Photo 23. Oak-hornbeam forest (Querco-Carpinetum) in the Forest of Bia§lowieza This forest type occurs on moist, eutrophic soils. (Photo: T. van Vuure)

partition seems also to have existed between the European bison (‘dry’) and the aurochs (‘wet’). In addition to this, there was also a division of their habitat in ‘high’ (European bison) and ‘low’ (aurochs) areas, as far as their height above sea level is concerned. Roughly speaking, the European bison inhabited the drier forest types in the lowlands and the higher mountains areas, while the aurochs frequented the wetter forest types in the lowlands and the lower mountain areas. In this way, the European bison with its dryarctic origin and the aurochs with its wet-tropical origin could both survive as grass-eaters in the same area, but different habitats, in Europe46. An ecological partition does not mean that the two animals lived strictly apart. They are quite likely to have met regularly as a result of

46 It is difficult to say whether and how the European bison, in the absence of the aurochs, changed habitat. Gilibert · European bison feed especially (1805, pages 429-438) reports (from hearsay) that in summer in the Forest of Bia§lowieza on sedges (‘carices’); modern research (Krasiński 1978, page 24) mentions that bison do not use unmown sedge meadows since they prefer the mown ones.

249


250


some overlap in their habitats. There must have been forest types in which both of them foraged for food during the summer period, possibly nutrient-rich linden-hornbeam forests (see Photo 23). Their meetings were probably more frequent during the winter period, when the aurochs, more so than in summer, visited the forest, since the quality of the grasses and graminoids in its wet habitats would become greatly diminished in winter (see Ch. 9.1.2). Thus, the aurochs is likely to have distinguished between its summer and its winter habitat. In summer, the emphasis may have been on grasses and graminoids in the wetter habitats; in winter, on bark, branches, tree leaves and bushes, as well as any edible remains of grasses and forbs in the drier, wooded habitats. Depending on the vegetation, the winter temperature, the geographical situation and the density of the population, the contrast between the summer and winter habitat will probably have been more or less marked. The marginal quantity of food in its winter habitat should be considered ‘emergency rations for survival’, but the food in its summer habitat provided the aurochs with most of the nutrients it needed. It is

· (East Poland). This forest Photo 24. Alder-ash forest (Circaeo-Alnetum) in the Forest of Bia§lowieza type occurs on wet, eutrophic soils, along small rivers. Periodic flooding may occur. (Photo: T. van Vuure).



hard to describe in detail, with reference to its winter habitat, which forest types the aurochs preferred, or how it competed with the European bison. Presumably, both preferred to stay in the more nutrientrich deciduous forests (see Ch. 9.1.2), but what each of them ate from which plant species and in what proportion cannot as yet be precisely ascertained. There is more information about the types of vegetation in the summer habitat of the aurochs. As was already explained, this habitat consisted of wetter vegetations, such as marshy forests and marshes. Marshy forests may include alder-ash forests along brooks and rivulets (see Photo 24) and willow forests along larger rivers. Such marsh forests occur on sand, clay and loam soils, on a basic soil that is solid enough to carry large animals like wild cattle. In addition to this, they are characterised by an often-lush vegetation of grasses and forbs (Borowski & Kossak 1972, Van der Werf 1991). The types of marsh forest mentioned here include some slightly wetter and drier varieties. Marsh forests that grow on peaty, non-mineral soils, such as many alder marsh forests, were

· Photo 25. Sedge marshes (Magnocaricion elatae) along the Le´sna River, in the Forest of Bia§lowieza (East Poland), square 602 (Photo: T. van Vuure). Such marsh vegetations probably played an important role in the ecology of the aurochs.

251


252


probably unsuitable, since the ground would have been too soft to carry bovine animals. Marshes included, first of all, salty coastal marshes, where the aurochs may have foraged on vegetations of salt plants in the tidal area. During certain periods in the prehistory of the Low Countries, there were very extensive tidal marshes along the coast. More inland there were other marshes, such as bogs and fens. The soil of the bogs, like the peat of the forests mentioned in the above, was too soft to carry bovine animals, which would have simply sunk down in it. Moreover, the scarcity of nutrients in bog vegetations is unlikely to have attracted the aurochs. Fens include many types of marshes that are under the direct influence of ground- or surface water and that are more or less treeless, due to the presence of water. The most relevant vegetations for this discussion of the aurochs are those that, like the ones just mentioned, occur on mineral, solid ground that heavy animals like wild cattle can walk on. They alone are likely to have been of use to them in their search for food. Fens, which probably belonged to the most important habitats of the aurochs, formerly occurred in Europe on a larger scale and in larger areas than today. They were increasingly forced back by drainage and cultivation. Rivers and marshes with a natural water regime that has not been influenced by man occur hardly, or not at all anymore in Europe. The groundwater used to be higher in many areas, and floods occurred more often. More or less extensive fen vegetations, dominated by sedges (Carex sp.; Association of Tall Sedges (Magnocaricion elatae)) occurred along brooks (formerly in Twente; Westhoff 1949) and small rivers (Drentse Aa; De Bruijn 1977, Reest; Dirkx et al. 1998) in the Netherlands, as well as abroad along larger rivers (Biebrza, Northeast Poland; Pa³czyñski 1984). They were nearly treeless due to their wet character. Nowadays, such sedge vegetations along small rivers may still be found in the Forest of Bia³owie¿a (see Photo 25). Large rivers, like the Missouri in North America (Weaver 1960) and the Pripyat in White Russia (Buskens et al. 1998), across large areas (hundreds of square kilometres) were accompanied by wet grass and sedge lands. Also in these areas, sedges dominate the treeless marshes. This is still evident in Wood Buffalo National Park (Central



Canada), where sedges figure large in the vegetations of the flood plains (see Photo 33) and form the main food of bison living there. Similar sedge marshes used to occur along the earlier Rhine, as may be seen in pollen diagrams from the vicinity of Wijk bij Duurstede (Holland) (see Fig. 31; Steenbeek 1990). Sedge marshes also existed along many rivers and in river deltas in the Low Countries at one time (see Fig. 41). The later sedge peat that originated from these may still be found in many places. The situation along the rivers at the time was totally different from today if one compares, for instance the river Rhine that has now been forced into a straightjacket of groynes and dikes. The said sedge marshes occur in

Fig. 41. The distribution of the various botanical types of peat formed during the Holocene in the northwest of the Province of Overijssel (Netherlands), which shows that locally, sedge vegetations could occupy extensive areas (after Pons 1992). Legend: 1. Sedge peat and reed-sedge peat 2. Sphagnum peat 3. Sedimentary peat 4. Open water 5. Peat brook and river courses (partly filled) 6. Boundary of the higher lying soils

253


254


very flat, low banks, which are flooded on a large scale at the slightest rise in the river water level. The flood plain of the Pripyat has a width of 8 kilometres, although the situation in this river valley can no longer be considered completely natural. In 1949, when it was still much more so, Kulczyñski described the flood plain of the Pripyat and its tributaries prior to the realisation of a large drainage plan. The sand and clay soils along the rivers, in combination with the layer of sedge roots, provide a solid basis that can carry heavy animals such as wild cattle. Thus, the marshes of the Zambezi delta in Mozambique are inhabited by thousands of African buffalo and even hundreds of elephants (Tinley 1974). The aurochs’ preference for river valleys as a summer habitat is still reflected in that of its domesticated descendants. During the growing season, present-day domestic cattle also prefer to live in riverside grasslands (Wallis de Vries 1994, Skovlin 1984, Roath & Krueger 1982a, 1982b, Pinchak et al. 1991, Goodman et al. 1989). The presence of water and the attractive food have been shown to be the main reasons for this preference. In relation to their surface areas, such grasslands are obviously much preferred over the drier and/or higher areas. Archaeological excavations may throw more light on the habitat preference of the aurochs. Though peaty soils may preserve animal bones well, the many bone finds of aurochs (especially of hunted ones) in peats can be illustrative of this preference in spite of this. Along the Tjonger River at Jardinga (Netherlands) a Mesolithic aurochs kill site was found at the very place the animals had been killed (Prummel et al. 2002, Prummel & Niekus 2005). The marshy vegetation of this valley was composed of alder (Alnus glutinosa) and sedges at the time (Baak et al. 2005, Bottema 2005). A comparable case was researched by Street (1999) and many more examples for Denmark and Germany are listed by Auler (1999). River valleys are also the habitat of an animal that may have been of great advantage to the aurochs: the beaver (Castor fiber). Both domestic cattle and beavers have a strong preference for the broad, flat basins of brooks and rivers that flow relatively slowly (Munther 1982). Lakeand river banks are the beaver’s habitat. It feeds on the marsh- and water plants, tree leaves and tree bark in these areas; the latter is mainly



Photo 26. Beaver meadow in North America in the beginning of the 20th century, which developed after the decline of the beaver dam and the emptying of the reservoir (after Dugmore 1914). Such meadows contain a large biomass of grasses and herbs that the aurochs in Europe probably used to take advantage of.

a winter food (Naiman et al. 1988). By regulating the water level, beavers are able to a certain extent to adapt their living area to their demands. They do this by constructing dams in brooks and rivers. These will make the water level upstream rise and result in shallow beaver ponds with marshy banks. In such areas, well protected against predators, beavers can find a lot of food. In rare cases, beaver dams in North America can reach a length of 80 m and a height of 5 m (Johnston 1994). Both for the construction of dams and lodges and for winter food, beavers have to fell trees. Along lake banks, fewer trees need to be felled than in river valleys, since no dams are built in the lakes. The impact upon the surrounding forest may vary quite considerably, depending on the length of the river, the number of beavers, their period of stay and the differences in the river level (Naiman et al. 1988; Johnston & Naiman 1987, 1990; Johnston 1994; Munther 1982). It will be greater

255


256


as the winter lasts longer, the beavers are more numerous, their stay more protracted and the decline of the river slighter. This is why the beavers in Louisiana (Southern United States) need far less timber than the beavers in Canada, where it is so much colder. The greater the impact on the forest, the more this is likely to have been advantageous to the aurochs in Europe at the time, since the thinning out and the destruction of pieces of forest must have provided more ground vegetation. Beavers used to occur both in the Great Wilderness (Mager 1941) and in the forests at Jaktorów (Twardowski & Kasperczyk 1992). The effects of beaver feeding not only include a decrease in the forest area along the banks, but also an increase of the area of marshland as a result of the rising water level. Research that Johnston (1994) has carried out in a part of Voyageurs National Park (Minnesota, United States) shows that through a fourfold increase of the number of beavers between 1940 and 1988, the area of wet grassland (mainly Calamagrostis canadensis) increased from 17 ha to 476 ha. The area of trees and bushes decreased by 50% in the zone that was influenced by beavers. Such effects are sometime found in pollen diagrams, as research by Garrison (1967) in Ohio (United States) shows (see Fig. 32). The felling of trees caused the proportion of forest to shrink, while that of marshland, especially sedges, increased. Such a transformation may work in favour of many species of fish, water birds and elk (see Photo 26). The amount of timber felled by beavers can sometimes be considerable. Elsewhere in Minnesota (United States), for example, it was ascertained that six beavers were able to ‘harvest’ nearly twice the quantity of vegetable biomass of a herd of ungulates in Serengeti (7,800 kg/ha/year versus 4,380 kg/ha/year; Johnston & Naiman 1990). Such an extraordinary effect is seen especially during the first years of establishment; it becomes less marked in later years. The effects of beaver feeding may thus vary greatly, both in their extent and their nature, also depending on the various other factors mentioned earlier. Another aspect of the influence of beavers is seen when these leave their living area, for whatever reason (e.g. lack of food). The beaver dam then falls into decay, the pond becomes empty and on the soil, which has become bare and often flat as a result of sand- and clay sediments,



a vegetation of grasses and sedges will develop (Naiman et al. 1988; Wilde et al. 1950; Dugmore 1914). Such an area is called a ‘beaver meadow’ (see Photo 26). In North America, such beaver meadows used to be eminently suited to settlers. The meadows, which often measured several hectares, were very suitable for pasturing cattle. The penetration of the North American primeval forests was highly facilitated in this way. In the beginning of the 20th century, old settlers spoke of the river valleys in which they lived as ‘beaver meadows’, a term that is reminiscent of the origin of these places (Ruedemann & Schoonmaker 1938). This same process is likely to have taken place in Europe. The ‘fields’ that are mentioned in the Route Descriptions for the Great Wilderness were mostly situated along rivers and at least part of them had presumably been effected by beavers (Mortensen 1938, page 23; Mager 1941, page 177, 1960, I, page 274). Not only man, but also aurochs probably took advantage of the activities of the beavers, which were very common in the Great Wilderness, among other places. At the aurochs kill site in the Tjonger valley at Jardinga (Netherlands) beaver was present at the time also (Prummel & Niekus 2005).

Seasonal migration To conclude that the aurochs possibly had a summer- and a winter habitat would be to suggest that these animals migrated in search of areas with better-quality food, both in spring and in autumn. Nothing has been passed down about such seasonal migrations, however. The last population in the Forest of Jaktorów was forced to stay inside the reserve. Animals that left the forest area were driven back and in winter, hay was brought to them inside the reserve. The European bison in the Forest of Bia³owie¿a are also kept inside the forest area with the aid of supplementary feeding in winter; seasonal migration is out of the question. European bison in the Caucasus, which are not provided with supplementary food, migrate to the valleys in autumn, and back again, higher into the mountains, in spring (Heptner & Naumov 1966). From the 19th and the early 20th century, they have

257


258


become known to cover often-long distances in this manner (Pfizenmayer 1929). The North American bison, which used to occur in great numbers, are known to have led rather a rambling life. Apart from the shifts the various subpopulations would engage in a southerly direction, and in spring to the north, during summer and winter extensive migrations into all kinds of directions could also be observed (Roe 1972). Some subpopulations hardly moved around at all, while others had migration routes that extended across hundreds of miles. The current bison population in Wood Buffalo National Park (Central Canada) has its summer- and winter residences right next to each other (Carbyn et al. 1993), or no more than 70 to 80 km apart at the most. With the help of the data about the habitat choices of the three grazers aurochs, European bison and wild horse, the occurrence of these three species may be clarified in a hypothetical diagram (see Fig. 42), in which that occurrence is related to the drier or wetter character of the various living areas, and the latter’s position above sea level.

Fig. 42. Hypothetical diagram representing the main occurrence in Europe of three large grazers, aurochs, European bison and wild horse, in relation to the height above sea level and the degree of moistness of the various rough vegetation types (Drawing: H.M. Klees).



Summary In this chapter, an attempt is made at reconstruction of the habitat of the aurochs, on the basis of archaeological data, old descriptions, comparison with related wild cattle species and the results of Chapters 9.1.1 and 9.1.2. Based on what was found in Ch. 9.1.1, the habitat of the aurochs must have been situated in a landscape that consisted of extensive forests alternated by several types of marshes. Bone finds and old descriptions suggest that there was a strong affinity between the aurochs and marshes and marshy forests, to which it owed its nickname ‘marsh walker’. River valleys, coastal salt marshes and other types of marshes are likely to have been among the aurochs’ favourite areas. For present wild cattle species, such as the forest buffalo in Africa and the wood buffalo in Canada, such areas, inside otherwise forested landscapes, were also preferred habitats. Apart from marshy forests, the aurochs is also likely to have lived in somewhat less wet forests. Between aurochs and European bison in the same area, a certain separation of habitats may well have existed, with aurochs in the somewhat wetter forests and bison in the somewhat drier ones, with some overlap. The hypothetical diagram in Fig. 42 represents the occurrence of the three large grazers, aurochs, European bison and wild horse.

9.2. Predation In its extensive distribution area, the aurochs had to contend with different species of predators. The occurrence of some of these has become known from descriptions and pictures, while that of some others derives from their presence inside the distribution area of the aurochs and from their choice of prey. Descriptions from Poland (Schneeberger in Gesner 1620; Œwiêcicki 1634) mention the wolf (Canis lupus) as a predator of the aurochs, probably the most important one there. The same was probably true almost

259


260


anywhere in Europe. Brown bear, lynx and possibly fox and wolverine will probably have been predators of aurochs calves in Europe, the first two also considered calves of the European bison in the Caucasus their prey (Heptner & Naumov 1966). In connection with aurochs, however, there is no written evidence about the bear and the lynx in. Yet another large predator occurred in the Balkans: the lion (Panthera leo). From the Middle East, lions are known to have hunted aurochs, and this is likely to have been the case in the Balkans, too. Their presence there has become known both from reports by writers such as Herodotus and Aristotle and from wall paintings, statues and bone finds in the area that stretches from Greece down to Hungary and Romania (Ninov 1999). The lion presumably became nearly extinct already in the 4th century BC in Greece and died in the 2nd century AD in Macedonia (Sallares 1991). The wolf and to a lesser degree the lion were the European predators; outside Europe, down to the Middle East, there was also the tiger. That the lion hunted the aurochs is shown in several pictures from Mesopotamia and Old Egypt (see Fig. 43), in which a lion may be seen attacking an aurochs (Von Lengerken 1955). This warrants the assumption that the aurochs had to contend with predators the size of a fox upward to brown bears and tigers, which would either eat calves or adult animals, depending on their own size. Reports that have been transmitted from the last living area of the aurochs, the forests near Jaktorów in Poland, mention predation. Schneeberger (in Gesner 1620) writes about the aurochs there: ‘They do not experience harm from the wolves, except when they, when they have just been born, wander around separately, because then they will be torn up by the wolves’. The last population of aurochs was guarded nearly constantly by hunters appointed by the Crown, which will definitely have prevented wolves having their way. The fact that Schneeberger mentions that the hunters had to go to the aurochs in the forest ‘every single day’ probably means that there was no supervision by night, and that wolves could get closer at these times. Schneebergers’ quote, in which he refers to the wolves, first of all shows that especially calves fell prey to wolves; moreover, adult aurochs apparently suffered little harm from wolves. Very young calves may



have been left lying in the forest to wait for their mother’s return, and have fallen prey to wolves as a result. Schneeberger writes about this: ‘When the time to give birth arrives, the cow will retire to the densest areas of the forest and remain there with the calf for about twenty days; as soon as she sees it has become stronger and jumps she will finally take it to the pasture, guarding it closely so that the calf is not caught by hunters or devoured by wolves. The hunters are unlikely to have been able to continually follow all the (roaming) bulls, since confrontations between aurochs and wolves took place now and then, as a description by Œwiêcicki (1634) shows: ‘More than once, a single huge aurochs bull has been seen to have gained the complete victory, at the cost of a number of wolves thrown on the ground and trampled to death’. The aurochs was probably not an animal that, like the horse, would seek safety in flight; rather, it would defend itself by butting with its horns and kicking with its legs. Such behaviour may still be seen in domestic cattle nowadays. 19th-century domestic cattle in Poland have been seen kicking their hind legs at wolves and standing in a circle to defend themselves (Pusch 1838). The first Heck cattle released in the wild at Rominten (former East Prussia), stood in a circle to defend their calves, which had been shot by the managers (Frevert 1957). This strategy has also been described in relation to the European bison in 19thcentury Bia³owie¿a, as has the protection of the calves against wolves by shielding them with their bodies during flight (Brincken 1826). Schaller (1967) reports that in India, gaur cows stood in a circle to defend their calf, which had been killed by a tiger. Whether ‘walking into the water’ was a strategy aurochs could use to shake off attackers cannot be said with certainty. Elk, which often occur in marshes, are definitely known to do this (Mech 1966, p.138). It so happens that for wolves, there is little sense in attacking an elk in deep water, since their hunting technique fails there. The 18 to 20 yearold aurochs bull that was hunted and hit by at least 9 arrows during the Boreal period in Denmark may have resorted to this strategy; it got into the water and perished there. In 1983, its complete skeleton was discovered in a former peat bog at Prejlerup (Aaris-Sørensen & Brinch Peters-

261


262


en 1986). Hartz & Winge (1906) described the find of the Vig bull, that died under the same circumstances. Which age groups of the aurochs were affected most by predation can possibly be concluded from the research of other large mammals. Many researches have revealed that especially young, old and diseased animals, the weaker individuals in general, therefore, are most likely to be killed or fall prey to predators (elk – Mech 1966, gaur – Schaller 1967). Of the population of bison in the Peace-Athabasca Delta in Wood Buffalo National Park (Central Canada), which was studied by Fuller (1966) during the sixties, wolves clearly killed three categories of animals in particular: young calves, old animals and those weakened by diseases or wounds. Healthy, adult animals, in the prime of their lives, are generally too alert, too fast or too strong to be killed by a predator. Schneeberger and Œwiêcicki seem to have agreed with this. Of the European bison that lived in the forests of Bia³owie¿a in the early 19th century, it was said that especially the young and ill animals were killed by wolves; by contrast, the healthy, strong animals did not suffer. In India, gaur calves in particular are killed by tigers, against few adult animals (Schaller 1967). The fighting spirit of the aurochs that has been so frequently praised by man, is still reflected to a certain extent by that of the Spanish fighting bulls. This became especially clear during the wild animals fights that were held in Spain up to the early 20th century, and in which large predators like lion, tiger or brown bear were baited against bulls (De Cossío 1943, Volume I). In these situations, although they were not quite natural, the bulls, which were between 4 and 5 years old, proved to be nearly always invincible. Besides predation by wolves, there were also other causes of death. In the bison population studied by Fuller, diseases like bovine tuberculosis formed a similarly important cause of death. In addition to this, many bison regularly perished by drowning. In the spring of 1958, nearly 500 bison drowned during a period of high water, and during the floods in the autumn and winter of 1960, some 3,000 bison died, a quarter of the population. According to Fuller, such regularly returning floods, and also diseases constitute much more important causes of bison death than predation by wolves. Cases of mass drowning were also described with regard to the bison still living on the prairies in the 19th century. While crossing



Fig. 43. Picture of a lion attacking an aurochs (Middle East, 2500 BC; after Von Lengerken 1955). In the Balkans, North Africa and the Middle East the lion was one of the predators of the aurochs.

rivers or stepping on unreliable ice in spring, each year large numbers of animals lost their lives through drowning (Roe 1972, p. 160 ff.). According to Jêdrzejewska & Jêdrzejewski (1998), research into the causes of fluctuations in the number of European bison during the past two hundred years in the forests near Bia³owie¿a has shown that the average annual temperature and the competition for food (with red deer) are the most important mortality factors. The amount of food that is available regulates the number of animals, therefore; not predation by wolves. The same is true for the elk and wild boar in the same area. For red deer and roe deer, the impact of predation by wolves and lynxes, respectively, is much larger than for the other ungulates. In these cases, predation is at least as important as the food factor. Comparable results were found by Messier (1991) for the elk in Isle Royale National Park (Michigan, United States) and the whitetail deer in the National Superior Forest (Minnesota, United States). There as well, the number of elk is regulated mainly by the amount of food; that of whitetail deer by both the food and predation by wolves. In Yellowstone National Park both wolves and the amount of food regulate wapiti deer (Ripple et al. 2001, Larsen & Ripple 2003, Ripple & Beschta 2005).

263


264


The limited aurochs information that is available suggests that what is true for the other wild cattle species also held for the aurochs. The remarks by Schneeberger and Œwiêcicki and the notes in the Inspection Reports (see Ch. 5.2.1 and 5.2.2) indicate little impact of predation by wolves and a dominant impact of the food factor. This latter may show from the abnormally great mortality among aurochs in severe winters (1556/ 1557 and 1598/1599) and the detrimental effects for the aurochs of the competition for food with the cattle, horses and pigs local farmers herded in the same area.

Summary With the help of early descriptions and pictures, it could be ascertained which animals considered aurochs as their prey. In Europe, this was mainly the wolf. This chapter explains how the aurochs are likely to have defended themselves against their predators. In the same way, early descriptions as well as comparison with wild bovine species still alive today allow the deduction that of this type of animal, the calves and old animals in particular fell prey to predators. Beside predation, other natural mortality factors such as diseases, death by drowning and malnutrition play a role in regulating the numbers of these animals. Various studies have shown that the amount of food that is available is the most important mortality factor among large ungulate animals, including bovine species. Written data from the 16th century suggest that this was the same for aurochs.

9.3. Social structure and reproduction Although there are only very few data about the social structure and reproduction of the aurochs, they may just suffice to provide us with a global impression of these aspects. In comparison with related Bovini members and with wild or feral cattle, this impression may add to, as well as elucidate matters.



Like so many other data, these stem from the last aurochs population at Jaktorów. About the group structure of the aurochs that lived there, Schneeberger reported: ‘In winter they run in herds and they rest in one place, quite like some army’. This may indicate the situation during the winter feeding period, when the animals were given supplementary hay. This probably happened at a central place; the population of aurochs, over 50 animals at the time according to Schneeberger, would concentrate there. The same phenomenon also occurs nowadays in the forests at Bia³owie¿a, during the winter feeding period of the European bison, when groups of about 100 animals will live together without showing any great aggressiveness (Pucek 1986). During the summer, they are in much smaller groups of 2 to 40 animals, or wander around alone. The size of a group of aurochs was also very different in summer than in winter: ‘In summer, however, they roam the wilderness separately47 and if one has strayed too far and not shown himself for a few days (because there are hunters that have to go to them every single day) he will be sought with very great diligence and forced by dogs to return’. Although this text is not unambiguous about who roam around solitarily, the difference with the winter situation does show. To be able to monitor the animals adequately, they were obviously not allowed to wander off too far. Perhaps to prevent them from leaving the forest area and grazing the farmland and pastures, they were retrieved with the help of dogs. The 1564 Inspection Report notes that not the animals wandered around separately during the summer (£ukaszewicz 1952): ‘In the forests at Jaktorów and Wiskitki, where the aurochs hide, during our travels we found a herd of 30 animals, including 22 old cows, 3 young bulls, 5 young calves. We were unable to find old bulls, since these had spread out in the forest on account of the thunder, but the hunters had told us that there were 8 of these old bulls’. This report must have been written during the summer (July or August), since ‘5 young calves’ are mentioned, as well as ‘a very old thin cow, which will have trouble getting through the winter’, further on in the report. 47 Since the word ‘singuli’ gives the impression that all the animals roam the forest separately in summer (contrary to what is to follow), Schneeberger must have either misunderstood his source, or have meant to indicate only the bulls.

265


266


The situation described in this Inspection Report is very similar to the group structure described for the European bison (Krasiñski 1978), the gaur (Schaller 1967) and the banteng (Halder 1976). As it happens, the social structure among those related bovine species consists of mixed groups of cows, calves and young bulls, separate from which the old(er) bulls roam around alone or in small groups. That the thunder had motivated the adult bulls to spread out through the forest is an argument that may have been brought forward as an excuse for not knowing where the bulls were, when this ought to have been known. There is no reason why the bulls would have been scared of the thunder while the other animals were not. The general idea about both the bovine species that live in the forest (European bison, gaur) and those that live in the steppes (bison, African buffalo) is that the bulls, as they grow older, become increasingly isolated. This may well have to do with a conscious avoidance of confrontations with other bulls. In larger groups of bulls, there are bound to be more unrest, mutual competition and fights, which would take a lot of time and energy of the bulls (Hoekstra & Vulink 1994). Such problems are less complex and easier to solve in smaller groups. Usually, only the old bulls live alone. Old banteng bulls, for example, are very rogue animals (Halder 1976). Among the bovine species that live in the forests, such as gaur, banteng and European bison, the mixed groups may include up to a few dozen animals; the groups of bulls are usually much smaller (Schaller 1967, Halder 1976, Pucek 1986): 2 to 6 gaur, 2 to 7 European bison. The 1- and 2-year old European bison bulls still live in the mixed groups; the 3- to 5-year olds may be found either in the mixed groups or in separate groups composed of bulls only. Older bulls either live alone (62%) or in groups of 2 or 3 (Krasiñski 1995). In the Oostvaardersplassen, the size of the groups of bulls is generally five; the herd that was studied only had one bull that lived alone (Hoekstra & Vulink 1994). The composition of both the mixed groups and the groups of bulls may change regularly, although the mixed groups usually have a fixed core of cows and calves, in particular. Unlike Bia³owie¿a, where the European bison receive supplementary food in winter, the group structure of these animals in the Caucasus,



where they do not, is virtually the same in the winter period as in the summer (Pucek 1986). Changes in the food situation apparently do not prompt any great changes in the group structure. Directly related to the group structure of bovine species is the remarkable difference in size (sexual dimorphism) between bulls and cows. A marked sexual dimorphism is characteristic for animals that live in groups part of the year, and alone the rest of the time (Guthrie 1990). Apart from bovine species, this is also a known fact about red deer and wild boar, among other animals. Animals that live in mixed herds all year round, such as the Przewalski horse (Equus przewalskii) and the oryx (Oryx beisa) show little sexual dimorphism. Bulls that live alone or in small groups do not just avoid a lot of stress in this way, but also have more food at their disposal (Prins 1987, Guthrie 1990). The greater risk of predation, in comparison with living in herds, is ignored in favour of this advantage. Around the mating season, a number of changes in the structure of the group take place among bovine animals that run wild; the same is likely to have been true of the aurochs. Both Œwiêcicki and Schneeberger report that aurochs bulls would mate with domestic cows in this period. Œwiêcicki describes how aurochs bulls, attracted by their smells, would go to the herds of domestic cows to lure away cows in heat to mate with. In the same way, aurochs bulls must have gone to the mixed (aurochs) groups to find the cows in heat, the way related bovine species still do. In the forests at Bia³owie¿a, the European bison bulls start to get restless around the start of the rutting season, which is in August and September there, and they begin to make more and longer tours. The groups of bulls dissolve gradually, and more and more bulls join the mixed groups (Krasiñski 1978). Not all the bulls join the cows in the growing mixed groups, however, so groups of bulls and rogue bulls may still be seen also during the mating season. Among the gaur in India and the banteng at Java, the groups of bulls also dissolve, and the bulls go to the mixed herds during the rutting season; rogue bulls may still be found during that season also here, however (Schaller 1967, Halder 1976). Among the Heck cattle in the Oostvaardersplassen and the feral white cattle of Chillingham Park (Northumberland, England), there are groups

267


268


of bulls in specific restricted living areas (home ranges) (Hall 1988, Hoekstra & Vulink 1994). In the Oostvaardersplassen, the mixed groups travel through these home ranges; any cows in heat are covered by the bulls that have remained there. Another cattle population that lives in a feral state in Europe is that of the Doñana National Park in South Spain (Lazo 1995). About 140 animals older than 1 year old live there in an area of 6,700 hectares, without supplementary feeding. Annually, 30 to 50% of the calves are harvested. The mixed groups there are very stable and have virtually separate living areas. The living areas of the oldest, dominant bulls hardly overlap; those of the less dominant, younger bulls show a lot of overlap. As with the gaur, the banteng and the European bison, prior to the mating season, the adult bulls start to get restless; they extend their living area by making longer tours, which bring them in touch with the mixed groups and with other bulls. During the mating season, these bulls live among the mixed groups; after this period, they once again retreat to their own living areas, separate from the mixed groups. The difference between the Oostvaardersplassen and the Doñana National Park, as regards the behaviour of the various social groups, may be related to the size of the area the animals have at their disposal. The 180 head of cattle in the study by Hoekstra & Vulink (1994) were in an area of 600 hectares. The individual home ranges of the cattle in the Doñana National Park varied from 160-2,500 ha (Lazo 1995). The average home ranges of 4- to 11-year old European bison bulls in the forests at Bia³owie¿a are around 2,288 hectares on average during the vegetation period; those of the older bulls measure about 1,406 hectares. For the gaur in the Kanha reserve (India), Schaller (1967) reports individual living areas of 6,000 hectares; for the Malaysian gaur, Conry (1989) mentions areas as large as 7,000 hectares. Both the gaur and the European bison have unlimited freedom to move about, as well as a choice of the various habitats that may be of greater or lesser importance to them, depending on the season. Obviously, the amount of food that is available also plays a part. In areas with plenty of grasses and graminoids, such as the steppes and the sedge marshes, much smaller living spaces will suffice for the bovine animals than in densely forested areas (see Ch. 9.4).



Photo 27. Scottish Highland cow with calf in the Imbos Forest (Central Netherlands) (Photo: J.M. Gleichman).

Schneeberger reports about reproduction: ‘In September they mate, while burning with lust, have numerous and very fierce fights and sometimes both fall down dead. But the one animal that they have noticed to be stronger, and constantly provoking others to fights, hunters will hunt under orders of the king, in a way that will be explained below. But they also hunt those that have been spotted covering domestic cows; for cows will become pregnant by them, but will either miscarry or give birth to non-viable calves. Finally, they give birth in May, some in September like domestic cows; this seldom happens, however, and the calves that are born in the autumn are weak and do not stay alive because of the fierce winter cold. When the time to give birth arrives, the cow will retire to the densest areas of the forest and remain there with the calf for about twenty days; as soon as she sees it has become stronger and jumps she will finally take it to the pasture,

269


270


guarding it closely so that the calf is not caught by hunters or devoured by wolves’ The months mentioned here by Schneeberger, September for mating and May for birthing, are unlikely to be correct. If conception was in September and birth in May, gestation could only have lasted 8 months. Since gestation in both domestic cows and related bovine species such as gaur (Schaller 1967) and European bison (Krasiñski & Raczyñski 1967) is about 9 months, a mating season in the months of August and September, with a birthing season in May and June, is more likely. In this way, the mating- and birthing seasons of the aurochs must have coincided with those of the European bison, who still realises his reproduction in the same period (Krasiñski 1978). Since, according to Schneeberger, the aurochs incidentally had its young later (in September), cows that had not become pregnant earlier, or young cows, could apparently sometimes get in heat (again) some 3 or 4 months after the regular mating season. It is likely, therefore, that reproduction of the aurochs, as of the European bison, took place during limited, fixed periods of the year. The calves, which were born in May or June, grew up in the period in which the grass grew most plentiful, which offered the best chances for survival. The reproduction of animals that live in captivity is not limited to fixed periods, but may take place throughout the year. Among European bison in captivity, births took place pretty much throughout the year (Krasiñski & Raczyñski 1967), and among the Chillingham cattle that are provided with supplementary food in winter, births are seen all through the year as well (Hall 1988). During the Imbos experiment with Scottish Highland cattle, which were not given any extra food in winter, births in the first year still took place in the period from April to September; after five years, however, they were concentrated mainly in the months of April, May and June (Van Wieren 1988) (see Photo 27). In the Doñana National Park (Spain), which has a milder climate, the mating season comprises June, July and August; births take place mainly in the period from February to August, though most occur in March, April and May (Lazo 1992). During the mating season, the bulls are more restless and adventurous than at other times, and behave to impress, by beating their horns against the bushes or into the ground (Schaller 1967, Schloeth 1961).



The throwing apart of mown corn by the aurochs in August, which was mentioned in Ch. 9.1.2, is likely to have been another expression of this behaviour. According to Schneeberger, aurochs bulls would fight together during the mating season, sometimes to the death. Schaller (1967) reports that this does not take place among gaur, and Hubback (1938) reports only a single case of a gaur bull having been killed. Schloeth (1961), who did research among the Camargue cattle, notes that this happened very incidentally.

Summary The information obtained from early descriptions and through comparison with wild cattle species still in existence suggests that the social structure among aurochs was similar to that of related bovine species. This social structure consisted of mixed groups of cows, calves and young bulls beside smaller groups of older bulls and single old bulls. The mating season and, consequently, also the season in which calves were born, were limited to certain periods of the year. The rutting season in Poland was in late summer (presumably August, September); calves were born in late spring (presumably May, June). Just before and during the mating season, the grown bulls would go to the mixed groups to mate with the cows there.

9.4. The impact of large herbivores on the forest growth

Introduction This chapter will try to cast light on the possible effect of large herbivores on the growth of trees and forests, and thus on the appearance of the landscape, through their consumption of branches, leaves and bark. The concept ‘large herbivores’ here does not only include the aurochs, but several other animals; all species from beaver to elephant, in fact,

271


272


which eat off trees incidentally or regularly, and thus have an impact on their growth. The search for examples and studies regarding this subject clearly is not limited to the aurochs or the areas in which it used to live; it extends into areas like North America and Central Africa. Interactions between large herbivores and trees may be seen in many areas on earth; when looking at such interactions it is important to assess man’s possibly dominant role either today or in the past. Only the long-term absence of all human interference, whether in the shape of hunting, supplementary feeding, cutting, burning, fighting predators, or other activities, can present a clear image of the real impact of large herbivores on the growth of the trees. The image of the natural Holocene landscape of Central Europe, which emerged from Ch. 9.1.1 as consisting of ‘forests and marshes’ can hardly be found in its natural, unharmed state any longer. Apart from a

· (East Poland) in the beginning of the 19th Fig. 44. Map of the central part of the Forest of Bia§lowieza · The forest vegetation is century (after Brincken 1826). In the lower centre the village of Bia§lowieza. rendered as a type of savannah with solitary trees. However, Brincken describes the forest area as one large continuous forest. The artist apparently depicted the forest landscape in his own artistic way. The dark vertical strip in the middle is a copying error.



few small, remote areas, where man with his agriculture and forestry has barely infringed on the natural processes, it is fair to say that everywhere on earth, the vegetation, the soil and the water balance have been tampered with. The Great Wilderness in the earlier East Prussia, the final large area to be called a ‘complete wilderness’, at least for the larger part, stopped existing as such around 1500, when the aurochs became extinct there. From the beginning of the 16th century its decline progressed, since it presented an obstacle to man at the time in every way. No single area in Central Europe deserved the epithet ‘complete wilderness’ from that period onwards, although some areas long afterwards retained the image of such a wilderness, in spite of their lack of a number of the original animal species. The forests at Bia³owie¿a constitute such an area, parts of which kept their appearance of an original wilderness for a long time. The descriptions of these forests that stem from the early 19th century, when the provision of the European bison population of over 700 animals with supplementary food had only just begun, evoke a picture that may still be seen reflected in many places in the area today. Although in that period, the original natural situation was clearly a thing of the past, a discussion of it may still prove useful to show what these forests, which hosted such a large population of European bison, really looked like at the time, and to dispel the misunderstandings about it, such as those caused by Vera (2000). Each winter, every European bison had about half a cartload of hay (‘charge de deux chevaux’; Brincken 1826, p. 108) at its disposal, which Brincken considered not a lot for a European bison. In addition to the hay, the animals would eat the branches and bark of deciduous trees and shrubs in winter (Brincken 1826, p. 57). Bia³owie¿a and the villages Teremiski and Pogorzelce were a lot smaller at the time than they are now. From these villages and the villages at the edge of the forest area, cows were pastured in the forest, but only in the fringe area, up to a certain boundary (Brincken 1826, p. 12). The same pattern of pasturing took place on a larger scale until way into the 20th century (Faliñski 1968, p. 203); it mainly occurred within 20 m from the forest road, not deep in the forest (author’s observation, 1972). Brincken, commissioned

273


274


by the Russian tsar, wrote a detailed description of the plant- and animal world of this forest. He describes the forests as follows: ‘the large forest……………constitutes an uninterrupted stretch of 52 km in length and 45 km in width, and 187 km around’ (p. 3). ‘Situated in the centre of the Imperial Forest, this village [Bia³owie¿a] is the first open area one encounters after half a day’s journey through the dark forests’ (p. 10). On page 18, he gives some more details about the appearance of the forest and its functioning: ‘The large forest that we have tried to sketch offers a vegetation on which (as has already been indicated in more general terms) the culture of Europe, with its attendant forestry economy, has not yet had the least effect. Everywhere around, the various species of indigenous trees can be seen growing in their specific stands, flourishing naturally, growing old, and falling over to make space for their successors. One will fail to find forests kept neat with the use of axes, dense regeneration or neatly ordered young trees, the things that are a forester’s pride elsewhere in the world. Indeed, a forest of this great size, growing under hard natural circumstances, has not only become rare in Europe, but also offers an opportunity for many interesting observations………………; the forester is interested to see how nature manages the forests without any influence by man, how it sows, grows and destroys and how the remains of dead trees form the beginnings of new generations’. Brincken describes the forest with the species of trees that can still be found there today, such as oak, linden, Scots pine and Norway spruce, alder, hornbeam, aspen, and, as one of the most common shrubs, hazel. There are no grassy areas or open, park-like landscapes. He describes the diverse character of the forest and the difficulties one may encounter passing through it. Locally, sometimes it is impossible to get through (pp. 29 and 30): ‘These forests are always massive and dense, and you can penetrate them only with difficulty. Besides the obstacles formed by the multiplicity of bushes you encounter thousands of obstructions formed by large tree trunks that have fallen and become overgrown by thorny brambles’. ‘The forest district of Nieznanów [in the southwest of the forest] looks as imposing as it is unpleasant, because of the enormous numbers of trees



that have been uprooted, which turn the forest into a truly inaccessible, and therefore unknown, area48. Crisscross through each other, as on a large battlefield, lie enormous numbers of uprooted and rotting trees, on which a new generation of trees already emerges; large numbers of young bushes grow between the dead and moss-covered branches of oak, poplar, linden and Norway spruce and thus grow on their remains; the kingdom of death and that of life are close together, connected, intermingled’. The map in Brincken’s book bears no relation to the reality of the forest he describes (see Fig. 44). It shows a kind of savannah landscape, with many single trees; entirely different from what Eichwald (1830) (also shown in Vera 2000, p. 272) printed. Apparently, it was left up to the artist to picture the forest in his own way, which resulted in an

· National Park (East Poland), square 256. This pollen record Fig. 45. Pollen diagram from Bia§lowieza roughly covers the last 800 years. During this period the area was densely forested, given the (darkcoloured) proportion of trees and shrubs in the diagram (after Mitchell & Cole 1998).

48

The name of the forest district, ‘Nieznanów’, derives from the word ‘ nieznany’, which means ‘unknown’.

275


276


artistic, rather than realistic depiction of the forest vegetation that was actually present.

Areas comparable to Europe Up to now, Europe and its natural Holocene landscape have been the focus of attention, since Europe belongs to the distribution area of the aurochs of which the most data have survived. To know the impact of large herbivores on forest vegetations, a central issue in the ecology of the aurochs, it is also interesting to ascertain which areas, comparable in climate and vegetation to Europe, may provide additional data about these aspects. The east of the United States and the southeast of Siberia are examples of such areas. Their climate is comparable to that of Central Europe, and they have many tree genera in common. The circumstances in both areas, as in Central Europe, may be considered favourable for tree growth. In addition to this, descriptions of the early appearance of the landscape of both areas have survived, relating to a period in which man exerted hardly any influence, and non on any large scale. In the period described, moreover, these areas still possessed their original range of large herbivores and their predators. During the Holocene period, the east of the United States from the Atlantic coast to the Mississippi basin naturally hosted four large herbivores: the whitetail deer (Odocoileus virginianus), the wapiti (Cervus elaphus), the elk (Alces alces) and the bison (Bison bison)49. Numerous descriptions of this area from the 16th century onwards mention its densely forested character (Roe 1972, pp. 842 - 852): ‘The thicknesse of the wood and greatnesse of Forrests doe hinder the sun from warming of the ground…’ ‘…a land of woods and bogs’. ‘…the whole country being but an interminable forest’. ‘The vast forest, which everywhere covers the country, and which in many places is impenetrable…’ ‘From Maine to Alabama, the woods were unbroken and impassable. The great Appalachian forest was in primitive days an exceedingly 49

In pre-Columbian times already bison were present in the eastern United States (Tankersley 1986).



dense tangle. At a few points the aborigines had worn narrow footways through it; but these trails were not adapted to pack-animals, the original means of transportation brought by the Europeans, but for the use of men who journeyed on foot, and could thus climb steeps inaccessible to a burdened beast…. The undergrowth of this forest country is far more dense than that which is commonly found in European lands’. ‘In the primeval forests it did not take long for a road to become impassable, if unused. Braddock’s Road over the Alleghanies, cut in 1755, was impassable in 1758… The Old Portage Road, cut in 1749, was cut out again in 1752’. Zoller & Haas (1995) mention similar descriptions of the area from many other sources, like: (Pennsylvania, 1743) ‘We observed the tops of the trees to be so close to one another for many miles together, that there is no seeing which way the clouds drive nor which way the wind sets, and it seems almost as if the sun had never shone on the ground since the creation’. (New Jersey and Pennsylvania, 1805): ‘Progress was painfully slow through an overshading forest’. Apart from these descriptions of forests that have often remained untouched, reports may also be found about local Indian tribes that changed the forest landscape with a view to hunting (Roe 1972, p. 850 ff.; Bonnicksen 2000, p. 267, 268). Using fire to create often-large open areas in the forest in order to attract wild animals and ensure a larger catch. In this way, Indians also played a large role in the continued existence of, for example, oak-chestnut forests, where maple and beech would otherwise dominate. The practice of burning was used throughout North America, and will be discussed in more detail in connection with the history of Wood Buffalo National Park (Canada)50. From the southeast of Siberia, the Ussuri area to be exact, descriptions have been passed down of a period in which man had only barely left his mark on the landscape there. The descriptions stem from Arsen50 On the basis of extensive sources research, Roe (1972, pp. 228–256) concludes that the numbers of bison in the forests east of the Mississippi valley must have been relatively small, and in marked contrast to the millions of bison that inhabited the prairies. Which areas of the forest the bison inhabited has as yet been the subject of little research. The records mentioned by Roe show that the larger part of the observations of bison stems from areas along rivers. Whenever they were seen in forests, their presence was strongly related to the occurence of salt licks, places where salt water comes to the surface. Such salt licks were frequented by bison and deer that would come from afar along the broad paths that had been trodden down in the forests, to drink salt water and eat salt soil. Like the bison in Wood Buffalo National Park, the early bison in the easterly United States seem to have been closely tied to the rivers

277


278


Photo 28. Forested hills in the Ussuri region (East Siberia) (Photo: J. Shibnev). Today, this forest landscape is still largely the same as the explorer Arsenjew (1924) found and described it in the beginning of the 20th century.

jew, who undertook expeditions through the Ussuri-area and SichoteAlin in 1902 and 1906, and included the plant- and animal world in his report (Arsenjew 1924). This area was still basically untouched at the time. Incidentally, hunters and fishermen would pass through it; they would stay in primitive shelters, as also used to be done in the Great Wilderness around the end of the Middle Ages. The forests consisted of deciduous as well as coniferous trees, and there were five species of large herbivores, the sika deer (Cervus nippon), red deer (Cervus elaphus), elk (Alces alces), roe deer (Capreolus pygargus) and the Siberian musk deer (Moschus moschiferus). The beaver (Castor fiber) was not naturally present here51. Of the large predators, the Siberian tiger (Panthera tigris) was, and still is, the best known. Arsenjew wrote the following description of the landscape he passed through on his long journeys (Part I, p. 126): 51 No indications were found of the presence of the beaver in these areas. Its distribution area does not begin until the Baikal region; from there it extends into Europe (Djoshkin & Safonow 1972).



‘The word ‘taiga’ used by the Siberian natives indicates the ‘virgin wilderness, in which there are no roads’. In contrast to the taiga of Central Siberia, which consists almost exclusively of coniferous forests and passes into the tundra in the North, the taiga at the Amur and the Ussuri usually consists of mixed and deciduous forest. Extensive areas are covered with these forests, usually still completely in their original state and never trodden by human feet. From the river valleys, forests stretch out across extensive plains, hills and mountainous areas up to the mountainsides. Beautiful timber forests, often consisting of veritable forest giants, are alternated by all kinds of marshes. Large quantities of tree trunks uprooted by storm, pieces of forest where fire has raged, full of halfcharred stumps overgrown by young wild trees together with the dense shrubbery form often completely impenetrable areas. All kinds of wild animals, the larger kind mainly represented by bears and tigers, find their impenetrable hiding places here. Only the narrow trails that the wild animals use, which are clearly trodden down under the twigs of the shrubs and the trees, may be seen like tunnels through the shrubbery’. Part I, pp. 129/130: ‘Sometimes, one suddenly sees an open area in the forest. The inexperienced forest hiker tends to go to such places, where he ends up in a mass of fallen trees. Large areas of the forest have been destroyed here by whirlwinds; the uprooted tree trunks lie crisscross through each other, often forming large obstacles. An open area in the forest indicates a marsh, a burning spot or trees uprooted by the wind. Such areas cannot always be avoided. If the trees that have fallen over are not too thick, they are removed with the help of axes; if a large tree blocks the road, however, it is hewn flat on the top and the sides, so that the horses may get over it with their packs. Naturally, this delays the transport quite a bit, so it is slow going to take horses through the taiga’. If these descriptions are compared to those from the earlier Europe, the similarity in terms of the problems people had to face in these jungles is remarkable. Particularly the leading along of pack animals, as was also done in the Great Wilderness, caused delays and turned such journeys into marathon sessions. Apparently, enough grass could be

279


280


found in the jungle for the few horses Arsenjew led along, since there were definitely no large grassy areas. These trips were also made with the help of a guide, who knew the terrain and was able to survive in the jungle. Arsenjew did not only record his trips in writing, but also on photographs. These provide a convincing rendition of his experiences in spite of their poor quality. Nowadays still, the Ussuri-area has retained a lot of its original character (see Photo 28).

Population density of wild cattle Information about the population density of the aurochs in their final living areas is scarce, and there is none at all about the Great Wilderness. Aurochs are known to have lived there but nothing was passed down about their numbers. About the situation in the forests at Jaktorów there are more data, as well as statistics for the final 70 years of the aurochs’ stay there. From his journeys to Poland halfway though the 16th century Von Herberstein (1557b) learnt that the number of aurochs, then still only living in Masovia, was ‘not large’. Schneeberger was the first to report their number, as ‘again over 50’, probably in 1559. The fact that he described it in this way probably indicates that the number of aurochs, as a result of the severe winter of 1556/1557, had been less in 1557, possibly less than 50. ‘Over 50’ is the largest number ever reported. Schneeberger’s report also shows that the population at the time was still able to recover from a period of decline. As far as is known, this proved no longer possible in the subsequent period. After this mention by Schneeberger, the number of aurochs decreased steadily. Around the year 1600 the population was dealt the finishing blow; only a few animals remained. The total forest area, i.e. the forests of Bolemów, Wiskitki and Jaktorów, measured about 25,000 ha. If the number of aurochs in Schneeberger’s time may be considered a normal, traditionally present number, the normal density would amount to approximately one aurochs on 500 ha. Not all parts of the area may have been equally suitable for foraging, so the actual, usual density is bound to have been higher. The



same is true, for example, of the European bison at Bia³owie¿a. There are about 250 European bison on the Polish 58,000 hectares of this forest area, which amounts to 1 animal on every 230 ha. Naturally, this is very global, since the number is regulated by man and affected by the winter provision of supplementary food and the summer ‘supplement’ in the shape of man-made pastures, clear-cut areas and verges. An additional part is played by the competitors for food, such as red deer. In the beginning of the 19th century, the number of red deer was very small here (Brincken 1826); in the beginning of the 20th century there were about 5,000 (Krasiñski 1978). Based on data about the past two hundred years, Jêdrzejewska & Jêdrzejewski (1998) conclude that the density of the European bison, for whose number food constitutes the main regulating factor, varied from 0.4 animals/ km2, with many red deer, to 1.5 animals/ km2, with few red deer52. By way of comparison, the density of European bison in the Caucasus (North Ossetia), which are not provided with supplementary food in winter, is 1 on every 60 ha (Kazmin & Smirnov 1992). About 12% of the (forest)area, in which the European bison live, consists of pastures and clear-cut areas, on which they obtain half of their food. Just before World War I, when the European bison lived in the poorly accessible forests of the central- and high mountains of the northwest Caucasus, an area of 524,700 ha, their population still numbered about 600 animals (Pfizenmayer 1929), which came to a single European bison on approximately 875 hectares. At the time, supplementary food was not provided either, predation occurred and poaching was discouraged. Grand Duke Michailowitsch, who was the single possessor of hunting rights in the area, seldom killed European bison. After the Russian Revolution in 1917, massive poaching obliterated this population of European bison in the twenties. For the gaur in the forested Kanha-reserve (India), a quarter of which consists of man-made pastures, Schaller (1967) noted a density of 1 animal per 100 hectares. Conry (1989) reported the density of the gaur in the Lepar River area (Malaysia). This area measures 68,000 hectares, 38% of which is secondary forest, grown after felling, and 23% agricultural land. These habitats are interesting for the gaur, unlike the prima52

The number of 1.5 European bison per km2 may be based on too high an estimate (Büchner 1895, Szalay 1938: page 75-76, Pucek 1991). However, Krasińska & Krasiński (2004) contradict this.

281


282


ry forests that cover the remainder of the area. Around 80 gaur live in this area on average, which amounts to about 1 animal per 850 hectares. In the Tai forest (Ivory Coast), the density of the forest buffalo varies depending on the degree of forestation (Hoppe-Dominik 1988). It is lowest in the primary forest, 1/500 ha, and highest in the large access zones, which have been cleared of trees by man and are overgrown with grasses, i.e. 1/15 ha. The above survey shows that the population density of large, herbivorous bovine animals is connected strongly to the openness of, i.e. the lack of trees in, a specific habitat. The more grass there is, the higher the density. This also shows the large role played by man, whose activities create grassy clear-cut areas, access roads, agricultural ground and secondary forests in areas that were almost entirely closed forests before he arrived. The number of bovine animals per surface area rises as a result. Habitats that were naturally open, such as steppes, river valleys and marshes, have the highest densities. The North-American prairies, which measured about 400 million hectares (Joern & Keeler 1995), used to be inhabited by approximately 30 million (McHugh 1972) to 40 million (Roe 1972) bison; a density of 1 animal on every 10 to 13 hectares. This density differs from that in Wood Buffalo National Park (Canada). Originally, a quarter of the Peace-Athabasca Delta there, some 367,000 ha, were suited as a foraging area for the bison, 75 % of whom use the area during some period of the year (Carbyn et al. 1993); the density in the Delta alone that may be calculated on the basis of this comes to 1 bison on 90 to 120 ha53. In comparison, the densities of the herbivores in the Oostvaardersplassen (in the year 2004) could be called very high, with ca. 3000 large herbivores on some 2,600 ha (J. Griekspoor pers. comm.), This may be possible because these are new, very nutrient-rich soils; moreover, in very cold winters, supplementary food was given. Bison are able to survive on exclusively grasses and sedges throughout the year, whereas the winter diet of Heck cattle mainly consists of grasses. It is uncertain whether the density of the aurochs at Jaktorów, 1/500 ha in 1559, was normal, or whether this was an already strongly declining population. Compared to other (present) bovine species that live in forests it does not seem unusual. According to the Inspection Reports, 53 Calculated for the entire Wood Buffalo National Park and based on bison numbers by Carbyn et al. (1993), Bradley (2004) and Bradley et al. (2004), bison density is 1 animal on 700-800 ha.



domestic cattle and horses were grazing the forests in which the aurochs grazed as well as early as 1564. The fact that many animals died during the severer winters (1556/1557 and 1598/1599) seems to indicate that as early as 1557, the health of the aurochs population was affected by the food competition with the domestic cattle during the summer. However, in view of the fact that it is not certain if the grasslands were all natural, or partly man-made, it remains difficult to ascertain the (natural) density of the aurochs there. High mortality during severe winters is not abnormal for a population of animals, however. As the hunting pressure exercised by the Polish kings was probably quite mild, and predation barely played a role, the aurochs population is likely to have been in accordance with the amount of food available; this latter was partly determined by man. In places where the aurochs used to live, greatly varying numbers of this animal’s bones have been found in the course of archaeological excavations. These varying numbers can be explained partly by the degree of conservation at the sites in question: bones will waste faster in dry or acid soils, but are well preserved in wet, oxygen-poor and/or calcareous soil. Another explanation may be found in the varying aurochs numbers that occurred in various places; finding few bones might indicate a slight occurrence and the other way around. Furthermore, the explanation may lie in the need or preference of earlier man to hunt this very animal. Though the number of bones found is not necessarily related directly to the degree of occurrence, the relative number of bones may constitute a measure for the former population density of an animal. The former low density of the North American bison east of the Mississippi River, compared to that of the Great Plains bison, is reflected in the numbers of bones found, for example. West of this river many bones were found, and to the east of it only a few (Tankersley 1986): this picture concurs with eyewitness accounts (Belue 1996, Roe 1972). Likewise, based on relative bone numbers it can be stated that during the last ice age in Northwest Europe red deer (Cervus elaphus) were very rare: during the Holocene, on the other hand, this animal was frequent-

283


284


ly present (Lister 1984). Conversely, during the last ice age the wild horse (Equus ferus) occurred in great numbers, but in the Holocene forested Europe it was rare (Uerpmann 1990). For the hare (Lepus europaeus) as well, the relative bone number may be a measure for the population density of this animal and at the same time a measure for the increase of landscape openness from the Neolithic period onwards (Söffner 1982, Hüster-Plogmann et al. 1999). The difficulty and real risks of aurochs hunting apparently have not been a hindrance to it. Both young animals and adult bulls were considered for this54. Nevertheless, based on bone finds of which a relation to the age of the animals in question could be ascertained, a preference to kill the least defensible ones appears to have existed. In Bedburg-Königshoven (Germany) a preference for young and female animals existed (Street 1999) and, exceptionally, in Ulkestrup Lyng Øst the remains of at least 19 young calves (and of at least 21 aurochs in total) were found (Richter 1982). Probably (young) calves were slaughtered in many more sites, but their remains are seldom found because of the great porosity of their bones and their presence during a short period of the year. All this does not alter the fact that if people were able to support themselves by catching fish, snaring beavers or keeping cattle or sheep, their need to hunt aurochs may well have been less great. In general, relatively few aurochs bones have been found, especially compared to the numbers of the much hunted red deer and wild boar (Sus scrofa)(Ekström 1993, Zeiler 1997, De Jong 1998, Gramsch 1987, Hüster-Plogmann et al. 1999, Kokabi 1994, Degerbøl & Fredskild 1970, Estevez & Saña 1999, Ervynck et al. 1999, Laarman 2001, Gehasse 1995). One exception to this rule is the archaeological site Bedburg-Königshoven (Street 1999). At this Early Mesolithic (Preboreal) kill site aurochs bones were by far the dominant category of material recovered. The explanation for this may be that during the Preboreal (and in the subsequent Boreal) the landscape was still rather open and not yet totally wooded, which may have offered more possibilities to the aurochs than the landscape of the subsequent Atlantic period; moreover, larger

54 Hunting for bulls is evident not only from many bone finds in human settlements, but also from remains of skeletons (of 3 adult bulls), in which arrow heads and/or bone wounds point to unsuccessful hunts (Prejlerup: Aaris-Sørensen & Brinch Petersen 1986, Vig: Hartz & Winge 1906, Önnarp: Ekström 1993); the Vig bull even turned out to be hunted at least twice (Aaris-Sørensen 1999).



numbers of aurochs made people more disposed to hunt them. Most aurochs finds from Denmark and South Sweden date from the Boreal period; far fewer are known from later periods (Degerbøl & Fredskild 1970, Ekström 1993). This also points to stronger afforestation resulting in a reduction of the aurochs number. Another case in which extensive aurochs hunting seems to have taken place is in Late Neolithic South Germany and Switzerland (Steppan 1999, Hüster-Plogmann 1999). Though man became more and more independent of hunting during the Neolithic (compared to the Mesolithic), nevertheless there were phases during the Neolithic period in which the growing human population faced crises (e.g. food shortage) as a result of which they had to apply themselves more intensively to hunting aurochs. The openness of the landscape at the time, which had been caused by man in the meantime, is likely to have contributed greatly to the larger number of aurochs that were obtained by hunting. The process of the disappearance of the aurochs from South Scandinavia may possibly throw more light on the earlier aurochs population density there and in other places. Because of the rise of the sea level that took place after the last ice age the South Scandinavian distribution area of the aurochs increasingly fell apart into isolated areas (Aaris-Sørensen 1999, Ekström 1993). This isolation probably started the ultimate disappearance of the aurochs from the various isolated areas (Aaris-Sørensen 1980); isolated small island populations are extra susceptible to disturbances by any natural cause. On the Island of Sjaelland the aurochs became extinct in 5900 BC, in South Sweden in 5400 BC and on the Island of Fyn in 3200 BC. Although the aurochs could still increase in number during the Boreal, in the early Atlantic period their numbers decreased (Ekström 1993)55. It is logical to relate this decrease to the increase of afforestation, since the human population density (and therefore the hunting pressure) was still very low at the time. Moreover, at the end of the Boreal alder (Alnus glutinosa) entered the area and probably took over large parts of the marshes, where the aurochs had been used to finding their food. In the long run, the remaining isolated aurochs population in South Sweden was probably too small to face threats like diseases, climate changes, etc. Any influx of aurochs populations from the European mainland was no longer possible. 55 Among others L. Larsson (cited by Ekström 1993, page 83) could show this process of decrease in the successive settlements of Ageröd, on the basis of the number of aurochs bones.

285


286


However, if the aurochs population had been large enough there would have been a sufficiently large enough buffer to resist threats. Generally speaking, the population density of the aurochs is likely to have varied strongly depending on the amount of food (grasses and graminoids) during the growing season. If its living area was largely forested, the density was relatively low; in areas with many sedge marshes it was probably relatively high, possibly close to that of the wood bison in North America. Table 14 gives a brief summary of the population densities of a number of bovine species in their living areas, as discussed in the above. Table 14. Survey of the population density of a number of bovine species and Heck cattle. More detailed explanations may be found in the text.

Bovine species/population Population density Area Aurochs Heck cattle

0.2 an./ km2 26 an./ km2 56

European bison

0.4 – 1.5 an./ km2 57 0.03 – 1.1 an./ km2 58

Bison

8 – 10 an./ km2 1 an./ km2 0.12 – 1 an./ km2 0.2 – 7 an./ km2

Gaur Forest buffalo

Forests at Jaktorów (Poland) Oostvaardersplassen (Netherlands) · (Poland) Forests at Bia§lowieza Caucasus Biosphere Reserve (Kavkaszky Zapovednik) Steppes in North-America Sedge marshes in Canada Forests in India Forests and marshes in Central-Africa

The effects of feeding: forest elephants, European bison and others If one were to compare the landscapes that could be found naturally, with hardly or no interference by man, in Europe, the easterly United States and southeast Siberia, they would turn out to consist mainly of 56 In 2004, the density of the total large herbivore fauna in the Oostvaardersplassen nature reserve was roughly one large herbivore per hectare (J. Griekspoor, pers. comm.). 57 See footnote 52. 58 T.P. Sipko, Severtsov Institute, Moscow (in litt. 2000).



almost uninterrupted forest. Where there was no forest, there were marshes, areas ravaged by fire or storm or beaver habitats. In the early days, when man still led a roaming existence and lived largely from the hunt, up to and including the Mesolithic period, his impact on the forest was basically non-existent. Not until he needed large areas for grass and agricultural crops (for his agricultural and cattle-raising pursuits), from the Neolithic period onwards, were the forests opened up on a large scale. This was done by burning and cutting. As soon as man stopped doing this, the forest would close up again; this was seen in the Great Wilderness as well as in the south of the Netherlands, in the period shortly after the Romans left this latter area. The process of opening up forest areas, carried out by man, may still be seen nowadays, with attendant favourable effects for specific animal species. Such effects promoted the occurrence of the forest buffalo in the Tai forest (Ivory Coast), which did not enter the primary forest area in large numbers until after man had interfered by felling. The same is true of the gaur in Malaysia, which was able to take advantage of the abandoned agricultural plots in the jungle (Wharton 1968), and for the forest elephants in the Gabon jungle, which strongly prefer the secondary forest that has shot up on the former agricultural plots (Barnes et al. 1991). In the same way, the European bison at Bia³owie¿a may often be found on the man-made grasslands and clear cuts (Krasiñska & Krasiñski 1995). Similarly, for the banteng in the Udjung Kulon jungle (Java, Indonesia) the grasslands that are being kept open by man are very important (Hoogerwerf 1970). All of these are examples that show man’s interference on a limited scale. Without that interference, the forest will quickly regain the terrain it has lost and the animals that initially took advantage of the opened-up areas will have to seek their food elsewhere. As the extent to which a forest is opened up grows, the original natural situation will be increasingly pushed to the background. An advanced stage of this is represented by the open, park-like landscapes of the Borkener Paradies in the west of Germany (Pott & Hüppe 1995); see Photo 36) and the New Forest in the south of England (see Photo 29). This latter area, with its very open character, impresses many as being natural, but is

287


288


Photo 29. New Forest (South England). Since the 11th century this area has been managed by man as a forest reserve and also used for cattle breeding and silviculture. Cattle, horses and several deer species graze there. The management has resulted in a half-open landscape of grassland, heath land, shrubs and forests (Photo: J.M. Gleichman).

actually managed almost completely by man. At first sight the area seems to be able to retain its open character only as a result of the feeding by various large herbivores (cattle, horses, deer). In reality, the regulations for its management support the natural effects of these large herbivores in such a way as to be barely noticeable. It so happens that every year, 1,000 hectares of New Forest heathland are burnt and mown to prevent the vegetation from becoming rougher, and young trees from growing up into large ones (Edlin 1969). Horses and cows are removed or added, supplementary food is provided, parts of the forest are fenced off; in brief, man has complete control of the area and ensures that it does not become forested all over (Tubbs 2001). The often very high population density of horses and cattle (locally up to 3 animals on 1 hectare) in many places has even caused thorny bushes such as hawthorn and blackthorn to disappear completely (Putman 1987).



In the forest- and heath area Wolfhezerheide, some 10 years of intensive grazing management with cattle (0.2 animals/ha) could not prevent the gradual closing up of the heath with Scots pine and birch (Bokdam & Gleichman 2000). Human interference in the shape of felling was necessary to keep the heath open. The Oostvaardersplassen nature reserve, with its seemingly natural management, is still a very open area even after 20 years, with still-increasing numbers of horses, cattle and red deer. An explanation for this, in the first place, is the fact that this area is very rich in nutrients, which allows for the long-term maintenance of a relatively high density of herbivores. Secondly, supplementary food was provided in some severe winters, effectively preventing the bottleneck situation caused by such winters in a natural situation. In addition to this, the surface area of the region is enlarged on a regular basis, to keep postponing the moment at which the maximum maintainable number of large herbivores will be reached59. In spite of the high grazing pressure, elder (Sambucus nigra) increasingly covers the earliest grazing areas (Cornelissen & Vulink 1996, p. 89) (see Photo 30). Because of its poisonous ingredients, elder is not or hardly eaten by horses and cattle, rather by red deer. The establishment of elder is promoted by moderate grazing; without grazing it would take hold at a much slower pace. Nevertheless, during recent years the number of red deer has increased exceptionally, which has caused a slow-down of the elder advance. By analogy with comparable situations (reviewed by Flueck 2000) it is to be expected that red deer numbers will decrease sharply in the near future, assuming that man will not interfere. After this decline of the deer (and the cattle and horses as well), elder will be able to continue its march. The bringing into action of wolves, the (natural) traditional predators of deer (,Jêdrzejewska & Jêdrzejewski 1998, Ripple et al. 2001), would undoubtedly accelerate this whole process60.

59

The last time this happened was in 1996 (P. Cornelissen, pers. comm..); during that year the grazed area was doubled. Since the Oostvaardersplassen reserve is in the situation of an island without large predators, it can be compared to real remote islands. The situation in question has been described by Sondaar (1977, 1985) for, e.g., the Island of Crete (Greece). Before the arrival of man, a particular morphologically adapted herbivore fauna (dwarf elephant, dwarf hippopotamus, deer) developed on this island, in the absence of predators. The herbivore fauna could not prevent the whole island from becoming overgrown by forests of mainly oak (Quercus sp.); tree growth was impossible only in extreme dry or salty spots (Bottema 1980). It is likely that not only the newly arrived herbivore fauna, but also the already adapted endemic fauna experienced a series of overpopulations, followed each time by massive starvation (Sondaar 1977, 1985). 60

289


290


As was shown from early descriptions and by pollen research, the natural landscape of Central Europe and comparable areas in North America and Southeast Siberia consisted of ‘forests and marshes’. This basically proves that, in areas where the circumstances for tree growth are favourable, in terms of temperature and moisture balance, for instance, forests will eventually grow naturally, without any interference by man. This tendency towards afforestation may be changed by man with the help of burning, felling and grazing cattle. Burning and felling were used to remove the forest, burning and grazing to maintain the open landscape that had been achieved. Superficial consideration of situations that showed many visible effects of the feeding by large herbivores has often led to the conclusion that such large herbivores would be able to open up forest vegetations and to keep them open. In the jungle, this ability is often ascribed to the elephant; in Europe, such an effect is sometimes attributed to the European bison. The forest elephant (Loxodonta cyclotis), a somewhat smaller species than the savannah elephant (L. africana), was allegedly perceived to

Photo 30. A part of the Oostvaardersplassen nature reserve. In 1984, this area was still an open plain; now it is increasingly growing thick with elder (Sambucus nigra) (Photo: T. van Vuure). Because of the current extremely high density of large herbivores the advance of elder is slowed down; after the future herbivore decline, elder will spread across the entire area.



be able to wreak a havoc of uprooted trees in the forest. Such cases are said to have been ascertained in the sixties of the 20th century in Liberia (Parren 1991). Research into the densities of the elephant populations and the causes for these specific incidents never took place in Liberia. If the tropical rain forests of Africa alone are considered for a comparison of areas favourable for tree growth, it turns out that elsewhere in Africa, forest elephants in situations in which they are not hunted, chased away or otherwise concentrated in specific areas actually have no impact whatsoever on the forest. The Tai forest (Ivory Coast), which hosts the largest permanent population of forest elephants in West Africa, is completely forested (Bousquet 1978, Merz 1986). 60% of this area consists of primary forest; the remainder is secondary forest created by felling. The density of elephants, which prefer to seek food in the secondary forest and along the roads that have provided access to the forest, is 2.3 animals per 1,000 hectares, on average for the whole forest. In the secondary forest, the density is 3 to 5 times as high as in the primary forest. In the open marshes of Azagny National Park (Ivory Coast), the density of elephants is approximately twice as high as in the Tai forest, i.e. 5 animals on every 1,000 ha. The densities in savannah areas, which are more open and contain more grass, are generally higher than in rain forests. In the Tai forest, nothing is known about the destructive influence of the animals on the forest (Bousquet 1978, Merz 1986). The same is true of the rain forests in Northeast Gabon (Barnes et al. 1991), which are still largely uninhabited, thus barely affected by man. Elsewhere in Gabon, the only, usual effect of the elephants on the forest turned out to be that they, like other animals, would make and maintain paths there (Campbell 1991). Outside those paths, barely any impact of the elephants was noticeable, and there was definitely no large-scale ‘destruction’. Since the forest elephant, on account of its foraging behaviour, may more aptly be called a culture follower (Barnes et al. 1991), in view of the fact that it likes to frequent secondary forests, agricultural grounds and access roads into the forest, the animal would not be likely to try to open up the forest locally by uprooting trees unless it was desperate. Elephants are frequently chased away from certain areas by humans, which may lead to concentrations of these animals elsewhere, causing

291


292


lack of food and subsequently, many uprooted trees as a result of the animals’ search for food. Such incidents have been frequently reported of the savannah elephant in East Africa (Buechner & Dawkins 1961, Glover 1963). If the animals are left at liberty to follow their own travel routes, their density will be regulated gradually by the amount of food available, and their effect on the forest will be barely noticeable as a result. Longterm research in the savannah forests of Lake Manyara National Park (Tanzania) has shown that in the long term, elephants and other large herbivores do not actually affect forest communities of Acacia tortilis (Loth 1999). The existing vegetation turned out to be a determining factor for the presence and composition of herbivore populations, instead of the other way around. By uprooting trees, elephants do not create open forest, but rather speed up the regeneration of the forest. The European bison in Europe, in view of its feeding on branches and bark and uprooting of trees, is said to effect the forest in such a way that it gets the character of an open, park-like forest with meadows. Such an effect has never yet been proved, however. It is inspired by assumptions, in part based on the research carried out by Borowski & Kossak (1972), in which the authors revealed that the summer food of the European bison consisted of one third branches, bark and leaves of trees. A later study, of the rumen content of European bison that had been shot, showed this proportion to be about 10%, and also made clear that the European bison mainly feeds on grasses (Gêbczyñska et al. 1991). It is true that large-scale stripping, feeding and sometimes uprooting of young trees by European bison has been ascertained, but it has never been observed that this leads to the eventual opening-up of the forest. The opposite fact has been proven: they actually have no influence on the forest. This evidence is supported not only by global observations of Polish and Russian populations of European bison, but also by research into the effects of feeding on tree bark on the height growth of trees and the development of forests made up of such trees. In view of suspicions held by forestry organisations with regard to the possible detrimental effects of the feeding on tree bark by European bison and red deer on the growth of oak and ash trees, in 1972 the



forests at Bia³owie¿a were checked for evidence of such an effect (Borowski & Van Vuure 1974). For this purpose, about 100 stripped and 100 unstripped oak trees, which grew in a moist, nutrient-rich linden-hornbeam forest (Tilio-Carpinetum), were compared. The same was done with ash trees that grew in a wet, nutrient-rich alder-ash forest (Circaeo-Alnetum). For both species of trees, the lengths of the annual shoots that had grown after the year in which large-scale stripping had taken place were measured. In the case of the oak trees these were three shoots, against four for the ash trees. The trees were 10 to 15 years old, with an average diameter of about 5 centimetres and a height of 6 to 7 m (oak) and 7 to 8 m (ash), respectively. Comparison of the length of the annual shoots of stripped trees with those of unstripped trees showed that largescale stripping had no visible detrimental effect on the height growth of the trees. The height growth of stripped trees did not visibly suffer as a consequence. Only trees whose trunks had been stripped all round died. For the oak trees, this amounted to only 5 % of the stripped trees. In the oak patch, where the young trees were fairly close together, only 2 trees had been uprooted. A visit of the plots in question that took place in 1998, 26 years after the research was carried out, revealed that both the oak and the ash trees had developed normally, entirely in accordance with the expectations of 1972 (see Photo 31). The restorative capacity of the trees could be called adequate, and stripping wounds were barely or no longer visible. Trees may have died at a later stage as a result of wound infections, but this has not had any effect on the structure of the remaining forest. The natural mortality rate of trees as a result of the competition for food and light is so high that the number of trees that die as a result of stripping is negligible. A larger-scale follow-up study, in which a total of 6,000 oak, ash, and to a lesser extent, alder, hornbeam, elm and linden trees were measured, was carried out in the same forests by Fruziñski et al. (1975). This study also showed that very few trees died, and also that stripping did not actually influence the structure of the forest: the structure of stripped stands was no different from that of unstripped ones. A mortality rate among stripped trees that is comparable to the one found in the two studies mentioned above was shown by the research car-

293


294


· (East Poland), square 396. 35 Photo 31. Oak stand in the cultural part of the Forest of Bia§lowieza Years ago European bison and deer heavily harmed the oaks in this forest section by their bark peeling, in spite of which the young oaks from those days have developed into a closed oak forest that is growing well. (Photo: T. van Vuure).

ried out by Van Wieren (1986). During this study, which was also carried out in the forests at Bia³owie¿a, observations were carried out of stripped trees no more than a few kilometres from feeding places, in the month of April. Of the 423 trees observed, 27 were dead, which amounts to about 6%. The effect of bark stripping on the growth of coniferous trees has also been ascertained. In the Netherlands, this was studied in cultivated forests on the Veluwe, for Scots pine (Pinus sylvestris) (Pels Rijcken 1965) and Douglas fir (Pseudotsuga menziesii) (Reijnders 1972). Both studies showed that stripping did not affect the height growth of these trees, nor cause any loss of increase. The mortality rate was negligible also here. Prior to this, research in Poland carried out by Szczerbiñski of Scots pine (cited in Fruziñski et al. 1975) had already shown that the effect of stripping on the structure of coniferous forests is negligible.



It has been proven that European bison are able to uproot trees, but the diameter of such trees does not exceed 10 cm (Pucek 1986). Not all of those trees were deliberately uprooted, moreover; most of them went down when the animals were running through the forest. The consumption of bark and branches by European bison may look disastrous to human standards, but the effect on the growth of the trees is marginal. Photo 32 shows a former meadow along the £utownia River in the Bia³owie¿a Forest. After the mowing of such meadows is stopped, young trees begin to appear. In spite of the feeding by European bison and red deer, analogous with comparable game meadows also this meadow will close up with trees after a few years. Observations in Russia, in areas where European bison live, show that these animals barely affect the natural rejuvenation of the forest (Borowski et al. 1967). This was also shown in the reserve in the Caucasus (North Ossetia), where no supplementary food is given in winter. Preferred plants such as Rubus caesius and Festuca montana grow and

· Formerly, this meadow was Photo 32. Part of the Lutownia River Valley in the forest of Bia§lowieza. mown by man and many bison grazed there. After mowing was stopped, the area was gradually taken over by trees and bushes; bison no longer graze there (Photo: M. Karczewska). §

295


296


rejuvenate quite well, and the main forest tree in the area, the beech, is hardly eaten at all (Kazmin & Smirnov 1992). Of the thousands of young trees originally planted on a clear-cut area in a cultivated forest, only a few dozen eventually grow to their full stature. In the natural process of forest rejuvenation, the starting material is even more extensive, since the seedling stage should also be taken into account. As was seen before, mortality as a result of large-scale stripping does not affect this. Clear cutting and subsequent cultivation by man may well have exacerbated the problem (for forestry) of stripping. By concentrating attractive food (leaves, branches, bark) in a certain area, the extent and the effect of stripping and feeding are increased. Research in the European part of Russia revealed that the impact of the elk is strongest in those areas where clear cutting and massive cultivation take place (Kuznetsov & Lozinov 1992). The thinning-out of young trees also turned out to yield more food for the elk. The increased amount of food in forests that were managed in this way resulted in a large increase of the number of elk. The increase of clear-cut areas and secondary forest (young trees) enlarges the possibilities of the animals that prefer to forage in these areas. The same was already seen in the jungle, with regard to elephants, gaur and banteng. The numbers of elk and their detrimental effects on the trees turned out to be much more limited in forests that had not been influenced by man, i.e. ecosystems that still retained their natural character (Kuznetsov & Lozinov 1992). Since the food, that is to say the young trees and shrubs, was spread out through the forest, the animals needed to search harder for it and the total amount of food per surface area was smaller than in a forest that had been opened up locally by man. The number of animals in need of that food became smaller as a result of this, so the population was more evenly distributed across the forest. This was already mentioned in the discussion of the density of bovine animals in forested areas. The less involvement by man, the more limited the supply of food available to bovine animals and deer is likely to be. Thus, in early Europe, not only the density of elk was lower than in the current cultivated forests, but also that of red deer (2-10 on 1,000 ha; Fröhlich 1955, Meister 1969) and roe deer (1-3 on 100 ha; Schwend 1950, Meis-



ter 1969). The same is true of North America, where the densities of the whitetail deer (Odocoileus virginianus) used to be lower than they are now: 2-4 per 100 ha against the current 10 per 100 ha (North Wisconsin, United States) (Alverson et al. 1988). Man ensured larger food supplies also there, and this deer took advantage of that. A herbivore that is undoubtedly able to have a great impact on the forest, be it locally, is the beaver. The ability of this rodent, which lives by lakes and along rivers, to temporarily or permanently open up its living area by felling trees and building dams, was already discussed in Ch. 9.1.2. No other animal is usually capable of bringing about such effects.

Wood Buffalo National Park (WBNP): A case study A representation and clarification of the position of large herbivores in a natural, largely forested ecosystem is best given on the basis of the actual situation in such an ecosystem. Wood Buffalo National Park, in the northeast of the Canadian state of Alberta, an area that gained its reserved status in 1922, is eminently suited for this purpose. In comparison with, for example, the surface area of the Netherlands, which is about 33,000 km2, this nature reserve is quite large, some 44,800 km2 (Carbyn et al. 1993). The whole area is inside a boreal coniferous belt, which mainly consists of Picea glauca, Picea mariana, Pinus banksiana, Populus tremuloides and Populus balsamifera (Schwarz & Wein 1997). Many rivers, the largest of which are the Peace and the Athabasca, transect it. The Slave River is its eastern boundary. Coniferous forests, alternated by lakes and marshes, dominate the landscape. The largest marshes are situated along the large rivers and in the so-called PeaceAthabasca Delta in the south. This is one of the largest inland freshwater deltas on earth, with a surface area of 1,533,000 ha. WBNP hosts the largest bison population of North America, which lives free and is neither given supplementary food nor hunted (Photo 22). As it happens, the management of these bison and also other animals is not interfered with by man, and entirely left to nature. Thus, in the case of the bison, the number of animals is regulated by the amount of food available in

297


298


the marshes (their main foraging areas), as well as by floods and the predation by wolves. The other large herbivores that live here are elk, whitetail deer and reindeer. Beavers also find their ideal biotope here. Two processes that may shed light on the relation between the large herbivores and the vegetation are currently taking place in this large nature reserve. In the first place, a development is taking place on the so-called ‘dry grasslands’ (Schwarz & Wein 1997, 1999). These grasslands, called dry to distinguish them from the wet grasslands along the rivers, occur in areas unaffected by river floods. In the period that Raup (1935) carried out his research of the vegetation, they covered many hundreds of hectares in various places. Their nature and origin never ceased to puzzle him even then, since they were entirely surrounded by coniferous forest. Although there was sporadic regeneration of aspen (Populus tremuloides) at the time, the grasslands still had a very open character, which for some mysterious reason they seemed able to retain. Studies carried out in 1996, however, revealed that all these areas had closed up with aspen timber (Schwarz & Wein 1997, 1999). Only a few small growth areas, in which the soil was too dry, had not yet closed up completely. The ‘prairies’ described earlier by Raup are now completely dominated by aspen, and barely anything remains of the grasslands. The forest, in the shape of a pioneer vegetation of aspen, has once again taken its original position; original, since closer inspection of the history of the area has uncovered an explanation for the earlier grasslands. Historically, local Indian tribes in the north of Alberta used to burn the forests in order to create grasslands and attract game. When the WBNP was founded in 1922, they had to stop their burning activities inside the reserve. Outside the reserve, the burning continued into the late forties, when it was stopped also here. Inside WBNP, man instigated no more burning, and the dry grasslands started to close up. Raup saw the first effects of this when he visited in the early thirties. The closing-up of the initially open grasslands shows that the local large herbivores under natural circumstances were unable to stop the advance of the forest. The forest dominates this area once again, and will continue to do so unless it is removed by fire (Schwarz & Wein 1997)61. Small, natural 61 Although Timoney (1999) is critical about some points of the research carried out by Schwarz & Wein (1997), he nevertheless agrees with the authors that soil factors such as moisture- and salt content actually determine the presence (or absence) of forest.



forest fires regularly occur in the coniferous forests of WBNP, as a result of lightning. These fires are invariably followed by forest rejuvenation with aspen.

Photo 33. The Peace-Athabasca Delta in Wood Buffalo National Park (Central Canada) is a vast marshy area in which sedge vegetations play an important role. These vegetations are of vital importance for the bison living there (Photo: Parks Canada, Wood Buffalo National Park).

The other process in WBNP, which has a lot to do with the functioning of large herbivores, is taking place along the large rivers and in the said delta. The issue in question is the water regimen of the area. The most important river is the Peace, which rises in the Rocky Mountains. The Peace-Athabasca Delta, which is fed mainly by this river, is a gigantic mosaic of lakes, fairways, grasslands, sedge lands and forests. The area is very flat, with differences in height of only a few centimetres (Healey 1994). The spring floods are essential for the vegetational character of the delta; they take place in May, June and July (their high point is in June) and their dynamics are preponderant to the nature of

299


300


the vegetation. The grass- and sedge lands of this delta constitute the most important foraging area for the bison (Carbyn et al. 1993) (see Photo 33). They comprise 24% of the delta, some 367,000 ha. Both in summer and in winter, the bison live here to eat grasses and sedges.

Photo 34. Because of a changing water regime the water level in the Peace-Athabasca Delta (Central Canada) is gradually falling. As a result of this large areas, originally wet, sedge vegetations, may be colonized by willows (Salix sp.) (Photo: Parks Canada, Wood Buffalo National Park).

Just how important the water regimen really is for the delta vegetation became apparent after a one-off intervention by man, outside the area. It so happens that in 1968, a dam upstream in the Peace River was finished, which caused a reservoir to fill up. This was to have dramatic consequences for the water balance in the delta. The construction of the dam reduced the frequency and scale of the floods, in particular; their dynamics became considerably less, as did the difference between high and low tides. The delta began to dry out as a consequence62. This had 62 Another explanation for the drying-up of the delta is given by Timoney et al. (1997). Based among other things on muskrat returns and flood frequency, they conclude that the delta is subject to a climatological alternation of wet and dry periods. These variations take place according to a sine-based model with periods of ca. 120 years. The dam in the Peace River was constructed at the very moment a climatologically drying trend set in.



immediate effects on the food supply of the bison, among others. The dehydration also caused receding water levels in the productive, formerly wet grass- and sedge lands. Favourable circumstances for the germination of willows ensued, and these began to take root on a massive scale (low willow shrubbery) (see Photo 34). This had not been possible before; it had been too wet. In the early nineties, as much as 47% of the areas interesting to bison had become overgrown by willows, and this process still continues (Carbyn et al. 1993)63. If predictions prove correct, about 85% of the original wet grass- and sedge lands will be covered with willow by the year 2031 (Jaques 1990). Former marshes overgrown with willow have lost their appeal to the bison, whose number decreases steadily partly as a result of this. Around 1970, the total number of bison was still about 12,000, 10,000 of which lived in the Peace-Athabasca Delta. The 2,000 animals north of the Peace river do not seem to decrease in number, but of the Delta population, which used to number 10,000 at one time, only 382 bison were left by 1998 (Carbyn et al. 1998), clearly only a fraction of the original number. Emergency measures to make the delta wetter again with the help of dams made of rock have as yet been of little avail (Healey 1994). Thus, a one-off intervention in the water regimen of a large river has had disastrous consequences for this population of bison. These developments, caused by a sudden, once-only action, shed more light on a number of fundamental processes in this area. It has turned out that the wet grass- and sedge lands were treeless because willows and other trees could not germinate and grow up in spring. If they get this opportunity, however, massive growth takes place. Secondly, this forest growth was revealed to be unstoppable by large herbivores; colonisation progresses in spite of feeding. The appearance of more willow forest makes the area more interesting for elk, for which the willow constitutes important food. Elk are not able to stop the eventual growth of willow forests either, however, just as they were unable to prevent the growth of aspen in the ‘dry’ grasslands. That such wet sedge vegetations along rivers owe their existence to the water regimen of those rivers was also shown in the basin of the Pripyat River (White Russia/Ukraine). Kulczyñski (1949) concluded that 63 As in WBPN, in the nearby MacKenzie Bison Sanctuary willow cover has actually increased as well over the past 30 years (Larter et al. 1994).

301


302


Photo 35. In the middle of this picture from Wood Buffalo National Park a light-coloured strip can be recognised that stretches to the horizon. At one time this zone was clear cut to accommodate a never-realised road. Afterwards the zone grew thick with wild shoots of trees in spite of the presence of large herbivores. In the centre of the strip is a trail that is used extensively by bison and infrequently by man (Photo: L.M. Brusnyk).

the drainage by man of the sedge marshes there resulted in their growing over with willow and birch. Judging from the soil profile (sedge peat), these marshes had been able to survive for centuries as sedge vegetation because the almost permanently saturated soil did not allow any tree growth.

Discussion In the above, it is argued that various studies have shown that the natural landscape of Central- and Western Europe, if it were allowed to return, would consist of almost completely closed forests alternated by marshes. Based on this and on the capacities of the current large herbivores, the effect of feeding by these animals has been concluded to be marginal. Various other opinions exist, however, which do consider these large herbivores to have a great impact on the tree growth and, thus, the appearance of the landscape. These opinions have resulted from the



increased use of cattle and horses in nature conservation projects and from ecological studies of populations of large herbivores in various European and African nature reserves.

The animal species The herbivores relevant to this discussion include a large variety of species, whose numbers and appearances have undergone numerous changes in the course of evolution. In historic times, the aurochs and the wild horse became extinct in Europe as a result of activities by man. Historically, the latest period in which many species became extinct was the transition of the last Ice Age to the warmer Holocene period, some 12,000 to 10,000 years ago. In this period, in Europe among other places, many large herbivores and their predators became extinct, including the mammoth, the steppe bison, the giant deer, the lion, the hyena and the sabre tooth tiger. Martin (1984) finds a cause for this massive extinction in the increase of the hunting pressure exerted by man with his greatly improved hunting techniques (‘overkill-theory’). Many large mammals were allegedly unable to deal with this and died. Also on other continents, many animal species died between 30,000 and 10,000 years ago; as in Europe, according to Beutler (1996) and Bunzel-Drüke (1997), this was caused by man. Others (Guthrie 1984, Vereshchagin & Baryshnikov 1984) deny man’s influence here and seek explanations in the climatologic and attendant vegetational changes. As it happens, between 12,000 and 10,000 years ago a few short, marked climatological changes took place which in a very short time changed the climate on the northern hemisphere from a dry and cold climate with little snow in winter to a warm and wet one with a lot of snow. These changes allegedly were so rigorous that many animal species were unable to adapt; they died and became extinct as a result. Periods like the one discussed here, in which many species became extinct, took place many times in the course of evolution (Stanley 1989), and often were larger-scale and had greater consequences than the one on the transition of the Pleistocene to the Holocene period. About 250

303


304


million years ago, in the Upper Permian period, 70 to 90% of all animal species on earth became extinct in a period of several million years. As far as can be traced, climatological changes, for whatever reason, were at the basis of many of these cases. The argument about the cause for the extinction of many mammals in Europe in the beginning of the Holocene period is part of the discussion about the effect of large herbivores. One of the issues is which species really naturally belong in Europe, if man’s influence is not considered; another is whether we can still speak of a natural range of mammals and a natural landscape if these lack the woolly mammoth, the woolly rhinoceros, the giant deer and other animals. According to some authors (Poortinga 1981, Bunzel-Drüke et al. 1994), we are very far removed from such a natural landscape, precisely because all those species are missing. Poortinga, as a ‘reasonable ecological alternative’ for the species that have disappeared, suggests that elephants and rhinoceros are introduced in Europe to aid the closest possible approximation of the natural landscape. It so happens that large herbivores have a lot of impact on the landscape in his vision; after their extinction, caused by man, the vegetation must have changed essentially, from a grassy steppe with little forest into a densely forested landscape. Zimov et al. (1995) propound that the disappearance of large herbivorous mammals from the arctic zone of the northern hemisphere, which also obliterated their effects of trampling and feeding, changed the dry, grassy steppe into a wet, mossy tundra. In their opinion, the reintroduction in the tundra areas of these large herbivores or their relatives may possibly reverse this process, changing the tundra back into a grassy steppe. To test their theory, an experiment (‘Pleistocene Park’) is being carried out in the Kolyma tundra (Northeast Siberia). Yakutian horses, reindeer, elk and musk oxen are currently living there on an area of 160 km2. Bison, and possibly Siberian tigers, will be added at a later stage (Stone 1998). Many mammals became extinct in a fairly short period on the transition of the Pleistocene to the Holocene period; in the course of the Pleistocene, however, species also became extinct on a regular basis, to be succeeded or replaced by others (Stanley 1989). Although it is quite well known which animal species lived during that period of two million years,



it is not always clear exactly which species lived in which period, and what their living conditions in that period were like. The reason for this is that warm and cold climates alternated during the Pleistocene period. The warmer periods cannot simply be compared to the Holocene, since they occurred in all kinds of gradations; the same is true of the fauna in those warmer periods and the fauna during the Holocene. If man were to have wiped out a number of mammals at the end of the Pleistocene and in the course of the Holocene period, it would be reasonable to assume that in Europe today, there are many vacant niches for large mammals. If this extermination was caused by climatological changes, the presence of such niches becomes very doubtful. The filling up, according to man’s own discretion, of suspected vacant niches is a hazardous affair, therefore. Species such as the aurochs, the wild horse and the Balkan lion, whose extinction by man has been proven, have left vacant niches that could be filled by cows, horses and (bredback) Berber lions. Whether animal species should be brought to Europe for the simple reason that they existed during the warmer period of the Pleistocene, and/or because there is no physical barrier for their distribution, is a topic for discussion. The fallow deer (Dama dama) is tolerated in the Low Countries for both these reasons, the second of which could warrant the introduction of the tiger, for example, in Europe, outside the Balkans. Since a vacant niche for a large feline is suspected in Europe, the tiger, which during its distribution in Asia had already progressed into the Caucasus (Mazak 1979), might be able to fill it. Opinions about the (re)introduction of animal species differ greatly; some scientists are very cautious, while others seek the completest possible filling-in of the niches. The information about Pleistocene species is far from complete; the early distribution of species that still exist may also be problematic on occasion, however. The exact European living area of the European bison is not completely clear, for example; the northern, western and southern boundaries of its distribution area cannot be ascertained with any certainty. Some use C.C. Flerov’s map (rendered in Pucek 1986, p. 292) to indicate this distribution area; he drew the border of the distribution area of the European bison through the south of Sweden, South

305


306


England and North Spain. The evidence for its occurrence in these areas is too dubious to be used, however; bone finds, toponyms or written records are practically lacking. In itself, this does not necessarily mean that the species did not occur in those areas; it may have become extinct there at a very early date. For instance, in Sweden all post-glacial Bison finds date from the Boreal period and the latest Bison finds from Denmark date from the Preboreal period; both are periods during which the landscape was not yet entirely overgrown with forests (Ekström 1993, Degerbøl & Iversen 1945). Its former distribution in the Netherlands, Belgium and France is more unclear. Apart from an occasional toponym and a few bone finds nothing is known about this, whereas bones of the (ice age) steppe bison (Bison priscus) are found in huge numbers. It is remarkable that nearly no European bison remains have ever been found in human settlements in Western Europe, unlike in Germany and Switzerland and even more often in Eastern Europe; only in France two (neolithic) sites are mentioned by Poulain (1980). There do not appear to have been any physical barriers to prevent the animal from going to Western Europe, but climatological or other factors may have limited its distribution in such a way that its occurrence, as a more continental species, was less obvious in these countries than in Centraland Eastern Europe.

The impact of large herbivores As much as there are different opinions about which large herbivores were indigenous to Europe, there is also discussion about the impact of the various species on the development of trees and shrubs. Up to 1975, when Van de Veen proposed to allow more natural processes to take their course in nature conservation, management of the latter had been determined mainly by human interference, and through the conservation of cultivated situations that had developed with time. Large herbivores and their predators especially are central to the processes indicated by Van de Veen, and the main issues are the effects of that predation and the impact of feeding on trees and shrubs.



The withdrawal of man’s interference and its replacement by the said processes would make nature more dynamic and allow the development of a truly natural landscape. The effects of the large herbivores, the European bison in particular, would create a ‘more openly structured forest’ (Van de Veen 1975). Van de Veen did not deem red deer capable of this, since he did not think that even population densities of 30 to 40 deer/100 ha would be able to prevent young Scots pine from growing up to constitute closed timber. At best, deer would be able to slow this process down somewhat. Moreover, according to the same author, the predation by wolves would keep the density of deer from ever becoming this high. In later years, Van de Veen substantiated his vision of a more open forest with findings of research carried out in savannah areas in East Africa. Through their feeding, the large herbivores in savannah areas (the Serengeti, for example) would be able to control the growth of trees and shrubs to such an extent that a variegated landscape of grasslands, thickets, pieces of forest and solitary trees could develop. Closure of the forest would be made impossible by these animals. This notion of the various large herbivores in their specific niches, giving the landscape the character of an openly structured forest, was later adopted and propagated by many authors and institutions: in the Netherlands, by Van der Lans & Poortinga (1986), Vera (1997) and the Stichting Kritisch Bosbeheer, in Germany by Remmert (1985), Hofmann & Scheibe (1997) and Bunzel-Drüke et al. (1994). Similar ideas in North America also bring to mind the discussion about the Serengeti: the discussions about the character of the natural landscape here are specifically concerned with the semi-arid landscape of Yellowstone National Park and the proven impact of feeding by the wapiti (Kay 1997). In recent studies it was shown that this impact has been greatly diminished since the reintroduction of wolves (Ripple et al. 2001, Larsen & Ripple 2003). These ideas about the appearance of the natural landscape in Europe, referred to in Germany as the ‘Megaherbivorentheorie’, refer to relatively large numbers of large herbivores, whose number, feeding and digestive specialisms would ensure that the natural landscape forms a

307


308


colourful variety of all kinds of vegetational structures, in which all herbivores will be able to exist optimally. Influencing of vegetational structures by one species to benefit another (‘facilitation’) plays an important role in this system. Browsers and intermediate feeders give the forest landscape an open structure, in which a lot of grassland develops (see Photos 29 and 36). In this grassy landscape, consumers of tall grasses will have a facilitating function for those that eat short grasses. The adherents of this theory disagree about the measure of openness of this forest landscape. Vera (2000) and Hofmann & Scheibe (1997) assume a very open, park-like landscape. They ignore the findings of modern palynology, which start from the almost closed European forest landscape revealed by pollen analyses. They also believe that the European bison in particular, by uprooting trees and feeding on bark and branches, is able to give the forest a very open structure (Vera 2000, pp. 355, 356; R.R. Hofmann pers. comm.). With reference to the openness of the park-like landscape (the percentage of grassland), Vera (in Borgesius & Van Tol 1998) quotes a proportion of ‘30-40% on average, and at least 50% on the impoverished Veluwe [Province of Gelderland, Netherlands]’. Van der Lans (pers. comm.) also propounds an open forest landscape, though less open than in the model proposed by Vera. In this rather more closed system, it is also the European bison that is supposed to be able to open up the forest (Van der Lans & Poortinga 1986, p. 51) and ensure more structure in the forest (‘fine tuning’, ditto p. 112). Bokdam, Cornelius & Krüsi (2001) also start from a landscape of vegetational mosaics and the impact of large herbivores on this landscape, but they emphasize that factors such as the availability of nutrients in the soil will determine the production of the grasslands and the proportion of feeding inhibitors in ligneous plants, and thus influence the intensity of feeding and the speed of succession. To test their Megaherbivorentheorie in practice, Hofmann & Scheibe (1997) have planned the ‘Multi-Spezies-Projekt’, in which as many different kinds of indigenous large herbivores as possible are brought together to show their actual ability to create an open, park-like landscape in Germany. Because of the obstinacy of the authorities, finding a



Photo 36. The Borkener Paradies reserve along the Ems River in Germany (Photo: J.M. Gleichman). Here, the age-long management with cattle and horses has created a half-open landscape with grasslands, bushes, solitary trees and stretches of wood.

large-scale location for this project in former East Germany has turned out to be impracticable (R. Cornelius pers. comm.). Vera (2000) assumes grasslands colonised by thorny thickets, which will eventually bring forth tall forest vegetation. As a consequence of feeding, in time this forest will be opened up and turned into grassland by large herbivores; later on, thorny thickets will once again establish themselves. Vera especially pays attention to tree species like oak (Quercus robur) and hazel (Corylus avellana). In his theory, the establishment and juvenile growth of these tree species in thorny thickets would be impossible without the feeding preferences of large herbivores like aurochs and wild horses. However, the apparently larger role of the oak in former times, in contrast with the present situation, that is emphasized by Vera, may be explained in several other ways. Besides a good growing site, two main factors determine whether oak will establish and

309


310


grow well: sufficiently light growing circumstances and little or no feeding damage by herbivores (among others: Palmer et al. 2001, Mountford et al. 1999, Borgesius & Van Tol 1998, Kelly 2002, Lüpke 1998, Lüpke & Hauskeller-Bullerjahn 1999, Linhart & Whelan 1980, Kuiters et al. 1997, Langbein 1997). This actually means that oak does not tolerate much shading by shadow trees like e.g. hornbeam (Carpinus betulus) or beech (Fagus sylvatica), or too high densities of herbivores. A period often referred to is the Mesolithic, during which hornbeam and beech had not yet arrived in Northwest Europe. According to pollenand plant macrofossil data, oak played an important role in the forests at the time. This major role can partly be explained by the absence of hornbeam and beech and partly by the low natural densities of herbivores inhabiting the forests (see the above in this Chapter). Besides this, man probably regularly burnt forest parts to provide more food for its game animals in this way (several papers in Scott et al. 2000; for the United States, see Bonnicksen 2000). It is likely that oak took advantage of the light growing circumstances that resulted from such burnings to extend its territory. To illustrate his theory, Vera describes the situation in the Forest of Bia³owie¿a, where there is allegedly hardly any natural regeneration of oak and very little hazel. On the other hand, Bobiec (2002) contradicts this, writing that ‘numerous anecdotal observations and photographic documentation’ confirm the regeneration of oak; in addition to this he mentions hazel as the ‘most common shrub’ in the forest. Earlier, Brincken (1826, page 22) had already ranged hazel among the most common shrubs there at the time. Pollenanalytical research in the Bia³owie¿a National Park, spanning the last 800 years, shows a continuous and evident presence of oak and hazel (Mitchell & Cole 1998). The possibly smaller number of oaks in this forest than in earlier centuries may be attributed to the excessive densities of bison and deer during the last centuries, and even today. The former situation in Ireland is also illustrative. Throughout the Holocene, Ireland lacked aurochs, bison, wild horse and (until recently) red deer (Woodman et al. 1997), while oak and hazel were very abundant regionally (Bradshaw et al. 2003). Beech did not reach Ireland till 3000 years ago, and hornbeam did not arrive in Ireland naturally (Godwin



1975, Rackham 1980). Apparently oak profited from the absence of the aforesaid shadow trees. Apart from this, large herbivores clearly did not play a key role in the establishment and growth of oak and hazel here; on the contrary, their absence was beneficial to these trees. Comparable conclusions are drawn by Mitchell (2005), who has found that Holocene pollen diagrams from Ireland closely resemble those in the rest of Europe. As was indicated before, many authors regularly refer to the savannah areas in East Africa to underline their ideas about the ability of large herbivores to shape the landscape, also in Europe. It is remarkable that all the large, closed African jungles are always largely ignored in these arguments, although large herbivores also live in those areas, in which, moreover, the influence of man is as yet barely noticeable. It is true that there is a more limited number of species of large herbivores; these basically possess the same feeding capacity, however. Elephants also occur in these closed jungles, beside wood buffalo, bongo (Taurotragus eurycerus) and okapi (Okapia johnstoni). Their population densities are relatively low, and they have been proven unable to really change the structure of the forest, in spite of their feeding. Their numbers and ecologies have adapted to the existing (forest) situation. The East-African savannahs host a multitude of herbivore species. Changes in the population densities of these animals and the attendant results for the vegetational structure have often been impressive. In particular the impact of elephants has often been recorded, with references to the destruction of forests (Buechner & Dawkins 1961, Glover 1963). In many cases, such destruction could be explained by temporary overpopulation caused by man, which had resulted in a shortage of food. Elephants will also resort to breaking off branches and uprooting trees in other cases, however. Research into the impact of elephants on the tree growth in a savannah forest used always to be based on short-term observation. More and more frequently, however, results are published of long-term observations, during which both the changes, and the causes for these changes, in the number and size of trees in the savannah areas of East Africa are considered (Van de Vijver et al. 1999, Loth 1999). Research by Van de Vijver et al. shows that both the precipitation factor

311


312


and the aging processes of trees are much more important for the structure of the savannah forests than the effects of elephants. Loth’s research shows that the uprooting of trees does not create permanent grasslands but rather accelerates the rejuvenation of the savannah forest. Germination of Acacia tortilis, one of the most important savannah trees, benefits from a lot of light and bare soil, free from competition with other plants. The removal of large trees, by aging processes or by the impact of elephants, creates such circumstances. The most important driving force behind this whole ecosystem is the precipitation factor. This eventually determines the extent of forestation, while the influence of large herbivores is merely marginal (Loth 1999). The situation in the Hungarian steppe area of Hortobagy proves that it is important to consider all factors that may affect the character of a landscape. The area in question is often seen as a natural steppe area inside the otherwise forested Central Europe. Closer consideration reveals that the true picture is a little different, however. In the first place, the amount of precipitation (527 mm/year) in the area is sufficient for successful forest growth; in the second, man has left a considerable mark on the landscape (Den Hollander 1947, Dunka et al. 2003). Man cut down the original forests and the rivers that transect this plain, which used to be slow and meandering, were all canalised in the 19th century. As a result of this, the great floods that used to be so common in the early days no longer occur. As a consequence of the dehydration and silting up of the soil, here a treeless landscape has developed in the course of time.

The European bison The European bison plays a large role in the discussion of the impact of large herbivores on the tree growth in Europe. A lot of research of this animal was carried out in the Bia³owie¿a Forest. Whether this forest may be called natural and, if not, whether any conclusions may yet be drawn from the research into the impact of the European bison, is very much the question. The current situation in this forest cannot be called completely natural. Apart from the strict nature reserve, the forest area



is subjected to a management with a view to timber harvesting, although this is done less rigidly than is common in the Low Countries. The management of the forest is not natural, therefore, and the composition of its animal species is no longer original, either. A number of species that used to live here, such as aurochs, bear, wolverine, wildcat and flying squirrel have disappeared64. A number of remaining species, including the European bison, are hunted by man and given supplementary food in winter. Man controls the number of European bison. Some have used this argument of quantity control to stress that, if man would no longer hunt and provide supplementary food, the numbers of animals initially ascertained ‘randomly’ by man would become increase, and the impact on the forest of, for example, the European bison would become greater as a result. In that case, the true size of feeding by large herbivores would be revealed; the forest would become increasingly open and the forest structure would clearly change. Research into the fluctuating numbers of the various animal species in this area during the 19th- and early 20th centuries and into the factors that influence these numbers nevertheless shows that the current numbers are not as random as they are sometimes made out to be (Jêdrzejewska et al. 1997). The subject of this research was the period 1802-1919; the numbers of large herbivores and their predators, as well as the management engaged in by man in this period, were revealed. Not only the size of the European bison population is known, therefore, but also the extent to which it was hunted and poached by man, how it was provided with supplementary food and which factors effected fluctuations in its population. On the basis of this information, Jêdrzejewska et al. (1997) have concluded that man’s influence, as well as the number of large herbivores, were determining factors for the number of European bison. Predation barely played a part in this. With regard to the 19th-century situation, apart from times of war, they ascertained a balanced density for the European bison (i.e., in balance with the amount of food available) of 0.5/ 100 ha. For red deer this number is 4, against 3 for roe deer and 0.3 for elk. Data for the period 1991–1995 (Jêdrzejewska & Jêdrzejewski 1998)

64 It is not known when the aurochs became extinct in this forest; Hedemann (1939), describing the history of the forest from ca. 1500 onward, does not mention it at all. It is doubtful whether the wild horse naturally occurred in this forest. Reports about ‘wild horses’ do not clarify whether these are horses that were originally wild, and in which areas they lived exactly (Brincken 1826, Mager 1941).

313


314


show that the density of the European bison population more or less concurred with this number, while that of the elk was lower (0.2) and those of roe deer and red deer higher (5.1 and 5.6, respectively). The population of European bison is more or less in balance with the current amount of food, therefore, but this is determined to a large extent by man’s management, which provides game meadows, clear cuts, verges, access to farmers’ fields and -meadows and supplementary food in winter. According to some authors (F.W.M. Vera, R.R. Hofmann, H.E. van der Lans), the pressure of feeding on the forest would increase, and the structure of the forest would change, if this management and the hunt on the European bison were stopped. The forest would even be opened up. Others assume that the removal of human interference will decrease the available amount of food (both in summer and in winter), which will result in a smaller carrying capacity, thus in fewer European bison, and a more limited impact. In a part of the Bia³owie¿a Forest, the National Park, hunting and silviculture were prohibited as early as 1921. There are no game meadows or clear cuts in this part. In spite of the fact that they have free access, the European bison hardly use this strict nature reserve, allegedly because it yields less food than the surrounding cultivated forests (Krasiñska & Krasiñski 1997). The population of European bison that inhabits the Bia³owie¿a Forest in the current circumstances hardly affects the development of the forest. Such, in any case, is the opinion that prevails among the Polish researchers of this forest area (Z. Pucek, L. Mi³kowski pers. comm.). Although some consider the influence of large herbivores a determining factor for the openness of the forest, others search for abiotic factors capable of achieving this. In Central- and West European forests, natural fires and feeding by insects have had little effect. Fire may play an essential part in steppe areas, steppe forests and boreal coniferous forests, however. Beside occasional, infrequent factors such as avalanches, volcano eruptions and floods, storms constitute a more regular and omnipresent factor. Storms may have a great impact and turn large forest areas into chaos. The question is whether large herbivores benefit from this. Tentative research (Reynolds 1969) shows that deer and cattle hardly benefit from such ravaged areas, rich in nutrients though they may be. Uprooted



trees impede their movement, while at the same time young trees use the protective circumstances to grow up. Clearly, storms may open up the forest, but also create circumstances for its rejuvenation. The number of European bison is not only determined by the competition for food among its own species, but also by the presence of other species that compete for food. The competition for food between the aurochs and the European bison in their time may be somewhat elucidated by considering the competition between the European and the domestic cow in the 19th century. Around 1860, the increase of the number of domestic cows caused a dramatic decrease in the number of European bison (Jêdrzejewska et al. 1997). Conversely, the number of European bison increased again between 1890 and 1900, when the pasturing of cattle in the forest became more limited. The extremely high number of red deer around 1910 did not affect the number of European bison, however.

The Swiss National Park The discussion about the impact of large herbivores on the forest growth and the resulting landscape currently focuses on developments in the Swiss National Park. This nature reserve in the east of Switzerland is about 169 km2, and has been a strict nature reserve since 1914. This means that there is no more hunting inside the area, cattle are no longer grazed there and no more forestry takes place. The starting point in 1914 was a very open forest landscape with (formerly) mown and grazed mountain meadows and pioneer forests of mainly mountain pine (Pinus montana), which had resulted from large-scale felling in the preceding centuries (Leibundgut & Schlegel 1985). Largely man, therefore, had determined the character of this area and it had become a cultivated landscape. After this became a strict nature reserve, the numbers of red deer and chamois increased greatly. The hunt for these species had been abolished, natural predators were lacking and the mountain meadows were free from grazing by cattle and supplied a lot of food. In the eighties, their numbers stabilised to the current number of around 2,000 deer

315


316


(in July and August) and the same number of chamois (Krüsi et al. 1995). During the growing season, the deer stay inside the national park, but 90% of them live outside in winter, in lower areas. In addition to this, in some parts of the reserve almost half the deer seek their food in areas outside the reserve at night (Krüsi et al. 1996). The Swiss National Park is no closed ecosystem, therefore, but part of a larger whole. The vegetation of the national park has changed gradually in the past 80 years. A continuous succession of grass- and forbs communities occurs on the mountain meadows, but large areas are still free of forest (Schütz et al. 1998). The current landscape shows the unmistakeable influence of large herbivores, whose feeding ensures that the mountain meadows do not close up with trees (Krüsi et al. 1995). Superficial consideration of this phenomenon might lead to the conclusion that the herbivores that live there are able to keep the forest open and to leave a lasting mark on the landscape. On the basis of this consideration, Krüsi et al. (1995) argue that naturally, there must have been a cycle from forest, to open area, to forest (‘Wald-Freiland-Wald-Zyklus’), in which large herbivores play an important part. More detailed consideration reveals the matter to be a little different, however, since many processes coincide and some (natural) processes are entirely lacking. Under the present circumstances, the large herbivores in this area affect tree growth in two ways. In the first place, they create favourable circumstances for the germination of mountain pine through their consumption of grasses and forbs. This tree species is dependent on sunny stands, with little competition of grasses and forbs, for its germination and development. The fact that grazing by red deer was able to ensure such circumstances was revealed in an experiment in which a positive correlation was found between the increasing number of deer and the growing number of small trees (< 150 cm) (Krüsi et. al 1996). This experiment also showed that the number of large trees hardly increased, in spite of the growth of the number of small trees. Not until the number of deer stabilised could many of the trees grow to their full height. This indicates another effect of the deer: their intensive feeding off young mountain pine. Both Krüsi et al. (1995) and Leibundgut & Schlegel (1985) believe that the large feeding pressure is the most important rea-



son the mountain meadows have remained open for such a long time. The tree growth-promoting impact of deer is limited to the germination and settlement of the trees, therefore; their intensive feeding subsequently keeps these trees short. In spite of the still-open character of the national park, slow reforestation of the mountain meadows is taking place. The open areas are expected to close up with mountain pine in the long term; still later, this pioneer species will be replaced in part by Norway spruce (Picea abies) and larch (Larix decidua) (Schütz et al. 1998). How the resulting forests in the ‘Wald-Freiland-Wald-Zyklus’ will be opened up by large herbivores is not explained by Krüsi et al. (1995). Apart from possible avalanches, there does not seem to be any indication of a factor that may affect such openness in the short term. As soon as a forest of mountain pine has settled, natural cyclical decline dynamics will ensure that locally, old trees disappear from the forest canopy. The germination and growth of young mountain spruce is related to the degree of openness of the forest canopy; the more open this is, the greater the increase of young trees will be. The germination-promoting impact of deer feeding is of considerably less importance here, and a far greater number of trees will be able to grow up undisturbed among the fallen trees. The rejuvenation of the forest keeps pace with its decay (Krüsi et al. 1995). The Norway spruce around the national park do not suffer from the high feeding pressure, either, and the effect of large herbivores on these trees is limited (Krüsi et al. 1996). In the Swiss National Park, a number of processes that are essential to a natural situation are lacking: the process of predation, and that of fluctuations in the amount of food available to herbivores. Large predators like wolf, bear and lynx are lacking, so that the number of herbivores is largely determined by the amount of food available. This factor may be very variable in nature, and have its effect on the size of the population; winter is a particular time of stress with regard to this. The deer in the current National Park are able to avoid stressful food situations by leaving the National Park area in summer or winter in order to seek food elsewhere, in the farmers’ meadows or in clearcut areas in cultural forests; deer who constantly stay in the central part of the Na-

317


318


Photo 37. The Stabelchod alp in the Swiss National Park, autumn 1998 (Photo: J. Bokdam). No agriculture, cattle breeding, silviculture or hunting has taken place in this nature reserve since 1914. This picture shows how forest renewal, progressing from the forest edge on the left, is taking place in the open field.

tional Park have a 10 % lower body weight than those who leave the Park temporarily (Blankenhorn 1977). In fact, this nature reserve is not a closed ecosystem. Unlike inside the National Park man can hunt deer outside the reserve, causing deer that take refuge inside the National Park to be culled when they leave it. Population crashes, such as occur regularly in ecosystems that are managed naturally, have not (yet) taken place in this area. In addition to this, the essential predators of red deer and chamois do not occur here. Research in the Bia³owie¿a Forest has shown that, apart from the food factor, predation by the wolf and the lynx are important mortality factors (Jêdrzejewska & Jêdrzejewski 1998). Recent research in Yellowstone National Park (United States) has shown that wolves, after an absence of 70 years, have not only decimated the population of wapiti deer (Cervus elaphus), but have also indirectly enabled the formerly heavily browsed willows and aspen to



develop into trees at last (Ripple et al. 2001, Larsen & Ripple 2003, Ripple & Beschta 2005a). If the Swiss National Park constituted a closed ecosystem, from which the deer and chamois would be unable to escape and predation would be present, the population size of these animals would probably fluctuate much more. Lesser feeding pressure will be beneficial to the advanced growth of mountain pine. If the current density of deer (17–26 animals/100 ha) were to decrease to one third (6–9 animals/100 ha), the forest would extend four times as quickly (Krüsi et al. 1998). If ideas about the reintroduction of the wolf, bear and lynx, also in the east of Switzerland, where the national park is situated, were to become reality, the situation there would become a lot more natural and more clarity would ensue. Beside the effects of large herbivores on the extension of the forest, the mountain pine itself also has a disadvantage with regard to its distribution. Apparently, this tree is unable to disperse across long distances, since forest extension takes place exclusively immediately adjacent to the forest edge (Krüsi et al. 1998) (see Photo 37). Seed dispersion and settlement, therefore, always take place from newly developed forest edges and at a limited distance from these. Such slow distribution has also been shown in other tree species, such as the white spruce (Picea glauca) in Canada (Timoney & Peterson 1996).

Summary This chapter deals with the possible impact of large herbivores (not just the aurochs, and not only in Europe!) on the growth of trees and forests. Wherever possible, situations have been sought in which man does not exert any influence on the ecosystem. To add to what was already written in Ch. 9.1.1 about the natural landscape of Europe, a description is given of the appearance of the forest landscape at Bia³owie¿a (East Poland) in the early 19th century. In addition to this, descriptions are given of the landscapes that prevail in areas comparable to Europe, the east of the United States and South-

319


320


east Siberia, from a period in which man still hardly exerted any influence there. The natural landscape in the latter two areas basically concurs with that of Europe. Since the population density of bovine animals may be directly connected to the impact of their feeding on the development of trees, the population density of the aurochs in its last living area in Central Poland was ascertained, and compared to the densities of a number of closely related, still-existing bovine species. In general, archaeological bone finds in Europe indicate a relatively low aurochs population density. The population density of the aurochs is likely to have varied a lot, depending on the openness of the forest. The large herbivores include in particular a few animals that are allegedly capable of shaping the forest landscape by way of their feeding. The supposed impact of two of these, the European bison in Europe and the forest elephant in Africa, are discussed in more detail. Research into their impact on the forest reveals this to be very slight under natural circumstances, and to have no disturbing effect on the development of the forest. What is true of the European bison and the forest elephant generally also applies to other large herbivores. Only the beaver is able locally to exert a great influence on the forest growth. The practical example of the natural ecosystem in Wood Buffalo National Park (Central Canada) was used to ascertain whether feeding by large herbivores affects the establishment of forest. Research in this area has revealed that the absence of trees in certain areas is not caused by the large herbivores (bison, deer, beavers), but by the water regimen of the large rivers and, to a lesser extent, by natural forest fires. The nature and scale of river floods cause extensive, almost treeless wet grassand sedge lands, which in summer and winter constitute the foraging area for the bison. The final paragraph, ‘Discussion’, highlights the conflicting opinions about the impact of large herbivores and deals in more detail with specific related issues.



Photo 38. Wall painting in the Lascaux Cave (France), depicting an aurochs cow with a reddish brown coat and a black head and neck.

Photo 39. This Jersey bull has a coat colour (blackish brown with a fairly coloured eelstripe) that closely resembles that of the original aurochs. The fair poll and ears are domestication features. Not only in so-called ‘primitive’ cattle can aurochs features still be seen, but also in modern breeds like this one (Photo: IVO ‘Schoonoord’, Zeist).

321


322


Fig. 46. A map from 1792 (by K. de Perthées) of an area in Central Poland. In this map the outlines of the last living area of the aurochs (see Fig. 10) are rendered in yellow. The dark grey areas are forests, the light grey ones are fields and meadows.

Photo 40. Spanish fighting bull in the surroundings of Santa Cruz de la Sierra (Extremadura, Spain). This bull has a number of features of the aurochs bull: a deeply blackish- brown coat with a pale eelstripe, a light snout and forward- and inward curving horns (Photo: M. Bunzel-Drüke).

Black Cyan Page 323

THE BREEDING-BACK EXPERIMENT OF THE HECK BROTHERS

10. THE BREEDING-BACK EXPERIMENT OF THE HECK BROTHERS 10.1. Cattle breeds In the course of the thousands of years following the domestication of the aurochs and as a result of the continuous selection in favour of milk yield, meat production and draught capacity, almost 1,000 cattle breeds and -varieties developed up to this day65. These breeds show a tremendous variety of fur colours, horn sizes, horn shapes, body sizes and udder sizes, although they are actually all descendants of animals that evolved from the aurochs some 9,000 years ago, in the Middle East and Pakistan. After their initial introduction, the genetic composition of the domesticated cattle that were imported to Europe by the first cattle breeders is very likely to have remained unchanged by crossbreeding with the wild congeners that occurred locally. In fact, there would have been little sense in crossbreeding the imported cattle, which had already undergone strenuous selection as to size, early maturity, tractability and production, with animals that might counteract those characteristics. In many places, occasionally domesticated cattle managed to escape in spite of their continuous tending by man, and returned to a feral state. There are numerous reports of this. In Europe, examples include the Mostrenca cattle at the Doñana National Park (Lazo 1995), the Betizu in the Basque Country (Van Vuure 1998), the Chillingham cattle in North England (Hall 1988) and the cattle of Swona (Orkney Islands; Hall & Moore 1986) and in the Camargue (Schloeth 1958). Outside Europe, such cattle occur in the National Park of El Kala (North Algeria; see Photo 41), on islands near New-Zealand, on an island of the Seychelles, on Hawaii, the Falkland Islands, the Aleutian Islands, in Colombia, Japan (the Island of Kushinoshima) and various areas in the United States (Hall & Moore 1986). The Island Amsterdam, in the Indian Ocean, also hosts such a population (Guintard & Tardy 1994).

65

The completest enumeration of cattle breeds, -populations and -varieties may be found in Mason (1996).

323

Black Cyan Page 324

324


Although not all these populations are equally wild and unattended, this does show that domestic cattle may easily turn wild and adapt to a natural food supply. Domestic cattle may vary strongly from their wild ancestors in appearance, but their digestions have probably remained largely unchanged (see Ch. 9.1.3). Almost all the foreign cattle populations mentioned here are of European descent, since European cattle breeds were transported all across the world. As a result of this, physical features of European cattle, and also possible aurochs characteristics such as the eelstripe, may be found anywhere in the world (Joshi & Phillips 1953; Joshi et al. 1957; French et al. 1966). The Criollo cattle of South- and Central America and the Texas Longhorns from the south of the United States, for example, originally came from Spain and Portugal, respectively. In the 19th century, the eelstripe of the Criollo cattle was reported to be ‘very characteristic’ (Dobie 1980, p. 22). The Heck brothers used exclusively European cattle breeds (the so-called ‘taurine’ breeds) for their breeding-back experiments; in view of that fact, a

Photo 41. Domestic bull gone wild in the El Kala National Park, North Algeria (Photo: K.J.M. De Smet). As has been shown for many countries, domestic cattle can run wild and survive on natural food without man’s help quite easily.

Black Cyan Page 325


global survey of these breeds is provided below. With the help of albumin analyses of domestic cattle that descend from the aurochs, i.e. taurine as well as zebu breeds, Baker & Manwell (1980) divided these into seven groups, based on their kinship. One of those groups, that of the European cattle breeds, may subsequently be subdivided into seven groups66. A. North-European: Scandinavian and English breeds, including Finnish cattle, Swedish and Norwegian red cattle, Fjäll cattle, Aberdeen Angus, Galloway, Scottish Highland cattle, White English Park cattle (e.g. Chillingham cattle) (see Photo 42), Shorthorn, Hereford and Longhorn. B. Pied Lowland: Dutch, Belgian, Danish, German, Polish and Russian Black Pied and Red Pied cattle, as well as Groningen Whiteheaded and Belgian Blues, among others. C. European Red Brachyceros: German, Danish, Polish, Latvian, White Russian and Bulgarian red cattle, including Angler and Rotvieh. D. Channel Island Brachyceros: Mainly Guernsey and Jersey. E. Upland Brachyceros: These breeds occur mainly in the Alps and Balkans and include Simmenthal, Braunvieh, Gelbvieh, Werdenfelser, Brown Swiss, Allgäuer, Montafoner, Buša, Rhodope, Corsican and Sardinian cattle. Breeds in North Africa (Algerian Atlas cattle) and the Middle East (Syrian and Egyptian cattle) are related to this group. F. Primigenius-brachyceros Mixed: These are crossbreeds of Steppe cattle and Upland cattle, including Charolais, Limousin, Piemontese, Bulgarian and Romanian brown cattle. G. Primigenius: Mainly Spanish, Italian and Balkan breeds, including Spanish and Portuguese fighting cattle, Chianina, Maremmana, Romagnola and Steppe cattle (Podolic cattle). The Criollo and the Texas Longhorn in America are related to the Iberian breeds. Fig. 47 shows where the two most important domestication centres (of the taurine and of the zebu breeds) used to be, as well as the kinship between the seven subgroups listed above. Of these seven groups, the Primigenius cattle are considered the most original, and seen as cattle that still show the most aurochs characteristics, such as late maturity, specific horn shape, fur colour, body size, udder size and sexual dimorphism. This does not mean that other breeds no longer show any au66

Nomenclature of these groups is based on Baker & Manwell (1980).

325

Black Cyan Page 326

326


Fig. 47. The domesticated descendants of the aurochs can be roughly divided into two groups, taurine cattle and zebu cattle. This picture shows where these originated and into which areas they distributed (after Manwell & Baker1980 and Baker & Manwell 1980, adapted). 1. Area of origin of taurine cattle. 2. Area of origin of zebu cattle. A. North European breeds B. Pied Lowland breeds. C. European Red brachyceros breeds. D. Channel Island brachyceros breeds. E. Upland brachyceros breeds. F. Primigenius-brachyceros mixed breeds. G. Primigenius breeds. ? A probable alternative domestication centre (Troy et al. 2001, Gautier 2002). (Please see the text for a more detailed explanation of A-G)

rochs characteristics. Highly upgraded cattle such as Frisian cattle (see Photo 12), Brown Swiss and Jersey (see Photo 39) also still display the horn shape and fur colour characteristic of aurochs. A short description of the main cattle breeds involved in the breeding-back experiment of the Heck brothers or in later experiments is given below. Some dairy breeds that were also involved will only be

Black Cyan Page 327


mentioned briefly in Ch. 10.2. Unless indicated otherwise, the data for the survey below stem from French et al. (1966) and Felius (1995), as well as from the author’s own observations. Scottish Highland cattle. (See Photo 27) This late-mature meat cattle was developed into a breed in the late 18th, early 19th century. It is longhaired and long-bodied. The colour is a solid light- to dark brown, sometimes black. The average withers height is 128 cm in bulls, 115 cm in cows. This cattle breed had long slim horns, which grow outward and up and sometimes backward; rarely in the shape of those of the aurochs. The udder is enlarged. Chillingham cattle67. The origin of this breed is unknown. It has been kept in a feral state in a fenced-off area in the north of England for centuries. The colour is white (sometimes with black spots), with black (sometimes red) ears and a black snout. The average withers height is 120 cm for bulls and 110 cm for cows. The curved horns are positioned high, not forward pointing. The udder is enlarged. Steppe cattle. (See Photo 16) This group of late-mature cattle owes its name to the Ukraine steppes. They probably came over from Central Asia in the Middle Ages. There are several types, from the Ukraine to Hungary. Their colour is light- to dark grey, sometimes black. The forehand is darker than the rest of the body. The calves are born a reddishbrown colour and change to grey at six months. The shoulder height of Hungarian Steppe cattle is 150 cm in bulls, 138 cm in cows. The horns of these draught- and meat breeds are long, protruding to the sides and often lyre-shaped. They are long-bodied. The udder is enlarged. Corsican cattle. The origin of these cattle, which live on Corsica, is unknown. It is quite similar to Sardinian and North African cattle. These days, crossbreeding with other cattle breeds has produced all kinds of colours (Quittet & Denis 1979), such as black, blackish brown, reddish 67 Beside Chillingham cattle, there are various other breeds of ‘White Park cattle’ in England, which have different characteristics.

327

Black Cyan Page 328

328


Photo 42. White English Park cattle in the former East Berlin Zoo. In several places in England, among others in Chillingham, white cattle (in several types) have been held in parks for centuries. (Photo: T. van Vuure).

brown, black- and red pied and grey. Eelstripes also occur. The shoulder height is 130 cm for bulls and 115 cm for cows. The curved horns of this draught- and dairy breed are positioned high. The udder is enlarged. Camargue cattle. The origin of this breed is unclear. It has either occurred from the Middle Ages onward (Felius 1995), or the population that lived there in the early 19th century was obliterated by a disease and replaced by a different cattle breed, the Salers (Moll & Gayot 1860). It lives in a feral state and is used for bullfights. The colour of both sexes is black or blackish brown. The height at the withers is 130 cm in bulls and 120 cm in cows. The curved horns are positioned high. The udder is small.

Black Cyan Page 329


Spanish fighting cattle. (See Photos 10 and 40) Up to the 18th century, bulls were selected from local populations to be used in the bullfights (De Cossío 1943). In the 18th century, specific breeding with selected animals started to take place. Selection and breeding were based exclusively on ‘fighting spirit’. As far as physical characteristics are concerned there is no really uniform breed of fighting bulls; rather, there are various breeding stocks. Colours vary widely among the different breeding farms (Rodríguez Montesinos 1994), though black is most common among the bulls, dark brown for the cows. Reddish brown, white, grey, red pied and black pied animals also occur, as do eelstripes, both light and dark. The height at the withers is usually between 120 and 130 cm but may range between 110 and 140 cm depending on the breeding farm. There are long-bodied as well as square-set animals. The shape of the horns may also vary strongly depending on the breeding farm. The horns, which are between 50 and 60 cm in length, occur in various shapes and positions. The aurochs horn shape, forward-pointed and curved inward (called ‘brocho’ in Spain), occurs regularly. The udder is small. Chianina. (See Photo 48) These draught- and meat cattle are among the earliest European cattle breeds, and allegedly existed already during the Roman period. Their build is square, their fur colour white. Initially, the calves are reddish brown. The shoulder height is 150-160 cm in cows and 160-175 cm (sometimes over 180 cm) in bulls. The horns are rather small and curved forward. The udder is enlarged.

Summary In the course of centuries, domestication of the aurochs resulted in hundreds of different domestic cattle breeds. Wild populations occur of some of these. European domestic cattle breeds are divided into seven groups on the basis of albumin similarities. In addition to this, short descriptions

329

Black Cyan Page 330

330


are given of the seven cattle breeds involved in the breeding-back experiments discussed below, as well as in other such experiments.

10.2. The breeding-back experiment Well before the extinction of the last aurochs in the forests at Jaktorów, this animal species had become largely forgotten elsewhere in Europe. After 1627, the memory of it disappeared completely. As a result of bone finds, discussions about the possible historic existence of the aurochs ensued at the end of the 18th century. In the beginning of the 19th century and in the course of that discussion, some felt the urge to recreate the animal in its original shape. In an article in 1835, the Polish zoologist Jarocki already argued in favour of an attempt to achieve this (£ukaszewicz 1952). The distribution of suitable bulls and cows in large forest areas, the first years in the absence of any wolves to give the cattle the chance to become accustomed, might possibly lead to the return of the original species in the long term. According to Jarocki, creating the correct, natural circumstances would turn domestic cattle into aurochs. As far as is known, this experiment was not carried out at the time. The idea to revive the ancestor of domestic cattle persisted, however. Around the transition of the 19th into the 20th century, when an increasing number of bone finds were done and more and more publications about the aurochs started to appear, a clearer image of the animal’s appearance emerged. Partly as a result of these developments, practical experiments to try to revive the aurochs actually started to take place. In early 20th-century Germany, the brothers Heinz and Lutz Heck carried out such experiments. Their father, who was the director of the Berlin zoo, had gathered an extensive collection of cattle breeds from many countries, which provided the brothers with a reasonable survey of the possible breeds and characteristics. In the 20s and 30s of the 20th century, with the use of cattle from this collection added to by animals obtained elsewhere in Europe, crossbreeding experiments took place which, according to the brothers, would lead to the ‘return’ of the aurochs.

Black Cyan Page 331


331

According to Heinz Heck (1951), the experiments were inspired mainly by the brothers’ curiosity to find out what the aurochs had actually looked like, and also to show this species to the public. In addition to this, the difference between the aurochs and the European bison could be clarified in this manner, and this was a matter of considerable interest at the time. In addition to this, revival of an extinct species might somehow make up for the loss of all the other species that had already been exterminated by man. As a basis for the crossbreeding and the selection of cattle, the brothers used their own individual images of the aurochs. The information for these images was obtained from early descriptions and pictures of the aurochs (rock paintings, descriptions by Schneeberger, Von Herberstein and others), as well as from observations and interpretations of characteristics of still-existing cattle breeds or individual animals. From these sources, both Lutz and Heinz Heck composed a description of the appearance of the aurochs. In Table 15, these are compared both with each other and with the conclusions drawn in Chapter 7 of this book. Table 15. Comparison of a number of physical characteristics of the aurochs, based on the writings of Heinz and Lutz Heck and the conclusions drawn in Chapter 7 of this book.

Characteristic The appearance of the aurochs accord ing to Heinz Heck (described in Heck, H. (1934), unless specified otherwise)

The appearance of the aurochs accord ing to Lutz Heck (described in Heck, L. (1934), unless specified otherwise)

The appearance of the aurochs on the basis of the results found in Chapter 7 of this book

Fur colour

†† Black. Thin short summer fur, of a lighter colour than the rougher winter coat.

†† Deep blackish brown to black. In winter, the fur is likely to have been longer and denser than in summer.

†† Shiny black in summer; the longer winter fur flowing and velvety.

Black Cyan Page 332

332


Table 15. Continued

68 69

Rather wide eelstripe along the back, white in the middle, yellowish and reddish yellow at the sides68

On the back, a more or less wide reddish marking (‘saddle’). Red eelstripe along the spine.

Along the length of the back, a narrow stripe of a lighter colour, probably greyish.

Longer, curly, reddish yellow hair on the forehead69.

–

Black curled hairs between the horns and on the forehead.

The area around the snout was white.

Lighter-coloured or white The area of the snout was light-coloured. area around the snout.

Light-coloured area around the eyes.

–

–

‡‡ Reddish brown, with a white belly and inside of the thighs. In winter, the sides of the neck were blackish (Heck 1980).

‡‡ Reddish. Varying from a light (grey) to a darker colour (black). Thin, short summer fur, of a lighter colour than the rougher winter coat.

‡‡ Reddish brown. Black in rare cases.The winter coat is likely to have been longer and denser than the summer fur.

The tail tip was black.

–

–

The area of the snout was white (more clearly so than in the bull; Heck 1980).

–

–

In 1980 he describes it as ‘a light, yellowish-white eelstripe’ (Heck 1980). In 1980 he gives this description: ‘between the horns there are often longer, brownish hairs’ (Heck 1980).

Black Cyan Page 333


333

Table 15. Continued

Horns

– Light-coloured area around the eyes (more clearly so than in the bull; Heck 1980).

–

– The head was lightcoloured (Heck 1980).

–

Calves Calves Red; they changed In the first 4 months, colour after 6 months. the calves of both sexes had the same colour (a dark reddishbrown).

Calves Reddish brown. Within half a year, the male calves would turn the colour of adult bulls. The female calves kept their reddish brown colour.

Outside Europe In Egypt, the aurochs had a whitish saddle on its back.

Outside Europe In Egypt, the aurochs had a reddish saddle on its back. In North Africa the aurochs was a lighter red, and of a lighter colour than in more northern areas.

Outside Europe There is no reason to assume that the colour of the aurochs outside Europe was clearly different than inside.

The thickest part was white, turning into a bluish colour and with a black tip.The horns grew sideways, then forward and then up. There was considerable variation in this aspect.

‘As they may be seen in Spanish fighting cattle, Camargue cattle, Corsican cattle, English Park cattle and Watussi cattle’.

The horns of the aurochs always showed a characteristic shape and position. The spirally curved horns were pointed forward and curved in. The horns were lightcoloured with dark tips.

Black Cyan Page 334

334


Table 15. Continued

Size

At least that of the larger cattle breeds, i.e. bulls of 800-900 kg and cows of 500-600 kg. The largest subspecies were almost 2 m in height (Heck 1980).

Between the smallest and largest domestic cattle breeds, i.e. bulls of 250-350 kg and cows of 200-250 kg70.

The bulls had a shoulder height between 160 and 180 cm.The withers height of cows was around 150 cm.Body weight possibly like that of European bison and banteng.

Udder

Small; disappeared after 6-8 months’ feeding.

As in the Spanish fighting bulls, the Corsican cattle and the Zebu cattle.

Small, hardly visible.

Body shape

Fairly short and high on – the legs. Straight back. Slightly raised pelvis and tail join.

Since the height of the withers was approximately equal to the length of the trunk, the animals were squarely built. The withers were barely or not at all raised.

Comparison of these descriptions shows that the brothers really did not agree on all the aspects of the extinct aurochs’ appearance. In the first place there was a discrepancy in the colours they attributed to the eelstripe and the ‘saddle’. Heinz assumed a white eelstripe with coloured sides of his own invention and lighter-coloured hairs between the horns. Lutz retained the notion of a red eelstripe, the way he had sometimes seen it in Spanish fighting bulls. For the European aurochs, Heinz did not assume a lightcoloured back (‘saddle’), though he does mention this for the Egyptian aurochs. Lutz, on the other hand, assumes a reddish saddle. For the NorthAfrican aurochs, he thought this would have been lighter than for animals 70 He appears to have gained a different opinion in 1952, when he writes that the aurochs weighed between 600-800 kg and that the shoulder height was between 170 and 180 cm (Heck 1952b). How this was established is not reported.

Black Cyan Page 335


from the more northerly areas. With regard to the colour of the cow the brothers did not quite agree either. Both mentioned a reddish/reddish brown colour, but Lutz assumed more variation, both to a grey and a black colour. How Heinz came up with the black tail tip for cows is a mystery; this is true of several aspects mentioned by the Heck brothers, however. With regard to the horn shape, Heinz gives a somewhat more detailed description than his brother, who indicates a more global shape, the way he saw it in certain cattle breeds. As to the weight, their opinions clearly varied in 1934: Heinz refers to a weight of at least the larger cattle breeds, while Lutz compares the aurochs to the medium-sized cattle breeds. He reports that heavier animals may have occurred, however, and illustrates this on the basis of the ‘Auerochs’ that was shot in Brandenburg in 1595, and which weighed 700 kg (’14 Zentner’). This ‘Auerochs’ was no aurochs, however, but a European bison (Szalay 1938, p. 45, Kühlhorn 1955). For his description of the aurochs and still-living cattle breeds, Lutz provided a survey in which he outlined to what extent a number of cattle breeds, as regards specific characteristics, were still similar to the aurochs (Heck, L. 1934, p. 289). In this survey, which is given below (Table 16), he has indicated for the various cattle breeds if their specific characteristics concur with aurochs characteristics (=), whether they have become smaller or less pronounced (‘Minusmutation’) (-), or have become larger or more pronounced (‘Plusmutation’) (+) Table 16. Survey composed by Lutz Heck (1934), indicating to what extent certain aspects of a number of cattle breeds were still like those of the aurochs (see text for explanation).

Aurochs Spanish fighting cattle Camargue cattle Corsican cattle Scottish Highland cattle English Park cattle Swedish Fjäll cattle

Horn

Body weight

Colour

Fur

Udder

= = = + —

= = = = = = -

= = + = + +

= = = = + = =

= = = + = = +

335

Black Cyan Page 336

336


Heck’s distribution of plusses and minuses is not always unambiguous: he considered the horns of Camargue cattle equal to those of the aurochs and those of English Park cattle smaller, although there is hardly any difference between the horns of these two breeds. That the colour of Camargue cattle would get a plus is understandable, since both the bulls and the cows are black. Why the colours of Scottish Highland cattle (light- to dark brown) and Swedish Fjäll cattle (white) were given plusses is less clear, however. A strange aspect of this survey devised by Heck is that it would seem to indicate that Spanish fighting cattle are like aurochs in every aspect. The horn shape, body size and colour, as well as the fur and the size of the udder of aurochs and Spanish fighting cattle are the same, according to Lutz. If Spanish fighting cattle were like aurochs in every respect, therefore, one might begin to wonder why Lutz still started his crossbreeding experiments to revive the aurochs. He might have spared himself the effort, since the animal was still alive in Spain anyway. The reason he did not just limit himself to Spanish fighting cattle may well be that, according to his own reasoning, there was no cattle breed that had retained all the characteristics, including the invisible ones, of the aurochs. Obviously, Spanish fighting cattle were not quite like aurochs; Camargue and Corsican cattle were also used, therefore, to be able to have as many characteristics as possible, also non-visible ones, at his disposal during selection. Genes that had disappeared in the one breed might still be present in the other. An excessive number of too different breeds would only confuse matters, however, and make selection more difficult. How Lutz managed to select in favour of such invisible characteristics during his crossbreeding experiment is unclear. If such selection is impossible or does not take place, however, it makes no sense to seemingly involve these invisible characteristics by using many different breeds. Initially, Lutz mainly used only three breeds: Spanish fighting cattle, Camargue cattle and Corsican cattle. According to him, these breeds had all the physical aspects and characteristics of the aurochs (Heck, L. 1934). Heinz started with a larger number of more diverse cattle breeds, to gather as many characteristics and features of the aurochs as possible in that way. He did not only use breeds that may be called ‘primitive’

Black Cyan Page 337


(Corsican cattle, Hungarian Steppe cattle) and possess certain aurochs characteristics, but also breeds whose appearance is far removed from that of their wild ancestor (Black Pied Frisian, Angler). Both Heinz and Lutz selected in favour of specific physical characteristics in their crossbreeding products, such as colour, horn shape and such. Characteristics such as able-bodiedness, resistance against disease, strong legs or size differences between bulls and cows, which could be found in aurochs, were not taken into account during the selection procedures. It looks as though the brothers thought such aspects would return ‘naturally’ in the final product. Heinz in particular assumed that the entire domestic cattle population still in existence had retained the complete genetic make-up of their wild ancestor. He described the experiment as ‘mixing cattle breeds in a melting-pot’, which process, after selection by man, would eventually yield the original, entire aurochs. In 1921, Heinz Heck began his version of the experiment in the Hellabrunn zoo (Munich) with the use of his father’s cattle collection. Characteristic for both this and Lutz’ experiment was the two brothers’ extreme vagueness about their reasons for crossing certain heads of cattle, and about the criteria they used to judge the prodigy possibly to be used for further breeding. Both indicate only globally the breeds that were used, and the moment at which, in their opinion at least, the aurochs returned. Heinz started mixing a ‘Black Pied Lowland bull’ with an Angler cow (Heck, H. 1934) (see Table 17). The Angler is a North German, reddish brown, slightly built dairy breed (Felius 1995). At the same time, a ‘Podolic Steppe’ bull was mixed with a Scottish Highland cow. Podolic Steppe cattle (which also include Hungarian steppe cattle) are grey with relatively large horns, and originally stem from the southern Ukraine (Felius 1995). The cow born from the first crossing was mixed with the bull from the second. The bull that resulted from this was mixed with a Hungarian steppe cow, which produced ‘Nordland’. This bull was crossed with a Corsican cow, yielding the bull ‘Primus’ that, according to Heck, ‘already clearly showed the desired aurochs characteristics’. At best, this may have indicated the light snout, since neither the small horns of

337

Black Cyan Page 338

338


the still young animal nor its rather drab fur colour, which betrayed its Hungarian steppe ancestor, would remind anyone of aurochs. ‘Primus’ was subsequently crossed with his mother, the Corsican cow, and the bull ‘Glachl’ was born as a result in 1932. This is the animal that Heinz considered the first bred-back aurochs, since it allegedly had the exact colour of aurochs, and its size and horns also seemed to be turning out right. More of a thorough description is not provided, and the accompanying photos do not give a clear idea, either. Primus was subsequently crossed with his mother, and a nameless female calf was born in 1933, the second ‘bred-back aurochs’. The survey below charts these crossbreeding procedures: Table 17. Crossbreeding as carried out by Heinz Heck (according to himself) in the 20s and 30s of the 20th century, in order to create a bred-back aurochs.

† Black Pied Lowland x ‡ Angler

† Podolic Steppe x ‡ Scottish Highland

Cow

x

Bull

Bull x Hungarian Steppe ‡ Corsican ‡ item Cow

x

x

† ‘Nordland’

† ‘Primus’ † ‘Glachl’

(Source: Heck, H. 1934)

If Heinz performed 5 crossing steps in about 12 years and 5 times 9 months are subtracted for the pregnancies, all the animals must have been about 2 years old. Animals of that age are not yet fully grown (this is usually at age 3 or 4), so they cannot really be judged for their eventual horn shape, udder size and bodily proportions, among other things. If these were selection criteria, the animals should have been older. Thus, the result desired by Heinz Heck was accomplished in a period of about twelve years, a result that had been achieved ‘incredibly

Black Cyan Page 339


quickly’, according to Heck himself. He considered it a miracle that all the aurochs characteristics, which had been distributed among the numerous cattle breeds from the time of domestication, had become concentrated in the two animals that had been created in the course of this experiment (Heck 1951). ‘Der Ur lebt wieder’ (‘the aurochs lives again’) was his enthusiastic reaction. Although he clearly seemed very satisfied with his bred-back aurochs, he still continued crossbreeding. Not only Primus and his relations were crossed; in 1934 Heinz also planned new crossings of English Park cattle, Steppe cattle and Scottish Highland cattle, in order to obtain a higher weight (Heck, H. 1934). These products of crossbreeding would subsequently be crossed with the slightly built Allgäuer71, probably because of the latter’s ‘deerlike’ character. In addition to this, he intended to exchange bulls with his brother Lutz, who was also carrying out an experiment around the same period. According to Lutz (Heck, L. 1952a), Heinz also used Werdenfelser72 cattle in his experiments, but Heinz himself makes no mention of this. From 1932/1933 onward, the progress of the crossbreeding carried out by Heinz is unclear, therefore, and can no longer be traced. In spite of all the praise, the breeding procedures had evidently not yet led to the desired result: from the moment the first so-called bred-back aurochs appeared the animals were continually tinkered with to improve them. Heinz did not report if any animals were culled between 1921 and 1932 because they did not meet the requirements. According to his own report (Heck 1951) there was not a single throw-back to the original cattle breeds used for the crossbreeding. Later on (Heck 1980) he had a different opinion, and seemed to consider such throw-backs as normal. As more generations of cattle grow up he expected fewer ‘imperfections’. A little later than his brother, and with an emphasis on different cattle breeds, Lutz Heck began his own experiment to breed back the aurochs in Berlin. From the start, he mainly concentrated on three breeds: Spanish fighting cattle, Camargue cattle (French fighting cattle) and Corsican cattle. He chose these breeds because he recognized in them the features and characteristics the aurochs had displayed, in his opinion. He took into account such features as horn shape, colour, build, 71 72

Grey-coloured dairy breed from South-German mountain areas (Felius 1995). The Werdenfelser is a brown dairy breed from Bavaria (Felius 1995).

339

Black Cyan Page 340

340


udder size and character traits (Heck, L. 1934) (see Table 16). The three breeds were to play principal parts in his version of the bred-back aurochs. How important he considered them is shown by the extensive descriptions he gave of his travels to Corsica, the Camargue and Spain. He reported on the cattle he observed there and stated his reasons for selecting certain animals in those areas. In 1927 he bought a bull calf and two female calves on Corsica, in 1931 a bull, two cows, a male- and a female calf in the Camargue. Finally, in 1932, from the breeding farm of Conradi in the area of Seville, he bought a bull and two cows. In the period 1929-1933, according to his own records (Heck, L. 1934, pp. 239, 266, 275), 27 calves were born, including a Camargue bull calf and a female calf, two Spanish fighting bull calves and three Corsican female calves and five bull calves of the same breed. As is shown in Table 18, the other calves were products of crossbreeding of the three breeds, as well as with other breeds. Table 18. Calves born in the period 1929-1933 as a result of crossbreeding experiments carried out by Lutz Heck.

Camargue † Spanish fighting Corsican † Camargue † Camargue † Spanish fighting Spanish fighting Spanish fighting Corsican † Corsican † Corsican † Corsican †

†

† † †

x x x x x x x x x x x x

Camargue ‡ Spanish fighting ‡ Corsican ‡ English Park ‡ Corsican ‡ Swedish Fjäll ‡ Corsican ‡ Corsican ‡ Swedish Fjäll ‡ English Park ‡ Steppe ‡ Montafoner ‡73

: 1 † calf, 1 ‡ calf : 2 † calves : 5 † calves, 3 ‡ calves : 1 † calf, 1 ‡ calf : 1 † calf, 1 ‡ calf : 1 † calf : 1 † calf : 1 ‡ calf : 4 calves : 2 calves : 1 calf : 1 calf

(Source: Heck, L. 1934)

In 1952 (Heck, L. 1952a, p. 118), Lutz reported that at the time, beside these cattle, he had also bred in ‘Angler, Allgäuer and other’ breeds. He did not clarify what the other breeds had been. They defi73

The Montafoner is a brown dairy breed from Austria (Felius 1995).

Black Cyan Page 341


nitely included Scottish Highland cattle, since his own records showed that 80% of the first 90 calves consisted of mixes of French and Spanish fighting cattle with Corsican cattle, in addition to which he obtained seven calves with English Park cattle, six with Scottish Highland cattle and four with Angler. He forgot to mention the calves he had obtained, according to his 1934 report, from Swedish Fjäll and Steppe cattle cows. In 1934, Lutz was still of the opinion that, apart from the three South-European breeds mentioned, no other breeds qualified for breeding. Swedish Fjäll cattle were unsuitable because they were dominantly hornless, English Park cattle because of their white colour, Scottish Highland cattle because of their long hairs and the position of their horns, which protrude sideways, and Steppe cattle because of their silver grey colour. His 1952 description (Heck, L. 1952a) of his version of the breeding-back experiment does not show whether the above-mentioned breeds that he had actually crossed had actually contributed to his bred-back aurochs, or indeed exactly which crossings had taken place. In addition to this, his brother and he also mutually exchanged cows and bulls, and nothing is known about the use of those animals (Heck, H. 1934, p. 15, Heck, L. 1934, p. 275). Whether any records were kept of the crossbreeding carried out by both brothers before and during World War II is not known. None have survived, in any case (M. Bunzel-Drüke in litt.), and since Lutz and Heinz have died, exact details about the extent to which any breeds contributed to the present Heck cattle will never be known. It is uncertain whether the brothers would have known this themselves, since their own inadequate description of their experiments, and the fact that no notes of these seem to have survived, prompt suspicion that they were not exactly strict about recording their findings properly. The two brothers agreed that their breeding results were very alike (Heck 1980, Heck, L. 1952a), which led them to conclude that ‘the fundamental principle of the breeding back was correct, since this led to the same result, albeit by different methods’. Lutz put this ‘great similarity’ into perspective, however, by stressing that specific differences were demonstrable up to that point (1952). He illustrated this with quotes from a letter by the English archaeologist Zeuner, who suggested that

341

Black Cyan Page 342

342


he knew what the two breeding stocks looked like. Also in a later publication, Zeuner (1963) himself did not clearly reveal how he knew where the animals of the Berlin stock were kept after World War II. In his opinion, the difference between the Berlin and Munich stock was clearly visible. Allegedly, the first was more like the Southwest-European aurochs, the other more like the Central-European one. He based this opinion on criteria that were rather vague and contestable, the more so since around that time, and still today, data about local varieties of the aurochs were lacking. The fact that Zeuner could distinguish several types of Heck cattle is not surprising. The genetic heterogeneity of the population offers every opportunity for that. The difference Werner (1976) makes between the two breeding stocks is not convincing either, since he based it on the dubious descriptions by Zeuner and the Heck brothers, and on the scarce photos from the 30s and 40s. To judge from those few published photos (in Heck, L. 1952a; see Photo 43) of animals in the breeding stock of Lutz Heck, there do seem to have been differences with the animals of Heinz, among others with regard to the size of their horns. The heterogeneity of the two breeding stocks and the unreliability of the eyewitnesses mentioned make comparison difficult, however.

Photo 43. A cow of the Heck cattle stock bred by Lutz Heck and released in the Schorfheide reserve (Germany). This pictures dates from 1944 (after L. Heck 1952a). It is unlikely that any of this stock has remained after the 2nd World War.

Black Cyan Page 343


Contrary to Heinz, who kept his animals exclusively in zoos and small wildlife reserves, Lutz chose to distribute his animals in larger wildlife areas, thus to regain ‘the characteristics of the wild animal’. In 1938, six years after the importation of the Spanish fighting cattle, a group of Heck cattle was distributed in the forest area of Rominten, in former East Prussia (Frevert 1957). The animals never really ran wild there, since they would go from one feeding place to another in winter. As this offered too much competition to the red deer, the pride of Rominten, in 1942 it was decided to take 16 of the 22 animals to the forests at Bia³owie¿a. There they multiplied without supplementary feeding in winter to 23 animals in 1944, the year in which the Russians conquered this area from the Germans. Another group of animals was distributed on the Schorfheide, near Berlin. All animals of the populations set out in the wild were lost as a result of the war situation (Heck 1952a, p. 123). Heinz Heck estimated the total number of Heck cattle that survived the war as between 37 and 41 animals in 1951; according to him, not a single animal of the Berlin stock survived (Heck 1951). This was contradicted by Lutz, however, who alleged that the larger part of the Heck cattle in 1952 consisted of animals from the Munich stock, with only a few Berlin animals (Heck 1952a). It remains unclear, therefore, whether the Berlin stock still lives on in the present-day Heck cattle, although it is certain that the animals of the Munich stock dominate. According to Lutz, after the war only the populations in West Germany remained, at Augsburg, Duisburg, Cologne, Munich, Munster and Neanderthal. As far as is known, neither Heinz nor Lutz practiced any more selection and breeding of Heck cattle after the war; this was left to the individual ideas of the then owners. The further progress of the post-war breeding is hard to trace, however, because of the inadequate records that were kept of this. Up to the late 50s, however, (Hungarian) Steppe cattle are still likely to have been bred in, and in 1952 a head of Watussi cattle, originally from Africa, was even allegedly bred in at the Duisburg zoo (M. Bunzel-Drüke in litt.). If the Heck brothers brought any consistency to their breeding experiments before the war, which is very doubtful, this was definitely entirely obliterated by the insights of the individual later owners. As it happens, because of a lack of good

343

Black Cyan Page 344

344


basic knowledge individual breeders have different ideas about the size, colour and horn shape of the original aurochs, and this is reflected in their breeding results. The influence of the Heck brothers in the world of the German zoos, which was so great before and during World War II, strongly diminished after this period. The parts that both, but particularly Lutz, had played during the nazi regime had done a lot of damage to zoological research in Europe and created a lot of ill will in scientific circles (Daszkiewicz & Aikhenbaum 1998). Many zoo owners removed the Heck cattle from their collections (Dathe 1980, Koehler 1952). An anonymous source suggests that around 1955, there were still about 50 head of Heck cattle in the Munich zoo (Anonymus 1955). According to the first official Heck cattle herd book, which was started in Berlin on 1 January 1980, the number of animals on 1 January 1979 was 88. This was probably too low an estimate, since a number of locations were missing from the list, such as the Antwerp and Han-sur-Lesse zoos. According to the present herd book, kept by the SIERDA74, on 1-1-2001 there were 597 registered Heck cattle in Belgium, Germany and France. The larger part of the present Heck cattle live unregistered in Dutch nature reserves. In 2004, their number was about 660 (J. Griekspoor pers. comm.).

Summary In the 20s and 30s of the 20th century, the brothers Lutz and Heinz Heck in Germany made attempts to revive the aurochs. They did this by crossbreeding several cattle breeds to which they ascribed aurochs characteristics, and by applying selection to the subsequent offspring. Each of the brothers used his own selection of breeds. According to their own reports, the desired result, the bred-back aurochs (Heck cattle), was speedily obtained and quite similar to the aurochs. Unlike Heinz, Lutz distributed his animals in the wild, in Germany and Poland. Lutz’ Heck cattle probably did not survive World War II, and only the animals bred by Heinz still exist. After the war, individual breeders applied their own ideas to the further breeding of Heck cattle. 74 ‘Syndicat International pour l’Elevage, la Reconnaissance et le Développement de l’Aurochs-reconstitué’ – International Association for the Breeding, Recognition and Development of the Bred-back Aurochs (formerly SIERDAH).

Black Cyan Page 345


Currently, Heck cattle are kept in many areas, including the Oostvaardersplassen in the Netherlands.

10.3. Evaluation of the breeding-back experiment The beginning and first development of the Heck cattle took place in Germany during the nazi regime (1933-1945). Outside Germany, these experiments were barely known. After the war, when the greatly diminished population of Heck cattle became more widely known and more criticism became feasible, the first critical noises were heard in Germany. They were aimed both at the way the basic data for the experiments had been acquired (Kühlhorn 1955, Koehler 1952) and at the appearance of the animals that had resulted from the experiments (Anonymus 1955, Herre 1953). The ease with which the Heck brothers had presented their evidence in support of their theory was denounced first of all, together with their careless use of historic sources and their incorporation of these in their descriptions of the original aurochs75. Criticism also focused on the carelessness, the ease and the speed with which they had carried out their experiments, and on the lack of clarity regarding the selection criteria they had applied. Their conceit and satisfaction with their ‘perfect’ end result, and the way in which they praised their two breeding stocks (‘Der Wildstier, der Ur, lebt wieder’ - ‘The wild bull, the aurochs, lives again’) failed to engender sympathy. The appearance of the Heck cattle the brothers had bred was another object for criticism. It was clearly different in many ways from that of their wild ancestor, although the brothers themselves were very content and seemed unable to notice any differences. The breeding-back experiments have obviously been an object of criticism for a long time. Oddly enough, neither of the brothers themselves ever expressed any criticism or doubt about their own experiment or that of the other, at least never in any written form. Both seemed undividedly satisfied and proud, although it would have seemed logical to have reasonable doubt about their methods and results. This is even 75

See Chapter 2 for a more detailed description of the use of historic sources by the Heck brothers.

345

Black Cyan Page 346

346


more so at present, with an increasingly clear idea of the aurochs and of the way in which the breeding-back was carried out at the time. If the experiment is subjected to a little closer scrutiny, first of all it becomes clear that the image the two brothers had of the original aurochs (as recorded in their descriptions, see Table 15) is actually an amalgamation of data from both old descriptions and personal observations of cattle breeds that still existed at the time. Using their own discretion, they combined these two sources into aurochs descriptions they deemed exact. Remarkably, however, their ‘exact’ descriptions differ. Heinz persists in describing a mainly white eelstripe, while Lutz describes the aurochs with a red eelstripe. Lutz mentions a red saddle, the way he assumed it looked on the Egyptian aurochs and in the fighting bulls he had seen in Spain. Heinz only mentions colour saddles from ancient Egypt, of a ‘whitish’ colour, as could allegedly be seen in some pictures (see Ch. 7.3). What prompted Lutz to attribute a red saddle to the Egyptian aurochs is a mystery. He does not give a source for his assumption, and ancient Egyptian pictures only ever show white, yellow and light brown saddles on what very probably were cattle that had run wild, which had all kinds of colours and markings anyway. His reasoning may have been that the Egyptian aurochs had a colour saddle Spanish fighting bulls sometimes have red colour saddles therefore the (European) aurochs had a red colour saddle. Heinz’ mention of a black tail tip, black neck sides, lighter-coloured hairs between the horns and on the forehead and a light-coloured head (in the cow) was prompted by his observation of existing cattle breeds, since old descriptions or pictures could not have inspired this. From Lascaux, a painting of a reddish brown cow with black head and neck (see Photo 38) is known, but to what extent this characteristic really existed and was widely distributed is unknown. The lighter hairs between the horns often occur in combination with a lighter colour saddle on the back. The characteristic ‘white belly sides and insides of the thighs’ was also probably observed in breeds that still existed. Old pictures do not show anything of the kind. As to the horn shape, neither of the brothers gives a correct description of the curves. Although clarity about the shape and position of the

Black Cyan Page 347


horns had been obtained already in the early 20th century (Von Leithner 1927, Ewalt and Laurer 1911, among others), neither Heinz nor Lutz apparently knew about this. The shape of the horns could not only be found in the literature; in the beginning of the 20th century, many skulls, including complete horn cores, had already been found, enabling anyone who wanted to see them and to conclude how aurochs horns were shaped. Neither description by the brothers mentions the typical forward-pointing, inward-curving horn shape. Heinz does mention the forward position, but not the inward curve. Lutz (1934) bases his description on the horn shapes depicted in cave paintings in the Abrigo de los Toros cave near Tormón (Valencia area, Spain), which he calls ‘true’ (see Fig. 48). The horns depicted there show all kinds of strange shapes,

Fig. 48. Drawings of aurochs in the Abrigo de los Toros Cave at Tormón (Spain) (after L. Heck 1934). Lutz Heck thought he might ascertain the shape of aurochs horns on the basis of such drawings. However, the curves of aurochs horns are difficult to render on a flat surface.

347

Black Cyan Page 348

348


however, and do not present a realistic image. Lutz actually remains rather vague about the horn shape, saying it appears in Spanish and Corsican cattle, Camargue cattle, English Park cattle and Watussi cattle. Of the horns of the latter, he says that they have ‘exactly the same curve and shape’ as those of the aurochs, only ‘very much enlarged’. Watussi cattle do have impressive horns, but the shape and position of these are nothing like those of the horns of their wild ancestor. The position of their horns is high, and they have a single curve that points outward and up. Also with regard to the size and weight, the Heck brothers differed in opinion at the time of the breeding-back experiment. Heinz focused on a weight that was twice as high as that of the animals Lutz had in mind, i.e. 500-800 kg against 200-350 kg. Later on, Lutz obviously changed his opinion; by 1952, he mentioned a weight of 600-800 kg (Heck 1952b). Lutz initially probably contemplated the three South-European breeds, since their sizes and weights are relatively limited. Later on he clearly adhered more to Heinz’ opinion. A reason was never given. The horns shape, colour and other aspects of the various animals used for the crossbreeding experiments were never exactly described by the brothers. Descriptions of the various breeds were given, but which individual animals were used and for what reason was never reported. The same is true of the selection procedures that took place during the breeding experiments: the brothers did not clarify why a certain animal would or would not suit, and why specific animals were used for further experiments. Since the two brothers initially started out with different cattle breeds and focused on different aurochs appearances, it is even more remarkable that their final products, according to themselves, looked exactly alike and exactly like the aurochs, and that no throw-backs took place to any of the bred-in ancestors. Photos from the 30s and 40s, which depict the Heck cattle of Lutz Heck, show that there were differences between the two breeding stocks, however. This is hardly surprising, since a different set of cattle breeds was used for each of these stocks. Closer contemplation of the still-existing Heck cattle, offspring of animals that resulted from the crossbreeding experiments carried out at

Black Cyan Page 349


the time, show that these are unlike that original aurochs in many aspects (see Table 19). The complacency of the two brothers may probably partly be explained by their possible unwillingness to admit failure of the experiment, which led them to consistently emphasise the professionalism of the initial argument in favour of the experiment, as well as the actual course of this experiment and the beauty of its result. For another part, it seems that their practical criteria for the desired appearance were very broad, so that in spite of the large variation in shapes and colours almost all products of their crossbreeding could be called ‘successful’. Yet another part could be explained by the brothers’ simple lack of thorough knowledge about the aurochs. Their reasoning about the appearance of the aurochs and about the correct methods to breed back such an animal is not exactly remarkable for its thorough scientific basis. Some data were haphazardly obtained from the literature, but the research was never very in-depth. What had been written or could be observed was often taken for granted without further study or critical contemplation. In spite of the fact that the horn shape of the aurochs had already become known from bone finds in the 20s, Lutz still derived this shape from the many forms that may be observed in the Abrigo de los Toros cave. Also Heinz reports that the position of the horns varied ‘not inconsiderably’, although he claimed to have seen a number of skulls with horn cores. He did not recognize, however, that aurochs horns, in spite of the ‘considerable variation’, as he called it, all had the same basic shape, be it with very small individual variations. A lack of basic knowledge, broad selection criteria and the complacency of the two brothers led to their excessive simplification of the breeding-back experiment, to a ‘simple mixing of a few breeds, selection, and the subsequent revival of the pure-bred, original aurochs’. It took Heinz 12 years at the most to revive the aurochs, according to his own accounts. Lutz took less than 11 years to distribute his bred-back aurochs at Rominten. Of those 11 years, the fighting bulls brought in from Spain only participated during the final 6. In such relatively short periods, it can never be possible to stabilise certain characteristics, which stem from such various breeds, together in one single breed (Koehler 1952), without abundant segregation of bred-in characteristics. As it happens, so many unde-

349

Black Cyan Page 350

350


sirable traits are mixed in together with the desired ones that it takes very long to remove these undesirables features and to stabilise only the desired ones in all the animals of the population homozygously. This is also apparent in the current Heck cattle, in which the traits of the breeds that were bred in during the 20s and 30s of the 20th century, such as their horn shapes, sizes and fur colours, are still clearly distinguishable. Lutz Heck must have known that not all the calves that resulted from the breeding of Heck cattle would have the desired colours, and that ‘throw-backs’ were bound to occur. It so happens that Frevert (1957) reports that shortly after the distribution of Heck cattle at Rominten (in 1938), calves were already being shot on account of their ‘wrong’ colours, i.e. ‘white and red spots’. Also today, after 70 years, Heck cattle still have such divergent colours (see Photo 44). In spite of all the severe criticisms one could make on the execution of these experiments by the Heck brothers, it may be argued in their favour that their ideas have greatly contributed to the return of the wild ox into the public consciousness, and that domestic- and wild cattle play a role in the conservation and management of particular nature reserves today. In a way, their thoughts and experiments were appealing and stimulating; the immediate cause for the aurochs research set down in this present book was a picture of a head of Heck cattle (see ‘Preface’). The Heck brothers tried to realise their fantasies, but did not take into account the necessary scientific framework.

Description of Heck cattle This section will provide a description of the Heck cattle that still exist, the offspring of probably only the breeding stock of Heinz Heck, to ascertain to what extent these current Heck cattle correspond to the image of the original aurochs; in other words, whether the breedingback experiment may be called a success on the basis of this. Descriptions of Heck cattle that take into account the exact numbers and percentages of the various horn shapes, fur colours, markings

Black Cyan Page 351


and body shapes of bulls and cows do not exist at present. Descriptions given by various observers only present a global image of the animals. As early as 1955, one observer concluded that the size of the aurochs as ascertained from skeleton finds was clearly different from that of the Heck cattle (Anonymus 1955). The difference in size between bulls and cows, characteristic of the aurochs and other originally wild cattle species, did not resemble that of their wild ancestors either. Herre (1953) concluded that the creation of the Heck cattle had not revived the aurochs, but that ‘scientifically worthless domestic cattle crossbreeds, by their similar fur colour and horn shape, only provide a vague impression of this impressive wild animal’. He calls their body shape ‘tonnig’ (‘tubby’), in the way of domestic animals. Moreover, he denies the wild nature ascribed to them by the Heck brothers. As to their being pure-bred, with regard to their physical features and the non-occurrence of throw-backs, he denies this by concluding ‘……that the pied colours of domestic cattle do not seldom reappear’. Herre also stresses the similarly small difference in size between the bulls and the cows, which is common in domestic cattle but not in aurochs. The abso-

Photo 44. Heck bull with a piebald coat in the Oostvaardersplassen reserve. After 70 years such a recessive feature may still become visible (Photo: H. Kampf).

351

Black Cyan Page 352

352


lute size of the Heck cattle, according to Herre, has no relation to that of the original aurochs either. He includes drawings of an aurochs bull and -cow, with respective shoulder heights of 200 and 180 cm. He compares them with Heck cattle in the Osnabrück zoo, whose withers height is 140 cm for bulls and 135 cm for cows. As to the size of the aurochs, Herre’s own estimate is actually rather high, since aurochs of such size probably never existed during the Holocene period. Guintard & Denis (1996; ‘Pour un standard de l’aurochs de Heck’) (see Table 19) take a somewhat more exact approach in their description of the Heck cattle. They provide a description of their morphology based on measurements and visual observations. Their report does not make clear of which animals in which nature reserves these observations were carried out. The characteristics described by them seem to represent the greatest common denominator, and to constitute the goal that breeders should focus on. With regard to the shoulder height, their measurements showed that of bulls to be 142 cm on average, and that of cows 131 cm76. The shape of the horns is described as ‘lyre-shaped high-positioned’ for the cow and ‘more widely spread and the tip pointed forward’ in the bull. The fur colour is basically reddish, ‘with a strong to very strong black colouring, with black legs’. The bull is generally darker than the cow and may have a blackish appearance. There is a whitish zone around the snout. Also according to these authors, the red colour, independent of the extent of the black colouring, remains visible in the hairs on the head and in the eelstripe, both in the bull and in the cow. As the reddish calves grow older, a progressive black colouring of the head, neck, shoulder, lower parts of the trunk, tail and legs appears, earlier in bulls than in cows. In this way, a reddish saddle remains on the back. The udder is little developed. Far from all Heck cattle fit this description. As a matter of fact, the Heck cattle population is generally very heterogeneous as regards their physical traits, and probably also with respect to their other bodily, non-visible characteristics, though these are less conspicuous. A more detailed description of the appearance of the current Heck cattle, which may be seen in various places in Europe, will be given below. 76 These sizes are a little smaller than those of the current dairy- and meat breeds in Europe, whose bulls and cows have withers heights of 150 and 140 cm, respectively (Frahm 1982).

Black Cyan Page 353


If the appearance of the Heck cattle is compared to that of the aurochs, the first thing that stands out is that their shoulder height is clearly less than that of their wild ancestors. The average height of the withers of the aurochs cow was 150 cm, that of the aurochs bull 170 cm. Heck cattle are about 20 (cows) to 30 cm (bulls) lower, respectively. The difference between the withers height of bulls and cows is less pronounced than in aurochs, whose cows’ shoulder height was 88% of that of its bulls, against 93% in Heck cattle. If there were the same difference in Heck cattle, the cows would have a withers height of 125 instead of 131 cm. Also among one of the largest domestic cattle breeds, the Chianina, the difference between the sexes is not as pronounced as in the aurochs. Chianina bulls attain an average withers height of 170 cm, and cows of 93% of that, i.e. 158 cm (Frahm 1982). If the horn shape of Heck cattle is considered, their horns are revealed to usually protrude sideways, pointing upward, sometimes with backward-curving tips. Occasionally, the tips curve forward slightly. Aurochs-type horns, i.e. forward-pointing and inward-curving at the same time, occur only very rarely in Heck cattle. The horn shape of Hungarian Steppe cattle may often be found in Heck cattle: widely protruding, lyre-shaped horns (see Photo 45). Guintard & Denis (1996)

Photo 45. Like this specimen, many Heck cattle show the characteristic lyre-shaped horns that originate from Hungarian Steppe cattle (Photo: J. Bokdam).

353

Black Cyan Page 354

354


report 51 and 48 cm for the average horn lengths of bulls and cows, respectively. Compared to those of the aurochs, these Heck cattle horns are clearly smaller. This is particularly true of the bulls. The sexual dimorphism, which in aurochs manifested itself clearly in the size of the horns, among other things, has all but disappeared in Heck cattle: the difference is only a few centimetres, against 20 to 30 cm in aurochs. Generally speaking, the udders of the Heck cattle may be termed small. Large udders occur regularly, however, clearly revealing the dairy breeds that were bred in at the time. Heck cattle are not only smaller than their wild ancestors; their physical proportions are also different. The shoulder height of the aurochs was almost the same as the trunk length, which gave the animal a more or less square build. In Heck cattle, the build is more rectangular, since the legs are shorter, relatively speaking. The average trunk length of the bulls is 169 cm, of the cows 151 cm (Guintard & Denis 1996); more or less the same as in aurochs and most other current cattle breeds (Matolcsi 1970). Since the legs are shorter, relatively speaking, the animals look rather long-bodied and short-legged, like most other domestic cattle (see Fig. 16). Occasionally, long-legged animals with a more square build occur. In addition to this, the difference between the (heavier) forehand and the (lighter) hindquarters, which may have been apparent particularly in the aurochs bull, is much less clear in Heck cattle, which gives the animals a more ‘squat’ appearance. The relatively short, broad head characteristic of domestic cattle may also be found in Heck cattle. The aurochs had a longer, relatively narrower head. The Heck brothers’ reasoning, that certain traits of the Heck cattle, such as ‘wild’ behaviour, resistance to diseases and certain aspects of social behaviour, as features that had ‘returned spontaneously’, argued in favour of the similarity of Heck cattle to the aurochs, underestimated the capabilities and hidden characteristics of domesticated cattle. Features such as the resistance to diseases, original social behaviour and wild characteristics, such as timidity and aggressiveness, may have been hidden in the various crossed-in breeds, to appear again at the crucial moment of a wilder existence. Such traits may reappear under the right circumstances. Clearly, therefore, this does not mean

Black Cyan Page 355


that they were created completely anew in Heck cattle. The fact that Heck cattle are relatively well able to survive in a feral state, and do not really suffer from diseases or ailments, proves the theory that many domestic cattle breeds are still able to cope with the food supply and extreme climatological circumstances that prevail in nature reserves. The use and functioning of Heck cattle in nature reserves will be discussed in more detail in Chapter 11. If, with the colours of the aurochs bull and -cow in mind (see Ch. 7.3), the colours of the current Heck cattle are considered, the conclusion is warranted that the difference in colour between the sexes is not anywhere near as clear and consistent as was the case with the aurochs. Heck cattle, both bulls and cows, may have very variegated colouring. In the cows, colours vary from light brown, through reddish brown and dark brown, to black. Black cows frequently occur (see Photo 46). Cows also regularly have eelstripes. The standard colour of the aurochs cow was reddish brown, rarely black. Eelstripes, narrow and grey in colour, only occurred in bulls, whose standard colour was a deep blackish brown.

Photo 46. Heck cow in the Ronde Put reserve (Belgium) (Photo: T. van Vuure). This cow has a black coat, an enlarged udder and the typical Heck cattle horns.

355

Black Cyan Page 356

356


In Heck cattle, the colour of the bulls varies from black/white, brown/ black or red/black pied, to completely black. Eelstripes may either be light (whitish or yellow) or dark (dark brown or reddish brown), and their width varies from a few centimetres to over 10 cm. They are often missing, however. Moreover, grey colouring of the hairs occasionally occurs in bulls, a trait that probably stems from the grey Upland cattle or Hungarian Steppe cattle that were bred in at the time. The grey colouring (discolouring) of the back, flanks and forehead also seems to stem from the Hungarian Steppe cattle. Discolouring of the back and flanks is also sometimes called a ‘colour saddle’, and occurs regularly. Such a saddle may vary in colour from grey to light brown and reddish brown (see Photo 47). The aurochs calf was usually reddish brown, rarely black. Some Heck cattle calves may be reddish brown, but their colours are very various, ranging from very pale brown to black. Heck cattle bulls as well as -cows may have a lighter-coloured area around the snout, which may vary in size and colour (white to reddish brown). It does not occur in many animals. The aurochs (bull) also had a light area around the snout, but its size is not known. In many Heck cattle, the hair on the forehead and between the horns is of a lighter colour, i.e. pale brown to reddish brown. As far as can be ascertained, the aurochs did not have a lighter-coloured area there; this area constitutes a discoloured zone that appeared at a later stage, of the kind that also occurred on the back and the flanks. On account of the absence of any marked similarity in size, colour and horn shape, among other aspects, Heck cattle cannot be considered to resemble the aurochs closely. Rather, they should be seen as a population of cattle in which a few aurochs characteristics may be found; a trait they share with many other cattle populations. In Chapter 11, more attention is paid to the question whether there is any difference in the functioning of Heck cattle and aurochs, and to the possible usefulness of the first species for purposes of grazing. For the sake of clarity, Table 19 once again compares the physical characteristics of Heck cattle with those of the aurochs.

Black Cyan Page 357


Photo 47. Heck cattle in the Slikken van Flakkee reserve (SW Netherlands). The left bull has an evenly black coat with a pale eelstripe, the right one has a so-called colour saddle of lighter-coloured hair (Photo: T. van Vuure).

Table 19. Physical characteristics of Heck cattle as compared with those of the aurochs. The data for the Heck cattle were taken from Guintard & Denis (1996) and personal observations. More details for Heck cattle may be found in Ch. 10.3, for the aurochs in Ch. 7.

Characteristic

Heck cattle

Aurochs

Withers height

††: 142 cm on average ‡‡: 131 cm on average

††: 170 cm on average ‡‡: 150 cm on average

Fur colour

The difference in colour between Clear difference in colour the sexes is often unclear. Bulls are between the sexes. generally darker than cows. ††: From black pied via red, ††: A deep blackish brown to brown and grey to black. Colour black. Along the back was a saddles. Lighter as well as darker narrow light-coloured eelstripe, eelstripes. Light-coloured zone probably greyish. Light zone around the snout, sometimes around the snout. also between the horns.

357

Black Cyan Page 358

358


Table 19. Continued

‡‡: From light brown via red, brown and grey to black. Lightcoloured zone around the snout and sometimes also between the horns. Eelstripes sometimes occur. Calf: Usually reddish brown, otherwise varying from light brown to black.

‡‡: Reddish brown, rarely black. Possibly a light zone around the snout

Horns

The (slightly) curved horns point upward strongly. Often lyreshaped. Very rarely as in the aurochs. Relatively short and thin.

Very characteristic and relatively stable as to shape and position. The horns curve forward and inward. Relatively long and thick.

Build

Because of the relatively short legs, the body is longish (rectangular) in shape.Little difference between the forehand and hindquarters.

Since the shoulder height more or less equals the trunk length, the body is squarely built.Clear difference between forehand and hindquarters.

Head

Relatively short and broad.

Relatively long and narrow.

Udder

Very variable in size. Both small ones and large ones occur.

Small. Hardly visible.

Calf: Reddish brown, rarely black.

Summary Quite soon after the development of Heck cattle, criticism on both the execution and the results of the breeding-back experiment began. It was aimed in the first place at the image the Heck brothers had formed of the aurochs, and which had a mixture of truth and (mainly) fantasy as

Black Cyan Page 359


its basis. Moreover, the brothers did not agree about this image. In the second place, there was criticism about the way breeding and selection had taken place, and about the speed with which the final results had been reached. The selection criteria were vague and rather broad, and no good records had been kept of the crossings, so that it is unclear which breeds contributed to the final product, and to what extent. The breeding-back experiment of the Heck brothers is thus characterised by an unprofessional and untransparent way of working. Comparison of the physical characteristics of Heck cattle and aurochs shows that there are few similarities between the two (see Table 19). Only the colour of the fur of some Heck cattle shows some similarity. Other traits, such as the horn shape and body size, do not concur.

359

Black Cyan Page 360

360


11. FINAL REMARKS, CONCLUSIONS AND RECOMMENDATIONS In the preceding chapters, on the basis of numerous data from several disciplines, an attempt was made to answer a number of questions about the aurochs and about Heck cattle. At the set-up of the Aurochs Project in the beginning of 1998, the main issues were formulated around the appearance of the aurochs and its way of living. Additional questions deriving from these main issues focus on the effects of feeding by large herbivores (including the aurochs), on the image presented by the natural Holocene landscape of Europe, and on an assessment of Heck cattle and their usefulness for nature management. Such questions are closely connected to the discussions currently being held in nature conservation circles in the Low Countries, with regard to the landscape to be aspired to and the cattle to be used. Although broadly speaking, the issues in question have been addressed, many details still remain unclear. The following discussion of the conclusions reached in the preceding chapters, and of the resulting recommendations, is divided into three parts. The first part will focus on the animal, the second on the landscape and the last on a combination of these two aspects.

11.1. The animal Characteristics The physical characteristics of the aurochs that could be reconstructed from the various sources are described in Chapter 7. In broad outlines, a relatively reliable picture could be composed of its body size, fur colour, horn shape and udder size. The horn shape and -position were accurately reconstructed on the basis of the many horn cores,

Black Cyan Page 361

FINAL REMARKS, CONCLUSIONS AND RECOMMENDATIONS

and a number of horns, that have survived. For lack of tangible evidence, the other three characteristics could only be ascertained with a little less precision. The physical aspects of the aurochs changed from the moment of its domestication, since man, instead of nature, started to apply selection criteria. Segregation into all kinds of horn shapes, fur colours and body sizes began to take place. Nowadays, no cattle breed that still displays all the physical characteristics of the aurochs remains, although some cattle breeds or -populations have retained more of those characteristics than others. The greatest number of such original aurochs characteristics may still be found in the populations of Spanish fighting cattle, to which the Portuguese and French fighting cattle are closely related. Heck cattle, contrary to the claims of the Heck brothers and their followers, are only vaguely similar to their wild ancestors. At best, some kinship may be discerned in their fur colour. Neither their horn shape nor their udder size show any similarity (see Table 19). A characteristic that Heck cattle, but probably also all other domestic cattle breeds, do have in common with the aurochs is their choice of (natural) food throughout the year, as well as its digestion. As far as can be ascertained, the digestive system, which is mainly geared to the digestion of grasses and graminoids, has remained unchanged in domestic cattle. This may not be the case for their assimilation of the digestive end products in physiological growth- and reproductive processes. At the time of domestication, these processes were modified by selection to be of greater benefit to man. Generally speaking, domestic cattle mature earlier and remain smaller than their wild ancestor; they fatten more readily and are able to reproduce earlier and more often. This has resulted in a cattle breed that produces a relatively large number of descendants, milk and fat, and retains a manageable size. In situations where domestic cattle are able to run wild, their early maturity has not yet constituted an impediment. In such cases, the main issue is the digestive system of domestic cattle, which adapts easily to a natural food supply. This has been shown to be true of various breeds, including Heck cattle.

361

Black Cyan Page 362

362


Generally speaking, domestic cattle have been shown to be able to run wild easily. This has happened in many areas in the world, under widely diverging circumstances, both in lowlands and in mountain areas. Such populations develop quite well without the help of man, and have no notable health problems. The successful development and relatively good health conditions of the Heck cattle in the Oostvaardersplassen are no exceptions to this. A characteristic that may also be different in domestic cattle, as compared to the aurochs, is that of temperament. Selection has supposedly made domestic cattle less aggressive and more easily manageable. This is certainly true of cows, but bulls are actually still rather unreliable individuals, which have to be controlled by certain measures so as to be useful. Aggressive behaviour may be a characteristic of certain breeds (Jersey bulls and Chianina bulls are known to be hard to handle), but the management of the cattle, for example with regard to the space available to them and the nature of their contacts with man, may also have an influence. The behaviour of the Spanish fighting bulls on the various breeding farms varies from aggressive, through easily approachable, to escapist. It is doubtful whether the aurochs was always as savage and aggressive, and as different of the current domestic cattle, as is often mentioned. The aurochs may have possessed certain mental characteristics that enabled man to turn it into a domestic animal more easily than related breeds, such as the European bison and bison, of which man never managed this. In judging the characteristics of the current European cattle breeds it is important to realise that their genetic material has derived mainly from cattle that stem from the Middle East. After the introduction of these cattle in Europe during the Neolithic period, these animals adapted to the climate, among other things. Any difference in winter hardiness between the various European cattle breeds is unlikely to be large. As was shown in the breeding-back experiment conducted by Lutz Heck, the South-European breeds he used were able to survive the northeastPolish winters without any problems. Whether these breeds still possess the fur structure of aurochs is unknown, and to what extent a changed fur structure plays a part in the heat insulation of other breeds remains unclear for now.

Black Cyan Page 363


Running wild One can only imagine what, in the long run, would have happened to domestic cattle forced to live in the same circumstances as the aurochs in its time, i.e. in a landscape of ‘forests and marshes’, with competitors for food, predators like the wolf, and seasonal and annual fluctuations in the climate and the food supply. Under such circumstances, a social structure comparable to that of the other species of the Bovini is likely to develop. Extensive acreage and large freedom of movement are essential to such a natural social structure. In view of the (relatively consistent) appearance of other animal species, the cattle that have run wild inside a specific area, for example Central Europe, will increasingly evolve in a certain direction, eventually to constitute a uniform type. In addition to this, domestic cattle will probably mature later in natural circumstances, which is a reversal of the process that takes place during domestication. They will grow to their full size, and become more muscular, later (particularly the bulls), and reproduce at a later age. Late maturity is a feature better suited to natural circumstances; we find it in all the other members of the Bovini. The difference in size between the bulls and the cows will become more marked. For reasons of greater ablebodiedness, manoeuvrability and speed, the animals’ build will become more square (equal shoulder height and trunk length) than is the case in many current domestic cattle, which have a more rectangular shape (greater trunk length than withers height). The udder will grow smaller, of a relative size known in all mammals. A relatively large udder is superfluous, vulnerable and energetically uneconomic. These are the processes likely to occur, since they take place generally among the Bovini. The same may be true of other anatomic and physiological processes, but whether these take place exactly as they do with the other Bovini obviously remains unclear. Although it is not yet clear as a result of which process (reduced benefit of the olfactory organ?) the head of domestic cattle has become shorter and, relatively speaking, broader, in the course of the domestication process, natural circumstances may cause a reversal of this process, possibly resulting in renewed elongation of the head. This may also cause an increase in

363

Black Cyan Page 364

364


brain content. Domestic cattle have smaller brains than the aurochs, a decrease that has affected adversely the quality of their senses, among other things77. Protracted periods under extreme circumstances will have a strong selective effect, in favour of the animals with the sharpest sensory perceptions. This concludes the processes that may also be found in other members of the Bovini. The fur colour and horn shape of the aurochs were not generally found inside this bovine group, but were more specific for the species. Whether this same fur colour and horn shape will ever reappear in their original manifestations is very much the question. The fact that both these features originally were very consistent, and apparently very functional, within the species, warrants the assumption that there were strong evolutionary and social forces that sustained them, and left little space for any great variation. It is very likely, therefore, that an animal may develop that is not like the original wild animal in every aspect, but which is definitely globally similar. The processes described in the above will take an extremely long time.

Breeding back These natural processes are unlikely to produce a perfect copy of the aurochs, man will not be able to achieve this, either, if only because the appearance of the animal is not known in every one of its details. Any approach will always remain global at best. Attempts to accelerate the process of ‘aurochs creation’, and obtain an aurochs-like creature from domestic cattle, have always held a strong appeal to the imagination78. Jarocki wanted this as early as 1835; the Heck brothers tried it in the 20s and 30s of the 20th century, and another attempt was made in the 80s in the United States. Currently, attempts to turn Heck cattle into animals more similar to the aurochs are made in several places, including the Lippe area (Soest, Westphalia) in Germany, where Spanish fighting cattle, Sayaguesas, Italian Chianina cattle and other breeds are being used 77 The hearing, smell and sight of such domestic animals as pigs, dogs, cats and rabbits is known to have deteriorated in comparison with the senses of their wild ancestors (Herre & Röhrs 1990). 78 There are similar ideas with regard to other animal species. Nowadays, attempts are also being made to breed back the Berber lion (Panthera leo leo) and the quagga (Equus quagga quagga), with the use of animals that still possess a number of characteristics of the extinct subspecies.

Black Cyan Page 365


to try and make the Heck cattle that are living there more like the aurochs in colour, horn shape and size. Why, really, should we breed back the aurochs? In fact, the animal did not play a dominant role in modelling the landscape. On the other hand, it did play such a role in the existence and experience of earlier people, who lived with and on them; it must have been an imposing animal. If other organisms (e.g. insects, fungi) depended on the aurochs, they can probably live on in domestic cattle. According to Heinz Heck (1934, 1949), a fair reason for breeding back the aurochs could be to correct a mistake (sc. extinction) made by man. He argued that the process would also make clear the differences between aurochs and European bison to the general public. For science, it would be not only interesting to see what the animal looked like, but also how it lived among other animals, in its niche of the (natural) landscape. The latter argument fits in well with the aspiration felt in some countries to restore the natural landscape with all its original participants. At the same time

Photo 48. Chianina cow in the flood plains of the Lippe River (Germany). Heck cattle are crossbred with such large white cattle there to increase the shoulder height of the former (Photo: T. van Vuure).

365

Black Cyan Page 366

366


such a landscape is indispensable for reshaping the aurochs, to refine the bred-back animal in accordance with the natural requirements. As was shown in Chapter 10, the breeding-back experiment of the Heck brothers was inadequate. On the basis of vague criteria and without proper knowledge of the appearance of the aurochs, the two brothers made inaccurate selections among the crossbreeding products of various cattle breeds. They did not use the knowledge about the aurochs that was available at the time, nor did they take advantage of the breeding techniques others were using to create new cattle breeds in the same period79. Their experiment can actually be described as a missed chance; if they had worked more meticulously, less led by their fantasies and complacency, a lot more would have been achieved today. The present situation is more or less the same as before their experiment. The aurochs is known to have been a consistent bovine species that was well adapted to its surroundings. To recreate a cattle breed with the same adaptability, and similar to the aurochs, the aurochs characteristics still present in the various domestic cattle breeds will need to be combined in a single breed. In principle, there are three ways to do this: through remodelling Heck cattle by making selections inside the population, by crossbreeding Heck cattle with other breeds that may contribute to a more similar appearance, or by making a ‘new’ attempt at selecting and crossing certain cattle breeds, without using Heck cattle. To find the desired features inside the existing Heck cattle population is no easy task. Animals with the right fur colour can be found, but the right udder size is more difficult, the correct horn shape is hardly to be found and animals of the right size are completely lacking. The population is very heterogeneous, which makes the realisation of stable desirable characteristics (homozygoty) difficult and time-consuming. The advantage of the Heck cattle in the Oostvaardersplassen, however, is 79 The experiments of the Heck brothers took place during a period when elsewhere in the world, the new dairy breed Jamaica Hope and the new meat breed Santa Gertrudis were being created (Rhoad 1949; Ministry of Agriculture, Jamaica 1972). These latter two experiments both took a period of about 40 years, 20 of which were used in search of, and for the selection of, good starting material; the remaining 20 to create a new breed with this. In both cases, combinations of ‘only’ two features were concerned: good milk yield and good heat tolerance in the first case, good meat production and good heat tolerance in the second. In both cases, inbreeding and crossing of inbred lines were applied.

Black Cyan Page 367


that during the past fifteen years, these have been selected, to a certain extent, for a life under difficult circumstances. A method that may have results sooner is the breeding in of cattle with characteristics that clearly do not occur in the Heck cattle population. In Germany near the Lippe, for example, Heck cattle are crossed with Chianinas, whose size resembles that of the aurochs. In itself, it should be possible to affect an increase in the size of Heck cattle. A disadvantage of such a crossing, however, is that together with the factor for a greater shoulder height, undesirable features are crossed in as well, such as white fur and small horns. Such undesirable traits all have to be bred out again later, which may take a long time. Against a single desirable feature that is bred in, therefore, there may be several undesirable traits that need to be bred out again. Faster results may be achieved by using cattle that already possess two or more desirable characteristics. In this way, there will be fewer undesirable traits that have to be removed later. If such cattle are used, however, crossbreeding them with Heck cattle is not advisable, since the latter breed has so few of the desired features themselves. As it happens, in such cases Heck cattle have more undesirable traits than the cattle they are to be crossed with, and it is better to seek other breeds to cross with. The Spanish and Portuguese fighting cattle form a cattle population that as a whole possesses more aurochs characteristics than the current Heck cattle. Not only does this heterogeneous population display more aurochs traits, these are also often combined in one and the same animal; besides, all cows of these cattle have small udders. Reddish-brown cows with small udders and aurochs horn shapes may be found along blackish-brown bulls with light eelstripes, light snouts and aurochs horn shapes (see Photo 40). Selecting and crossbreeding such animals may bring the desired target closer more quickly than the previous methods. Inbreeding will be an unavoidable means to achieve stabilisation of desirable features; this will hopefully ward off segregation in the shape of diverging individual animals. The first step could consist of collecting male and female animals with desirable physical traits (small udders, aurochs horn shape and -coat colours) in not too small numbers. These animals should then be bred in a

367

Black Cyan Page 368

368


system of crossbreeding and backcrossing till all of them possess the mentioned traits; the desired coat colour becomes visible already half a year after birth, udder size and horn shape not until some years later. The second phase would be to breed these (still rather small) animals for size (larger body, longer legs) and late maturity, since these are two strongly related factors. This will be no short-term experiment, but one which will take at least 30 to 40 years in total; one should beware of excessive optimism. That last part of such an experiment will be relatively time-consuming; breeding for late maturity means that the selected animals become fertile at an increasingly advanced age, so that they have calves later in life. The extent to which modern genetic research and genetic techniques might be helpful for the realisation of this goal could also be considered. To test the animals’ physiological properties they must be fed on grass and straw in the meantime, and supplemented with branches and leaves in winter. Distribution of the animals in a natural situation (including predators) will eventually refine the end product as to the capacities required of it; e.g. the skull will elongate to the original aurochs skull length. In this situation throw-backs and animals with pathological defects can still be eliminated by man. Also for veterinary reasons, it would be wise to eventually create a bovine species that is to a large extent different from domestic cattle. In the same way that the wild boar is physically very different from the domestic pig, and takes up a unique position in nature reserves, a bovine animal that is very similar to the aurochs, and very clearly different from domestic cattle, might also gain a separate status.

Conclusions and recommendations 1. The physical characteristics of Heck cattle barely resemble those of the aurochs. 2. If the effects of feeding by aurochs are to be copied in nature reserves, this might well be done with the help of domestic cattle. Many cattle breeds would suit, including Heck cattle.

Black Cyan Page 369


3. Heck cattle are of little value for the breeding of a bovine animal that looks like an aurochs. As starting material for such breeding experiments, it would be better to use cattle that are more similar to aurochs to begin with.

11.2. Effects and landscape Effects The current nature management in various European countries attributes great value to the effects of the feeding of large herbivores, such as horses and cattle. From the 70s of the last century, in certain nature reserves in the Low Countries conservation has taken place the way it has been done for centuries in areas such as the New Forest in England, i.e. by distributing domestic animals as a cheap mowing method and a way to keep the landscape open. The latter aspect means that excessive rejuvenation of the forest is counteracted, so that an open, park-like landscape is retained. At a later stage, many people started adding an extra dimension to this ‘management with the help of animals’. Especially and initially by Stichting Kritisch Bosbeheer (see Ch. 1) and later also Vera (2000) brought forward the theory that large herbivores were not only able to retain a beautiful landscape in an inexpensive manner; they could even do this without man’s help. By their feeding alone, large herbivores were allegedly able to counteract tree growth. The animals could be creators of the landscape, and large herds of horses, aurochs and European bison could inhabit it. Among these large herbivores, the European bison in particular was assumed to have been able to counteract the growth of trees, by eating tree bark and branches and by uprooting trees. Also in Polish forestry circles, such ‘disastrous’ effects are stressed time and again. Research into the alleged effects of European bison on the forest shows, however, that in the long term, the activities of these animals barely affect the development of the forest (see Ch. 9.4). The sight of many trees stripped,

369

Black Cyan Page 370

370


fed on and sometimes uprooted by European bison has played an important part in the notion that this would be detrimental to the forest as a whole. It has been shown, however, that the forest continues to grow in spite of such feeding, that open areas close up again eventually and that basically, the essential growing power of the trees is not affected adversely. Also in a natural, self-regulating ecosystem like Wood Buffalo National Park (Canada), large herbivores such as bison and deer turned out to be unable to counteract forest growth; in this area, only the effects of water and fire could achieve that. The local closing up with forest of dehydrated sedge marshes and open areas affected by fire could not be counteracted by the said herbivores. Elephants in African rain forests and savannah areas by nature also proved barely able to affect tree growth. The occurrence and growth of trees in these savannah areas turned out to be determined, not by elephants or other large herbivores, but by basic requirements such as moisture and light. Under certain circumstances, however, large herbivores are indeed able to counteract forest growth. This has been shown in the New Forest and in various nature reserves in the Low Countries, to name but a few. Human support is required under such circumstances, however. Relatively high population densities of cattle and horses during the growing season of the vegetation can only be realised by supplementary feeding of these animals in winter, either locally or in stables. Only a density thus manipulated to remain high may sustain the extreme feeding pressure on the vegetation. The research described in this book portrays large herbivores like the aurochs, not as creators of the landscape, but as obliged to adapt to existing vegetations or any changes thereof. Besides feeding, trampling must have been another aurochs effect. This should be seen as the making of trails and the rooting around in the soil by the bulls. The extent of this is likely to have been limited. In addition to this, by its consumption of grasses and graminoids the aurochs probably sped up the cycle of plant material, which caused faster mineralisation. Its droppings and eventually its mortal remains may have constituted the means of existence for numerous more or less specialised organisms (‘reducents’). Only speculation is possible about the function

Black Cyan Page 371


of the aurochs with regard to the distribution of plant seeds. As it happens, cattle are known to favour the consumption of wild apples (Malus sylvestris), if these are available. The seeds of this plant find an excellent germination medium in the excrement of the cattle. Whether this is also true of acorns, which often leave the body intact, is unknown.

Landscape The discussions about the impact of large herbivores on the growth of the forest, which have been going on in the past decennia, are inextricably connected to the image people have of the resulting landscape. Those who attribute great influence to these animals assume that the natural landscape had an open, park-like character. Others, who consider this effect as limited, see a mainly forested landscape. In many current nature reserves in the Low Countries, the management goal is to retain the (semi) open landscape, with or without the underlying thought that this is the natural situation. An important argument in favour of openness is that such a semi-open landscape hosts more plant- and animal species than an entirely forested environment. An open landscape that is partly forested does not only contain forest-specific species, but also many plant- and animal species from steppe areas. The latter are almost or entirely lacking in forested areas. In addition to this, a semi-open landscape has a greater variety of vegetational structures (forest, grassland, shrubs, solitary trees) per surface area than a forested landscape. Many attribute great aesthetic and conservational value to such variety. Apart from a few, all nature reserves in Europe may be called seminatural, since interference by man has been a determining factor in their appearance. Ever since the Great Wilderness (in former East Prussia) ceased to exist, around 1500, there have been no completely selfregulating land ecosystems in Central- and Western Europe. Large untouched forest areas, cut by small and larger rivers, with flood plain forests and marshes, are no longer available. Nor, for that matter, is their original fauna. The current forests are almost all cultivated, made and maintained by man. Man regulates the number of wild large herbi-

371

Black Cyan Page 372

372


vores, and distributes his domestic animals throughout the open landscape in high densities. From year to year, approximately the same density of cattle is maintained, a phenomenon unknown in nature. Man also controls the water balance, so that floods hardly ever occur and many soils become dehydrated. As was already shown in the Discussion (see Ch. 9.4), many researchers attribute a key role in the shaping of the landscape to the large herbivores. The research discussed here (Ch. 9.1) reveals a different image. Beside the already mentioned slight effect of large herbivores, it has also been concluded on the basis of numerous studies that the landscape of Central- and Western Europe during the Holocene period consisted mainly of forests. Entomological research, pollen research, reports by Roman writers and descriptions of the Great Wilderness all evoke an image of a landscape that consisted of extensive forests, alternated by marshes. The only open areas in this forested landscape could be found in wet areas such as salt marshes, peat bogs and fens (along rivers and in seepage areas), since tree growth was impossible there. As the research results presented here indicate that such a landscape existed naturally, this also means that the above-mentioned, semi-open landscape would not have existed in natural circumstances. That this was the landscape that appeared at some stage in large areas of Europe was a direct result of man’s interference. By cutting, burning and the creation of cattle pastures, man created this semi-open, cultivated landscape that may still be seen in many European nature reserves today. These human activities have become conditions for the existence and continuation of that landscape. The population densities of large herbivores in forest areas that were closed by nature were revealed to have been much lower than in the current (usually open) nature reserves, in which seasonal grazing by horses and cattle takes place on a regular basis.

Conclusions and recommendations 1. In natural circumstances, in areas favourable to tree growth large herbivores like aurochs were unable to stop the growth of the forest;

Black Cyan Page 373


at best they could slow it down somewhat. The aurochs was not able to open up the forest; neither were other large herbivores (with the exception of the beaver) in such circumstances. 2. To retain an open, park-like landscape, an unnaturally high population density of large herbivores in needed, kept by man in winter. The latter may sometimes have to help by burning and felling. 3. In the current climatological circumstances, the natural landscape in by far the larger part of (Central and Western) Europe would be covered almost entirely with forests. Areas that will not allow tree growth are marshy areas (along the coast), peat bogs and fens (along rivers).

11.3. The animal in the landscape Depending on the way in which they are kept, the deployment of cattle in nature reserves may make specific demands of the animals. Circumstances may vary, from situations in which cultivated landscapes are managed with the help of seasonal pasturing, to conditions that approximate the original natural circumstances. In the first case, many cattle breeds may suit, since the animals are stabled in winter, do not have to contend with any predators, are able to calve under supervision and remain under veterinary inspection. Moreover, because of such monitored circumstances the animals can be kept in high densities. The situation changes as soon as the cattle also have to spend the winter outside. Additional requirements include their resistance to the winter cold and, if no supplementary food is provided, an ability to make do with the scarce natural food supply. In addition to this, medical checkups become less easy and, if bulls are let in with the cows, the latter need to be able to calve without appreciable difficulty. If circumstances are created to approximate the natural situation, the demands made on the animals are even higher. In such a case, they would lack veterinary care, and should also be able to get through the winter, avoid or fend off predators such as wolves, function in a completely developed social structure and manage to reach and utilise the various foraging areas. The latter situation does not (yet) occur in the

373

Black Cyan Page 374

374


Low Countries, where seasonal or year-round grazing takes place in smaller areas, with at least a very little supplementary feeding in winter. Life is hardest for the cattle in the Oostvaardersplassen, who lack veterinary care. In addition to this, their natural food supply may show strong variations through the year. The Heck cattle mortality rate of 70 animals in 1999, which is about 14% of the population, may seem high according to human standards, but is actually quite normal for a naturally regulated population. At present, adaptation need not go beyond an animal’s suitability to such semi-natural circumstances. Even in the most extreme situation in the Low Countries, the Oostvaardersplassen nature reserve, animals never need to run far and fast to search for food or to fend off wolves. The fact that many domestic breeds do not do too badly in semi-natural circumstances may indicate that they still seem well able to survive in such circumstances. In many areas of the world, domestic cattle may be seen to be easily able to run wild, and to adapt to the local food supplies. Also there, bottleneck periods such as dry spells or winter are successfully withstood. Diets can be adapted in such periods, and adaptation to the cold is equally feasible. Indications exist that even the presence of wolves and bears need not constitute an impediment to the cattle’s running wild (Dobie 1980). There is little clarity as to the resistance against disease. It is not know whether, and to what extent, the aurochs was resistant against certain diseases. Heinz Heck’s claim (1980, p. 13) that Heck cattle were resistant against foot- and mouth disease and catarrhal fever should be investigated. If this is really the case, that resistance must stem from one of the breeds bred in at the time, since it is well known that domestic cattle have a certain measure of resistance against both these diseases. The Heck cattle in the Oostvaardersplassen, who are seen as non-kept animals’ and do not need to be inoculated, may generally be considered a healthy population80. The Oostvaardersplassen are surrounded by a buffer zone of 500 metres, inside which no domestic cattle is allowed. In North America, buffer zones are usually a lot larger, to avoid contacts of bison with domestic cattle and conflicts with cattle breeders about such contacts, with regard to the possible transfer of brucellosis, anthrax and cattle tu80 The exemption from the Destruction Act initially granted for the Heck cattle in the Oostvaardersplassen was later withdrawn. In principle, dead animals have to be removed if possible.

Black Cyan Page 375


berculosis. In itself, the occurrence of diseases among wild cattle need not be a reason for concern, as long as their effects remain limited. Diseases imported from elsewhere, for example the rinderpest in Africa, may lead to disastrous consequences in a very short period. According to the “Leidraad Grote Grazers” (guidelines with regard to large herbivores)81, which is completely in keeping with the ‘ethical guidelines’ of the Dutch Forestry Commission (11-11-99), dead cattle have to be removed, if possible, on account of the possible dangers of contamination. How large the actual veterinary risk of dead cattle in nature reserves really is is still not known, however. This aspect is being researched. Leaving (large) dead animals where they are might prove enrichment to nature conservation; in this way, high-quality food is not removed, which is important particularly in nutrient-poor areas. In addition to this, cadavers are an (often) indispensable source of food for many organisms, ranging from fungi and insects to birds and mammals. If there were plenty of carrion eaters (such as white-tailed eagles, raven, wild boar) in the large nature reserves, cadavers could be cleared away quickly which would probably greatly reduce the possible risk of contamination by diseases. This risk would also be less if natural winter deaths as a result of food shortages were allowed to occur. Interference by man, for example in the shape of ‘timely’ culling based on human standards, may constitute a disturbing factor in ecological processes (Prins 1998). The plan for the future replacement of the Heck cattle in the Oostvaardersplassen by European bison, because of the veterinary problems, is not really a solution. Obviously, European bison may contract the same diseases as Heck cattle. The fact that present European bison have a small genetic basis and are not always free of disease in the wild does not make their presence any less of a risk.

The aurochs habitat Swamp forests and marshes are likely to have played important parts in the life of the aurochs. Causes for the decline and extinction of the aurochs may be found in both the hunting and the ousting of this ani81

Drawn up by the Dutch Ministry of Agriculture, Nature Management and Fisheries, 19-1-2000.

375

Black Cyan Page 376

376


mal from its most significant food supply, the nutrient-rich forelands along rivers. Man took up the wet grass- and sedge lands there as cattle pastures. In addition to this, the forests along rivers were also felled to accommodate meadows. Also elsewhere, man occupied the more productive soils, leaving the less productive ones in current nature reserves. In the larger part of Europe, the combination of a humid summer habitat and a drier winter habitat was of vital importance to bovine animals such as aurochs. Before man started to construct ditches and dikes, the landscape was very much wetter than nowadays, with marshes and floods on a scale that is hard to imagine today. If a population of wild cattle is to exist in the best possible way, the presence of both wet and dry habitats is essential. It would be interesting to see how much of a division of habitats between Heck cattle and European bison would result from the latter animal’s introduction in the Oostvaardersplassen (see Ch. 9.1.3). The water management conducted in the Low Countries has caused many soils to dehydrate. Beside the extraction of underground water for the drinking-water supply, for agricultural reasons this management aims to drain off superfluous surface water as quickly as possible. This is done by canalising rivers and brooks, and by using dikes and groynes to keep large rivers in narrow, deep basins. Fortunately, things are already starting to change, and brooks are once again allowed to meander in some places. Also large rivers may possibly regain a more natural character in some areas, as a result of greatly increased dynamics. In this connection, the plans with regard to the IJssel River (Central Netherlands) link up well to the creation of a natural habitat for the wild descendants of the aurochs82. The enlargement of the IJssel River Basin proposed in these plans (the moving or even removal of main dikes, particularly in its northern half) will greatly increase the dynamics of the river. The sedge marshes near the river, which constituted the aurochs’ foraging grounds at the time, are connected to very flat areas with little fall, which makes them marshy almost throughout the year. Adding dynamics to the IJssel River may possibly restore such sedge vegetations. In addition to this, connections may be made from the IJssel River forelands to the Veluwe, to combine the wet and the drier habitats. Linking up nature reserves makes the living areas larger as well as more 82 Provisional plans for this have been drawn up in the report ‘De Rijn op termijn’, which was published in 1999 by WL Delft Hydraulics at Delft (the Netherlands).

Black Cyan Page 377


complete. In the long run, cattle that live in dry, nutrient-poor areas, such as in parts of the Veluwe, may well develop a lack of important nutritive elements such as calcium, phosphorus and sodium (Groot Bruinderink et al. 1997, Wallis de Vries 1994). Access to more nutrient-rich areas, such as riverbanks and brook sides, may alleviate these shortages. Relatively dry areas like the Veluwe may also become more interesting as a result of more moisture. If all the Veluwe brooks, free of cultivating measures, could resume their original meandering courses, and less drinking- and irrigation water was to be pumped up, the area would greatly increase in value. In forests managed by man, the proportion of grasses and forbs is usually larger than in natural forests. By his regular thinning, the clearcuts, the many verges and game meadows, man contributes to the more open structure of cultivated forests, and increases the proportions of grasses and forbs. If one considers the food supply, however, natural forests also have their advantages. Particularly in the West- and Central European situation, the proportion of deciduous trees is greater, which means that there is more food for ungulates. The average age of the trees is higher as well, which results in more tree fruits83. In addition to this, natural forests host a very large proportion of dead trees, which provide food for, among others, many kinds of insects and fungi, of a richness and variety that no longer occurs in the cultivated forests. The scarcity of grasses and forbs in the first growing phases of the forest is counterbalanced by an increase in the later stages, when large trees are lost and more small open areas develop. In the Central-European natural forests, large to very large open areas could only result from storms. In this respect, fire and insects play negligible parts (see Ch. 9.1.3). Small- as well as large-scale uprooting of trees by wind may stimulate forest rejuvenation. If there are naturally low densities of herbivorous ungulates, moreover, tree seedlings, which often grow up protected by fallen tree crowns, may ensure sufficient and timely rejuvenation. Light-loving tree species that constitute popular food, such as oak trees, may also retain their proportion of the forest stock in that way. Whether the oak will once again become as important as during the 83 · the largest population density of wild boar may be found in the strict nature reserve north In the forests at Bia§lowieza, of the village. The tree stock of oaks is oldest there and yields the most acorns (J¸edrzejewska & J¸edrzejewski 1998). In addition to this, there is a lot of dead wood.

377

Black Cyan Page 378

378


Atlantic period is very much the question. The circumstances for growth were very different at that time than they would be now. The temperature was a little higher than at present and important tree species such as beech and hornbeam did not yet occur. As a model of reference for future natural forests, therefore, it might be better to use the situation of around 2000 years ago, when a composition of tree species had developed that is comparable to the current one. The prominent part played by the oak in the European Medieval landscape was inextricably connected to the semi-open cultivated landscape created by man. Without man’s interference, its role would have been much more modest.

Population density and area size In its time, the population density of the aurochs is likely to have been very diverse. Depending on the season (summer or winter) and the productivity of the living area (open marshy riverbanks or forests), this density could vary quite a bit. If cattle are distributed in nature reserves with the aim to create circumstances that are as natural as possible, a minimum population size should definitely be adhered to. Although cattle do seem able to withstand a large degree of inbreeding84, it is still advisable to use ample margins, for example a minimum of 100 animals. If this is impossible, and the population remains too small, exchanges between cattle populations that live in similarly small areas may provide a remedy against excessive inbreeding. The same goes for predators. If one assumes a population density like the one calculated for a grazer like the European bison at Bia³owie¿a, i.e. 1 animal on every 200 ha in the presence of red deer, such a population of 100 head of cattle would need a minimum of 20,000 ha of living space with nutrient-rich 84 In the past three centuries, the population of feral Chillingham White cattle (Northern England) varied between 13 and 80 animals, with an average of about 40 animals (Blokhuis 1982). As a result of this, there is a lot of inbreeding and the animals may be called almost genetically uniform (Visscher et al. 2001). The degree of homozygoty is even greater than among natural populations of wild cattle. Extensive inbreeding is also characteristic for Spanish fighting cattle and Heck cattle that are kept by small breeders. By nature, populations of wild cattle (for example African buffalo during rinderpest epidemics) may become very small temporarily (‘bottleneck’), which causes inbreeding. If such a period is only brief, with a subsequent increase of the population, the genetic diversity will grow again. The current global population of the European lowland bison stems from a group of 13 animals. Moreover, a single bull of the Caucasus population was at one time involved in the population of the European upland bison. On 31-12-98, the total population of European bison consisted of about 2,850 animals (European Bison Pedigree Book 1998).

Black Cyan Page 379


forests. If the latter were nutrient-poor, the living area would have to be a lot larger. If a forest area were to be combined with nutrient-rich river forelands, however, a much smaller area would suffice for such a population. To ensure a well-balanced social structure of the population, the animals should have the space and the freedom to deal adequately with social relations and tensions. If cattle are kept in relatively small nature reserves for purposes of grazing, the requirements for life in a (nearly) natural situation do not apply. Animals in such a situation are completely managed by man, and problems such a food shortages, inbreeding and physical health will be much easier to keep in check. In a comparatively large nature reserve like the Oostvaardersplassen, nature conservation focuses on the maintenance of an open landscape with short grazing vegetations for the geese. If man really were to stop interfering, the populations of Heck cattle, horses and red deer would (probably) decrease considerably in the near future. The landscape would become even more densely forested. Eventually, if the area would become almost entirely forested, with the exception of excessively marshy areas, only a few dozen horses and cattle would remain. Any dead animals are difficult to find, let alone remove, in such a densely forested landscape.

Conclusions and recommendations Nature management is torn between two extremes, the cultivated situation and the natural situation. The choice for either of the two, or for a hybrid form, depends on the goals and possibilities. In the cultivated situation cultivated forests and (semi-) open cultivated landscapes are conserved with a view to the vegetations and plant- and animal species that occur there. The cattle here are used as cheap mowers and pruners. The cattle need to be manageable and community-minded. Many domestic cattle breeds may be used for this, not necessarily of the aurochs type. The animals are stabled in winter (sea-

379

Black Cyan Page 380

380


sonal grazing). Where their grazing does not result in the conservation of desired vegetations, man should resort to mowing, burning or felling.

In the natural situation, man no longer interferes, original plantand animal species occur and the natural landscape in Central- and Western Europe would consist of forests and, in places where forests cannot grow, marshes. This is a situation that no longer occurs in by far the larger part of Europe. There, as in the Low Countries, however, possibilities to approximate that situation do exist. For a wild bovine animal that descends from the aurochs, this could signify large living areas, in which river forelands are combined with forests, with the presence of wolves as predators, a dependence on the natural food supply, and freedom from veterinary limitations. If such a bovine animal is to survive under those circumstances, its appearance and physiology should be as similar as possible to the bovine animal that was best adapted to those circumstances in its time: the aurochs; the animal that constituted an essential part of the original ecosystem.

Summary of conclusions and recommendations The animal 1. The physical characteristics of Heck cattle barely resemble those of the aurochs. 2. If the effects of feeding by aurochs are to be copied in nature reserves, this might well be done with the help of domestic cattle. Many cattle breeds would suit, including Heck cattle. 3. Heck cattle are of little value for the breeding of a bovine animal that looks like an aurochs. As starting material for such breeding experiments, it would be better to use cattle that are more similar to aurochs to begin with.

Black Cyan Page 381


Effects and landscape 1. In natural circumstances, in areas favourable to tree growth, large herbivores like aurochs were unable to stop the growth of the forest; at best they could slow it down somewhat. The aurochs was not able to open up the forest; neither were other large herbivores (with the exception of the beaver) in such circumstances. 2. To retain an open, park-like landscape, an unnaturally high population density of large herbivores in needed, kept by man in winter. The latter may sometimes have to help by burning and felling. 3. In the current climatological circumstances, the natural landscape in by far the larger part of (Central and Western) Europe would be covered almost entirely with forests. Areas that will not allow tree growth are marshy areas (along the coast), peat bogs and fens (along rivers).

The animal in the landscape Nature management is torn between two extremes, the cultivated situation and the natural situation. The choice for either of the two, or for a hybrid form, depends on the goals and possibilities. In the cultivated situation cultivated forests and (semi-) open cultivated landscapes are conserved with a view to the vegetations and plantand animal species that occur there. The cattle here are used as cheap mowers and pruners. The cattle need to be manageable and comm.\unityminded. Many domestic cattle breeds may be used for this, not necessarily of the aurochs type. The animals are stabled in winter (seasonal grazing). Where the grazing does not result in the conservation of desired vegetations, man should resort to mowing, burning or felling. In the natural situation, man no longer interferes, original plantand animal species occur and the natural landscape in Europe would consist of forests and, in places where forests cannot grow, marshes or steppes. This is a situation that no longer occurs in by far the larger part

381

Black Cyan Page 382

382


of Europe. There, as in the Low Countries, however, possibilities to approximate that situation do exist. For a wild bovine animal that descends from the aurochs, this could signify large living areas, in which river forelands are combined with forests, with the presence of wolves as predators, a dependence on the natural food supply, and freedom from veterinary limitations. If such a bovine animal is to survive under those circumstances, its appearance and physiology should be as similar as possible to the bovine animal that was best adapted to those circumstances in its time, the aurochs: the animal that constituted an essential part of the original ecosystem.

Black Cyan Page 383

APPENDIX

APPENDIX Passage from Gesner (1602, pp.141 and 142) in which the letters by Anton Schneeberger and Johann Bonar were published (translation from Latin by M.J.T. Verhallen): ‘Since mention has just been made of a Sarmatian85 or Masovian ‘Thurus’ or ‘Turo’, I will here add the things that, after what was written and published in the old days, have been written to me about the animal by scientists. To begin with, Lord Anton Schneeberger in a letter to me writes the following: ‘The native hunters obviously mistakenly call the ‘Thuri’ bison and buffalo: some, however, more simply and correctly call them ‘wild forest cattle’. Others think they should be called ‘Pyrrhic cattle’86, since a more accurate name is preferable. They look a lot like domestic cattle, but they are much larger and covered with longer hairs, and have two horns, which are curved forward and slim. The forehead, because of the curly, frizzy hair, makes them terrible to behold. Their hoof is cloven, more hollow than in domestic cattle. The cows are smaller than the bulls and not as long. When a bull is born, its hair is a chestnut colour (blackish brown, dark brown or the colour of black ash; ‘plowy’87 as the Polish call it), within half a year they become completely black, however, while the line over the spine, some two fingers wide, remains a lighter colour. The cows always keep the colour described in the above, and black cows are found very rarely. They live in a part of the Hercynian Forest, which is situated five miles from Warsaw, the capital of the province of Masovia, near the villages of Sochaczew and Kaski. Most of them occur in the very dense parts of the wood, the same forest has red deer, roe deer and wild boar, but only very few of these. In the autumn they feed on acorns (it so happens that the forest has enormously large oak trees, they say some are bigger than 9 m around), and then they are fatter and more shiny than at other times. In winter they eat branches of shrubs and trees; however, 85 86 87

Sarmatia indicates East Poland and a part of Russia. In Classical Italy, ‘Pyrrhic cattle and ‘Lucanian cattle’ were names for the elephants introduced there from Africa. Here, the Polish word ‘p§lowy’ means ‘reddish brown’ or ‘fox-coloured’.

383

Black Cyan Page 384

384


they are also supplied with hay that the farmers of the above-mentioned villages have gathered for the aurochs in summer; in summer they leave the forests and go into the fields and devour mown grain and when they are satisfied they throw the rest apart with their horns, unless they can be chased away by dogs. It is said that in the 57th year88, several animals perished from the fierce cold, but now it is suspected that there are once again over 50, since the accurate number is being concealed, they do this led by some kind of superstition. In winter they run in herds and they rest in one place, quite like some army. In summer, however, they roam the wilderness separately89 and if one has strayed too far and not shown himself for a few days (because there are hunters that have to go to them every single day) he will be sought with very great diligence and forced by dogs to return. The animal is very fast, but not long-living; it is said that only a few have lived beyond the 15th year, they do not experience harm from the wolves, except when they, when they have just been born, wander around separately, because then they will be torn up by the wolves. An aurochs is not afraid of humans and will not flee when a human being comes near, it will hardly avoid him when he approaches it slowly. And if someone tries to scare it by screaming or throwing something, this will not scare it in the least, but while it stays in its place it will actually open its mouth, widen it and close it again quickly, as if it is making fun at the human for his attempt. When it is standing in the road or somewhere else, one must go around it, even if one is driving a carriage, since it will not move off the road by itself. When challenged they become very hot-tempered, but if the person who has provoked it stretches out on the ground, nothing bad will happen to him, since they spare those who are stretched out, just like lions, with remarkable kindness, but if he does not stop challenging them, they will attack with their horns and throw him up into the air with them. In September they mate, while burning with lust, have numerous and very fierce fights and sometimes both fall down dead. But the one animal that they have noticed to be stronger, and constantly provoking others to fights, hunters will hunt under orders of the king, in a way that will be explained below. But they also hunt those that have been spotted covering domestic cows; for cows will become pregnant by them, but will either miscarry or give birth to non-viable calves. Finally, they 88 89

This is the year 1557. See Ch. 9.3.

Black Cyan Page 385

APPENDIX

give birth in May, some in September like domestic cows; this seldom happens, however, and the calves that are born in the autumn are weak and do not stay alive because of the fierce winter cold. When the time to give birth arrives, the cow will retire to the densest areas of the forest and remain there with the calf for about twenty days; as soon as she sees it has become stronger and jumps she will finally take it to the pasture, guarding it closely so that the calf is not caught by hunters or devoured by wolves. More than once, they took calves away from the aurochs and brought them to domestic cows to be raised by them; this attempt failed, however, because they all died. They only hunt by order of the king; very many riders and footmen go hunting with arrows, blunderbusses90 and with many dogs. Soon they will force the one animal they have decided to hunt, by shouting and barking of the dogs to separate from the others, in the meantime the riders and those that carry the blunderbusses etc. hide partly in ditches, partly behind the larger trees, so that the animal does not become furious at seeing such a large group of people. As soon as it has been separated and set apart it will be bombarded with the blunderbusses, nevertheless it will still run away, even though it has been hit by very many bullets, sometimes for a whole day or longer, until it is struck in the chest, since it will then fall very quickly. Farmers will continuously split pieces of wood, which they will throw against, slantwise and straight on top of it, thus surrounding the animal while standing firmly on it, so that it will no longer move to resist, and while it is held thus and is still alive, they will pull off the skin which covers its forehead between the two horns, and which by its curly hair makes the animal horrible to behold; when this has been ripped off they will finally kill it: immediately after it has been killed they will take out the heart, when they have cut this in half they will find a small bone in the shape of a cross, which they will send to the king together with the skin that has been torn off the head, and they greatly value both. Some say that this bone is advantageous to pregnant women, and to those who suffer greatly when giving birth. They send chunks of meat in the skin to the king, sometimes fresh, sometimes salted. I have heard that Sigismundus, a king which is remembered with a lot of respect, several times sent salted pieces of meat of an aurochs to emperor Charles V as a big gift’. 90

A blunderbuss is a type of fire arm.

385

Black Cyan Page 386

386


To this, the renowned Lord Baron Bonar has added the following: ‘I cannot agree at all to calling them ‘Pyrrhic cattle’, because Pyrrhus was absolutely the first of the foreign kings to show elephants in Italy, and because, moreover, several poets called them ‘Pyrrhic cattle’, while they used this word in a meaning, contrary to its normal meaning and since they were first seen in Lucania, some dared to call them ‘Lucanian cattle’, but everybody knows how much elephants are different from cattle. That is why it seems wrong to me to call them ‘Lucanian cattle’ or ‘Pyrrhic cattle’, since they are not. And this [the aurochs] is quite a different kind of animal, different from the elephant, and otherwise similar to a bovine animal. We might as well call him a Sarmatian- or Masovian bovine animal, since it is certain that this animal is found nowhere else but in Sarmatia. And repeatedly when I thought about this, and what kind of animal this was, I could not imagine anything else than that these animals by some coincidence stem from some outstanding cattle and have increased to this number, or that the European bison that also lives in Masovia, by mating with a domestic cow has produced this breed which, when afterwards their number had increased and they were seen, by order of kings and royals of that place were caught to be guarded (and are still guarded to this day), in such a way that these wild forest cattle developed and should not be considered domestic cattle, they were then given the name ‘Thur’ [aurochs] since we, when a horse or any other animal walks proudly or fervently, according to our Polish vocabulary will say: he walks like an aurochs. But since, on account of the sloppiness of the people and the lack (of knowledge) of the authors, it is not quite clear and also cannot be found out when or under which king these animals originate, so it is hard to show whether the name ‘Thur’ was given to them because of their fierceness or the etymology of the word is determined by the pride of these animals also with regard to other animals. When the skin of this animal has been cleaned it is covered with very fine black hairs, the skin is then torn into belts, girded with these pregnant women can give birth with miraculous ease. The ‘corrigium’91, however, is very thick, so that on account of its hardness it is not fit for anything but horse’s bridle and -saddle, since it does not tear at all’.

91

With ‘corrigium’, most likely the skin on the back is indicated. Also see Ch. 7.4.

Black Cyan Page 387

REFERENCES

REFERENCES Aaby, B. (1994): NAP percentages as an expression of cleared areas. In: B.Frenzel (Hrsg.). Evaluation of land surfaces cleared from forests in the Roman Iron Age and the time of migrating Germanic tribes based on regional pollen diagrams. Paläoklimaforschung 12: 13 - 27. Aaris-Sørensen, K. (1980): Depauperation of the mammalian fauna of the island of Zealand during the Atlantic period. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i København 142: 131 - 138. Aaris-Sørensen, K. (1999): The Holocene history of the Scandinavian aurochs (Bos primigenius Bojanus, 1827). In: G.-C. Weniger (Hrsg.). Archäologie und Biologie des Auerochsen.: 49 - 57. Neanderthal Museum. Aaris-Sørensen, K. & Brinch Petersen, E. (1986): The Prejlerup aurochs - an archeozoological discovery from boreal Denmark. Striae, 24: 111 - 117. Acker Stratingh, G. (1844): Over eenige wilde dieren, welke vroeger in ons Vaderland geleefd hebben. Mededeelingen uit het gebied van Natuur, Wetenschappen en Kunst. Deel I, nrs. 18 en 19. Adametz, L. (1898): Studien über Bos brachyceros, die wilde Stammform der Brachycerosrassen in Europa. Journal für Landwirtschaft 46: 17 - 31. Adametz, L. (1925): Kraniologische Untersuchungen des Wildrindes von Pami¹tkowo. Arbeiten der Lehrkanzel für Tierzucht an der Hochschule für Bodenkultur in Wien, III: 1 - 25. Adametz, L. (1933): Die Bedeutung der ‘Abzeichen’ des Banteng (Bibos banteng Raffl.) und des Urs für das Abstammungsproblem des Hausrindes. Biologia Generalis 9: 33 - 38. Allen, J.R.L. (1997): Subfossil mammalian tracks (Flandrian) in the Severn Estuary, S.W. Britain: Mechanics, preservation and distribution. Philosophical Transactions of the Royal Society of London. B, Biological Sciences 352 (1352(: 481 - 518. Alverson, W.S., Waller & D.M., Solheim, S.L. (1988): Forests too deer: edge effects in Northern Wisconsin.. Conservation Biology 2(4): 348 - 358 . Ammann, B., Birks, H.J.B., Brooks, S.J., Eicher, U., Von Grafenstein, U., Hofmann, W., Lemdahl, G., Schwander, J., Tobolski, K. & Wick, L. (2000): Quantification of biotic responses to rapid climatic changes around the Younger Dryas: A synthesis. Palaeogeography, Palaeoclimatology, Palaeoecology 159(3-4): 313-347. Anderson, P.M. & Brubaker, L.B. (1986): Modern pollen assemblages from Northern Alaska. Review of Palaeobotany and Palynology 46(3-4): 273-291.

387

Black Cyan Page 388

388


Andreyev, V.N. (1971): Horse herding for meat in Yakutskaya ASSR. Polar Record 15: 931 - 933. Andrieu, V., Field, M.H., Ponel, P., Guiot, J., Guenet, P., Beaulieu, J.-L. de, Reille, M. & Morzadec-Kerfouw, M.T. (1997): Middle Pleistocene temperate deposits at Dingé, Ille-et-Villaine, northwest France: pollen, plant and insect macrofossil analysis. Journal of Quaternary Science 12(4): 309-331. Anonymus (1955): Rückzüchtung des Auerochsen missglückt?. Natur und Landschaft 30 (3): 48. Anstadt, M. (1965): Polen - land, volk, cultuur. Het Wereldvenster, Baarn. Arsenjew, W.K. (1924): In der Wildnis Ostsibiriens; Forschungsreisen im Ussurigebiet. Band 1 und 2. Scherl, Berlin. Baak, P.J., Bottema, S., Cappers, R.T.J. & Woldring, H. (2005): Vegetatiehistorie van Jardinga. Paleo-Aktueel 14/15: 46-50. Baales, M. & Street, M (1996): Hunter-gatherer behaviour in a changing Late Glacial landscape: Allerød archaeology in the Central Rhineland, Germany. Journal of Anthropological Research 52: 281-316. Badam, G. L. (1984): Holocene faunal material from India with special reference to domesticated animals. In: J. Clutton-Brock & C. Grigson. Animals and Archaeology 3. Early herders and their flocks: 339 - 353. BAR International. Baker, C.M.A. & Manwell, C. (1980): Chemical classification of cattle. 1. Breed Groups. Animal Blood Groups and Biochemical Genetics 11: 127 - 150. Baker, R.G. & Drake, P. (1994): Holocene history of prairie in midwestern United States: Pollen versus macrofossils. Ecoscience 1(4): 333-339. Banerjee, S. & Saha, U. (1976): On the occurrence of Bos namadicus Falconer, District Bankura, West Bengal. Current Science 45 (5): 186 - 187. Barber, K.E. & Clarke, M.J. (1987): Cranes Moor, New Forest: palynology and macrofossil stratigraphy. In: K.E. Barber (ed.). Wessex and the Isle of Wight: Field guide. Quaternary Research Association, Cambridge: 33-44. Barnekow, L., & Sandgren, P. (2001): Palaeoclimate and tree-line changes during the Holocene based on pollen and plant macrofossil records from six lakes at different altitudes in northern Sweden. Review of Palaeobotany and Palynology 117(1-3): 109-118. Barnes, R.F.W., Barnes, K.L., Alers, M.P.T. & Blom. A. (1991): Man determines the distribution of elephants in the rain forests of northeastern Gabon. African Journal of Ecology 29: 54 - 63. Bartosiewicz, L. (1985): Interrelationships in the formation of cattle long bones. Zoologischer Anzeiger 215 (3/4): 253 – 262. Bartosiewicz, L. (1997): A horn worth blowing? A stray find of aurochs from Hungary. Antiquity 71: 1007 - 1010.

Black Cyan Page 389

REFERENCES

Bartosiewicz, L. (1999): Aurochs (Bos primigenius Bojanus, 1827) in the Holocene of Hungary. In: G.-C. Weniger (Hrsg.). Archäologie und Biologie des Auerochsen: 103 - 117. Neanderthal Museum. Bauer, R. & Haupt, H. (1976): Das Kunstkammerinventar Kaiser Rudolfs II, 1607 - 1611. Jahrbuch der kunsthistorischen Sammlungen in Wien, Band 72. Beck, H. (1973): Auerochse - III Philologisches.. In: J. Hoops. Reallexikon der Germanischen Altertumskunde, Band 1, Lieferung 4. W. de Gruyter, Berlin - New York. Behre, K.-E. (1988): The role of man in European vegetation history. In: B. Huntley & T. Webb III (eds.). Vegetation history. Handbook of vegetation science, Vol. 7: 633 - 672. Belue, T.F. (1996): The long hunt: death of the buffalo, east of the Mississippi. Mechanicsburg, Stackpole Books. Bendz, G. (1963): Kriegslisten/ Sextus Iulius Frontinus. Schriften und Quellen der Alten Welt, Band 10. Berlin. Benecke, N. (1994a): Archäozoologische Studien zur Entwicklung der Haustierhaltung in Mitteleuropa und Südskandinavien von den Anfängen bis zum ausgehenden Mittelalter.. Akademie Verlag. Benecke, N. (1994b): Der Mensch und seine Haustiere: die Geschichte einer jahrtausendealten Beziehung. Theiss, Stuttgart. Bergström, P.L. & Wijngaarden-Bakker, L.H. van (1983): De metapodia als voorspellers van formaat en gewicht bij runderen. IVO-rapport B-206, and IPP-publikatie 320. Beutler, A. (1996): Die Grosstierfauna Europas und ihr Einfluss auf Vegetation und Landschaft.. Natur- und Kulturlandschaft 1: 51 - 106. Birks, H.H. & Birks, H.J.B. (2000): Future uses of pollen analysis must include plant macrofossils. Journal of Biogeography 27(1): 31-35. Bjor, K. & Graffer, H. (1963): Beiteundersøkelser på skogsmark. Forskning og Forsøk i Landbruket 14: 121–365. Blake, S. (2002): Forest buffalo prefer clearings to closed-canopy forest in the primary forest of Northern Congo. Oryx 36(1): 81-86. Blankenhorn, H.J. (1977): Angewandte Wildforschung und das Hirschproblem im Nationalpark und seiner Umgebung. Schweizer Naturschutz 43(6): 7-9. Blokhuis, H.J. (1982): De wilde witte runderen van Chillingham. Zeldzaam Huisdier 7(2): 58-60. Bobiec, A. (2002): „Grazing ecology” from the Bia³owie¿a Primeval Forest perspective. Acta Theriologica 47(4): 509-511. Bökönyi, S. (1956): Trinkbecher aus Urhörnern in Ungarn. Säugetierkundliche Mitteilungen 4: 145 - 150.

389

Black Cyan Page 390

390


Bökönyi, S. (1962): Zur Naturgeschichte des Ures in Ungarn und das Problem der Domestikation des Hausrindes. Acta Archaeologica Hungarica 14: 175 – 214. Bökönyi, S. (1974): The history of domesticated mammals in Central and Eastern Europe.. Budapest. Boessneck, J. (1957): Funde des Ures, Bos primigenius Boj. 1827, aus alluvialen Schichten Bayerns. Säugetierkundliche Mitteilungen 5: 55 - 69. Bohlin, B. (1938): Einige jungtertiäre und pleistozäne Cavicornier aus Nord-China. Nova Acta Regiae Societatis Scientiarum Upsaliensis, Ser. IV, Vol. 11, nr 2. Bohlken, H. (1958): Vergleichende Untersuchungen an Wildrindern (Tribus Bovini Simpson 1945). Zoologische Jahrbücher, Abt. Physiologie 68: 113 - 202. Bohlken, H. (1961): Der Kouprey, Bos (Bibos) sauveli Urbain 1937. Zeitschrift für Säugetierkunde 26(4): 193 - 256. Bohlken, H. (1962): Probleme der Merkmalsbewertung am Säugetierschädel, dargestellt am Beispiel des Bos primigenius Bojanus 1827. Morphologisches Jahrbuch 103: 509 - 661. Bohlken, H. (1964): Vergleichende Untersuchungen an den Schädeln wilder und domestizierter Rinder. Zeitschrift für Wissenschaftliche Zoologie 170: 323 - 418. Bojanus, L.H. (1827): De uro nostrate eiusque sceleto commentatio. Verhandlungen der Kaiserlichen Leopoldinisch-Carolinischen Akademie der Naturforscher, Bonn 13(2): 411 - 478. Bokdam, J., Cornelius, R. & Krüsi, B.O. (2001): A conceptual model for nutrient-mediated successional grazing mosaics. In: B. Gerken, & M. Görner. Neue Modelle zu Masznahmen der Landschaftsentwicklung mit groszen Pflanzenfressern und praktische Erfahrungen bei der Umsetzung. Natur- und Kulturlandschaft, Band 4: 122 – 131. Höxter/Jena. Bokdam, J. & Gleichman, J.M. (2000): Effects of grazing by free-ranging cattle on vegetation dynamics in a continental NW-European heathland. Journal of Applied Ecology 37: 415 - 431. Bollongino, R., Burger, J. & Alt, K.W. (2003): Import oder sekundäre Domestikation? Der Ursprung der europäischen Hausrinder im Spiegel molekulargenetischer Analysen an neolithischen Knochenfunden. Beiträge zur Archäozoologie und Prähistorischen Anthropologie IV: 211-217. Bonnicksen, T.M. (2000): America’s ancient forests – from the ice age to the age of discovery. John Wiley & Sons, Inc. Borchardt, L. (1913): Das Grabdenkmal des Königs Sahu-re. Band II: Die Wandbilder, V: Die Tierdarstellungen. Berlin. Borgesius, J.J. & Tol, G. van (1998): Wilde ideeën, eenzijdige visie. Metaforen die te ver voeren. Nederlands Bosbouw Tijdschrift 70, May/June: 98 - 101.

Black Cyan Page 391

REFERENCES

Borowski, S. & Kossak, S. (1972): The natural food preferences of the European bison in seasons free of snow cover. Acta Theriologica 17: 151 - 169. Borowski, S., Krasiñski, Z. & Mi³kowski, L. (1967): Food and role of the European bison in forest ecosystems. Acta Theriologica XII, 25: 367 - 376. Borowski, S. & Vuure, T. van (1974): Impact of bark peeling on growth height in oak and ash. Sylwan 118(2): 30 - 36. Bos, J.A.A., Dickson, J.H., Coope, G.R. & Jardine, W.G. (2004): Flora, fauna and climate of Scotland during the Weichselian Middle Pleniglacial – palynological, macrofossil and coleopteran investigations. Palaeogeography, Palaeoclimatology, Palaeoecology 204(12): 65-100. Bosscha Erdbrink, D.P. (1986): De keratine oeros-hoorn uit de collectie Stolzenbach: nog een paar opmerkingen. Cranium 3(1): 35 - 39. Bottema, S. (1980): Palynological investigations on Crete. Review of Palaeobotany and Palynology 31: 193-217. Bottema, S. (1987): De invloed van de vegetatie op de fauna in Nederland gedurende het Laat Quartair. Nederlands Bosbouw Tijdschrift 59 (9/10): 287 - 294. Bottema, S. (2005, in press): Vegetation history of Mesolithic Jardinga: a palynological study. Palaeohistoria. Boule, M., Breuil, H., Licent, E. & Teilhard, P. (1928): Le Paléolithique de la Chine. Archives de l’Institut de Paléontologie Humaine, Mémoire 4. Bousquet, B. (1978): Un parc de forêt dense en Afrique: le Parc National de Taï (Côte d’Ivoire).. Bois et Forêts des Tropiques 179: 27 - 46. Bradley, D.G., MacHugh, D.E., Cunningham, P. & Loftus, R.T. (1996): Mitochondrial diversity and the origins of African and European cattle. Proceedings of the National Academy of Sciences, USA 93: 5131 - 5135. Bradley, M. (2004): Wood Buffalo National Park Bison Survey, March 2003. Unpublished Parks Canada Report 29 pp. Bradley, M., Handle, A., Sargent, P. (2004): Wood Buffalo National Park Bison Survey, March 2002. Unpublished Parks Canada Report, 30 pp. Bradshaw, R.H.W., Hannon, G.E. & Lister, A.M. (2003): A long-term perspective on ungulatevegetation interactions. Forest Ecology and Management 181: 267-280. Bremen, A. von (1072): Gesta Hammaburgensis ecclesiae pontificum. Boek IV. Quellen des Neunten und Elften Jahrhunderts zur Geschichte der Hamburgischen Kirche und des Reichs. Ausgewählte Quellen zur Deutsche Geschichte des Mittelalters, 1961, deel 11: 160 - 503. Wissenschaftliche Buchgesellschaft, Darmstadt.

391

Black Cyan Page 392

392


Brentjes, B. (1967): Die Tierwelt von Chatal Hüyük. Säugetierkundliche Mitteilungen 15(4): 317 – 332. Brincken, J. (1826): Mémoire déscriptif sur la Forêt de Bia³owie¿a en Lithuanie. N. Glücksberg, Varsovie. Brockmann-Jerosch, H. (1918): Das Lauben und sein Einfluss auf die Vegetation der Schweiz. Jahresbericht der Geografisch-Ethnografischen Gesellschaft in Zürich: 3 - 21. Brockmann-Jerosch, H. (1936): Futterlaubbäume und Speiselaubbäume. Berichte der Schweizerischen Botanischen Gesellschaft 46: 594 - 613. Brombacher, C., & Jacomet, S. (1997): IV. Ackerbau, Sammelwirtschaft und Umwelt: Ergebnisse archäobotanischer Untersuchungen. In: J. Schibler, H. Hüster-Plogmann, S. Jacomet, C. Brombacher, E. Gross-Klee & A. Rast-Eicher: Ökonomie und Ökologie neolithischer und bronzezeitlicher Ufersiedlungen am Zürichsee. Ergebnisse der Ausgrabungen Mozartstrasse, Kanalisationssanierng Seefeld, AKAD/Pressehaus und Mythenschloss in Zürich. Monographien der Kantonsarchäologie Zürich 20: 220-299. Fotorotar AG, Zürich und Egg. Brugal, J.P. (1987): Cas de ‘nanisme’ insulaire chez l’aurochs. Actes du Congrès National des Sociétés Savantes. Section des Sciences 112(2): 53 - 66. Bruijn, O. de (1977): De zeggen in het stroomgebied van de Drentse A. Doctoraalscriptie, RU Groningen. 2 Volumes. Brussaard, L. (1985): A pedobiological study of the dung beetle Typhaeus typhoeus (Coleoptera, Geotrupidae). Thesis. Wageningen. Büchner, E. (1895): Das allmähliche Aussterben des Wisents (Bison bonasus (Linn.)) im Forste von Bjelowjesha. Mémoires de l’Académie Impériale des Sciences de St.-Petersbourg. Série 8, Classe physico-mathématique. Vol. 3(2). Buechner, H.K. & Dawkins, H.C. (1961): Vegetation change induced by elephants and fire in Murchison Falls National Park, Uganda. Ecology 42: 752 - 766. Bulas, G., Thomas, L.L. & Whitfield, F.J. (1961): Polish-English, Volume II. Mouton & Co., The Hague. Buntjer, J.B. (1997): DNA repeats in the vertebrate genome as probes in phylogeny and species identification. Thesis Utrecht. Buntjer, J.B., Otsen, M., Nijman, I.J., Kuiper, M.T.R. & Lenstra, J.A. (2002): Phylogeny of bovine species based on AFLP fingerprinting. Heredity 88: 46-51. Bunzel-Drüke, M. (1997): Klima oder Übernutzung - Wodurch starben Grosstieren am Ende des Eiszeitalters aus?. Natur und Kulturlandschaft 2: 152 - 193. Bunzel-Drüke, M., Drüke, J. & Vierhaus, H. (1994): Quaternary Park - Überlegungen zu Wald, Mensch und Megafauna.. ABU info 17/18 (4/93 & 1/94): 4 – 38.

Black Cyan Page 393

REFERENCES

Burczak-Abramowitz, N.I., Meladze, G.K. & Czikowani, G.Cz. (1991): Tur, Bos primigenius Boj. z pó¿nego holocenu Kaukazu. Przegl¹d Zoologiczny XXXV (1-2): 123 - 131. Buskens, R., Erve, F. van, Moller Pillot, H. & Straaten, J. van der (1998): De Pripyat in WitRusland - Verslag van een excursie in 1997. Stichting Saxifraga, Tilburg. Buttenschön, J. & Buttenschön, R.M. (1978): The effect of browsing by cattle and sheep on trees and bushes. Natura Jutlandica 20: 79 - 94. Buttenschön, J. & Buttenschön, R.M. (1985): Grazing experiments with cattle and sheep on nutrient poor acidic grassland and heath. Natura Jutlandica 21(7): 117 - 140. Caesar, G.I. (58 - 52 v. Chr.): Commentarii de Bello Gallico, Libri I - VII. (in various translations). Campbell, D.G. (1991): Gap formation in tropical forest canopy by elephants, Oveng, Gabon, Central Africa.. Biotropica 23(2): 195 - 196. Carbyn, L.N., Lunn, N.J. & Timoney, K. (1998): Trends in the distribution and abundance of bison in Wood Buffalo National Park. Wildlife Society Bulletin 26(3): 463 - 470. Carbyn, L.N., Oosenbrug, S.M. & Anions, D.W. (1993): Wolves, bison....and the dynamics related to the Peace-Athabasca Delta in Canada’s Wood Buffalo National Park. Circumpolar Research Series nr 4. University of Alberta, Edmonton. Cerilli, E. & Petronio, C. (1991): Biometrical variations of Bos primigenius Bojanus 1827 from middle Pleistocene to Holocene. In:’Proceedings of the international symposium ‘Ongulés/ungulates’’, Toulouse, France: 37 - 42. Chaix, L. (1994): L’aurochs d’Étival & les aurochs de Franche-Comté. In: Aurochs, le retour aurochs, vaches & autres bovins de la Préhistoire à nos jours. Centre Jurassien du Patrimoine, Lons-le-Saunier: 67 - 75. Chauvet, J.-M., Deschamps, E.B. & Hillaire, C. (1998): De grot Chauvet - de oudste grotschilderingen ter wereld. Jan van Arkel, Utrecht. Christy, C. (1929): The African buffaloes. Proceedings of the Zoological Society of London: 445 - 462. Churcher, C.S. (1972): Late Pleistocene vertebrates from archaeological sites in the plain of Kom Ombo, Upper Egypt. Life Sciences Contributions, Royal Ontario Museum 82. Clason, A.T. (1961): Twee oerosschedels uit Drente. Varia Bio-Archaeologica 14: 183 - 188. Clason, A.T. (1965): De oeros in Groningen en Noord-Drente. Groninger Volksalmanak: 167 - 178. Clason, A.T. (1986): Het voorkomen van het wilde paard Equus ferus Boddaert 1785 in Nederland vanaf het Laat-Glaciaal. Lutra 29(2): 303 - 306. Clason, A.T. & Es, L. van (1992): De oeros - Bos primigenius - van Britsum (Fr.). Paleo-aktueel 3: 81 - 83. Clason, A.T. & Es, L.J.M. van (1993): De oeros - Bos primigenius - van Britsum (Fr.) gedateerd. Paleo-aktueel 4: 110.

393

Black Cyan Page 394

394


Clason, A.T., Laarman, F.J. & Vries, L.S. de (2000): Oeros en eland.. Cranium 17(1): 15 - 16. Clavel, M. (1967): La forêt en Gaule d’après les sources littéraires. In: Actes du Colloque sur la Forêt, Besançon, 21-22 octobre 1966. Cahiers d’Etudes Comtoises 12. Annales Littéraires de l’Université de Besançon, Vol. 88. Clot, A. & Duranthon, F. (1990): Les mammifères fossiles du Quaternaire dans les Pyrénées.. Museum d’Histoire Naturelle de Toulouse. Clutton- Brock, J. (1986): New dates for old animals: the reindeer, the aurochs and the wild horse in prehistoric Britain. Archaeozoologia, Mélanges: 111 - 117. Conry, P.J. (1981): Habitat selection and use, movements and home range of Malayan gaur (Bos gaurus hubbackii) in Central-Pahang, Malaysia. M.S. Thesis, University of Montana, Missoula, Montana, USA. Conry, P.J. (1989): Gaur Bos gaurus and development in Malaysia. Biological Conservation 49: 47 - 65. Conwentz, H. (1897): Moorbruecken im Thal der Sorge auf der Grenze zwischen West- und Ostpreussen. Danzig. Cornelissen, P. & Vulink, J.T. (1995): Begrazing in jonge wetlands. Flevobericht nr. 367. Rijkswaterstaat, Directie IJsselmeergebied. Cornelissen, P. & Vulink, J.T. (1996): Grote herbivoren in wetlands - evaluatie begrazingsbeheer Oostvaardersplassen. Flevobericht nr. 399. Directoraat-Generaal Rijkswaterstaat. Cossío, J.M. de (1943): Los toros: tratado técnico e histórico. Deel I. Espasa-Calpe, Madrid. Cremers, J. (1927): Bos primigenius Bojanus, oeros. Natuurhistorisch Maandblad 16(4): 44 - 45. Daszkiewicz, P. (1997): Nieznany list Baltazara Hacqueta do Georges Cuviera.. Kwartalnik Historii Nauki i Techniki 42 (3-4): 139 - 141. Daszkiewicz, P. & Aikhenbaum, J. (1998): Aurochs, retour d’un animal préhistorique....ou manipulation scientifique?. Le Courrier de l’Environnement de l’INRA 33, avril: 73 - 79. Dathe, H. (1980): Begründung der Einrichtung eines Zuchtbuches für das urähnliche Rind. In: Internationales Zuchtbuch für Auerochsen: 5 - 6. Berlin. Degerbøl, M. & Fredskild, B. (1970): The urus (Bos primigenius Bojanus) and neolithic domesticated cattle (Bos taurus domesticus Linné) in Denmark. With a revision of Bos remains from the kitchen middens. Det Kongelige Danske Videnskabernes Selskab, Biologiske Skrifter 17(1): 1 – 234. Degerbøl, M. & Iversen, J. (1945): The bison in Denmark. A zoological and geological investigatin of the finds in Danish Pleistocene deposits. Danmarks Geologiske Undersøgelser. II Række, nr. 73. Dementiev, G.-P. (1958): Quelques notes sur l’aurochs. Mammalia, 22: 161 - 167.

Black Cyan Page 395

REFERENCES

Dinnin, M.H. & Sadler, J.P. (1999): 10.000 Years of change: the Holocene entomofauna of the British Isles. Journal of Quaternary Science 14(6) : 545 - 562. Dio Cassius (± 200): Historiae/ Dio’s Roman history. Translation E. Cary. The Loeb Classical Library. 9 Volumes 1968-1970. Dirkx, G.H.P., Smeerdijk, D.G. van & Haaster, H. van (1998): Historische ecologie van het Reestdal – Een onderzoek naar historische referentiebeelden voor natuur. Assen. Het Drentse Landschap/ Wageningen. DLO-Staring Centrum. Djoshkin, W.W. & Safonov, W.G. (1972): Die Biber der alten und neuen Welt. Die Neue Brehm-Bücherei nr. 437. A. Ziemsen Verlag, Wittenberg-Lutherstadt. Dobie, J.F. (1980): The longhorns. University of Texas Press, Austin and London. Döhle, H.J. (1990); Osteologische Untersuchungen am Ur (Bos primigenius Bojanus 1827) von Neumark-Nord. In: D. Mania, M. Thomae, T. Litt & T. Weber: Neumark – Gröbern. Beiträge zur Jagd des mittelpaläolithischen Menschen:177-191. Deutscher Verlag der Wissenschaften, Berlin. Domecq y Diez, A. (1986): El toro bravo. Espasa-Calpe, Madrid. Doroszewski, W. (1958 - 1969): S³ownik jêzyka polskiego. 10 Volumes. Polska Akademia Nauk, Warszawa. Driesch, A. von den & Boessneck, J. (1974): Kritische Anmerkungen zur Widerristhöhenberechnung aus Längenmaszen vor- und frühgeschichtlichen Tierknochen. Säugetierkundliche Mitteilungen 22: 325 - 348. Driesch, A. von den & Boessneck, J. (1976): Zur Grösze des Ures, Bos primigenius Bojanus 1827, auf der Iberischen Halbinsel. Säugetierkundliche Mitteilungen 24(1): 66 - 77. Duerst, J.U. (1899): Die Rinder von Babylonien, Assyrien und Ägypten und ihr Zusammenhang mit den Rindern der alten Welt. Commissions-Verlag von G. Reimer, Berlin Duerst, J.U. (1931): Grundlagen der Rinderzucht. Springer, Berlin. Dugmore, R.A. (1914): Romance of the beaver. J.P. Lippincott Company, Philadelphia. Duncan, P. (1992): Horses and grasses; the nutritional ecology of equids and their impact on the Camargue. Springer. Dunka, S., Fejér, L. & Papp, F. (2003): A Közép-Tiszántúl vízi története. Vízügyi Történeti Füzetek 16. Budapest. Dusburg, P. von (1679): Chronik des Preussenlandes. Ausgewählte Quellen zur Deutschen Geschichte des Mittelalters. Band XXV, 1984. Wissenschaftliche Buchgesellschaft, Darmstadt. Vertaling: K. Scholz, D. Wojtecki. Edlin, H.L. (1969): New Forest: Forestry Commission Guide. Her Majesty’s Stationery Office, London.

395

Black Cyan Page 396

396


Edwards, C.J., MacHugh, D.E., Dobney, K.M., Martin, L., Russell, N., Horwitz, L.K., McIntosh, S.K., MacDonald, K.C., Helmer, D., Tresset, A., Vigne, J.-D. & Bradley, D.G. (2004): Ancient DNA analysis of 101 cattle remains: limits and prospects. Journal of Archaeological Science 31(6): 695-710. Eichwald, E. (1830): Naturhistorische Skizze von Lithauen, Volhynien und Podolien in geognostisch-mineralogischer, botanischer und zoölogischer Hinsicht. Wilna . Ekman, J, (1972): The urus female (Bos primigenius Boj.) from Slågarp, Southern Sweden. Zoologica Scripta 1(5): 203 - 205. Ekström, J. (1993): The Late Quaternary history of the urus (Bos primigenius Bojanus 1827) in Sweden. Lundqua Thesis, volume 29. Lund University. Ellenberg, H. (1986): Vegetation ecology of Central Europe. Cambridge University Press. Empel, W. (1962): Morphologie des Schädels von Bison bonasus (Linnaeus 1758). Acta Theriologica VI, 4: 53 - 112. Erny-Rodmann, C., Gross-Klee, E., Haas, J.N., Jacomet, S., & Zoller, H. (1997): Früher ’human impact’ und Ackerbau im Übergangsbereich Spätmesolithikum/Frühneolithikum im schweizerischen Mittelland. Jahrbuch der Schweizerischen Gesellschaft für Ur- und Frühgeschichte 80: 27-56. Ervynck, A., Neer, W. Van, Lentacker, A. (1999): Introduction and extinction of wild animal species in historical times: the evidence from Belgium. In: N. Benecke (ed.). The Holocene history of the European vertebrate fauna: 399-407. Verlag Marie Leidorf. Es, L.J.M. van (1990): De oeros in Noord- en Midden-Nederland. Doctoraal onderzoek, Groningen. Escherich, G. (1917): In den Jagdgründen des Zaren. In:’Bialowies in deutscher Verwaltung’. Militärforstverwaltung Bialowies: 192 - 218. Estévez, J. & Saña, M. (1999): Auerochsenfunde auf der Iberischen Halbinsel.. In: G.-C. Weniger (Hrsg.). Archäologie und Biologie des Auerochsen: 119 – 131. Neanderthal Museum. Ewald, R. & Laurer, G. (1911): Über die Hornentwicklung des Bos primigenius Boj.. Zentralblatt für Mineralogie, Geologie und Paläontologie 21: 684 - 687. Faliñski J.B. (1968): Park narodowy w Puszczy Bia³owieskiej. Warszawa. Faliñski, J.B. & Faliñska, K. (1986): Vegetation dynamics in temperate lowland primeval forests; ecological studies in Bia³owie¿a forest. Dordrecht, Junk. Felius, M. (1995): Cattle breeds: an encyclopedia. Misset, Doetinchem. Flechier, M. (1695): La vie du cardinal Commendon; ou l’on voit ses voyages, ambassades........ Amsterdam. Translation of the work by A.M. Graziani: De vita Joannis Francisci Commendoni cardinalis. Parisiis 1669.

Black Cyan Page 397

REFERENCES

Flueck, W.T. (2000): Population regulation in large northern herbivores: evolution, thermodynamics, and large predators. Zeitschrift für Jagdwissenschaft 46: 139-166. Fleming, H.F. von (1719): Der vollkommene teutsche Jäger. Leipzig. Forcellini, E. (1940): Lexicon totius latinitatis. 6 Delen. Patavii. Typis Seminarii. Frahm, K. (1982): Rinderrassen in den Ländern der europäischen Gemeinschaft. Ferdinand Enke Verlag, Stuttgart. Francfort, H.- P., Sacchi, D., Sher, J.A., Soleilhavoup, F. & Vidal, P. (1993): Art rupestre du bassin de Minusinsk: nouvelles recherches franco-russes. Arts Asiatiques XLVIII: 5 - 52. Franz, L. (1967): Was war der Schelch?. Zeitschrift für deutsches Altertum und deutsche Literatur 96 : 74 - 78. Fraser, A. (1972): The bull. Osprey Publishing. Fremery, N.C. de (1831): Over eenen hoorn en gedeelte des bekkeneels van Bos primigenius, in february 1825 bij de Eembrugge gevonden. C.G. Sulpke, Amsterdam. French, M.H., Johansson, I. & Joshi, M.R. (1966): European breeds of cattle - Vol I en II. FAO Agricultural Studies nr. 67. Frevert, W. (1957): Rominten. BLV-Verlag, München - Bern - Wien. Fröhlich, J. (1955): Wald und Wild im Urzustand. Allgemeine Forstzeitung 66: 192 - 193. Fruziñski, B., £abudzki, L. & Wlaze³ko, M. (1975): Debarking impact upon silvicultural values of deciduous thickets. Polish Ecological Studies 1(2): 51 - 60. Fuller, W.A. (1966): The biology and management of the bison of Wood Buffalo National Park. Ottawa. Fullton, J. (1971): Bullfighting. The Dial Press, New York. Gaillard, M.-J., Birks, H.J.B., Emanuelsson, U., Karlsson, S., Lagerås, P. & Olausson, D. (1994): Application of modern pollen/land-use relationships to the interpretation of pollen diagrams; reconstructions of land-use history in south Sweden, 3000 - 0 BP. Review of Palaeobotany and Palynology 82: 47 - 73. Garrison, G.C. (1967): Pollen stratigraphy and age of an early postglacial beaver site near Columbus, Ohio. The Ohio Journal of Science, 67(2): 96 - 105. Garry, C.E., Schwert, D.P., Baker, R.G., Kemmis, T.J., Horton, D.G. & Sullivan, A.E. (1990): Plant and insect remains from Wisconsinan interstadial-stadial transition at Wedron North Central Illinois, USA. Quaternary Research Orlando 33(3): 387-399. Gautier, A. (1968): Mammalian remains of the northern Sudan and Southern Egypt. In: ‘The prehistory of Nubia’, Wendorf, F. (ed.) Vol I: 80 - 99. Gautier, A. (1988): The final demise of Bos ibericus?. Sahara 1: 37 - 48.

397

Black Cyan Page 398

398


Gautier, A. (1993a): Mammifères holocènes du Sahara d’après l’art rupestre et l’archéozoologie. Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, Vol. XXVI, fascicolo II: 261 - 267. Gautier, A. (1993b): Fossiele zoogdieren in de Vier Ambachten. In: ‘Over de Vier Ambachten, 750 jaar Keure, 500 jaar Graaf Jansdijk’. A.M.J. de Kraker, H. van Royen, M.E.E. De Smet (red.): 71 - 78. Gautier, A. (1996): Les restes animaux de la Grotte d’Escoural.. Recherches Préhistoriques à la Grotte d’Escoural, Liège, ERAUL, Vol. 65: 323 - 336. Gautier, A. (2002): The evidence for the earliest livestock in North Africa: or adventures with large bovids, ovicaprids, dogs and pigs. In: F.E. Hassan (ed.): Droughts, food and cultureecological change and food security in Africa’s later prehstory. Kluwer Academic/ Plenum Publishers. Gêbczyñska, Z., Gêbczyñski, M. & Martynowicz, E. (1991): Food eaten by the free-living European bison in Bia³owie¿a Forest. Acta Theriologica 36: 307 - 313. Gedda, B., Lemdahl, G. & Gaillard, M.-G. (1999): Late glacial and early Holocene environments inferred from a tufa deposit at Fyledalen, S. Sweden. GFF121: 33-41. Gehasse, E.F. (1995): Ecologisch-archeologisch onderzoek van het Neolithicum en de Vroege Bronstijd in de Noordoostpolder met de nadruk op vindplaats P14. Thesis, Amsterdam. Geist, V. (1996): Buffalo nation: history and legend of the North American bison. Voyageur Press Inc. Gentry, A., Clutton-Brock, J. & Groves, C.P. (1996): Case 3010 - Proposed conservation of usage of 15 mammal specific names based on wild species which are antedated by or contemporary with those based on domestic animals. The Bulletin of Zoological Nomenclature 53(1): 28 - 37. Gentry, A., Clutton-Brock, J. & Groves, C.P. (2004):The naming of wild animal species and their domestic derivatives. Journal of Archaeological Science 31(5): 645-651. Germonpré, M. (1993): Osteometric data on Late Pleistocene mammals from the Flemish Valley, Belgium. Studiedocument van het Koninklijk Belgisch Instituut voor Natuurwetenschappen, 72. Germonpré, M. & Ervynck, A. (1988): Pleistocene zoogdierresten te Uitbergen (Oost-Vlaanderen, België). Cranium 5(1): 5 – 7. Gesner, C. (1602): Historia animalium. Liber I. De quadrupedibus viviparis. Francofurti. Gesner, C. (1620): Historia animalium. Liber I. De quadrupedibus viviparis. Francofurti. Giffen, A. E. van (1913): Die Fauna der Wurten. Teil I. Dissertatie, Groningen. E.J. Brill, Leiden.

Black Cyan Page 399

REFERENCES

Gilibert, J.E. (1805): Abrégé du système de la nature de Linné, histoire des mammaires ou des quadrupèdes et cétacées. Lyon, Paris. Gilmore, L.O., Fechheimer, N.S. & Baldwin, C.S. (1961): Inheritance of black hair pattern in cattle lacking the extension factor for black (E). IV. Partitioning phenotypes by castration. The Ohio Journal of Science 61(5): 273 - 277. Girling, M.A. (1989): Mesolithic and later landscapes interpreted from the insect assemblages of West Heath Spa, Hampstead. In: D. Collins & D. Lorimer (eds.). Excavations at the Mesolithic Site on West Heath, Hampstead 1976 – 1981: 72 - 89. BAR British Series 217. Gispen, W.H., Oosterhoff, B.J., Ridderbos, H.N., Unnik, W.C. van & Visser, P. (1975): Bijbelse encyclopedie. 2 Volumes. J.H. Kok - Kampen. Glover, J. (1963): The elephant problem at Tsavo. East African Wildlife Journal 1: 1 - 10. Gobet, E., Tinner, W., Hochuli, P.A., Van Leeuwen, J.F.N. & Ammann, B. (2003): Middle to late Holocene vegetation history of the Upper Engadine (Swiss Alps): the role of man and fire. Vegetation History and Archaeobotany 12: 143-163. Godwin, H. (1975): The history of the British flora; a factual basis for phytogeography. London. Goetcheus, V.G. & Birks, H.H. (2001): Full-glacial upland tundra vegetation preserved under tephra in the Beringia National Park, Seward Peninsula, Alaska. Quaternary Science Reviews 20: 135-147. Goetz, H.-W. & Welwei, K.-W. (1995): Altes Germanien. Auszüge aus den antiken Quellen über die Germanen und ihre Beziehungen zum römischen Reich. 2 Volumes.. Wissenschaftliche Buchgesellschaft, Darmstadt. Goodman, T., Donart, G.B., Kiesling, H.E., Holeczek, J.L., Neel, J.P., Manzanares, D. & Severson, K.E. (1989): Cattle behavior with emphasis on time and activity allocations between upland and riparian habitats. In: R.E. Gresswell, B.A. Barton & J.L. Kershner (eds.). Practical approaches to riparian resource management; an educational workshop: 95-102. US Bureau of Land Management. Billings, Montana. Górski, A. & Gru¿ewski, M. (1994): A contribution to the knowledge of the quaternary fauna of NE Poland. Przegl¹d Zoologiczny XXXVIII (1-2): 131 - 134. Gould, G.N. & Morgan, K.G. (1934): Acorn poisoning. The Veterinary Record 14(2): 33 - 35. Gramsch, B. (1987): Betrachtungen zum Ur-fund am Schlaatz bei Potsdam. Veröffentlichungen des Museums für Ur- und Frühgeschichte Potsdam, Band 21: 69-74. Greig, J. (1989): From lime forest to heathland - five thousand years of change at West Heath Spa, Hampstead, as shown by the plant remains. In: D. Collins & D. Lorimer (eds.). Excavations at the Mesolithic Site on West Heath, Hampstead 1976 – 1981: 89 - 99. BAR British Series 217.

399

Black Cyan Page 400

400


Grigson, C. (1973): The comparative craniology of Bos taurus L., B. indicus L., B. primigenius Boj. and B. namadicus Falc.. Ph.D. Thesis, London University. Grigson, C. (1974): The craniology and relationships of four species of Bos. 1. Basic craniology: Bos taurus L. and its absolute size. Journal of Archaeological Science 1(4): 353 - 379. Grigson, C. (1976): The craniology and relationships of four species of Bos. 3. Basic craniology: Bos taurus L. Sagittal profiles and other non-measurable characters. Journal of Archaeological Science 3(2): 115 – 136. Grigson, C. (1978): The craniology and relationships of four species of Bos. 4. The relationships between Bos primigenius Boj. and B. taurus L. and its implications for the phylogeny of the domestic breeds. Journal of Archaeological Science 5(2): 123 - 152. Grimm, J.L. & Grimm, W.K. (1854 - 1954): Deutsches Wörterbuch. Hirzel, Leipzig. Groenman-Van Waateringe, W. (1978): The impact of Neolithic man on the landscape in the Netherlands. In: S. Limbrey & J.G. Evans (eds.). The effect of man on the landscape: the lowland zone. Council for British Archaeology. Research Report nr. 21 : 135-146. Groenman-Van Waateringe, W. (1993) : The effects of grazing on the pollen production of grasses. Vegetation History and Archaeobotany 2: 157-162. Gromova, V.I. (1930): Note préliminaire sur le Bos primigenius Boj. de l’URSS.. Comptes Rendus de l’Académie des Sciences de l’URSS: 85 - 90. Gromova, V.I. (1931): Contribution à la connaissance de l’ure (Bos primigenius Boj.) de l’Europe orientale et de l’Asie septentrionale. Annuaire du Musée Zoologique de l’Académie des Sciences de l’URSS, Leningrad 32(3): 293 - 364. Groot Bruinderink, G.W.T.A., Van Wieren, S.E., Hazebroek, E., Den Boer, M.H., Maaskamp, F.J.M., Lamers, W., Slim, P.A. & De Jong, C.B. (1997): De ecologie van hoefdieren. In: S.E. van Wieren, G.W.T.A. Groot Bruinderink, I.T.M. Jorritsma & A.T. Kuitert (red.). Hoefdieren in het boslandschap: 31 - 69. Backhuys Publishers, Leiden. Groves, C.P. (1981): Systematic relationships in the Bovini (Artiodactyla, Bovidae). Zeitschrift für zoologische Systematik und Evolutionsforschung 19: 264 - 278. Grüß, J. (1931): Zwei altgermanische Trinkhörner mit Bier- und Metresten. Praehistorische Zeitschrift 22 : 180-191. Guintard, C. (1988): L’aurochs et ses essais de reconstitution. Thèse pour le Doctorat Vétérinaire, diplôme d’état, Nantes. Guintard, C. & Denis, B. (1996): Pour un standard de l’aurochs de Heck. Ethnozootechnie 57: 25 - 30. Guintard, C. & Tardy, F. (1994): Les bovins de l’Ile Amsterdam - un exemple d’isolement génétique. In: ‘Aurochs, vaches & autres bovins de la préhistoire à nos jours’: 203 - 209. Centre Jurassien du Patrimoine, Lons-le-Saunier.

Black Cyan Page 401

REFERENCES

Guthrie, R.D. (1984): Mosaics, allelochemics and nutrients: an ecological theory of the late Pleistocene megafaunal extinctions. In: P.S. Martin & R.G. Klein (eds.): Quaternary extinctions - a prehistoric revolution: 259 - 298. The University of Arizona Press. Tucson, Arizona. Guthrie, R.D. (1990): ‘Frozen fauna of the Mammoth Steppe - The story of Blue Babe’. The University of Chicago Press, Chicago and London. Gysseling, M. (1960): Toponymisch woordenboek van België, Nederland, Noord-Frankrijk en West-Duitsland (vóór 1226). Deel II. Belgisch Interuniversitair Centrum voor Neerlandistiek. Hahne, J. & Melles, M. (1997): Late- and post-glacial vegetation and climate history of the south-western Taymyr Peninsula, Central Siberia, as revealed by pollen analysis of a core from Lake Lama. Vegetation History and Archaeobotany (6): 1 - 8. Halder, U. (1976): Ökologie und Verhalten des Banteng (Bos javanicus) in Java. Mammalia Depicta, Beihefte zur Zeitschrift für Säugetierkunde. Heft 10. Verlag Paul Parey, Hamburg und Berlin. Hall, S.J.G. (1988): Chillingham Park and its herd of white cattle: relationships between vegetation classes and patterns of range use. Journal of Applied Ecology 25: 777 - 789. Hall, S.J.G. & Moore, G.F. (1986): Feral cattle of Swona, Orkney Islands. Mammal Review 16(2): 89 - 96. Hannon, G.E. (1999): The use of plant macrofossils and pollen in the palaeoecological reconstruction of vegetation. Thesis. Acta Universitatis Agriculturae Sueciae 106. Harrison, D. L. (1964): The mammals of Arabia. Vol. I. Ernest Benn Ltd., London. Hartz, N., & Winge, H. (1906): Om uroksen fra Vig. Saaret og dræbt med flintvaaben. Aarbøger for Nordisk Oldkyndighed og Historie. II. R., 21: 225-236. Hasselt, G. van (1805): G. van Hasselt’s Bijdragen voor den Burg van Nijmegen, tevens, onder d’oude dieren daar van die auer-ox gegischt, wiens kop bij den doorbraak des Banddijks te Weurd in dit jaar aldaar te voorschijn kwam. J.H. Meuleman, Jr., Arnhem. Hawley, A.W.L. (1987): Bison and cattle use of forages. In: H.W. Reynolds & A.W.L. Hawley, (eds.): Bison ecology in relation to agricultural development in the Slave River Lowlands, NWT: 49 - 52. Healey, M.C. (1994): Effects of dams and dikes on fish habitat in two Canadian river deltas. In: W.J. Mitsch (ed.). Global wetlands: old world and new: 385 - 398.. Elsevier Science. Heck, H. (1934): Der Ur. Das Tier und Wir, Monatszeitschrift für alle Tierfreunde. SonderNummer, März. Heck, H. (1949): Die Rückzüchtung ausgestorbener Tiere. Orion 4: 401 - 410. Heck, H. (1951): The breeding-back of the aurochs. Oryx 1(3): 117 - 122.

401

Black Cyan Page 402

402


Heck, H. (1980): Der neue Auerochse. In: H. Dathe (Hrsg.). Internationales Zuchtbuch für Auerochsen: 7 - 14. Berlin. Heck, L. (1934): Über die Neuzüchtung der Ur oder Auerochs. Berichte der Internationalen Gesellschaft zur Erhaltung des Wisents 3(4): 225 - 294. Heck, L. (1952a): Über den Auerochsen und seine Rückzüchtung. Jahrbücher des Nassauischen Vereins für Naturkunde 90: 107 - 124. Heck, L. (1952b): Tiere - mein Abenteuer. Erlebnisse in Wildnis und Zoo. Im Verlag Ullstein, Wien. Hedemann, O. (1939): Dzieje Puszczy Bia³owieskiej w Polsce przedrozbiorowej (w okresie do 1798 roku). Instytut Badawczy Lasów Pañstwowych. Rozprawy i sprawozdania. Seria A, nr. 41. Heikkilä, R. & Mikkonen, T. (1992): Effects of density of young Scots Pine (Pinus sylvestris) stand on moose (Alces alces) browsing. Acta Forestalia Fennica nr. 231. Hengeveld, G.J. (1865): Het rundvee, zijne verschillende soorten, rassen en veredeling. 2 Delen.. De Erven Loosjes, Haarlem. Heptner, V.G. & Naumow, N.P. (1966): Die Säugetiere der Sowjetunion. Band I: Paarhufer und Unpaarhufer. VEB Gustav Fischer Verlag, Jena. Herberstein, S. von (1557a): Moscovia der Hauptstat in Reissen. Michael Zimmerman, Wien. Herberstein, S. von (1557b): Rerum Moscoviticarum commentarii. In aedibus Ioannis Steelsii, Antverpiae. Herlin, A.H. (1994): Effects of tie-stalls or cubicles on dairy cows in grazing or zero-grazing situations: studies on behaviour, locomotion, hygiene, health and performance. Thesis. Sveriges Landbruksuniversitet Uppsala. Herodotus (440 BC/ 1954): The Histories. Translation: A. de Sélincourt. Penguin Books. Herre, W. (1953): Wie sah der Auerochse aus?. Kosmos 49(11): 504 - 507. Herre, W. & Röhrs, M. (1990): Haustiere - zoologisch gesehen. Gustav Fischer Verlag. Heybroek, H.M. (1963): Diseases and lopping for fodder as possible causes of a prehistoric decline of Ulmus. Acta Botanica Neerlandica 12: 1 - 11. Heymanowski, K. (1972): The last mainstay of aurochs and the organization of their protection in the light of contemporary documents. Sylwan 116(9): 9 - 28. Hilzheimer, M. (1910): Wie hat der Ur ausgesehen?. Jahrbuch für Wissenschaftliche und Praktische Tierzucht 5: 42 – 93. Hilzheimer, M. (1917): Der Ur in Ägypten. Studien und Forschungen zur Menschen- und Völkerkunde 14: 9 – 16. Hilzheimer, M. (1937): Die Geschichte eines Wisenthorns und ein neuentdecktes Horn vom Ur (Bos primigenius). Zeitschrift für Wissenschaftliche Zoologie 150(1): 97 – 106. Hirsch, Th. (1863): Die littauischen Wegeberichte. Scriptores Rerum Prussicarum 2: 662 - 711.

Black Cyan Page 403

REFERENCES

Hoekstra, A. & Vulink, J.T. (1994): De sociale organisatie van een kudde Heckrunderen - het ontstaan van stiergroepen. Flevobericht nr. 353.. Rijkswaterstaat, Directie IJsselmeergebied, Lelystad. Hölzer, A, & Hölzer, A. (1988): Untersuchungen zur jüngeren Vegetations- und Siedlungsgeschichte im Blindensee-Moor (Mittlerer Schwarzwald). Carolinea 46: 23-30. Hofmann, R.R. (1989): Evolutionary steps of ecophysiological adaptation and diversification of ruminants. Oecologia 78: 443 - 457. Hofmann, R.R. & Scheibe, K. (1997): Überlegungen zur Rekonstruktion der natürlichen Grosstierfauna Mitteleuropas auf der Grundlage ihrer morphophysiologischen Differenzierung und ihrer potentiellen ökologischen Nischen.. Natur- und Kulturlandschaft 2: 207 - 214. Hofmann, R.R. & Stewart, D.R.M. (1972): Graser or browser: a classification on the stomach structure and feeding habits of East African ruminants. Mammalia 36(2): 226 - 242. Hollander, A.N.J. den (1947): Nederzettingsvormen en problemen in de Groote Hongaarsche laagvlakte: een Europeesch ‘Frontier’ gebied. Amsterdam. Hoogerwerf, A. (1970): Udjung Kulon; the land of the last Javan Rhinoceros. Brill, Leiden. Hoppe-Dominik, B. (1988): Habitatpräferenz und Nahrungsansprüche des Waldbüffels, Syncerus caffer nanus, im Regenwaldgebiet der Elfenbeinküte. Dissertation. Universität Braunschweig. Hryniewiecki, B. (1938): Anton Schneeberger, 1530-1581: ein Schüler Konrads Gesners in Polen. Veröffentlichungen des Geobotanischen Institutes Rübel in Zürich, Heft 13. Hu, F.S. & Davis, R.B. (1995): Postglacial development of a Maine bog and palaeoenvironmental implications. Canadian Journal of Botany 73(4): 638-649. Hubback, T. (1938): Malayan gaur or seladang (Bibos gaurus hubbacki). Journal of the Bombay Natural History Society 40(1): 8 - 19. Huddard, D., Gonzalez, S. & Roberts, G. (1999): The archaeological record and mid-Holocene marginal coastal palaeoenvironments around Liverpool Bay. Journal of Quaternary Science 14(6) : 563 - 574. Hüster-Plogmann, H., Schibler, J. & Jacomet, S. (1999): The significance of aurochs as hunted animal in the Swiss Neolithic. In: G.-C. Weniger (Hrsg.). Archäologie und Biologie des Auerochsen: 151 - 160. Neanderthal Museum. International Commission on Zoological Nomenclature (2003): Opinion 2027 (Case 3010). Usage of 17 specific names based on wild species which are pre-dated by or contemporary with those based on domestic animals (Lepidoptera, Osteichthyes, Mammalia): conserved. Bulletin of Zoological Nomenclature 60(1) March 2003: 81-84.

403

Black Cyan Page 404

404


Isberg, O. (1962): Uroxen (Bos primigenius) i Sverige. Geologiska Föreningens Forhandlingar 84(4): 416 - 518. Jackson, S.T., Overpeck, J.T., Webb, T., Keattch, S.E. & Anderson, K.H. (1997): Mapped plant-macrofossil and pollen records of late Quaternary vegetation change in Eastern North America. Quaternary Science Reviews 16: 1-70. Janis, C.M. (1986): An estimation of tooth volume and hypsodonty indices in ungulate mammals, and the correlation of these factors with dietary preference. In: D.E., Santoro & J.-P., Sigogneau-Russell, D. (eds.). Teeth revisited. Proceedings of the VIIth International Symposium on Dental Morphology, Paris. Russell : 367 - 387. Mémoires du Muséum National d’Histoire Naturelle, Serie C, Sciences de la Terre 53. Janis, C.M. (1988): Correlation of relative muzzle width and relative incisor width with dietary preference in ungulates. Zoological Journal of the Linnean Society 92: 267 - 284. Jánossy, D. & Vörös, I. (1981): Pleistozäner Skelettfund des Ures (Bos primigenius Bojanus) von Nagybajom (Ungarn). Fragmenta Mineralogica et Palaeontologica 10: 79 - 96. Janssen, C.R. (1974): Verkenningen in de palynologie. Utrecht. Jaques, D.R. (1990): Vegetation habitat types of the Peace-Athabasca Delta: 1976 - 1989. Final Report. Parks Canada. Jarman, P.J. (1971): Diets of large mammals in the woodlands around Lake Kariba, Rhodesia. Oecologia 8: 157 - 187. Jêdrzejewska, B. & Jêdrzejewski, W. (1998): Predation in vertebrate communities: the Bia³owie¿a Primeval Forest as a case study. Springer. Jêdrzejewska, B., Jêdrzejewski, W., Bunevich, A.N., Mi³kowski, L. & Krasiñski, Z.A. (1997): Factors shaping population densities and increase rates of ungulates in Bia³owie¿a Primeval Forest (Poland and Belarus) in the 19th and 20th centuries. Acta Theriologica 42(4): 399 - 451. Jiu Lampo & Wei Qi (1980): Some animals from the Holocene of N.China. Vertebrata Palasiatica, Vol. XVIII, 4: 327 - 333. Joern, A. & Keeler, K.H. (1995): Getting the lay of the land: Introducing North American native grasslands. In: A. Joern & K.H. Keeler (eds.). The changing prairie - North American grasslands: 11 - 24. Oxford University Press. Johnston, C.A. (1994): Ecological engineering. In: W.J. Mitsch (ed.). Global wetlands - old world and new: 379 - 384. Johnston, C.A. & Naiman, R.J. (1987): Boundary dynamics at the aquatic-terrestrial interface: the influence of beaver and geomorphology. Landscape Ecology 1(1): 47 – 57. Johnston, C.A. & Naiman, R.J. (1990): Browse selection by beaver: effects on riparian forest composition. Canadian Journal of Forest Research 20: 1036 - 1043.

Black Cyan Page 405

REFERENCES

Joly, D.O. & Messier, F. (2001): Limiting effects of bovine brucellosis and tuberculosis on wood bison within Wood Buffalo National Park. Final Report, March 2001. University of Saskatchewan, Sasketoon. Jong, C.B. de, Cornelissen, P. & Vulink, J.T. (1997): Grote grazers in de Oostvaardersplassen - dieetsamenstelling op basis van faecesanalyse. Werkdocument 97.114 X. DirectoraatGeneraal Rijkswaterstaat. Jong, T. de (1998): Topstukken uit de diepte - Prehistorische dierenresten uit Noord-Brabant. Cranium 15(2): 84 - 110. Jonstonus, J. (1657): Historiae naturalis de quadrupedibus libri. Ioannus Iacobi Fil. Schipper, Amstelodami. Joshi, N.R., McLaughlin, E.A. & Phillips, R.W. (1957): Types and breeds of African cattle. FAO Agricultural Studies, nr. 37. Rome. Joshi, N.R. & Phillips, R.W. (1953): Zebu cattle of India and Pakistan. FAO Agricultural Studies, nr. 19, Rome. Kahlke, H.D. (1994): Die Eiszeit.. Urania Verlag. Kalis, A.J. & Meurers-Balke, J. (1998): Zur pollenanalytischen Untersuchung neolithischer Brunnensedimente - ein Zwischenbericht. In: ‘Brunnen der Jungsteinzeit’. Materialien zur Bodendenkmalpflege im Rheinland, nr. 11: 247 - 260. Rheinisches Amt für Bodendenkmalpflege. Rheinland Verlag, Köln. . Kar³owicz, J., Kryñski, A. & NiedŸwiedzki, W. (1919): S³ownik jêzyka polskiego. Tom VII. Warszawa. Kay, C.E. (1995): An alternative interpretation of the historical evidence relating to the abundance of wolves in the Yellowstone ecosystem. In: L.N. Carbyn, S.H. Fritts, & D.R. Seip. Ecology and conservation of wolves in a changing world. Canadian Circumpolar Institute, Occasional Publication nr. 35: 77-84. Kay, C.E. (1997): Testimony before the U.S. House of Representatives Subcommittee on National Parks and Public Lands. Oversight hearing on science and resource management in the national park system. February 27. Kazmin, V.D. & Smirnov, K.A. (1992): Winter nutrition, food resources and trophic influence of the European bison on forest phytocoenoses in the Central Caucasus. Byulletin Moskovskogo Obshchestva Ispytatelei Prirody Otdel Biologiceskii 97(2): 26 - 35. Kêdzierska, Z. (1959): Lustracje Województwa Rawskiego 1564 i 1570. PAN, Instytut Historii, Warszawa. Kêdzierska, Z. (1965): Lustracje Województwa Rawskiego XVII wieku. Instytut Historii PAN, Warszawa, Wroclaw.

405

Black Cyan Page 406

406


Keller, C. (1902): Die Abstammung der ältesten Haustiere. Zürich. Kelly, D.L. (2002): The regeneration of Quercus petraea (sessile oak) in southwest Ireland: A 25-year experimental study. Forest Ecology and Management 166(1-3): 207-226. Kendall, R.L. (1969): An ecological history of the Lake Victoria Basin.. Ecological Monographs 39: 121 - 176. Kinnaird, J.W., Welch, D. & Cummins, C. (1979): Selective stripping of rowan (Sorbus aucuparia L.) bark by cattle in North-east Scotland. Transactions Botanical Society of Edinburgh 43: 115 - 125. Klatt, B. (1912): Über die Veränderung der Schädelkapazität in der Domestikation. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, nr. 3: 153 - 179. Klaus, F.E.S. (1969): Im Urwald von Bialowies. Wild und Hund 72(16): 384 - 386. Klös, H.-G. & Wünschmann, A. (1968): Die Rinder. In: B. Grzimek (Hrsg.). Grzimek’s Tierleben: Enzyklopädie des Tierreiches. Teil XIII: 368 – 436. Kindler Verlag, Zürich. Kneller, M. & Peteet, D. (1999): Late-glacial to early Holocene climate changes from a central Appalachian pollen and macrofossil record. Quaternary Research Orlando 51(2): 133-147. Kobryñ, H. & Lasota-Moskalewska, A. (1989): Certain osteometric differences between the aurochs and domestic cattle. Acta Theriologica 34(4): 67 - 82. Kobryñczuk, F. & Kobryñ, H. (1975): Growth rate of bones in the postembryonic development of the European bison.. Acta Theriologica 20 (13): 151 - 157. Koch, W. (1927): Über angebliche Anzeichen von Domestikation am Skelett von Rindern. Zeitschrift für Morphologie und Ökologie der Tiere 7(3): 444 - 471. Koehler, O. (1952): Besprechung von ’Heck, L.: Über den Auerochsen und seine Rückzüchtung. Jahrb. d. Nass. Vereins für Naturkunde 90, 1952, 107-124’. Zeitschrift für Tierpsychologie, 9: 498-499. Koenigswald, W. von (1999): Palökologie und Vorkommen des pleistozänen Auerochsen (Bos primigenius BOJANUS 1827) im Vergleich zu den grossen Rindern des Pleistozän. In: G.C. Weniger (Hrsg). Archäologie und Biologie des Auerochsen: 23 - 33. Neanderthal Museum. Kokabi, M. (1994): Osteologische Untersuchung der Knochenfunde aus der Freilandstation von Rottenburg-Siebenlinden I. In: M. Kokabi & J. Wahl. Beiträge zur Archäozoologie und prähistorischen Anthropologie. Forschungen und Berichte zur Vor- und Frühgeschichte Baden-Württemberg 53: 197-211. Kooistra, L.I. (1996): Borderland farming. Possibilities and limitations of farming in the Roman Period and Early Middle Ages between the Rhine and Meuse. Thesis, Leiden. Krasiñska, M. & Krasiñski, Z.A. (1995): Composition, group size and spatial distribution of European bison bulls in Bia³owie¿a Forest. Acta Theriologica 40(1): 1 - 21.

Black Cyan Page 407

REFERENCES

Krasiñska, M. & Krasiñski, Z.A. (1997): Utilization of the Bia³owie¿a National Park by European bison. Parki Narodowe i Rezerwaty Przyrody 16(2): 63 - 77. Krasiñska, M. & Krasiñski, Z.A. (2004): ¯ubr – monografia przyrodnicza. SFP Hajstra. Warszawa - Bia³owie¿a. Krasiñski, Z.A. (1978): Dynamics and structure of the European bison population in the Bia³owie¿a primeval forest. Acta Theriologica 23: 3 - 45. Krasiñski, Z.A. & Raczyñski, J. (1967): The reproduction biology of European bison living in reserves and in freedom.. Acta Theriologica 12, 29: 407 - 444. Krüsi, B.O., Schütz, M., Bigler, H.C., Grämiger, H. & Achermann, G. (1998): Huftiere und Vegetation im Schweizerischen Nationalpark von 1917 bis 1997. In: R. Cornelius & R.R. Hofmann (Hrsg.). Extensive Haltung robuster Haustierrassen, Wildtiermanagement, Multi-Spezies-Projekte. Neue Wege in Naturschutz und Landschaftspflege? :62 - 74. Workshop des Instituts für Zoo- und Wildtierforschung (IZW), 25 - 26 März, Berlin. Krüsi, B.O., Schütz, M., Grämiger, H. & Achermann, G. (1996): Was bedeuten Huftiere für den Lebensraum Nationalpark. Eine Studie zu Nahrungsangebot und Waldverjüngung.. Cratschla 4(2): 51 - 63. Krüsi, B.O., Schütz, M., Wildi, O. & Grämiger, H. (1995): Huftiere, Vegetationsdynamik und botanische Vielfalt im Nationalpark. Ergebnisse von Langzeitbeobachtungen. Cratschla 3(2): 14 - 25. Kruytzer, E.M. (1949): De vindplaatsen van het oerrund Bos primigenius Boj. in Limburg in het bijzonder het moerasveen van Voerendaal. Natuurhistorisch Maandblad 38(3): 27 - 29. Kühlhorn, J. (1955): Der Auerochs von 1595. Zeitschrift für Säugetierkunde 20(2/3): 186. Kuiters, A.T., Slim, P.A. & Hees, A.F.M. van (1997): Spontane bosverjonging en hoefdieren. In: S.E. van Wieren, G.W.T.A. Groot Bruinderink, I.T.M. Jorritsma & A.T. Kuiters (red.). Hoefdieren in het boslandschap: 99-129. Backhuys Publishers, Leiden. Kuhn, W. (1977): Preseka. In: H. Weczerka (Hrsg.). Handbuch der historischen Stätten. Schlesien: 416 - 417. Stuttgart. Kulczyñski, S. (1949): Peat bogs of Polesie (Torfowiska Polesia). Mémoires de l’Académie Polonaise des Sciences et des Lettres. Classe des Sciences Mathématiques et Naturelles. Serie B: Sciences Naturelles, nr. 15: 1 - 356. Kuznetsov, G.V. & Lozinov, G.L. (1992): The estimation of the impact of moose (Alces alces) on the succession of vegetation of broad-leaved and taiga formations of the European part of the USSR. In: Proceedings of the International Symposium ‘Ongulés/Ungulates 91’. Sept. 2 - 6, 1991, Toulouse.

407

Black Cyan Page 408

408


Kwang-Chih Chang (1986): The Archaeology of Ancient China. Yale University Press, New Haven and London. Laarman, F.J. (2001): Archeozoölogie: aard en betekenis van de dierlijke resten. In: J.W.H. Hogestijn & J.H.M. Peeters (red.). De mesolithische en vroeg-neolithische vindplaats Hoge Vaart-A27 (Flevoland). Deel 16. Rapportage Archeologische Monumentenzorg 79. ROB Amersfoort. La Baume, W. (1944): Ostpreußische Funde vom Ur (Bos primigenius). Altpreußen 9: 28 - 32. La Baume, W. (1947): Diluviale Schädel vom Ur (Bos primigenius Bojanus) aus Toscana. Berichte der Schweizerischen Paläontologischen Gesellschaft 40(2): 299 - 308. Lagerås, P. & Bartholin, T. (2003): Fire and stone clearance in Iron Age agriculture: new insights inferred from the analysis of terrestrial macroscopic charcoal in clearance cairns in Hamneda, Southern Sweden. Vegetation History and Archaeobotany 12(2): 83-92. Landelijke Werkgroep Kritisch Bosbeheer (1980): Bosbeheer in opspraak 1975-1980. Documentatiemap 1. Kritisch Bosbeheer, Boxtel. Langbein, J. (1997): The ranging behaviour, habitat use and impact of deer in oak woods and heather moors of Exmoor and the Quantock hills. The British Deer Society. Lans, H.E. van der & Poortinga, G. (1986): Natuurbos in Nederland - een uitdaging.. IVN, Amsterdam. Larsen, E.J. & Ripple, W.J. (2003): Aspen age structure in the northern Yellowstone ecosystem: USA. Forest Ecology and Management 179(1-3): 469-482. Larter, N.C. & Gates, C.C. (1991): Diet and habitat selection of wood bison in relation to seasonal changes in forage quantity and quality. Canadian Journal of Zoology 69(4): 2677 - 2685. Larter, N.C., Sinclair, A.R.E. & Gates, C.C. (1994): The response of predators to an erupting bison, Bison bison athabascae, population. Canadian Field-Naturalist 108(3): 318-327. Lasota-Moskalewska, A. & Kobryñ, H. (1990): The size of aurochs skeletons from Europe and Asia in the period from the Neolithic to the Middle Ages. Acta Theriologica 35(1 - 2): 89 - 109. Laurent, J.M. & Wattenbach, W. (1986): Chronik der Slaven/ Helmold. Phaidon. Lauwerier, R.C.G.M. (1988): Animals in Roman times in the Dutch eastern river area. Nederlandse Oudheden 12. Project Oostelijk Rivierengebied 1. ROB - Amersfoort. Lawrence, M.J. & Brown, R.W. (1974): Mammals of Britain: their tracks, trails and signs. Blandford Press, London. Lazo, A. (1992): Caracteristicas poblacionales del ganado bovino asilvestrado de la Reserva Biológica de Doñana (S.O. de España). Doñana Acta Vertebrata 19(1-2): 85 - 96. Lazo, A. (1995): Ranging behaviour of feral cattle (Bos taurus) in Doñana National Park, S.W. Spain. Journal of Zoology, London 236: 359 - 369.

Black Cyan Page 409

REFERENCES

Leclant, J. (1979): Ägypten. Band I: Das Alte und das Mittlere Reich. Verlag C.H. Beck, München. Lehmann, U. (1949): Der Ur im Diluvium Deutschlands und seine Verbreitung. Neues Jahrbuch für Mineralogie, Geologie und Palaeontologie, Abhandlungen Abt. B, 90; 163 – 266. . Leibundgut, H. & Schlegel, J. (1985): Waldbauliche Untersuchungen in Bergföhrenbeständen des Schweizerischen Nationalparkes. Schweizerische Zeitschrift für Forstwesen 136(11): 945 - 955. Leithner, O. von (1927): Der Ur. Berichte der Internationalen Gesellschaft zur Erhaltung des Wisents, Band II, Heft 1 und 2: 1 - 139. Lengerken, H. von (1955): Ur, Hausrind und Mensch - Versuch eines Überblicks. Deutsche Akademie der Landwirtschaftswissenschaften zu Berlin. Wissenschaftliche Abhandlungen nr 14. Li, J.Q., Yang, G.F., Wu, X.H., Zhang, X.Q. & Qiu, L. (1991): A study of blood protein polymorphism of yellow cattle in South China. Journal of South China Agricultural University, Supplement: 5 – 10. Liberto, T.J. De & Urness, P.J. (1994): Comparative digestive physiology of American bison and Hereford cattle. Proceedings of 1st International Bison Conference, LaCrosse, WI. July 1993. Linhart, Y.B. & Whelan, R.J. (1980): Woodland regeneration in relation to grazing and fencing in Coed-Gorswen North Wales UK. Journal of Applied Ecology 17(3): 827-840. Linnaeus, C. (1758): Systema naturae: per regna tria naturae. I. Regnum animale. Editio decima. Holmiae. Linseele, V. (2004): Size and size change of the African aurochs during the Pleistocene and Holocene. Journal of African Archaeology 2(2): 1-21. Lister, A. M. (1984): Evolutionary and ecological origins of British deer. Proceedings of the Royal Society of Edinburgh 82 B:205-229. Litt, T. (2000): Waldland Mitteleuropa – die Megaherbivorentheorie aus paläobotanischer Sicht. In: Bayerische Landesanstalt für Wald und Forstwirtschaft (LWF). Grosstiere als Landschaftsgestalter – Wunsch oder Wirklichkeit? Berichte aus der Bayerischen Landesanstalt für Wald und Forstwirtschaft. Nummer 27: 49-64. Liu, H., Cui, H., Pott, R. & Speier, M. (1999): The surface pollen of the woodland-steppe ecotone in southeastern Inner Mongolia, China. Review of Palaeobotany and Palynology 105(3-4): 237-250. Loftus, R.T., MacHugh, D.E., Bradley, D.G., Sharp, P.M. & Cunningham, P. (1994): Evidence for two independent domestications for cattle. Proceedings of the National Academy of Sciences, USA 91(3): 2757 - 2761.

409

Black Cyan Page 410

410


Lorblanchet, M. & Welté, A.-C. (1994): Aurochs et bison dans l’art paléolithique. In: Aurochs, le retour - aurochs, vaches & autres bovins de la Préhistoire à nos jours: 27 - 65. Centre Jurassien du Patromoine, Lons-le-Saunier. Loth, P.E. (1999): The vegetation of Manyara - Scale-dependent states and transitions in the African Rift Valley. Proefschrift Wageningen. Lucanus, M.A. & Duff, J.D. (1928): The civil was (Pharsalia): Books 1-10. The Loeb Classical Library, 220. London, Heinemann. Lüpke, B. von (1998): Silvicultural methods of oak regeneration with special respects to shade tolerant mixed species. Forest Ecology and Management 106(1): 19-26. Lüpke, B. von, Hauskeller-Bullerjahn, K. (1999): Kahlschlagfreier Waldbau: Wird die Eiche an den Rand gedrängt? Forst und Holz 54: 563-568. £ukaszewicz, K. (1952): The ure-ox. Ochrona Przyrody 20: 1 - 33. Lydekker, R. (1898): Wild oxen, sheep, goats of all lands, living and extinct. Rowland Ward Ltd, London. Lydekker, R. (1912): The ox and its kindred. Methuen, London. Lynch, J.A., Hollis, J.L. & Hu, F.S. (2004): Climatic and landscape controls of the boreal forest fire regime: Holocene records from Alaska. Journal of Ecology 92(3): 477-489. Maciej z Miechowa (1521): Tractatus de duabus Sarmatiis Asiana et Europiana et de contentis in eis. In: S.A. Anninskij (1936). Traktat o dvuch Sarmatijach. Moskou-Leningrad. Mager, F. (1941): Wildbahn und Jagd Altpreußens, im Wandel der geschichtlichen Jahrhunderte. Verlag J. Neumann, Neudamm und Berlin. Mager, F. (1960): Der Wald in Altpreußen als Wirtschaftsraum. Band I und II. Böhlau Verlag, Köln, Graz. Manwell, C. & Baker, C.M.A. (1980): Chemical classification of cattle. 2. Phylogenetic tree and specific status of the Zebu. Animal Blood Groups and Biochemical Genetics 11: 151 - 162. Martin, P.S. (1984): Prehistoric overkill: the global model. In: P.S. Martin & R.G. Klein (eds.). Quaternary extinctions - a prehistoric revolution: 354 - 403. The University of Arizona Press. Tucson, Arizona. Martin, P.S. & Szuter, C.R. (1999): War zones and game sinks in Lewis and Clark’s west. Conservation Biology 13 (1): 26-45. Martin, T. (1990): Jungpleistozäne und holozäne Skelettfunde von Bos primigenius und Bison priscus aus Deutschland und ihre Bedeutung für die Zuortnung isolierter Langknochen. Eiszeitalter und Gegenwart 40: 1 - 19. Mason, I.L. (1984): Evolution of domesticated animals. Longman, London.

Black Cyan Page 411

REFERENCES

Mason, I.L. (1996): A world dictionary of livestock breeds, types and varieties. Wallingford: CAB International. Mathiasson, S. (1988): Uroxen (Bos primigenius) i västsverige - ett nytt subfossilfynd från Göteborg. Arstryck Göteborgs Naturhistorisk Museum: 16 - 34. Matolcsi, J. (1970): Historische Erforschung der Körpergröße des Rindes auf Grund von ungarischen Knochenmaterial. Zeitschrift für Tierzüchtung und Züchtungsbiologie 87: 89 - 137. Matuszkiewicz, W. (1966): Potencjalna roœlinnoœæ naturalna kotliny Warszawskiej. Materia³y Zak³adu Fitosocjologii Stosowanej U.W. Warszawa - Bia³owie¿a nr. 15. Mayrhofer, M. (1986): Etymologisches Wörterbuch des Altindoarischen. Heidelberg, Winter. Mazak, V. (1979): Der Tiger.. Die Neue Brehm-Bücherei 356. A. Ziemsen Verlag. McBurney, C.B.M. (1960): The stone age of northern Africa. Penguin Books, Harmondsworth. McHugh, T. (1972): The time of the buffalo. Knopf, New York. Mech, L.D. (1966): The wolves of Isle Royale. U.S. Government Printing Office, Washington. Meister, G. (1969): Ziele und Ergebnisse forstlicher Planung im oberbayerischen Hochgebirge. Forstwissenschaftliches Centralblatt 88: 97 - 130. Mela, P. (± 40): De chorographia libri tres; Kreuzfahrt durch die alte Welt. 2-Talige uitgave door K. Brodersen. Wissenschaftliche Buchgesellschaft, Darmstadt, 1994. Mellaart, J. (1975): The neolithic of the Near East. Thames and Hudson, London. Mertens, A. (1906): Der Ur, Bos primigenius Bojanus, mit besonderer Berücksichtigung der im städtischen ‘Museum für Natur- und Heimatkunde’ zu Magdeburg befindlichen Reste. Abhandlungen und Berichte. Museum für Natur- und Heimatkunde zu Magdeburg 1(2): 45 - 119. Merz, G. (1986): Counting elephants (Loxodontha africana cyclotis) in tropical rain forests with particular reference to the Tai National Park, Ivory Coast. African Journal of Ecology 24: 61 - 68. Messier, F. (1991): The significance of limiting and regulating factors on the demography of moose and white-tailed deer. Journal of Animal Ecology 60: 377 - 393. Ministry of Agriculure, Jamaica (1972): The development of the Jamaica Hope breed of dairy cattle. Animal Husbandry Bulletin nr. 2. Minkema, D. & Rooij, J. de (1982): Kleuren en aftekeningen bij runderen. Zeldzaam huisdier 7(2): 52 - 57. Mitchell, F.J.G. (2005): How open were European primeval forests? Hypothesis testing using palaeoecological data. Journal of Ecology 93(1): 168-177. Mitchell, F.J.G. & Cole, E. (1998): Reconstruction of long-term successional dynamics of temperate woodland in Bia³owie¿a Forest, Poland. Journal of Ecology 86: 1042-1059.

411

Black Cyan Page 412

412


Moerman, H.J. (1956): Nederlandse plaatsnamen - een overzicht. E. J. Brill, Leiden. Moll, L. & Gayot, E. (1860): La connaisance générale du boeuf.. Paris. Mortensen, H. und G. (1938): Die Besiedlung des Nordöstlichen Ostpreußens bis zum Beginn des 17. Jahrhunderts. Teil II: Die Wildnis im östlichen Preußen, ihr Zustand um 1400 und ihre frühere Besiedlung. Verlag von S. Hirzel, Leipzig. Mortensen, H. (1963): Zum Landschaftsbild des mittelalterlichen Urwaldes im nordöstlichen Mitteleuropa. Zeitschrift für Ostforschung 12(1): 97-99. Mott, R., J. (1978): Populus in late-Pleistocene pollen spectra. Canadian Journal of Botany 56: 1021 - 1031. Mountford, E.P., Peterken, G.F., Edwards, P.J. & Manners, G.J. (1999): Long-term change in growth, mortality and regeneration of trees in Denny Wood, an old-growth wood pasture in the New Forest (UK). Perspectives in Plant Ecology, Evolution and Systematics 2(2): 223-272. Munther, G.L. (1982): Beaver management in grazed riparian ecosystems. In: J.M. Peek & P.D. Dalke (eds.). Proceedings of the wildlife-livestock relationships symposium: 234 241. Forest, Wildlife & Range Experiment Station, University of Idaho. Muzzolini, A. (1983): L’art rupestre du Sahara Central. Classification et chronologie. Le boeuf dans le préhistoire Africaine. 2 Parts.. Toulouse. Naiman, R.J., Johnston, C.A. & Kelley, J.C. (1988): Alteration of North American streams by beavers. Bioscience 38(11): 753 - 762. Nederlof, L.-J. (1995): De vondst van de maand: Onverwachts een zeldzaamheid...over de vondst van een oeroshoorn. Cranium 12(2): 64. Nehring, A. (1888): Über den Schädel eines Franqueiro-Ochsen aus Brasilien. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 19 Juni: 91 – 99. Nehring, A. (1898a): Das Augsburger Urstierbild. Globus 74: 79 – 81 Nehring, A. (1898b): Über Ur und Wisent nach dem ‘Treßlerbuche’ des deutschen Ordens 1399 bis 1409. Globus 74(3): 44 - 47. Nehring, A. (1900): Das Horn eines Bos primigenius aus einem Torfmoore Hinterpommerns. Sitzungsbericht der Gesellschaft Naturforschender Freunde zu Berlin 16 Januar: 1 - 10. Nehring, K. (1965): Laub- und Reisigfutterstoffe. In: M. Becker, K. Nehring (Hrsg.). Handbuch der Futtermittel. Zweiter Band: 1 - 27.. Verlag Paul Parey. Nelson, S. M. (1990): The neolithic of Northeastern China and Korea. Antiquity 64: 234 - 248. Newberry, P.E. (1894): Beni Hassan, Part II. Archaeological Survey of Egypt. The Egypt Exploration Fund, London. Nickel, R., Schummer, A. & Seiferle, E. (1975): Lehrbuch der Anatomie der Haustiere. Band IV: Nervensystem, Sinnesorgane, endokrine Drüsen. Paul Parey.

Black Cyan Page 413

REFERENCES

Ninov, L.K. (1999): Vergleichende Untersuchungen zur Jagd und zum Jagdwild während des Neolithikums und Äneolithikums in Bulgarien. In: N. Benecke (ed.). The Holocene history of the European vertebrate fauna - modern aspects of research: 323 - 338. Nobis, G. (1954): Zur Kenntnis der ur- und frühgeschichtkichen Rinder Nord- und Mitteldeutschlands. Zeitschrift für Tierzüchtung und Züchtungsbiologie 63: 153 - 194. Nobis, G. (1993): Zur antiken Wild- und Haustierfauna Kretas - nach Studien an Tierresten aus den archäologischen Grabungen Poros bei Iraklion und Eleftherna bei Arkadhi. Tier und Museum 3(4): 109 - 120. Olson, T.A. & Wilham, R.L. (1982): Inheritance of coat coloration and spotting patterns of cattle: a review. Agriculture and Home Economics Experiment Station, Iowa State University of Science and Technology, Research Bulletin 595, Ames. Osborne, P.J. (1978): Insect evidence for the effect of man on the lowland landscape. In: S. Limbrey & J.G. Evans. The effect of man on the landscape: the lowland zone. Council for British Archaeology. Research Report, nr 21: 32 - 34. Oswald, W.W., Anderson, P.M., Brubaker, L.B., Hu, F.S. & Engstrom, D.R. (2003): Representation of tundra vegetation by pollen in lake sediments of Northern Alaska. Journal of Biogeography 30: 521-535. Paaver, K. (1965): The origin of the mammal fauna and the variability of mammals in the East Baltic area during the Holocene (in Russian). Estonian Academy of Sciences, Institute of Zoology and Botany. Tartu. Pa³czyñski, A. (1984): Natural differentiation of plant communities in relation to hydrological conditions of the Biebrza valley. Polish Ecological Studies 10: 347 - 385. Palmer, S.C.F., Truscott, A.-M., Mitchell, R.J. & Welch, D. (2001): Grazing in Atlantic oakwoods: Final report. Centre for Ecology and Hydrology, Banchory Research Station. Project C00342. Paravicini, W. (1989-1995): Die Preussenreisen des Europäischen Adels. 2 Teile. Jan Thorbecke Verlag Sigmaringen. Parren, M.P.E. (1991): Forest elephant (Loxodonta africana cyclotis Matschie) messenger-boy or bulldozer?: the possible impact on the vegetation, with special reference to 41 tree species of Ghana. Student Report. Department of Forestry, AV nr. 90/51. Partridge, L. & French, V. (1996): Thermal evolution of ectotherm body size: why get big in the cold? In: I.A. Johnston & A.F. Bennett (eds.). Animals and temperature - phenotypic and evolutionary adaptation: 265 - 292. Cambridge University Press. Payne, W.J.A. (1991): Domestication: A forward step in civilization. In: C. G. Hickham. Cattle genetic resources. World Animal Sciences 87: 51 - 72. Elsevier, Amsterdam.

413

Black Cyan Page 414

414


Peden, D.G., Van Dyne, G.M., Rice, R.W. & Hansen, R.M. (1974): The trophic ecology of Bison bison on short grass plains. Journal of Applied Ecology 11(2): 489-497. Pels Rijcken, P.H. (1965): Schilschade van edelherten aan de groveden.. Nederlands Bosbouw Tijdschrift 37(2): 30 - 65. Peters, H.F. & Slen, S.B. (1964): Hair coat characteristics of bison, domestic bison hybrids, cattalo and certain domestic breeds of cattle. Canadian Journal of Animal Science 44: 48-57. Petrie, W.M.F. (1920): Prehistoric Egypt. British School of Archaeology in Egypt, London. Pfizenmayer, E.W. (1929): Biologische und morphologische Notizen über den Kaukasuswisent. Abhandlungen der mathematisch-naturwissenschaftliche Abteilung der Bayerischen Akademie der Wissenschaften. Supplement-Band 11 - 14 Abhandlung: 497 – 504. Phoebus, G., Schlag, W. & Thomas, M. (1994): Das Jagdbuch des Mittelalters. Ms. fr. 616 der Bibliothèque Nationale in Paris. Glanzlichter der Buchkunst, Bd. 4. Graz/Austria. Pinchak, W.E., Smith, M.A., Hart, R.H. & Waggoner, J.W. (1991): Beef cattle distribution patterns on foothill range. Journal of Range Management 44(3): 267 - 275. Plinius, C. Secundus (± 70): Naturalis historiae, Libri XXXVII. (in various translations) Pöschl, J. & Hasenhüttl, E. (1995): Jagdmusik - Kontinuität in der europäischen Geschichte. Zeitschrift für Jagdwissenschaft 41(1): 43 - 51. Pomel, A. (1894): Paléontologie-Monographies. Boeufs-Taureaux. Carte Géologique de l’Algerie: 1 - 106. Ponel, P. (1995): Rissian, Eemian and Würmian Coleoptera assemblages from La Grande Pile (Vosges, France). Palaeogeography, Palaeoclimatology, Palaeoecology 114:1-41. Ponel, P., Matterne, V., Coulthard, N. & Yvenec, J.H. (2000): La Tène and Gallo-Roman natural environments and human impact of the Touffreville rural settlement, reconstructed from Coleoptera and plant macroremains (Calvados, France). Journal of Archaeological Science 27(11): 1055-1072. Ponel, P., Parchoux, F., Andrieu-Ponel, V., Juhasz, I. & Beaulieu, J.L. de (2001): A Lateglacial-Holocene fossil insect succession from Valleé des Merveilles, French Alps, and its palaeoecological implications. Arctic, Antarctic and Alpine Research 33(4): 481-484. Pons, L.J. (1992): Holocene peat formation in the lower parts of the Netherlands. In: J.T.A. Verhoeven (ed.). Fens and bogs in the Netherlands: Vegetation, history, nutrient dynamics and conservation: 7 – 79. Kluwer, Dordrecht. Poortinga, G. (1981): Stichting Tarpan. 2e druk. Groenwinkel Groningen. Portis, A. (1907): Di alcuni avanzi fossili di Grandi Ruminantia principalmente della Provincia di Roma. Palaeontographia Italica, 13.

Black Cyan Page 415

REFERENCES

Pott, R. & Hüppe, J. (1995): Die Bedeutung der Extensivbeweidung für die Pflege und Erhaltung nordwestdeutscher Hudelandschaften am Beispiel der NSG Borkener Paradies im Emstal bei Meppen (Emsland). Carolinea 53: 99-111. Poulain, T. (1980): La faune du Camp de la Vergentière. In: Association d’études préhistoriques et protohistoriques de Champagne-Ardenne, Châlons-sur-Marne. Actes du colloque interrégional sur le Néolithique de l’Est de la France, Châlons-sur-Marne, 24-25 mars 1979: 152-165. Pringle, W.L. (1987): Forage potential for livestock production. In: H.W. Reynolds & A.W.L. Hawley (eds.). Bison ecology in relation to agricultural development in the Slave River Lowlands, NWT: 53 - 62. Prins, H.H.T. (1987): The buffalo of Manyara. Thesis, Groningen. Prins, H.H.T. (1998): Sterfte van runderen in natuurgebieden. De Levende Natuur 99(3): 108 - 114. Prschewalski, N. v. (1877): Reisen in der Mongolei, im Gebiet der Tanguten und den Wüsten Nordtibets in den Jahren 1870 bis 1873. Jena. Prummel, W. & Niekus, M.J.L.Th. (2005): De laatmesolithische vindplaats Jardinga (Fr.): de opgravingen in 2002 en 2003. Paleo-Aktueel 14/15: 40-45. Prummel, W., Niekus, M.J.L.Th., Gijn, A.L. van & Cappers, R.T.J. (2002): A late Mesolithic kill site of aurochs at Jardinga, Netherlands. Antiquity 76 (292): 413-424. Pucek, Z (1986): Bison bonasus (Linnaeus, 1758) - Wisent. In: Niethammer, J. & Krapp, F. Handbuch der Säugetiere Europas: 278 - 315. Pucek, Z. (1991): History of the European Bison and problems of its protection and management. In: B. Bobek, K. Perzanowski & W.L. Regelin (eds.). Global trends in wildlife management. 18th IUGB Congress, Jagiellonian University Kraków, Poland; August 1987. Transactions Vol. I. Œwi¹t Press, Kraków-Warszawa. Pusch, E.G. (1838): Polens Paläontologie. Stuttgart. Pusch, E.G. (1840): Neue Beiträge zur Erläuterung und endlichen Erledigung der Streitfrage über Tur und Zubr (Urus und Bison). Archiv für Naturgeschichte. 6e Jahrgang, Band 1: 47 - 137. Putman, R.J. (1987): Invloed van grote grazers in het New Forest. In: S. de Bie, W. Joenje & S.E. van Wieren (red.). Begrazing in de natuur: 187 - 199. Pudoc Wageningen. Pyle, C.M. (1994): Some late sixteenth-century depictions of the aurochs (Bos primigenius Bojanus, extinct 1627): new evidence from Vatican MS Urb. lat. 276. Archives of Natural History 21(3): 275 - 288. Quittet, E. & Denis, B. (1979): Races bovines françaises. La Maison Rustique, Paris. Ralls, K. & Harvey, P. (1985): Geographic variation in size and sexual dimorphism of North American weasels. Biological Journal of the Linnean Society 25: 119 - 167.

415

Black Cyan Page 416

416


Rackham, O. (1980): Ancient woodland – its history, vegetation and uses in England. Edward Arnold. Rasmussen, P. (1989): Leaf-foddering of livestock in the Neolithic: archaeobotanical evidence from Weier, Switzerland. Journal of Danish Archaeology 8: 51 - 71. Rasmussen, P. (1993): Analysis of goat/sheep faeces from Egolzwil 3, Switzerland: evidence for branch and twig foddering of livestock in the Neolithic. Journal of Archaeological Science 20: 479 - 502. Raup, H.M. (1935): Botanical investigations in Wood Buffalo Park. Biological Series nr 20, Bulletin nr 74. National Museum of Canada, Ottawa. Rees, W. (1974): Preliminary studies into bush utilization by cattle in Zambia. Journal of Applied Ecology 11: 207 - 214. Regnell, M., Gaillard, M.-J., Bartholin, T.S. & Karsten, P. (1995): Reconstruction of environment and history of plant use during the late Mesolithic (Ertebölle culture) at the inland settlement of Bokeberg III, Southern Sweden. Vegetation History and Archaeobotany 4(2): 67-91. Reijnders, P.J.H. (1972): Schilschade door edelherten aan de douglas. RIN Arnhem. Remmert, H. (1985): Was geschieht im Klimax-Stadium? Ökologisches Gleichgewicht durch Mosaik aus desynchronen Zyklen. Naturwissenschaften 72: 505 - 512. Requate, H. (1957): Zur Naturgeschichte des Ures (Bos primigenius Boj. 1827) nach Schädelund Skelettfunden in Schleswig-Holstein. Zeitschrift für Tierzüchtung und Züchtungsbiologie 70: 297 - 338. Reynolds, H.G. (1969): Aspen grove use by deer, elk and cattle in southwestern coniferous forests. USDA Forest Service. Rocky Mountain Forest and Range Experiment Station. Research Note RM - 138. Reynolds, H.W., Hansen, R.M. & Peden, D.G. (1978): Diets of the Slave River Lowland Bison Herd, Northwest Territories, Canada. The Journal of Wildlife Management 42(3): 581 - 590. Rhoad, A.O. (1949): The Santa Gertrudis breed - the genesis and the genetics of a new breed of beef cattle. Journal of Heredity 40: 114 - 126. Richter, J. (1982): Adult and juvenile aurochs, Bos primigenius, from the Maglemosian Site of Ulkestrup Lyng Øst, Denmark. Journal of Archaeological Science 9(3): 247 - 259. Ripple, W.J. & Beschta, R.L. (2005a): Willow thickets protect young aspen from elk browsing after wolf reintroduction. Western North American Naturalist 65(1): 118-122. Ripple, W.J. & Beschta, R.L. (2005b): Refugia from browsing as reference sites for restoration planning. Western North American Naturalist 65(2): 269-273.

Black Cyan Page 417

REFERENCES

Ripple, W.J., Larsen, E.J., Renkin, R.A. & Smith, D.W. (2001): Trophic cascades among wolves, elk and aspen on Yellowstone National Park’s northern range. Biological Conservation 102(3): 227-234. Ritchie, J.C. & Haynes, C.V. (1987): Holocene vegetation zonation in the eastern Sahara. Nature, vol. 330: 645 - 647. Roath, L.R. & Krueger, W.C. (1982a): Cattle grazing influence on a mountain riparian zone. Journal of Range Management 35(1): 100 - 103. Roath, L.R. & Krueger, W.C. (1982b): Cattle grazing and behaviour on a forested range. Journal of Range Management 35(3): 332 - 338. Rodrigues, L.J. (1992): Anglo-Saxon verse runes. Llanerch Publishers, Felinfach. Rodríguez Montesinos, A. (1994): Pelajes y encornaduras del toro de lidia. Consejo General de Colegios Veterinarios de España, Madrid. Roe, F.G. (1972): The North American buffalo; a critical study of the species in its wild state. David & Charles: Newton Abbot. Roes, A. (1940): A drinking horn of the Viking Period. Mededelingen Koninklijke Nederlandse Akademie van Wetenschappen, Afd. Letteren, Nieuwe Reeks, deel 3, nr 3. Rokosz, M. (1995): History of the aurochs (Bos taurus primigenius) in Poland. Animal Genetic Resources Information 16: 5 - 14. Ropeid, A. (1960): Skav; ein studie i eldre tids fôr-problem. Universitetsforlaget, Oslo-Bergen. Rouse, J.E. (1970): World Cattle II. Cattle of Africa and Asia. University of Oklahoma Press, Norman. Ruedemann, R. & Schoonmaker, W. J. (1938): Beaver-dams as geologic agents. Science 88: 523 - 525. Ruprecht, A.L. (1976): Mammals of the Bia³ystok region Quaternary. Rocznik Bia³ostocki 13: 261 - 282. Rutten, L.M.R. (1909): Die diluvialen Säugetiere der Niederlande. Proefschrift Utrecht. Ryder, M.L. (1976): Hair. Studies in Biology nr. 41. Arnold. Ryder, M.L. (1980): Hair remains throw light on early British prehistoric cattle. Journal of Archaeological Science 7: 389 – 392. Ryder, M.L. (1984): The first hair remains from an aurochs (Bos primigenius) and some medieval domestic cattle hair. Journal of Archaeological Science 11: 99 - 101. Sallares, J,R. (1991): The ecology of the ancient Greek World. Duckworth. Schaefer, A.L., Young, B.A. & Chimwano, A.M. (1978): Ration digestion and retention times of digesta in domestic cattle (Bos taurus), American bison (Bison bison), and Tibetan yak (Bos grunniens). Canadian Journal of Zoology 56: 2355-2358.

417

Black Cyan Page 418

418


Schäfer, E. (1937): Der wilde Yak (Bos (Poëphagus) grunniens mutus Prze.). Der Zoologische Garten, Leipzig 9: 26 - 34. Schäfer, H. (1896): Neue Altertümer der ‘new race’ aus Negadeh. Zeitschrift für Ägyptische Sprache und Alterthumskunde 34: 158 - 161. Schaller, G.B. (1967): The deer and the tiger - a study of wildlife in India. The University of Chicago Press. Schaller, G.B. (1998): Wildlife of the Tibetan steppe. University of Chicago Press. Schild, R. & Wendorf, F. (1986): The late Pleistocene Nile in Wadi Kubbaniya. In: F. Wendorf, R. Schild & A.E. Close (eds.). The Prehistory of Wadi Kubbaniya, Volume 2: 15 - 100. Dallas. Schloeth, R. (1958): Cycle annuel et comportement social du taureau de Camargue. Mammalia 22 : 121 - 139. Schloeth, R. (1961): Das Sozialleben des Camargue-Rindes. Zeitschrift für Tierpsychologie 18: 574 - 627. Schlüter, O. (1921): Wald, Sumpf und Siedelungsland in Altpreussen vor der Ordenszeit. Niemeyer. Schreiber, A. & Göltenboth, R. (1990): The haemoglobins of wild cattle (Bovini Simpson 1945). Zeitschrift für Säugetierkunde 55: 276 - 283. Schütz, M., Wildi, O., Krüsi, B.O., Achermann, G. & Grämiger, H. (1998): Sukzession über 585 Jahre. Modell für die subalpinen Weiden im Schweizerischen Nationalpark.. Informationsblatt des Forschungsbereiches Landschaftsökologie 40. Schwarz, A.G. & Wein, R.W. (1997): Threatened dry grasslands in the continental boreal forests of Wood Buffalo National Park. Canadian Journal of Botany 75(8): 1363 - 1370. Schwarz, A.G. & Wein, R.W. (1999): Dry grasslands in Wood Buffalo National Park: reply. Canadian Journal of Botany 77: 918-921. Schwend, C. (1950): Wildstand und Lebensgemeinschaft Wald. Forstwissenschaftliches Centralblatt 69: 348 - 352. Scott, A.C., Moore, J. & Brayshay, B. (2000): Fire and the palaeoenvironment. Palaeogeography, Palaeoclimatology, Palaeoecology 164(1-4). Sinclair, A.R.E. (1977): The African Buffalo - a study of resource limitation of populations. University of Chicago Press, Chicago/London. Sisson, S. & Grossman, J.D. (1953): The anatomy of the domestic animals. W.B. Saunders Company, Philadelphia and London. Skorupka, S. (1968): S³ownik frazeologiczny jêzyka polskiego. Tom drugi. Warszawa. Skovlin, J.M. (1984): Impacts of grazing on wetlands and riparian habitat: A review of our knowledge. In: National Research Council/National Academy of Sciences. Developing strategies for rangeland management. Westview Press, Boulders and London: 1001 - 1103.

Black Cyan Page 419

REFERENCES

Smith, C.H. (1827): Order Ruminantia - The Taurine Group. In: E. Griffith et.al.. The Animal Kingdom. Vol. 4: 411 - 428. Smoliak, S. & Peters, H.F. (1955): Climatic effects on foraging performance of beef cows on winter range. Canadian Journal of Agricultural Science 35(3): 213 - 216. Söffner, W. (1982): Über die Grossäugerfauna Mitteleuropas im Postglazial; ein Beitrag zur Kenntnis der Beziehungen zwischen Wild und Vegetation. Zulassungsarbeit Institut für Botanik der Universität Hohenheim. Solounias, N., Moelleken, S.M.C. & Plavcan, J.M. (1995): Predicting the diet of extinct bovids using massateric morphology. Journal of Vertebrate Paleontology 15(4): 795 - 805. Sondaar, P.Y. (1977): Insularity and its effect on mammal evolution. In: M.K. Hecht, P.C. Goody & B.M. Hecht (eds.). Major patterns in vertebrate evolution: 671-707. NATO advanced Study Institutes Series. Ser, A. Life Sciences, Vol. 14. Plenum Press, New York. Sondaar, P.Y. (1985): Verdwenen leven van de Middellandse-Zeeëilanden. Cranium 2(2): 44-55. Splunder, I. van, Voesenek, L.A.C.J., Coops, H., Vries, X.J.A. de & Blom, C.W.P.M. (1996): Morphological responses of seedlings of four species of Salicaceae to drought.. Canadian Journal of Botany 74: 1988 - 1995. Stanley, S.M. (1989): Uitsterven - Rampen markeren elk nieuw begin. Wetenschappelijke Bibliotheek 17. Natuur & Techniek. Stechow, E (1929): Über die einstige Hege des Wisents im Urwalde von Bialowies. Abhandlungen der mathematisch-naturwissenschaftliche Abteilung der Bayerischen Akademie der Wissenschaften. Supplement-Band. Abteilung 11 – 14: 505 - 507. Steehouwer, K.J. (1987): De schofthoogteparadox - een nieuwe theorie ter verklaring van de schofthoogte-evolutie van het (pre-)historische rund. IPP - eindscriptie. Steenbeek, R (1990): On the balance between wet and dry - Vegetation horizon development and prehistoric occupation; a palaeoecological-micromorphological study in the Dutch river area. Thesis, Amsterdam. Steffen, H. (1931): Vegetationskunde von Ostpreußen. Jena, Verlag von Gustav Fischer. Steinmetz, R. (2004): Gaur (Bos gaurus) and banteng (Bos javanicus) in the lowland forest mosaic of Xe Pian Protected Area, Lao PDR: abundance, habitat use, and conservation. Mammalia 68(2-3): 141-157. Stella, E. (1518): De Borussiae antiquitatibus. Libri duo. Basiliae. Steppan, K. (1999): The significance of aurochs in the food economy of the Jungneolithikum (Upper Neolithic) in South-west Germany. In: G.-C. Weniger (Hrsg.). Archäologie und Biologie des Auerochsen: 161 - 171. Neanderthal Museum.

419

Black Cyan Page 420

420


Stolzenbach, L. (1984): Een keratinehoorn van de oeros (Bos primigenius Bojanus 1827) uit de Noordzee. Cranium 1(2): 63 - 65. Stone, P. (1961): Some famous drinking-horns in Britain. Apollo, a journal of the arts. Part I: 102 - 104. Part II: 143 - 145. Stone, R. (1998): A bold plan to re-create a long-lost Siberian ecosystem. Science, 2 October, 282: 31-34. Street, M. (1999): Remains of aurochs (Bos primigenius) from the Early Mesolithic site BedburgKönigshoven (Rhineland, Germany). In: G.-C. Weniger (Hrsg.). Archäologie und Biologie des Auerochsen: 173 - 194. Neanderthal Museum. Svenning, J.-C. (2002): A review of natural vegetation openness in north-western Europe. Biological Conservation 104(2): 133-148. Œwiêcicki, A. (1634): Descriptio topographica Ducatus Masoviae. Varsaviae. Szalay, A.B. (1915): Der Wisent in Ortsnamen. Zoologische Annalen 7(1): 1 - 80. Szalay, A.B. (1917): Wisente im Zwinger. Zoologischer Beobachter LVIII. Szalay, A.B. (1918a): Wisente im Zwinger. Zoologischer Beobachter LIX. Szalay, A.B. (1918b): War der Ur wild? Archiv für Geschichte der Naturwissenschaft und Technik VIII: 106 – 123. Szalay, A.B. (1930): Die Farbe des Ures. Der Zoologische Garten 3(9): 255 - 263. Szalay, A.B. (1938): Hundert irrige Wisentbelege. Verlag J. Neumann, Neudamm und Berlin. Szper, F. (1913): Nederlandsche nederzettingen in West-Pruisen gedurende den Poolschen tijd. Thesis, Universiteit van Amsterdam. Tacitus, P.C. (98): Germania, sive de origine, situ, moribus ac populis Germaniae. (in various translations). Tacitus, P.C. (117): Annales ab excessu divi Augusti. (in various translations) Tagányi, K. (1921): Alte Grenzschutz-Vorrichtungen und Grenz-Ödland: gyepü und gyepüelve. Ungarische Jahrbücher 1: 105 - 121. Tamboer-van den Heuvel, G. & Janssen, C.R. (1976): Recent pollen assemblages from the crest region of the Vosges mountains (France). Review of Palaeobotany and Palynology 21(3): 219 - 240. Tankersley, K.B. (1986): Bison exploitation by late Fort Ancient peoples in the Central Ohio river valley. North American Archaeologist 7(4): 289-303. Tetzner, F. (1902): Die Slawen in Deutschland. Braunschweig. Teunissen, D. (1990): Palynologisch onderzoek in het oostelijk rivierengebied; een overzicht. Mededelingen van de afdeling Biogeologie van de Discipline Biologie van de Katholieke Universiteit van Nijmegen. Mededeling nr. 16.

Black Cyan Page 421

REFERENCES

Thenius, E. (1980): Grundzüge der Faunen- und Verbreitungsgeschichte der Säugetiere. Eine historische Tiergeographie. V.E.B. Gustav Fischer Verlag, Jena. Thesaurus Linguae Latinae (1900 - …): Editus auctoritate et consilio academiarum quinque Germanicarum Berolinensis Gottingensis Lipsiensis Monacensis Vindobonensis. Lipsiae, Teubner. Vol. 1 – 10/2; A – prodeo. Thill, R.E. (1983): Deer and cattle forage selection on Louisiana pine-hardwood sites. USDA Forest Service. Southern Forest Experiment Station. Research Paper SO-196. Management 53(3): 540 - 548. Tietz, O. (2004, in press): The first sure evidence of an aurochs discovery (Bos primigenius Bojanus 1827) from the Weichselian Lateglacial in Germany. Zeitschrift für Geologische Wissenschaften, Berlin 32. Timoney, K., Peterson, G., Fargey, P., Peterson, M., McCanny, S. & Wein, R. (1997): Spring ice-jam flooding of the Peace-Athabasca Delta: evidence of a climatic oscillation. Climatic Change 35: 463-483. Timoney, K.P. (1999): Threatened dry grasslands in the continental boreal forests of Wood Buffalo National Park: commentary. Canadian Journal of Botany 77: 913-917. Timoney, K.P. & Peterson, G. (1996): Failure of natural regeneration after clearcut logging in Wood Buffalo National Park, Canada. Forest Ecology and Management 87(1 - 3): 89 - 105. Timpe, D. (1989): Wegeverhältnisse und römische Okkupation Germaniens. In: H. Jankuhn, W. Kimmig & E. Ebel (Hrsg.). Untersuchungen zu Handel und Verkehr der vor- und frühgeschichtlichen Zeit in Mittel- und Nordeuropa. Teil V: Der Verkehr, Verkehrswege, Verkehrsmittel, Organisation: 83 - 107. Abhandlungen der Akademie der Wissenschaften in Göttingen. Philologisch-Historische Klasse, Dritte Folge, nr. 180. Tinley, K.L. (1974): Marromeu wrecked by the big dams. African Wildlife 29 (2): 22 – 25. Tratz, E.P. (1958): Noch ein Urgehörn. Säugetierkundliche Mitteilungen 6(1): 27. Troy, C.S., MacHugh, D.E., Bailey, J.F., Magee, D.A., Loftus, R.T., Cunningham, P., Chamberlain, A.T., Sykes, B.C. & Bradley, D.G. (2001): Genetic evidence for Near-Eastern origins of European cattle. Nature 410: 1088 – 1091. Tubbs, C.R., (2001): The New Forest – history, ecology and conservation. New Forest Ninth Centenary Trust, Lyndhurst. Tustin, J.R. (1975): Grazing livestock among young Radiata Pine. What’s New in Forest Research nr. 22. Forest Research Institute, Rotura. Twardowski, T. & Kasperczyk, B. (1992): Reintrodukcja bobra Castor fiber w województwie œkierniewickim. Chroñmy Przyrodê Ojczyst¹ 48(2): 105 - 109.

421

Black Cyan Page 422

422


Udo, H.M.J. (1978): Hair coat characteristics in Friesian heifers in the Netherlands and Kenya. Thesis. Veenman & Zonen, Wageningen. Uerpmann, H.P. (1987): The ancient distribution of ungulate mammals in the Middle East. Beihefte zum Tübinger Atlas des Vordern Orients. Reihe A (Naturwissenschaften) nr. 27. Uerpmann, H.P. (1990): Die Domestikation des Pferdes im Chalkolithikum West-und Mitteleuropas. Madrider Mitteilungen 31: 109-153. Unger, M.F. & Ramey, R.F. (1966): Unger’s Bible Dictionary. Moody Press, Chicago. Van Soest, P.J. (1982): Nutritional ecology of the ruminant. O & B Books, Corvallis, Oregon, USA. Vance, R. E. (1986): Pollen stratigraphy of Eaglenest Lake, northeastern Alberta. Canadian Journal of Earth Sciences 23: 11 - 20. Veen, H.E. van de (1975): De Veluwe natuurlijk?: Het Veluwemassief:: ‘behouden’, ‘behouten’ of ‘woekeren met natuurlijke ontwikkelingsmogelijkheden’. Arnhem. Veen, H.E. van de (1985): Natuurontwikkelingsbeleid en bosbegrazing. Landschap, 2(1): 14 – 28. Vera, F.W.M. (1997): Metaforen voor de wildernis - eik, hazelaar, rund en paard. Proefschrift. Landbouw Universiteit Wageningen. Vera, F.W.M. (2000): Grazing ecology and forest history. CABI Publishing. Vereshchagin, N.K. & Baryshnikov, G.F. (1984): Quaternary mammalian extinctions in northern Eurasia. In: P.S. Martin & R.G. Klein (eds.). Quaternary extinctions - a prehistoric revolution: 483 - 516. The University of Arizona Press. Tucson, Arizona. Vergilius Maro, P. (± 30 B.C.): Georgica. Libri IV. (in various translations). Vijver, C.A.D.M. van de, Foley, C.A. & Olff, H. (1999): Changes in the woody component of an East African savanna during 25 years. Journal of Tropical Ecology 15(5): 545 - 564. Vink, I., Wolbers, H. (1997): Handboek melkveehouderij. Praktijkonderzoek rundvee, schapen en paarden, Lelystad. Visscher, P.M., Smith, D., Hall, S.J.G. & Williams, J.A. (2001): A viable herd of genetically uniform cattle. Nature 409, 6818: 303 - 304. Vlerk, I.M. van der (1942): Kwartaire Bovidae van Nederland. De schedels en hoornpitten, welke zich bevinden in het Rijksmuseum van Geologie te Leiden. Leidsche Geologische Mededelingen. Volume XIII: 1 - 28. Vörös, I. (1985): Early medieval aurochs (Bos primigenius Boj.) and his extinction in Hungary. Folia Archaeologica XXXVI: 193 - 219. Vörös, I. (1987): An aurochs (Bos primigenius Boj.) skeleton from the mesolithic peatbogs at Kecel-Rózsaberek. Folia Archaeologica XXXVIII: 65 - 86. Vries, J. de (1987): Nederlands etymologisch woordenboek. Bewerkt door F. de Tollenaere. E.J. Brill, Leiden.

Black Cyan Page 423

REFERENCES

Vries, L.S. de (1996): Karolingische faunaresten uit Wijk bij Duurstede-De Geer. Cranium 13(1): 58 - 64. Vulink, J.T. & Drost, H.J. (1991): Nutritional characteristics of cattle forage plants in the eutrophic nature reserve Oostvaardersplassen, Netherlands. Netherlands Journal of Agricultural Science 39: 263 - 272. Vuure, T. van (1998): Betizu, het wilde rund van Baskenland. Nieuwe Wildernis 4, zomer 1998: 12 - 15. Wallis de Vries, M.F. (1994): Foraging in a landscape mosaic: Diet selection and performance of free-ranging cattle in heathland and riverine grassland. Thesis Landbouwuniversiteit Wageningen. Watson, W. (1961): China before the Han dynasty. Thames and Hudson, London. Watts, W.A. & Winter, T.C. (1966): Plant macrofossils from Kirchner Marsh, Minnesota – a paleoecological study. Bulletin of the Geological Society of America 77: 1339-1360. Weaver, J.E. (1960): Floodplain vegetation of the Central Missouri Valley and contacts of woodland with prairie. Ecological Monographs 30: 37 - 64. Webb, S.D. (1977): A history of savanna vertebrates in the new world. Part I: North America. Annual review of Ecology and Systematics 8: 355-380. Werf, S. van der (1991): Natuurbeheer in Nederland. Deel 5: Bosgemeenschappen.. Pudoc Wageningen. Werner, A.R. (1976): Berlin cattle and Munich cattle. The Ark, III (2): 47 - 52; III (3): 76 - 82. Westhoff, V. (1949): Beken en beekdalen in Twente. In: A.F.H. Besemer, K. Hana, N. Tinbergen & J. Wilcke (red.). In het voetspoor van Thijsse: 36 - 64. H. Veenman & Zonen, Wageningen. Wharton, C.H. (1968): Man, fire and wild cattle in Southeast Asia. Proceedings Annual Tall Timbers Fire Ecology Conference 8: 107 – 167. Wiegels, R. & Woesler, W. (Hrsg) (1995): Arminius und die Varusschlacht; Geschichte Mythos - Literatur. Ferdinand Schöningh. Wieren, S.E. van (1986): Verslag studiereis Polen, 25/3 – 6/4 1985. Een verkennend onderzoek naar de schilaktiviteiten van de wisent. Unpublished. Wieren, S.E. van (1987): Het grazende dier. In: S. de Bie, W. Joenje & S.E. van Wieren (red.). Begrazing in de natuur: 34 - 53. Pudoc Wageningen. Wieren, S.E. van (1988): Runderen in het bos - begrazingsproef met Schotse Hooglandrunderen in het natuurgebied De Imbos - Eindrapport. Instituut voor Milieuvraagstukken, Amsterdam. Wieren, S.E. van (1992): Factors limiting food intake in ruminants and non ruminants in the temperate zone. In: F. Spitz, G. Janeau, G. Gonzalez, J. Janeau & C.S. Mann. Ongulés/ Ungulates 91. Proceedings of the International Symposium, sept. 2 - 6 1991, Toulouse.

423

Black Cyan Page 424

424


Wieren, S. E. van (1996): Digestive strategies in ruminants and nonruminants. Proefschrift Landbouw Universiteit, Wageningen. Wiesner, E. (1967): Ernährungsschäden der landwirtschaftlichen Nutztiere. VEB Gustav Fischer Verlag, Jena. Wijngaarden-Bakker, L.H. van (1997): Aurochs and Heck cattle. Anthropozoologica, nr. 2526: 193-199. Wilde, S.A., Youngberg, C.T. & Hovind, J.H. (1950): Changes in composition of ground water, soil fertility and forest growth produced by the construction and removal of beaver dams. Journal of Wildlife Management 14(2): 123 - 128. Wilson, D.E. & Reeder, D.M. (eds.) (1993): Mammal species of the world - a taxonomic and geographic reference. 2e Druk. Smithsonian Institution Press, Washington and London. Woodman, P., McCarthy, M. & Monaghan, N. (1997): The Irish Quaternary fauna project. Quaternary Science Reviews 16(2): 129-159. Woordenboek der Nederlandsche Taal (1893): Part X: O-ooilam. Edited by: M. de Vries, E. Verwijs & A. Kluyver. Martinus Nijhoff, A.W. Sijthoff. Wrzeœniowski, A. (1878): Studien zur Geschichte des polnischen Tur. Zeitschrift für Wissenschaftliche Zoologie 30, Supplement 45: 493 - 555. Zazula, G.D., Froese, D.G., Schweger, C.E., Mathewes, R.W., Beaudoin, A.B., Telka, A.M., Harington, C,.R. & Westgate, J.A. (2003): Palaeobotany: Ice-age steppe vegetation in east Beringia. Nature 423 (6940): 603-605. Zeiler, J.T. (1997): Hunting, fowling and stock-breeding at Neolithic sites in the Western and Central Netherlands. Thesis, Groningen. Zeuner, F.E. (1963): A history of domesticated animals. Hutchinson of London. Zhang Minghua (1984): A preliminary study of mammalian fauna in the neolithic sites of South China. Acta Theriologica Sinica 4(3): 177 - 185. Zheng Yi Li & Cao Cheng Xiu (1984): A new English-Chinese Dictionary. The Commercial Press and John Wiley and Sons Inc. Zimov, S.A., Chuprynin, V.I., Oreshko, A.P., Chapin III, F.S., Reynolds, J.F. & Chapin, M.C. (1995): Steppe tundra transition: a herbivore-driven biome shift at the end of the Pleistocene. The American Naturalist 146(5): 765-794. Zoller, H. & Haas, N. (1995): War Mitteleuropa ursprünglich eine halboffene Weidelandschaft oder von geschlossenen Wäldern bedeckt?. Schweizerische Zeitschrift für Forstwesen 146(5): 321 - 354. Zong Guanfu (1984): Fossils of Bos primigenius from Apa Tibetan Autonomous Prefecture, Sichuan. Vertebrata Palasiatica 22(3): 239 - 245.

Black Cyan Page 425

REGISTER

REGISTER Aberdeen Angus cattle 149, 174, 325 acorns 69, 75, 217, 222, 225, 231, 371, 377, 383 Africa 35-37, 40-43, 50, 53, 54, 78, 85, 118, 129, 132, 150, 158, 167, 219, 232, 233, 242, 248, 259, 263, 272, 286, 291, 292, 307, 311, 320, 325, 333, 343, 375, 383 African buffalo (Syncerus caffer caffer) 37, 42, 101, 122, 150, 219, 233, 242, 254, 266, 378 Alces alces 155, 276, 278. See also ‚elk’. alder (Alnus glutinosa) 72, 178, 183, 184, 186, 228, 236, 238, 239, 243, 250, 251, 254, 274, 285, 293 Algeria 42, 43, 216, 323, 324 Allgäuer cattle 325, 339, 340 Alnus glutinosa 254, 285. See also ‚alder’. Angler cattle 325, 337, 338, 340, 341 area size 280, 378 ash (Fraxinus excelsior) 228-230, 236, 239, 243, 250, 251, 292, 293 Asia 35, 37, 40-49, 52, 78, 101, 132, 150, 166, 167, 173, 183, 188, 215, 228, 305, 327 Association of Tall Sedges 252 Atlantic period 178, 184, 186, 187, 235, 284, 285, 378 Augsburg aurochs 117, 119, 124, 126, 142, 143, 153-155,161 aurochs-European bison confusion 83, 88 Austria 11, 24, 102, 340 backcrossing 368 banteng (Bos javanicus) 23, 37, 39, 40, 86, 99, 108, 116, 117, 119, 135, 149, 150, 151, 153, 154, 160, 165, 220, 244, 248, 266268, 287, 296, 334 basal length (aurochs skull) 104, 169, 215 beaver (Castor fiber) 60, 186-188, 254-257, 271, 278, 284, 287, 297, 298, 320, 373, 381 beaver meadow 255, 257

beech (Fagus sylvatica) 184, 186, 190, 208, 225, 235, 277, 296, 310, 378 Belgian Blue cattle 149, 325 Belgium 24, 58, 192, 194, 306, 344, 355 belt (aurochs skin) 92, 93, 142, 144, 145, 147, 154, 161, 164, 386 Berber lion (Panthera leo) 305, 364 Betizu cattle 16, 95, 323 Bhagwari cattle 149 birthing season (aurochs) 270 bison (American) (Bison bison) 37, 39, 101, 108, 134, 150-152, 220, 221, 244, 245, 248, 253, 258, 262, 266, 276, 277, 282, 283, 286, 297, 299, 300-304, 320, 370, 374 Bison bison 37, 220, 244, 276. See also ‘bison (American)’. Bison bonasus 37, 38, 116, 202, 221, 243. See also ‘European bison’. Bison priscus 244, 306. See also ‘steppe bison’. Black Pied cattle 129, 133, 141, 149, 159, 163, 325, 328, 329, 337, 338, 356, 357 body proportion/ -shape (aurochs) 99, 118, 165, 338, 351 body size (aurochs) 99, 103, 109, 118, 166, 171, 174, 323, 325, 336, 359-361 body weight (aurochs) 109, 119, 158, 165, 334, 335 Boreal period 176, 186, 187, 234, 235, 261, 285, 306 Borkener Paradies 287, 309 Bos acutifrons 35, 36 Bos arnee 37, 243. See also ‘water buffalo’. Bos brachyceros 86, 87 Bos gaurus 37, 220, 244. See also ‘gaur’. Bos javanicus 37, 86, 99, 220, 244. See also ‘banteng’. Bos primigenius 31, 32, 36, 37, 40, 48, 88

425

Black Cyan Page 426

426


Bos sauveli 37 Bos taurus 31, 32, 40 Bos trochoceros 88 Bovini 36-40, 45, 116, 150, 153, 213, 215, 218, 241, 243, 264, 363, 364 brain content (aurochs, cattle) 171 Braunvieh (Brown Swiss) cattle 110, 148, 149, 154, 325, 326 breeding-back (experiment) 15, 109, 110, 138, 323-344, 362, 366 brown bear (Ursus arctos) 233, 260 Brown Swiss cattle: See ‘Braunvieh’. browser 34, 183, 213, 214, 308 Bubalus depressicornis 37 Bubalus mephistopheles 37 Bubalus mindorensis 37 Bubalus quarlesi 37 Bulgaria 199, 325 bullfight 89, 94, 128, 139, 141, 328, 329 Camargue cattle 150, 271, 328, 333, 335, 336, 339, 348 Canada 37, 221, 244, 245, 247, 253, 256, 258, 259, 262, 277, 282, 286, 299, 300, 319, 320, 370 Canis lupus 258. See also ‘wolf’. Capreolus capreolus See ‘roe deer’. Carpinus betulus 190, 310. See also ‘hornbeam’. Castor fiber 186, 254, 278. See also ‘beaver’. cattle breeds 15, 18-20, 23, 53, 86, 118, 127, 128, 133, 139, 147-149, 152, 158, 159, 162, 166, 169, 170, 172, 174, 323-339, 344, 346, 348, 353-355, 361, 362, 366, 368, 373, 379-381 Caucasus 83, 202, 221, 243, 257, 260, 266, 281, 286, 295, 305, 378 Cervus elaphus 276, 278, 283, 318. See also ‘red deer’. Chianina cattle 149, 172, 174, 325, 329, 353, 362, 364, 365, 367 Chillingham cattle 162, 267, 270, 323, 325, 327, 328, 378 China 37, 45-48, 50, 129, 167, 182

Chinese water buffalo (Bubalus mephistopheles) 37 coastal marsh 236, 247, 252 coat-/ fur colour (aurochs/ cattle) 99, 135155, 165, 173, 174, 321-326, 329, 331, 338, 350-352, 357, 360, 361, 364, 366 colour genes (aurochs) 147-150 colour saddle (cattle) 138, 139, 141, 164, 332334, 346, 352, 356, 357 conversion factor (for shoulder height) 108, 111-113, 115, 171 Corsican cattle 325, 327, 333-341, 348 Corylus avellana 309. See also ‘hazel’. cranium. See ‘skull’. Crete, Island of 51, 89, 128, 139, 141, 289 crossbreeding aurochs/ domestic cattle 167, 168, 323 cultural-/ cultivated landscape 191, 239, 315, 372, 373, 378, 379, 381 curly-/ frizzy hair (aurochs) 92, 383, 385 decline (aurochs) 26, 53-78 Denmark 29, 52, 59, 60, 64, 107, 117, 134, 198, 235, 254, 261, 285, 306 dew claws (aurochs) 157, 158 Dexter cattle 172 disappearance/ extinction (aurochs) 53-78, 84, 285 disease (cattle) 20, 71, 75-78, 262, 264, 285, 328, 337, 354, 355, 374, 375 distribution area (aurochs) 36, 37, 41-52, 53, 61, 81, 102, 104, 129, 132, 133, 135, 175, 232, 234, 236, 259, 276, 285, 305 DNA research 29, 37, 39, 168 dodo (Raphus cucullatus) 72 domestication (aurochs) 48, 86, 88, 100, 107, 128, 131, 141, 147, 149, 151, 153, 162, 166-174, 215, 321, 323, 325, 326, 329, 339, 361, 363 Doñana National Park 268, 270, 323 early maturity (aurochs, cattle) 268, 270, 323 East Prussia 24, 61-64, 73, 75, 85, 123, 144, 192, 195, 199, 200-212, 237, 261, 273, 343, 371

Black Cyan Page 427

REGISTER

ecological partition 248, 249 ecology (aurochs) 174-322 edge- and hedgerow communities 190-192 eelstripe (aurochs) 138, 145-155, 321, 322, 324, 328, 329, 332, 334, 346, 352, 355358, 367 Egypt 43, 44, 54, 89, 126-128, 132, 138, 138141, 260, 325, 333, 334, 346 elder (Sambucus nigra) 190, 289, 290 elk (Alces alces) 19, 26, 46, 50, 60, 79, 106, 147, 155, 202, 213, 256, 261-263, 276, 278, 296, 298, 301, 304, 313, 314 elm (Ulmus sp.) 72, 180, 184, 222, 228-230, 235, 238, 293 England 29, 49, 55, 64, 103, 107, 108, 118, 122, 157, 162, 178, 179, 181, 190, 212, 225, 236, 237, 267, 287, 288, 306, 323, 327, 328, 369, 378 English Park cattle 149, 325, 328, 333, 335, 336, 339, 341, 348 Equus ferus 202, 284. See also ‘wild horse’. etymology 79-81, 386 European bison (Bison bonasus): Confusion with aurochs 83-88. Distribution area 305, 306. Food 221, 222. Habitat 243, 244, 248-251. Impact on vegetation 292297. Population density 248, 281, 286. Size 116, 117. Weight 119. evolution 31-40, 103, 303, 364 extinction (aurochs) 55, 60, 64, 72, 74, 78, 85, 202, 203, 330, 365, 375 facilitation 308 Fagus sylvatica 190, 310. See also ‘beech’. femur (aurochs) 108, 111, 116, 169, 174 fen 57, 209, 235, 245, 252, 372, 373, 381 fighting bull 94, 95, 97, 125, 134, 153, 262, 322, 329, 334, 340, 346, 349, 362 fighting cattle 16, 24, 125, 150, 159, 160, 174, 325, 329, 333, 335, 336, 339, 341, 343, 361, 364, 367, 378 fire 206, 245, 277, 279, 287, 298, 299, 314, 320, 370, 377 Fjäll cattle 172, 325, 335, 336, 340, 341

foliage (food) 197, 218, 228, 229, 231 food/ feeding (aurochs) 19, 21, 67, 72-75, 78, 93, 109, 117, 175, 213-218, 231, 239, 265, 363, 368, 380 food/ feeding (cattle) 18-23, 29, 34, 74, 174, 189, 213-217, 223-232, 236, 237, 246, 268, 270, 282, 288, 290, 324, 343, 355, 361, 363, 369, 370, 374-377, 379, 380, 382 food strategy 213-218 forest buffalo (Syncerus caffer nanus) 220, 222, 240, 242, 259, 282, 286, 287 forest elephant (Loxodonta cyclotis) 242, 286, 287, 290, 291, 320 Forest of Bia³owie¿a 15, 24, 75, 82, 96, 221, 222, 243, 249-252, 257, 272, 294, 295, 310, 312, 314, 318 Forest of Jaktorów 16, 24, 26, 65, 74, 76, 78, 79, 124, 216, 238, 239, 257 forest type 205, 236, 243, 249, 250, 251 France 24, 50, 59, 64, 95, 127, 137, 159, 192, 234, 237, 306, 321, 344 Franqueiro cattle 129, 172 Fraxinus sp. 190, 228. See also ‘ash’. Frisian cattle 158, 224, 326, 337 frontier forest 198, 199, 202 Fyn, Island of 285 Galloway cattle 149, 172, 226, 325 Gaul 192-194, 199, 212 gaur (Bos gaurus) 37, 39, 40, 45, 80, 108, 150, 151, 153, 154, 220, 240, 244, 248, 261, 262, 266-268, 270, 271, 281, 282, 286, 287, 296 Germania 26, 31, 55, 192-198, 212 Germany 11, 21, 24, 28, 29, 31, 36, 59, 61, 62, 64, 80, 84, 85, 97, 102, 106-108, 118, 121, 122, 142, 143, 167, 190, 192, 194, 218, 234, 236, 240, 241, 254, 284, 285, 287, 306309, 330, 342-345, 364, 365, 367 grazer 34, 35, 181, 183, 187, 213-217, 221, 227, 232, 236, 241, 242, 246, 247, 258, 259, 375, 378 Great Wilderness (‘Grosse Wildnis’) 62, 63, 65, 73, 156, 192, 195, 199, 200-212, 237,

427

Black Cyan Page 428

428


238, 241, 245, 246, 248, 256, 257, 273, 278280, 287, 371, 372 Greece 260, 289 habitat (aurochs) 45, 49, 50, 69, 70, 72, 74, 82, 97, 101, 117, 173, 175-259, 375-378 hair (aurochs) 25, 92, 93, 100, 145-147, 149, 151, 154, 160-165, 332, 334, 341, 346, 383, 385, 386 hare (Lepus europaeus) 284 hazel (Corylus avellana) 180, 184, 228, 230, 274, 309, 310, 311 heart bone (aurochs) 92, 94 Heck brothers 20, 29, 30, 109, 138, 323-359, 361, 364, 366 Heck cattle 11, 18, 20-24, 95, 96, 115, 148, 162, 173, 224, 261, 267, 282, 286, 341359, 360-369, 374-380 Hercynian Forest (‘Silva Hercynia’) 106, 194, 195, 198, 383 Hereford cattle 149, 226, 248, 325 Holocene period 28, 32, 37, 41-53, 58, 59, 87, 88, 97, 100-108, 115, 117, 120, 132, 166, 169, 175-179, 181-183, 185, 188, 216, 235, 253, 272, 276, 283, 284, 303-305, 310, 311, 352, 360, 372 hoof (aurochs) 116, 118, 155-158, 164, 236, 382 horn (aurochs): Colour 123, 124. Core 3537, 43, 47, 56, 57, 102-105, 107, 108, 120124, 128-135, 168, 347, 349, 360. Position 131-135. Shape 124-131. Sheath 57, 98, 120, 121, 129. Size 121-123. hornbeam (Carpinus betulus) 184, 190, 205, 206, 222, 235, 238, 239, 243, 249, 250, 274, 293, 310, 378 horse (domestic) 34, 35, 53, 67, 69, 75, 81, 91, 94, 109, 115, 135, 141, 164, 202, 203, 206-208, 211, 216, 217, 230, 234, 247, 248, 264, 279, 280, 283, 284, 288, 289, 303305, 309, 310, 313, 369, 370, 372, 379, 386 Hortobagy 312 humerus (aurochs) 108, 111-116, 120, 121, 171, 174

Hungarian Steppe cattle 110, 122, 159, 173, 174, 325, 327, 337-339, 341, 343, 353, 356 Hungary 29, 61, 63, 64, 106, 110, 117, 123, 129, 167, 199, 260, 327 hunting 19, 52-55, 59, 63, 65, 72, 74-76, 78, 79, 88-98, 106, 120, 124, 128, 138, 140, 143, 156, 166, 167, 174, 193, 197, 199, 203, 216, 235, 254, 260, 261, 269, 272, 281, 283285, 287, 303, 313-315, 318, 375, 384, 385 hypsodonty 214, 215 ice age 28, 35, 36, 41, 42, 49-52, 97, 100, 102, 183-185, 212, 234, 244, 283-285, 303, 306 impact (on forest growth) 29, 245, 246, 255, 256, 271-320, 369-373 inbreeding 76, 366, 367, 378, 379 incompatibility (genetic) 168 India 35-37, 40, 44, 45, 53, 104, 105, 133, 149, 150, 158, 167, 220, 244, 261, 262, 267, 268, 281, 286, 298, 323 insects 177-181, 191, 212, 245, 246, 314, 365, 375, 377 Inspection Report (Forest of Jaktorów) 26, 6771, 75-77, 83, 216, 218, 238, 264-266, 282 intermediate feeder 213-215, 221, 308 Ireland, Island of 49, 310, 311 Italy 50, 51, 55, 64, 79, 103, 104, 234, 383, 386 Jaktorów See ‘Forest of Jaktorów’. Japan 29, 48, 51, 323 Jersey cattle 110, 148, 149, 153, 174, 277, 321, 325, 326, 362 kouprey (Bos sauveli) 37 Lakenvelder cattle 149 landscape (natural) 175-212 Lascaux (cave) 23, 101, 117, 124, 126, 137, 153-155, 158, 159, 321, 346 late maturity (aurochs, cattle) 174, 325, 363, 368 Latvia 82, 325 Lepus europaeus 284. See also ‘hare’. Limousin cattle 149, 325 linden (Tilia sp.) 178, 180, 184, 205, 206, 222, 228, 230, 235, 238, 239, 243, 250, 274, 275, 293

Black Cyan Page 429

REGISTER

lion (Panthera leo) 42, 93, 138, 139, 233, 260, 262, 263, 303, 305, 364, 384 Lithuania 81-83, 199-202, 211 lowland anoa (Bubalus depressicornis) 37 Loxodonta africana 290. See also ‘savannah elephant’. Loxodonta cylotis 242, 290. See also ‘forest elephant’. lynx (Lynx lynx) 52, 202, 260, 263, 317-319 Mackenzie Bison Sanctuary 221, 301 Magnocaricion 251, 252 mammoth (woolly) (Mammuthus primigenius) 35, 50, 58, 137, 183, 215, 234, 303, 304 mantle- and fringe vegetations 189, 191 ‘marsh walker’ 240, 259 marshy forest 72, 239, 243, 251, 259 mating season (aurochs) 109, 163, 217, 267, 268, 270, 271 mauretanicus 36 Megaherbivorentheorie 307, 308 Mesolithic period 157, 158, 180, 190, 254, 284, 285, 287, 310 metacarpus (aurochs) 102, 108, 110-116, 169, 171 metatarsus (aurochs) 102, 108, 110, 111, 115, 116, 169 migration (seasonal) 257 molluscs 182 Montafoner cattle 325,340 ‘morstapa’ 240 mountain anoa (Bubalus quarlesi) 37 MRIJ (Maas-Rijn-IJssel) cattle 149, 158, 224 namadicus 36, 37, 44-46, 53, 104, 105, 133 nature (aurochs) 88-97, 362 nature conservation 12, 13, 17, 303, 306, 361, 375, 379 Neolithic period 46, 48, 107, 157, 162, 167169, 180, 183, 189, 190, 212, 223, 228230, 284, 285, 287, 306, 362 Netherlands 11, 17-24, 56-58, 64, 81, 97, 162, 177, 183, 187, 191, 192, 208, 212, 223, 224, 226, 237, 247, 252-254, 257, 269, 286, 287, 294, 297, 306-308, 345, 357, 376

neural spines (aurochs, European bison) 116, 117 New Forest 287, 288, 369, 370 Niata cattle 170 Norway 49, 228, 230 oak (Quercus robur) 69, 178, 205, 206, 208, 209, 217, 222, 228, 229, 238, 239, 249, 274, 275, 277, 289, 292-294, 309-311, 377, 378, 383 Oostvaardersplassen 11, 20, 24, 95, 224, 225, 266-268, 282, 286, 289, 290, 345, 351, 362, 366, 374-376, 379 ousting (aurochs) 72, 74, 75, 235, 375 pace length (aurochs, cattle) 158 Pakistan 45, 149, 167, 323 palynology/ pollen research 22-25, 175, 177, 180-192, 212, 245, 290, 308, 372 Panthera leo 260, 364. See also ‘lion’. Panthera tigris 278. See also ‘tiger’ peat bog 124, 175, 196, 212, 235, 245, 261, 372, 373, 381 Pinus sylvestris 205, 294. See also ‘Scots pine’. plant macrofossils 182, 190 Pleistocene period 28, 32, 35-37, 41-53, 58, 86, 88, 100-105, 108, 115, 117, 120, 126, 132, 133, 137, 166, 187, 244, 303-305 Poephagus mutus 37, 220, 243. See also ‘yak’. Poland 15, 19, 23-29, 31, 55, 60-66, 71-78, 81-84, 90, 94, 97, 98, 102, 123, 124, 147, 152, 161, 200, 205, 218, 231, 236, 237, 249252, 259-261, 271, 272, 275, 280, 286, 294, 319, 320, 322, 344, 383 pollen diagram 182-191, 208, 253, 256, 275, 311 poplar/ aspen (Populus sp.) 78, 185, 187, 188, 205, 208, 222, 228, 229, 235, 274, 275, 298, 299, 301, 318 population density (aurochs, cattle) 280-286, 288, 320, 373, 377-381 Populus sp. 185, 187, 246, 297, 298. See also ‘poplar/ aspen’. Portugal 118, 148, 324, 325, 361, 367

429

Black Cyan Page 430

430


Preboreal period 234, 235, 284, 306 predation 199, 203, 259-264, 267, 281, 283, 298, 306, 307, 313, 317-319 primigenius 31, 32, 36, 37, 40, 44, 46, 48, 88, 104, 105, 129, 325, 326 Prunetalia spinosae 189 quagga (Equus quagga quagga) 364 Quercus robur 309. See also ‘oak’. radius (aurochs) 108, 111-116, 169 red deer (Cervus elaphus) 48, 60, 74, 95, 109, 118, 119, 136, 147, 176, 202, 213, 233, 263, 267, 278, 281, 283, 284, 289, 292, 295, 296, 307, 310, 313-319, 343, 378, 379, 383 Red Pied cattle 141, 149, 325, 328, 329 reem 85, 89 reproduction (aurochs) 174, 264-271 rhinoceros (woolly) (Coelodonta antiquitatis) 35, 48, 50, 58, 183, 215, 234, 304 roe deer (Capreolus capreolus) 60, 119, 147, 202, 213, 263, 278, 296, 313, 314, 383 Romania 106, 144, 194, 195, 198, 229, 260, 325 Romans (writers) 24, 26, 56, 79, 89, 106, 125, 143, 192-198, 208, 210, 212, 372 Rominten/ Rominter Heide 96, 205, 261, 343, 349, 350 ruminants 33, 34, 213-215 rune (aurochs) 239 running wild/ feral (cattle) 18, 20, 24, 59, 95, 139, 167, 223, 264, 267, 268, 323, 327, 328, 355, 363, 374, 378 Russia 27, 36, 37, 48, 49, 63, 64, 71, 75, 85, 89-91, 123, 132, 141, 144, 182, 183, 200, 216, 221, 239, 243, 274, 281, 292, 295, 296, 325, 383 sacred forest 197 Sahara 42-44, 233 Salix sp. 186, 190, 221, 224, 225, 233, 300. See also ‘willow’. salt lick 277 Sambucus nigra 190, 289, 290. See also ‘elder’. savannah elephant (Loxodonta africana) 290, 292 Schorfheide 342, 343

Scots pine (Pinus sylvestris) 205, 223, 224, 229, 238, 274, 289, 294, 307 Scottish Highland cattle 17, 21, 149, 223, 224, 269, 270, 325, 327, 335-341 sedge marsh 245, 247, 251, 253, 268, 286, 302, 370, 376 sedge peat 186, 253, 302 senses (aurochs, cattle) 171, 364 Serengeti 42, 219, 233, 256, 307 sexual dimorphism (aurochs, cattle) 99, 108, 109, 112, 116, 136, 161, 167, 171, 174, 267, 325, 354 SIERDA 344 silva 193, 194 Silva Hercynia 194. See also ‘Hercynian Forest’. Simmenthal cattle 149, 325 Siwalik Hills 35 Sjaelland, Island of 52, 60, 285 skeleton (aurochs) 28-32, 61, 62, 86, 88, 102, 106, 112-119, 261, 284, 351 skin (aurochs) 27, 92, 93, 116, 142, 144-147, 154, 160, 161, 164, 385, 386 skull (aurochs, cattle) 35-39, 43, 44, 47, 56, 87, 88, 102-105, 124, 129-135, 168-171, 174, 215, 216, 347, 349, 368 snout (aurochs, cattle) 138, 149-151, 153, 155, 165, 322, 327, 332, 337, 352, 356358, 367 social structure (aurochs, cattle) 95, 109, 264-271, 363, 373, 379 Spain 36, 55, 94, 125, 134, 137, 160, 234, 262, 268, 270, 306, 322, 324, 329, 336, 340, 346, 347, 349 steppe bison (Bison priscus) 35, 58, 131, 183, 215, 234, 244, 303, 306 storm 19, 194, 246, 279, 287, 314, 315, 377 Sus scrofa 284. See also ‚wild boar’. Sweden 49, 59-61, 97, 102, 118, 228, 235, 241, 285, 305, 306 Swiss National Park 315-319 Switzerland 59, 117, 190, 230, 236, 243, 285, 306, 315, 319 symbolism (aurochs) 88-97

Black Cyan Page 431

REGISTER

Syncerus caffer caffer 219, 242. See also ‘African buffalo’. Syncerus caffer nanus 220, 222, 242. See also ‘forest buffalo’. systematics 36-40 Tai Forest 242, 282, 287, 291 tamarau (Bubalus mindorensis) 37 taurine cattle 166, 168, 326 teeth/ molars 34, 108, 109, 214-216 Texas Longhorn cattle 148, 324, 325 tibia (aurochs) 108, 111, 116, 169 tiger (Panthera tigris) 260-262, 278, 279, 304, 305 Tilia sp. 228. See also ‘linden’. toponym 79, 81-83, 97, 306 trail/ track (aurochs) 118, 157, 158, 236, 370 Trajan’s column 195, 198 trunk length 118, 170, 354, 358, 363 tur 63, 65, 73, 82-85, 91, 93, 106, 142, 144, 145, 156, 383, 386 Turkey 89, 128, 131, 166, 234 udder 99, 137, 158-160, 164, 165, 174, 323, 325, 327-329, 334-336, 338, 340, 352, 354, 355, 358, 360, 361, 363, 366-368 Ukraine 50, 301, 327, 336 Ulmus sp. 228. See also ‘elm’. ulna (aurochs) 108 United States 37, 188, 207, 211, 225, 226, 244, 256, 263, 276, 277, 286, 297, 310, 318, 319, 323, 324, 364 ur 62, 79-85, 97 Ursus arctos See ‘brown bear’.

urus 31, 32, 79, 141 Ussuri region 203, 211, 277-280 vegetation openness 181-184, 186, 191, 282, 284, 285, 308, 314, 317, 320, 371 wapiti (Cervus elaphus) 263, 276, 307, 318 water buffalo (Bos arnee) 37, 45, 243 Watussi cattle 172, 333, 343, 348 weight (aurochs) 119 White Park Cattle 327 width of the forehead (aurochs) 103, 104 wild boar (Sus scrofa) 60, 74, 109, 118, 143, 202, 233, 263, 267, 284, 368, 375, 377, 383 wild horse (Equus ferus) 176, 200, 202, 247, 258, 259, 284, 303, 305, 309, 310, 313 willow (Salix sp.) 178, 183, 186, 188, 190, 221, 224-227, 230, 233, 235, 240, 251, 300302, 318 withers-/ shoulder height (aurochs, cattle) 104, 109-120, 158, 165, 166, 169-172, 174, 327329, 334, 352, 353, 357, 358, 363, 365, 367 Witrik cattle 149 wolf (Canis lupus) 18, 60, 133, 166, 199, 202, 259-264, 270, 289, 298, 307, 318, 330, 372, 374, 380, 382, 384, 385 Wood Buffalo National Park 220, 244, 245, 252, 258, 262, 277, 282, 297-302, 320, 370 yak (Poephagus mutus) 37, 39, 40, 108, 150, 151, 173, 220, 243, 247 Yellowstone National Park 263, 307, 318 Zamoœæ 71 zebu cattle 37, 45, 53, 148, 149, 153, 154, 166-168, 226, 325, 326, 334

431

Black Cyan Page 432

432


Retracing the Aurochs: History, Morphology & Ecology of an Extinct Wild Ox

Ox

Ox

The Extinct

The Extinct

Ox

Ox

Ox

Ox

An Anthology of Structural Morphology

The Extinct

The Extinct

Ox

Ox

Ox

Ox

Living Extinct

Living Extinct

Living Extinct

Dictionary of Environment & Ecology

The Handbook of Morphology

The Waters & the Wild

The Ox-Bow Incident

An Introduction to English Morphology

The Evolution Of Morphology

Living Extinct

Yearbook of Morphology 2002 (Yearbook of Morphology)

Crystals: Growth, Morphology, & Perfection

An Introduction to English Morphology

An Introduction to English Morphology

Deleuze Guattari & Ecology

Retracing the Aurochs: History, Morphology & Ecology of an Extinct Wild Ox

Ox

Ox

The Extinct

The Extinct

Ox

Ox

Ox

Ox

An Anthology of Structural Morphology

The Extinct

The Extinct

Ox

Ox

Ox

Ox

Living Extinct

Living Extinct

Living Extinct

Dictionary of Environment & Ecology

The Handbook of Morphology

The Waters & the Wild

The Ox-Bow Incident

An Introduction to English Morphology

The Evolution Of Morphology

Living Extinct

Yearbook of Morphology 2002 (Yearbook of Morphology)

Crystals: Growth, Morphology, & Perfection

An Introduction to English Morphology

An Introduction to English Morphology

Deleuze Guattari & Ecology

Recommend Documents