Solnhofen: A Study in Mesozoic Palaeontology

Solnhofen '" A study in Mesozoic palaeontology "

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K. W. Barthel N.H.M. Swinburne S. Conway Morris

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Solnhofen A study in Mesozoic palaeontology

K. W. Barthel Form erly of InstilUle for Geology alld Pa/aeom %gy Technical University , Berlin

N. H. M. Swinburne Departmem of Earth Sciences The Open Uni versity , Millon Keynes

S. Conway Morris Department of Earth Sciences University of Cambridge

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Published by the Press Syndicate of Ihe University of Cambridge The Pitt Building. Trumpington Street. Cambridge CB2 l RP 40 West 20Ih Streel. New York, NY 10011-4211. USA JO Stamford ROlId, Oakleigh, Melbou rne 3166. Australia Originally published in German as So/nhoJm: Ein BUck in dit Ergrschichft by Ott Verlagl, Thun, 1978 and 0 Ott Verlag Thun 1978 This revised translation first published in English by Cambridge Universit y Press 1990 First paperback edition (wit h corm:tions) 1994 English translation 0 Cambridge University Press 1990 Printed m Great Britain at the University Press. Cambridge I1mis" Librur), ralilloRuillS ill publicu/ioll du/o

SMthe!. K. W. (K. Werner) 11./978 Solnhofen. - Rev. cd. I . West Ge rm any. Jurassic Strata . Fossils I. Ti tle 11 . SWinburne. N. H. M . (Nicola H. M .) Il l. Co nway Morris. S. 560' . 1'7640943 Librory of CongrtsJ cora/oguillg in publicOliOlI dOlO

Barthel. K. Werne r. !Solnhofc n. English] Solnhofen: a stud y in MeSOl:oie pa!ocon tology/K. W. 8arthel; [translated and revi$Cd by] N. H. M. Swinburne: [edited by] S. Conway Morris. p. em. Translation of: Solnhofen . Includes bibliographical references. ISBN 052 1 33344 X I. Palaeontology-Ju rassic. 2. Pal;leo1l1ology-Germany (West)Solnhofen. I. Swinburne. N. H. M. (Nicola Helga Marga ret). 1962. [I. Conway Morris, S. (Simon) Ill. Tille. 0E.733. 83713 1990 560' . I'7640943-dc20 1'9·25440 CIP

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Contents

I', d"cc hhrcv iations for museums

Tht Soln hofen limestone Introduct ion: T he limestone from Solnhofen IlislOry of plattenkalk exploitation

Cullections of foss ils

a

vi i

ix

4

9

( ;t'ological history and stratigraphy G eological history of the Southe rn Fra nconian A lb Pulacogeography and fac ies dist ribution in Late J urassic li mes

17 17 24

IIi'lI'UJ,:nl,)hy of the Solnhofen Plattenkalk

38 38

t ,'hv logy inlhe Solnhofen-Eichstatt a rea ()ccurrcnce of macrofossils ( ;l'l1 in ~ and microfossi ls llill,!!c nc tic altera tion of the sediment Ih'd' slrihut ion of e lements

1',lllu'ocnvironmcn( a nd sedimenta tion I'u lucucnvironmenl 111\' I'c~ tri c l ed basin model r h ~' mi " lr y or the Solnhofen waters and special preservation IIloml1" of microorgan isms I h\' i:ya no hactc rial mal t\ ilc pu:o.lliona l illodc l (Itill" (\e pi)',ilion,l1lhco ries

1'"lnl'\H'voluIo;Y 1' ,.I ,ll'udlllUtlc 1 II! III 1he lago o n 1(\'1' 111 1 ~'il1l11nIlI111i c~ 11'11 v"11 i" j CC()"Y' iC!11 '

40 41

49 52 56 56 56

59 64 64 65

67 71 71

73 79

84 v

Colltellts

6 Taphonomy Introduction Biostratinomy of the marine biota Biostratinomy of the terrestrial biota Fossi l diagenesis Chemistry of fossil preservation 7

The Fossils Int roduction Monerans and prot ists ~_

Non-vascular plants - brown algae Vascular plants Gymnosperms - seed ferns, Bennettitales. ginkgos , conifers Animals Invertebrates Sponges Cnidarians - jellyfish , hydrozoans , corals Annelid worms Bryozoans Brachiopods Molluscs - bivalves, gastropods, cephalopods Arthropods - crustaceans, chelicerales, insects Echinoderms - sea-lilies, starfish , brittle stars, sea-urch ins, sea-cucumbers Vertebrates Fish Reptiles Birds

89 89 89 93 96

100 102 102 102

lID

103 103

103 112 11 2 112 112 117 117 118

119 129 153 160 160 173 191

8 Conclusions: The Solnhofen Plattenkalk and comparisons to other plattenkalk lagerstattcn Summary of the characteristics of the Solnhofen Plattenkalk Olher plattenkalks wil h exceptionally preserved fossils

202 202 203

Appendix: Faunal and ftora ll ist Bibliography Systematic index Gener;!1 index

206 217 231 234

Preface

S ~'lhtll e n t ary rocks usually teem with fossils, but palaeontologists have long Il'I.'ognized that as a sa mple of o riginal life they are grossly in adeq uate . Th is is Im ply because in most circumstances only the resistant she lls and bones can IIIV IVt.! \0 fossilize. Very occasionall y, howeve r, the curtain of preservatio n is illt l' d slightly highe r and we begin to o btain a glimpse into the extraordinary '''Ilure of former life. Such deposits , where delicate and soft-bodied creatures m ' fossilized , are gene rally known as fossil1agershitten. Of these the Solnho10"11 IlIllcstonc is the most renowned Jurassic example. Over the centuries this '("1)("11 has produced a diverse collection of supe rbly preserved fossils, includ* 111 ", !lome anim als showing soft -part preservation . The Solnhofen organi sms are thl1l1t1oht to have been washed in to hypersali ne lagoona l basins and buried t 'l li dl y by fin e micritic mud , by which means they have been exceptio na lly

l't ~·'lC rved.

I ht ~ book com mences with a short history of the scient ifi c and commercial , _plll rlltion and explo.itation o f the limestone. The fo llowing chapter then p uts (ill' Solnhofen limestone into the context o f south G erm an stratigraphy and v. Illogical history . It next focllses upon events in La te J urassic times and the o",l uho fcn area. T he third cha pter presents a precise petrological description of Ih, Solnhofen limestone fro m its appearance at outcrop to elect ron microII l ullh~ of the microfa cies. Lit hological detai ls and geochem ical information are II • It 10 reco nstruct the palaeoenvironment at the site of depositio n, and , uIIl! lIl ll11S necessary fo r special preservation . The remai ning sections o f t he [< • •ilk pe rtain to the foss ils themselves: the o riginal palaeoecology o f t he ' " 1(11 111" 111 5, the short bursts (or last gasps) of life in the lagoonal basins and t he 1,11'hullomic processes affecting the fossils from buri al unti l their final exhumaI (un In the Im;1 chapler Ihe fossils are presented taxono mically , accompanied 11\ II "Inking co llection o f plates . Willi.. , preserving Ihe spirit of Barthel's book , this version is intended fo r a . 11. , . ~' udvanccd readershi p . I, is considerably reordered and more than half is 'I' w t1I (tic rial. Th is accommodates a change in emphasis towards discussio n of III! Solnhofen pa laeoe nvironment and the condit ions required for special I'" ~I'rv llti {) n . Th us it incl udes import ant research material not previously jtl'llllIhlc in an ea ~ jl y OI(;ccss iblc for m . In particular, emphasis is given to l it hll llt Kcupp's dCll1i led dcscription of the microfacies and discovery of

vii

Prcf(/c~

pu t.Hive cyanobacterial spheres. togethe r with his resultant stromatolite depositional theory. Meyer & Schmidt- Kaler's excellent book concern ing the stratigraphy lind palaeobiogeography of the area. published in 1984, has also been an im porta nt source o f material. K. Werner Barthe l slldly died in 1978, the yea r afler publ ication of his book Solllllofell : £ill Blick i1l (lie £r iI ' transitluIlU! h)"~I " 11lthnugh he

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Collections of fossils

thought that Archaeopteryx had evolved from the long-tai led pterosaur Rhamphorhynclllls (see Gou ld 1987). Nowadays we think that small running dinosaurs , relatives of CompsoglIathus, could have given rise to Archaeopteryx some time in the Late Jurassic. Both Archaeopteryx and Compsogllathus are described in chapter 7, where the rcader can see just how similar the creatu res are. The evolution of Archaeopleryx, the acquisition of feat hers and the origin of flight are subjects still bci ng actively debated. (The reader is referred to the symposium volume of the Archaeopteryx conference, held in Eichstatt, in 1984, edited by Hecht el al. 1985.) Recently , Archaeopteryx has again been in the public eye because of allegations by the physicist Fred Hoyle and mathematician C ha ndra Wickramasinghe (1986) that the fossils are the result of clever ni neteemh-century rorgeries. They claim that skeletons of the small dinosaur Compso811atlwswere take n, areas around the bones excavated and then infi lled by a paste made of ground-up limestone together with a bind ing agen t. The feathe rs of a modern bird were then pressed into the cement to make the impressions around the !>ke leton. Hoyle and Wick ramasinghe claim a major conspiracy involving Haberlei n , Owen (who alleged ly commissioned the forgery in order to undermine Darwin and Huxley) and a succession of curators at the Natural History Museum . Hoyle and Wickramasi nghe's allegations have been shown elsewhere (Charig et af. 1986, Swi nburne 1988, amongst others) to be mischievous and utterly without foundation. In 1877 a second Archaeopteryx was fou nd. It possessed even finer features as well as a complete skull and was sold to the museum of Berlin by Dr lliibcrlein's son, Ernst Haberlein . This Haberlein, together with a nother doctor, Dr Redenbacher [rom the local town of Pappe nheim, assembled a fi ne collection of Solnhofen fossi ls in the middle of the nineteenth century. Specimens in many museums sti ll bear the labels saying ' Haeber1ein'sche Samml ung' (from H abe rlein 's col lection) or ' Redenbachersche Sam mlung'. Archaeopteryx was but one of the splendid foss ils from the Solnhofen IOC;l lities and interest in the e ntire range o f fossi ls was stimulated . Major collections were established and Ihese encouraged a consideration of the So lnhofen fauna a nd flora as a whole. The first list of fossils was compiled by Ludwig Frischmann in 1853. In 1904 Johan nes Walt her published a comprehensive work e ntitled Die FalilUl tier SolnhoJeller Plattenkalke. This sum marlied Frischmann's list into a total of 650 known species, now thought to be an liver-estima te as it includes many incomplete a nd juvenile specimens as \cparate species. A more accurate figure is thought to be around 600 (0. Viohl , pen•. comm. 1986). Walther was also first to deal with the relat ionsh ip of the nrgilnisms to the sediment in which they were buried. In pa rticular he observed that almost all !>pecime ns were loc!>i b halo ensured that important fossils are either incorporated into the collect ions of the quarry owners or find their way into the hands of de;llers. To acquire a decent specimen of, say. a pterosaur would involve thousands of deutschmarks and an Arclweopteryx would be almost unpriceable! As fossils may even be an important part of the economy of the quarry, the working quarries exclude fossil hunters. There are various disused quarries , but fre sh rock is covered by debris piles which have been tho roughly scoured for fossils over the centuries. However, a few quarries have been especially designated as fossil quarries and here fresh sections are accessible and may be worked by amateurs. Although it is certai nl y less easy to come across fossils by accident than it once was this does not seem to have affected the popularity o f fossil hunting. In fact , this activity is so popular that entire German families will come to the region on fossil collecti ng holidays. For visitors to the area the magnificently situated Jura-M useum in Willi baldsburg , Eichstiill , houses the largest of the public collections. Also in the Eichstall area , at Harthof, is the private collect ion of the Museum Berg(!r . In Solnhofen itself is the Burgermeister-Mliller-Muscum as well as a large private collection open to the public at the Maxberg Quarry (Museum des Solnhofen Aktienvcreins). The Bavarian State Museum (Bayerische Staatssamm lung fUr Palaontologie und historische Geologie) in Munich is also well worth a visit.

2

Geological history and stratigraphy

Geological histor y of the Southern Franconian Alb rhe stratigraphy of Bavaria in southern Germany is relatively simple. The hcdding dips gently towards the southeast causing Jurassic rocks to outcrop over most of the Southern Franconian A lb (figs. 2.1-2.5). To the north and

northwest these pass downwards ;nlo Triassic sediments, whilst to the south ,lI1d southeast they are overla in by deposits laid down in the Tertiary Period. The o lder Pa laeozoic rocks, which arc metamorphosed and ridd led wi th igneous intrusions, underlie the area, but in the east are brough t to t he surface 111 Ihe fault blocks of the Bo hemian Massif. In Palaeozoic times these rocks formed parts of a series o f ancient mountain ranges which ran east-west across \()uthern Germany. These mountains were formed during a tectonic episode known as the Hercynian (Variscan) orogeny wh ich culminated late in th e l ';.rbonife rous Period at around 300 Ma. The mountain s were subject to t'rusion so that by the beginni ng of the succeeding Permian Period they had hee n worn down to expose their metamorphic and gra nite core. In the lu llowing periods these harder rocks persisted as a topographic high , known as the Vindelicisch Land , the position of which dominated regional palaeogeography until the e nd oflh e middle part of the Jurassic Period (see I1g2.6). Through tl nte the Vi ndclicisch Land subsided and began to be covered by sediments. In I arly Permian times conti nental sediments accumulated in the initial deprc\sions. Thereafter , as the land san k, the sea gradually transgressed southI·,j ... twards. To the northwest of the Southern Franconian Alb ma rine sedimC llts, such us Ihe well-known Middl e Triassic Muschelkalk, were deposited, " ll\: losing foss ils such as ce rati te ammonoids and oysters. In contrast. Triassic n lullc nts of the Southe rn Franconian Alb arc mostly non-marine sha les and ;lIllhlones, laid down in fluviatile and delta ic settings. lull y marine deposition over the entire area did not begin unlillhe Jurassic Pe riod when the Southern Francon ian Alb Illy under the shallow waters o r a WIde shelf sea . Sediments of Early Jurassic age (Lias) are dark shales and UI):ll1l1ceous (clayey) limestones and the period is often referred to as the Black I\IIU\..,;c . In the succeeding middle part of the Jurassic (Dogger) Period -I nd , lones prcdominate . and this interval is ofte n called the Brown Jurassic. \ .. t hI.' ,ca rldvanced du ring Middle J unNic times . Ihe coasHi ne migra ted 10 I he

17

Geological hislOfY fllld slrtlligmpity

Fig. 2.1 Outcrops of Jurassic rocks (stippled) in northern Europe. The inset shows the situation in the Southern Franeonian Alb and the section line of fig. 2.2. Adapted fronl Meyer. in Meyer & Schmidt-Kaler ( 1984) and Viohl (1985).

south and east but the palaeogeography was complicated by the developme nt of a northwesterly projecting spur of resistant land around Landshut with a corresponding indentation known as the Regensburg embayment. Conti nued incision of the Regensburg embayment into the coastline eventually breached the Vindelicisch Land ba rrier. The northern waters became connected to the sou therly Tethys Ocean and the Vindelicisch Land became an island . So it was th;11 by Late Ju rassic (MaIm) times, where the rock .. nrc often called t h e White Jurassic because of the llbunclancc of lir11(.' .. tOIli:'I, ll1l' \:on lin uing

Geological history of Ihe SOli/hem FrallCOlliuli Alb

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ma ri ne transgression had comple tely flooded the Vindelicisch Land . The tn lluence of land now came from the north from the uprising ' Mitte ldeutsehe Sl.:hwelle' . For the fi rst time the Southern Franco nian Alb could exchange walcrs directly with the Tethys Ocean . However, evide nce from recentl y lh"i llcd boreholes to the south o f Munich (used in the palaeogeographic ICl.:onstruction of Meyer, see fig. 4. 1, p. 57) suggests that such an interchange of Willers would have been impeded by carbonate shoals which lay under o nly a h'w me tres of water on the northernmost margin of Teth ys. In this t ranquil 'I'l1 l11g orga nically mediated carbo nate mounds and reefs spread across the I ,u ho nate platform of the Southern Franconian Alb. Between the reefs lay "1l1ll 11 basins which were infi lled by light co loured limestones and argillaceous Ihllc'tOlles. These incl uded the Solnho fen Plattenka lk. luwards the ve ry end of the J urassic, deposition over the Sout hern FrancoIIl,tll Alh came to an end, as land emerged in the northwest and east and t he Il'Ihys sca regrcsscd to the soulh . Exposed until Upper C retaceous times , the Mn cllOic lIcdimcnts were then subjected to the actio ns of wind and weather. I hl'cl'lI fI Cr, there was on ly one , short further period of widespread marine ~, ,IU tlcnlnt io ll which took place at the beginning of the Upper Cretaceous (in 1111 (\:nomanian- Turonill n stages, around 95 Ma). The sea advanced fro m the "Il Il H: II ~ I , nooding the area from Ries to Regc nsburg and laying down I ttd ~ \( lII c~ li nd impure limcstones. Around Rege nsburg Ih is sedi ment is now

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Nomenclature for subdivision o f the Jurassic Period.

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hg. 2.5 Seclion Ihrough strata of the Solnho ren area. The besl exposures oceur in valleys sueh as Ihose now occupied by the Danube and A)tmilh\.

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exposed , but in the Solnhofe n area , whe re it was once deposited over the plullcn kalk , it has since been eroded away. Evidence for its presence remains only in the Neuburg area, in thc form of a few silicified knolls, which resisted erosion. and sediments which survived by having been piped or channelled into Cllves benea th the previolls c rosion surface. Si nce the last rct reat of the seas, the Southern Francon ian Alb has remained

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Gcological history and stratigraphy

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limestones. Deepest of all is the Altmuhl valley wh ich runs betwecn Dollnstein and Kel heim. Although the present-day A ltmuh l river is too small to be responsible for such largc-scaleerosion, unt il the late lee Age the valley used to guide the waters which now flow furt her south down the Danube valley. Throughout the Quaternary Period deep ground fros ts furt her weathered the rock. At the end of the Ice Age men began to colonize the valley. Theold caves in the valley sides provided protection and the A ltmuhl region is very well known for the many fi nds of prehistoric artifacts.

Palaeogeography and facies distribution in Late Jurassic times LaI C in the Jurassic Period the Southern Fra nconian Alb WU'I co ve red by a broad shelf sea which beca me progressively shallower [wet 1110 ,,' ]IInlll l cd (sec

Pa{aeogeogmplly alld iades distribution

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lig. 2. 7, regional palaeogeography for Late Jurassic times). To the northeast lily the Bohem ian Massif, to the north and northwest the eme rge nt ' Mitteldeulsche Schwelle'. and to the south a shallow-water platform , represe nti ng Ihe submerged remnants of the Vindelicisch Land , which limited exchange of wuh,:r with the Tethys Ocean . Channels con nected the waters of the Tethys Ocean with those of the Southern Franconian Alb and along these condu its (a mc Tethyan-type marine sediments and animals. The positions of these pu!>,agcs are uncertain but they are genera lly placed to the west and thus ubscured under recent cover in the Ries Meteorite Cratcr. TI1 C Solnhofen wa ter mass must also have been connected to the northern Boreal Ocean, hccllUse there are also rare boreal representatives in the Solnhofen fauna (Zc i~s 19(4). Fvcnls leading to the formati on of the plattenkalk basi ns began in Oxfordian times (~cc fig , 2. 901) . In the west the deposition of regularly bedded argill aceous lill\c~toncs. similar 10 tho~c of Midd le Jurassic times conti nued , and was

Geological history a"d stratigraphy

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Fig. 2.7 General palaeogeography of northern Europe in Late Jurassic times. The shaded areas represent landmasses from a shallow continental sea on {he m.ugins of the Tethys Ocean. The small box shows the position or the Southern Franconian Alb. From Viohl ( 1985).

followed by purer limestones, wh ilst in the east a differen t facies was developed (a facies refers to an association of lithologies which are the deposits of a certain sedimentary environment). Slight submarine highs were colonized by an association of plate-like siliceous sponges and encrusted by cyanobacteria, between which were trapped peloids (amorphous lumps of calcium carbonate , some of wh ich were perhaps originally faecal pellets), whilst at the same time lime mud (micrite) was deposited on adjacent areas of lower relief(fig. 2.8). On the highs, in the sponge-alga l beds, not only was calcium carbonate produced at a faster rate , but this facies was cemented at an early stage making it resistant to compaction relative to the adjacenl bedded micrites. A primary difference in relief between the sponge mound highs and intervening basins was thus acce ntuated. In most places the organically constructed mounds formed gently rolling submarine hills but where the mounds protruded more abruptly from the seafloor (as in the Kelheim area) chunks of debris occasional ly fell from the edge of the reef mass into the su rrounding bedded facies . Tn the Kimmeridgian stage the sponges proliferated (figs 2.9h. c & d and 2. 10) and the barriers o rientated along a NW- SE trcllll liprCu\1 ~h'w l y north-

26

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Fig. 2.8 Lithostratigraphy of the Southern Franconian Alb. Note how the growth of the sponge-algal mounds controlled deposition of the bedded facies. From Vionl

( 1985),

Geological history (l/id straligra"hy

OXFOROIAN

(ox,. 21

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" Fig. 2.9 Palaeogeography of the Late Jurassic. The Southern Franconian Alb was a shelf area on the northern margin of the Tethys Ocean submerged under shallow water. During Late Jurassic times sponges colonized original highs in the underlyi ng topography, building mounds which accentuated the relief of the seafloor. Protected basins were formed between the sponge-algal mounds and in thcse bedded areas limestones wcredepositcd . (a) Oxfordian (OXl +2) - start of sponge-algal mound growth in the east, probably as a result of shallowing. (b) Early Kimmeridgian (kil ) - sponge-algal mounds spread westwards. (c) Middle Kimmeridgian (ki 2}- spongegrowth reaches a climax and adjaccnt sponge-algal mounds fu se 10 leave intervening basins. (d) Late Kimmeridgian (kil ) -watcr shallows further. SpongesspreaJ to the bottom orthe basins and appear in the bedded facies. Plattenkalk starts to bc dcposited in the east. (e) Early Tithonian (t~) - Plallenkalk is dcposited over most of the Southern Franconian Alb. (f) Latc Tithonian (tiJ) - sea starts to retreat and land emerges to the north. Planenkalk is still deposited in the south. From Meyer & Schmidt-Kaler (1984) and from Meyer in Meyer & SchmidtKaler (1984). wards. The chai ns fused into a network e nclosi ng small basins which gradually sh rank in size as they were overgrown by the rapidly accreting sponge mounds. The basins were first filled with bankkalk ('Bankkldk ') , i.e. thick (dm to m) regularly bedded limestones , often rich in ammonites and with slight ly undulose beddi ng pla nes. Bankkalk , like plattenkalk, has occasional th in intercalations o f argillaceous limestones. The sponges spreecb bn::lk wilh II l'lluchoidal 1

Lithology in th e Soltrh oJen- Eichstiilt arera

h g. 3.1 Eichstan plattenkalk facie s at Neumayer quarry, 500 m NE of Schernfeld . I hc beds arc typically 0.5-1.0 em thick and separated from cach olher by clayey p.!ftings. From Meyer & Schmidt-Kaler ([984).

frac ture to display a darker, beige-grey interior. Sometimes beddi ng planes I1 lay show other colou rs, such as concentric bands of yellow , brown o r grey , or the beaut iful fern-like markings called dendrites. Despite the plant-li ke llppcant nCe, these dendrites arc inorganic fea tures caused by a solutio n of the \ 111i1 11 amou nts of iron o r manganese from inside the beds and their reprecipiIII lion , FAUlE

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Fig. 3.5 Size frequenc), diagram of partide size of disaggregated plattenkalk . (a) F1inz from Maxberg quarry , Solnhofen; (b) fiiule from Schcrnfcld. From Kcupp (l977a).

Fig. 3.6 Plattenkalk foraminifera. (3) GUlidryina bllkowiensis Cushman & Glazcwski ( X 42) ; (b) Nodosaria ellglyplw Schwager (x22); (e) Margilllllina

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dislorla Kusnetzowa ( x 40) ; (d) Quillqileioclilino. egmonle1lsis Lloyd (X45) ; (e) Patellina [eifeli (Paalzow) from

above (left) , from below (right) and from the side (above) . From Groiss (1967).

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58

Chemistry 0/ the Sol/Iho/en wmers

the bod ies fell through the wate r to come to lie o n the sediment surface. Normally corpses lying o n th e seafloor are not left undisturbed because they .lte ut ili zed as a food source by scavenging ani ma ls. They would be d ismemhaed and partially consumed , prod ucing a large surface area for che mica l degradation by microbes and eventua lly resulti ng in the recycling of the o rganic molecules for the growth and construction of ot her o rganisms. But this was no t Ihe case in the Solnhofen environmenL The origin o f the exceptional preservatIon of the Solnhofen fossils is lin ked to the exclusion of macrobenthos from Ihe si te of burial , which ensured that the bodies were not ripped to pieces. I urt hermo re, the inacti vation of a large sectio n o f the no rmal microbial l'\lfllm unity resulted in a very slow rate of decay. Together these factors he lped to ensu re the remarkable preservation of the Solnhofen fossils.

Chemistry of the Solnhofen waters and special preservation HYPERSALINITY? The most convincing explanatio n fo r the poisono us properties o f the stagnant wate rs. that so favoured the exceptio nal prese rvation of the fossils. appears to he an excessive concent ratio n o f sa il. Regrettably, the evidence for hypersalililly continues to be indirect. On pp . 71-3. the evidence for th e climate "ffect ing the Solnhofen area is discussed . Arguments point to hot and dry w nditions, with no indications of any substantial runoff from adjacent landm:lsscs. Unde r such conditions, evaporatio n would be intense and , as the ~!llini t y increased , the dense brine would collect in pools at the bottom o f the IW'IlIls. The brine would have a range of conseque nces fo r the biota and its preservation. Because elevated salinities are fatal to most normal marine nrg:misms. macrobenthos wo uld be excluded from inhabiting the lagoon . !'hese ,mimals SlOp functioning in hypersaline solutions because wate r is withd rawn osmoticall y fro m the tissues. The shrivell ed appearance of ma ny "'iolnhofen jellyfi sh (especially those from the Gungolding-Pfalzpaint area) is qu ite consistent with this idea of hypersalinity. Ano ther consequence o f hypersalinit y is the preservatio n of organic material. ' Pickling' in salt solutio ns 1\ an effecti ve method of culinary preservation, because many of the decompll~11lg microbes arc inactivated and the decay processes greatly slowed . In 11;lt urlll hypersaline environme nts there are documented cases of exceptiona lly \low org:lIl ic decay whk h has resulted in specia l preservation of o rganisms. Fo r ~'·(i111lp l c. in the extreme ly ho t environ ment of Death Valley. Ca lifo rnia, there Me c~ampJcs of cx q ui~itc preservatio n o r insects in the sa lt-rich sedime nt (W. Ucrger. l>cr\. corn m .. 1989). "I he analogy !1elween the ... c.: extre mely salty c.:nvironments and the Solnho fe n lag(xlIlnl ba~i n ~ cl1n not he carried ve ry far. hecause salt concentral ions in the

l~a{aeoeIIVi'OI!",el!l mId sedimcll/mioll

Solnhofen waters never reache d very high values and th e evaporating lagoonal waters must have been constantly diluted by an influx of normal marine water. Certainly the salinity of the lagoonal waters nearly always re ma ined below the level o f saturation with respect to the common dissolved sa lls, as there are no evaporite beds in the plattenkalk sequences. This consideratio n lim its the salinity in the bottom waters to a maximum of 117 ppt , wh ich is the approx imate sali nity at which the first salt, gypsum (CaS04.2 H 20), would sta rt to precipitate. Sometimes the traces of sail crystals, lo ng si nce dissolved away a nd infilled by calcite (ma king salt pseudomorphs) are found on beddi ng planes, but they probab ly did not form on the lagoon fl oor and attest o nl y to loca l cond itions within the sediment. A more revea ling modern analogue to these salini ty-stratified lagoonal basins could perhaps be the Orca basin . Gu lf of Mexico (e .g. Trabant & Presley 1978). T hi s lies in a holl ow in the contine ntal she lf unde r fairl y deep water of a bout 1700 m (and in that it is very dissimilar to the relative shallow plattenkalk basi ns). A layer of brine has formed at the botto m of this depression as a result of the dissolutio n of some of the underlying evapori te deposits. During many centuries organic matte r a nd ot he r terrestrial sediments from the North American contine nt have been washed into the stagnati ng basin. Decaying o rgan ic maile r has totally depleted the brine layer of oxygen. In the anox ic and hypersaline waters , not hing, a pllrt from the microbial community, is able to survive , and without macrobe nthic organisms to bioturba te th e sedime nt , a fine lami nation is preserved. The remains of dead pla nk ton ic o rganism s wh ich fell into the basin. such as the tests of calca reous foraminifera and siliceous radiolaria ns, are exquisitely preserved. The occasio nal fragments of seaweed , which fo und the iT way into the sediment , show remarkable preservation in that the details of cell walls are still visible (Kenne tt & Pe nrose 1978). In the Orca basin a nox icity is the most important fllctor in retarding the rate of decay and promoting exceptio nal preservatio n . Hypersalinit y most like ly has a secondary effect in inhibiting the action of the sulphate-reducing bacte ria which are the first bacte riH to oxidize organic matter under anoxic conditions. This is inferred from measure men ts o f the act ivity of these bacteria which is abnormall y low in comparison with those measure ments from exclusive ly a noxic water such as the Black Sea (Wiesen berg et af. 1979), ANOXIC lTV? AnoxiH has developed in the Orca basi n not me re ly because it has been stagnant for a very lo ng time (carbon 14 calculations g ive a value of 79000 years) , but also because it has a high o rgan ic input a nd is too dee p for any ill sill l oxygen production by pho tosynthesis. Using the O rca ba .. in (I S a broad analogue for the pla tte nkalk basins, we can e xam in e the Vltrll)U S fru; to rs upon which the oxygen conte nt depends. Afte r a slOrm-inthu:l'd l' "dtILt1 /t1.: with the

60

Chemistry of the Solnhofen waters

I d hys Ocean , the water body begins with a ce rtain oxygen concent ration, 1I10st probably a pproaching the maximum amount of dissolved oxygen which may be held in waters of normal marine sa linity (5.4 mVI at salin ity 35 ppt and IX "C, see fig. 4.2). During stagnation the oxygen concentration wil l fallon two .It·counts. Firstly , with degradation of organic matter oxygen is used faster than It r.:; an be produced photosynthetically in the lagoon. Secondly, it will fall slowly .I~ the salini ty of the water rises and is able to hold less oxygen in sol utio n (see l(raph o f salinity vs oxygen concentrat ion , fig. 4.2). If a state of almost total anoxia in the boltom waters has been reached, the terrest riall y derived o rga nic matter is no longer broken down by aerobic microorganisms and accumulates 111 the sedi ment. (The presence of organic matter does not necessarily mean that the bottom waters were anoxic. If a large amoun t of organ ic material is huried in a ra pidly accumulati ng sediment , pore wate rs become anox ic because they have lost contact with the overlying bottom wate rs, even though the latter may still be oxic). T he U ppe r Solnhofen Plattenkalk docs not , in general, have a high proport ion of preserved o rga nic carbon, compared to some o ther Illatten kalks. H uckel (1974b) me ntions a measuremen t of 0.2% Iota I organic \." arbon (T OC) from a flinz bed and 0.9% T Oe fro m a fii ule bed. T his contrasts with an average value o f 1.2% fo r micritic limesto nes (while the very organicrich platten kalk o f Hjoula in Lebano n reaches 2.4%). Certain ly, the underl ylIlg beds ( Lower Solnhofen Pl attenkalk, maim zeta 2a at EichsUitt , and pl atten kalk from the maim epsilo n at Painten) and the overlying beds (M orn, heim beds in the Solnho fen-Eichstatt region) are more o rganic rich . T he Ilrganic matter imparts a brown-black ralherthan a grey-yellow colour to fresh \ urfaces and produces a sulphide smell on fract uring. The absence of preserved organic carbon is most like ly indicative o f a very low fa llo ut of organic material 10 the sediment , in othe r words lagoonal productivity was negligible . U nder these circumstances it is less likely that anoxia would be reached in t he time available before mixi ng and exchange o f the lagoona l waters.

,

_,L - -_ __ I

E

__ _

5




- - __

--- ---- ---

,

~ 1

-

HVposaline

Norma l marine

Hypersa li ne _

~~---o,~o----C,"o----C'"OC-'--."OC---~,oc---~,oo---~,~o----~,o Salinily 1%.)

hg. 4.2

Oxygen cQnccrm:lrion in walers of varying salinity demonstrating how. with

l m;rCn\lng sI11inil),.

oxygen ,(Jlubi!ll ), decreases . From Caq>enter (1966) .

Pa/aeocll1 1irOllmel1l1llld se(/ill/elltation

Other argumcnts for a general oxicity of the plattcnka lk waters have also been presented. Yeizer (1977) measured the iron and manganese contents of calcium carbona te from the plattenkalk flinz beds and compared his results with those from other south Ge rman Jurassic deposits. The elements Fe and Mn were introduced into the plattenkalk sediment , bound inside the lattices of clay and oxide minerals. iron predominantly in the valency state III and manganese in IV. Under reducing condit ions these elements gai n electrons from other atoms and acquire a valency state of II. Fe 2 + and Mn2+ are water-soluble ions and move out of their origi nal minerals into solution. Accordingly. calcite which precipi tated from anoxic waters would incorporate 11 relati vely high cont ent o f soluble Fe and Mn, in relation to a calcite prccipitflted frolf' oxic wate rs. Yeizer obtained average values of 100 ppm Fe and 70 ppm Mn for thc conccn tnH ions of these elements in the fli nz carbonate. Thcse results arc consistent with vil lucs obtained fro m carbonates wh ich formed in normal marine rather than those from anoxic env ironments. However. given the probable mode o f dcposition of these carbonates the resul ts arc perhaps not very surprisi ng. If (as described on pp. 65-7), the flin z beds are made out of carbonate formed in normal marine environments, deposited rapidly and lithified early , then they shou ld show a normal marine signature. To test if the bottom wate rs were anoxic the Fe and Mn contents of the well preserved foram inifera (which presumably suffered very little diagenetic exchange) should be measured. As the foraminifera almost certainly lived in the lagoon we might then havc a sounder basis for comment on the oxicity of the lagoonal waters. The two other arguments generally cited in favour of low oxygen concentration in the plattenkalk waters we find similarly inconclusive. Firstly, there is th e argument based o n pyrite. Pyrite (FeS2) forms under anoxic waters or in anoxic sediments. The su lph ide ion is supplied by the bacterial reduction of sulphate, which is derived mainly from seawa ter and the Fe 2 + from the reduction of Fe(lII) minerals. Pyrite is not a common constituent of the Solnhofen PllIltenkalk . although some finely disseminated pyrite does cause a bl ue-green tinge to some of the Solnhofen lithographic stones. But in compnrison wi th known anoxic deposits it is noticeably lacking. There arc a number o f possible explanations to account for the absence of pyrite, some of which ca n be dismissed. The suggest ion that pyrite fo rmed but has since bee n redissolved is addressed by Keupp ( 1977a, b). He argues that this is not the case, because there are no 'etching' struct ures on the ca lcite particles which would be made by an acidic solution which dissolved the pyrite. An explanat ion for the 1I0nforma tion of iron sulphides cou ld be in the inactivity o f sulphatc-reduci ng bacteria because they arc inhibited in the salt y watc rs or because the water... were not anox ic and ot her bacte ria preferentially reduced the organic matter. One of man y ot her explanations is a deficiency of iron. hC(;lH''>C the ... upply o f iron-containing minerals was too low .

62

Chemistry of tile Soll/llofel/ waters

A fina l argument advanced concerning the ox icity of the bottom waters is !lased upon the preservation of the siphunc1e of ammonites. The siphuncular lube is a paper-thin , weak ly phosphatic structure which runs inside the margin Illt he ammon ite shell to the initial chambe r from wh ich the animal commenced )!nlwth. In black shales deposited under anoxic conditions, although the spt.!cimens are crushed, the siphuncle is preserved as a distinct dark-coloured IlIIe. Siphunc1e preservation is normal for ammonites in the Mornsheim beds uve rlyi ng the Solnhofen Plattenka lk but , according to Keupp (1977a, b), the siphuncle in the plattenkalk ammonites is not preserved and this is evide nce iI!!a inst widespread bottom water anoxicity. Others, notably O. Viohl o f the Jura-Museum, Eichstiitt , would disagree with this statement of Keupp 's, d;lIming that siphunc1e preservation in the platten kalk ammonites is not unw mmon and making redundant this argument for plattenkalk waterox icity. In all probability, oxygen concen trations varied from normal marine to ulmost anox ic, this depe nding upo n th e period o f salin ity strati fication and the Icsultant degree of stagnation. OTHER POSSIBILITLES? A different explanation of the hostile conditions prevalent in the lagoona l wlltcrs was put forward by Paul de Buisonje of Amsterdam (1972, 1985). de Jiuisonje proposed thaI an upwelling curren t (presumably off the shelf edge) produced seasonal blooms of coccolithophorids in the lagoonal waters. As I he~c microorgan isms died and fell through the water column , toxi ns produced In the decay of their bodies soo n poisoned the bottom waters . Upwell ing mixed tll l\ pu trid water throughou tlheentire lagoon e nsuring an environment hostile ttl any marine macroorgani sms which the currents had swept into the area. I heir corpses sank to the bottom to be buried by a rain of coccoliths from the nvcrlying waters. de Buisonje's speculations are based on what appear to be vcry insecure I'lcmises. The original idea that the plattenkalk 'lagoon' was subject to an upwelling current is clearly based on the superficial similarity between pa laeot\~'og reriodically, storms stirred up this sediment (rom the seafloor and it heeame suspended in the turbulent water (fi g. 4.3b). The sediment-laden water \\ ;li-hcd over the coral patch reefs and fl ooded into the lagoon , the coa rse r lr"aclioll of the sedi ment , the 'reefal debris' , being dropped after a short ,h\ tance, whilst only the finest particles were transponed the long distance 10 the Solnhofen area . Some reefal organ isms were a lso drawn into the lagoonal waters. Their bodies sank to the lagoon floor to be covered by the more slowly 'C lIllllg, fi ner sed iment (fi~ . 4.3c) . Thh '" u~pen,io n Ihenry' of \cd illlent transport was firs t propou nded by the

Pulaeoe"virOlllllelll and sedimellllltiotl

Soulh

NOIlh

SEA

---

FAULE

Sallnlly-"rlllflcatiOtl wilh hyp ..... lln. bollom ...11 ..... Slow but con tinual doIrpo.lllon 01 I.r>d-d.lved ell,.

a

.c;:--.....:.. ,

REEF

"' ~---

Umlted pch,n!!" 01

IIIgoon,' ....1.' wil h open .... Some Ii ... _bon,I, brought In .

l"gill,,,IKI'" mlc.We llmel'

"

------Mlnne carbon.l. 00...

.

", -------_. ~ e- "'~)"-: brlnglllll 11m. mud '.om ..ound tI., 'HI .

Storm.... uh .,.... Ih. . . ., ",,"m,I, ••• """,Ined In 1M Ilow.

b

c

to

~

,,_.d try

,~",..,t. 51'11",,101' . ......... with .. ~ni IV -""l l f lclllon.

Fig. 4.3 Barthel's theory of deposition of the Solnhofen Plallcnkalk . The carbonate in both ninz and faule is regarded as allochthonous and of shallow marine origin.

Dutch sedimentologist van Stntatcn ( 1971). De position out of suspension explains how the particles rained down vertically upon the dead organ isms and shells. The different pulses of scdimCnI each formed a lamina draping around the fossil. This is an unusua l phcnomenon: us ually potential fo\\i! .. are buried by a unidirCCl ionlll . sediment -Iadcn curre nt which will , nlOolh lhe ..cdimcnt surface over the si te of burill I. In contrast . the depOSitIOn of Ih l' 111;lth:nkalk W:I!o ;{,

Other depositional theories

Ilot accompa ni ed by currents over the basin floor. In the Soln hofen-EichsUitt ,t rea this is clcarly shown by the saucer-shaped Inoceramus and Os/rea shells dnd ammon ite aptychi. 90% of which lie convex down, in an o rientat ion unstable in currents. Some bulbous ammonite she lls such as Aspidoceras and the occasiona l belemnite are embedded vertically in the sediment and this is the position in which they would first have settled. Other indications of very quiet wa ter are the resting impressions found adjacent to the body fossi ls formed as the sin king animal touched down. The blanket cove r of fine sediment also provides an explanation fo r the correlation of beds for many kilometres within somc basins (correlation of Oinz beds between dilTerent basins being a little more problematic). Barthel saw no essential difference between deposition of flin z and mule. rhe ftinz was a concentrated deposit brought into the lagoon by the stronger .. torms, whi lst the faule ca rbonate drifted in on gentle curren ts (too weak to l.'n lrain animal bodics), accumulating slowly over a long period of lime and diluted by land-derived clay (fig. 4.3a). The now well-preserved coccoliths and hent hic foraminifera , which (accordi ng to Barthel) form only a small pe rcentage of the fiiule carbonate. grew in the lagoon at times when the lagoonal wilters (wh ich were never in complete isolation from the sea) were of lowest ..alin ity. Barthel also suggested tha t the foraminifera might have grown on the .. ides of the sponge-algal mou nds and been washed into the more hype rsali ne depths . The time taken to deposit a ftin zlfiiule cou plet can be estimated o nly l'x lrcmely rough ly and by the following calculations: the amount of ammo nite l.'vtJlution between the top and bottom of the Solnhofen Plattenkalk is equivall'll! to about half an ammonite zone and, in Upper Ju rassic rocks, o ne ill11monite zone must last for on average a million years (given the number of ,I tllmonite zones and the tota l time to deposit the Upper Jurassic rocks as ,,, Iculated from radiomctric datcs). Therefore, the Sol nhofen Platte nk alk (U pper + Lower) may have taken around 500 000 years to be deposited. In this lillie some 500-2500 fl inzlfiiule couplets were formed (as calcu latcd by the ilvt::rage th ick ness of the beds and max imum thickness of Ihe seq uence), so 111:11, on average, each ftinz + faule cycle took of I he o rder of 200-1 (X)() yea rs to Ill.' deposited.

Other depositional theories Wh i]I' lh

seA

1.0.1 very much 1I~ if it fell

74

Life ill 'he lagoon

". , .,

-.-~""...,..,

'.

'.

, "

',.

.. , , ..

I·

I..

"

,

'.I "

':

'"

Itg. 5. 1 Trace fossil ; Llllllbricaria ;lIIeslimmr GoJdfuss, Schcrnfeld bei Eichsliilt; nla.~i mum

diameler of fossi l 102 mm ; BSPHGM 1964 XXIII 161.

direct ly from an o rga nism wh ich li ved in the l.. goon rathe r than being b ro ught III fro m elsewhere . A rarer type of LumbriclIrill. referred to as a separate trace fossil species, L. r('clll. probably has a different o rigin . L. reCf(l is an e longate worm-like trace ~h()rte r than thc marc COOlmon form , generally straight or sligh tly curved and l'rc~c rved wholl y or part ly in a phosphatic materi aL sometimes accompanied \1y ~'llcite (fig. 5.2). The phosphatic preservation suggests the coprolite con"'Icd mainly of fish fragment s with a secondary echinoderm compone nt. The Iragmcnl s which it contai ns are relatively large and presumably incompletely du.:wed . T his, together with its short length . suggests its mak er was prob'lbl y a li~ h . Cough balls (Jan icke 1970b) are anot her type of ejecta, but spat o ut o f the nlnul h of an animal. An elliptical ball. 12.5 by 15 em, which contained the ,cnn:cly chewed or digested rcmnants of a 45-50 em long fish is shown in fig. ~ .1, The di,ta'tcfu l morscll1lucrate dimes with their needle or sca le-like leaves. They generally have both male and

Fig. 7.s

Gymnosperm: Spht'llOzlIIIIiles Itlllfolius (Brongnillrl) Sapo.HIII ,

length 192 mm : BSPIIGM AS I Ris II .

Ei c h~tlill

TIlt! fossils

Fig. 7.6 Gymnosperm; 7Gillkgo jfabellallls (Unger). Solnhofcn; leaf lcngth 36 mm; BS PHGM AS V 39.

fe male cones. The woody female cones surround and protect the ovules which occur in pairs at the base of the cone scales and in d ue course give rise to seeds. The male cones are much smaller and expose the pollen sacs. HrachyphyJifllll is the commonest of the plal1cnkalk conifers and its leaf anatomy suggests it is related to the present-day monkcy-puzzle tree (Araucar· iaceae fami ly). A dense covering o f small sca le-like leaves is arranged in a spiral around the twigs and branches (fig. 7.7a). Cross-scctions o f the branches shO\I. an unusual feature; unlike o the r gym nosperms, which have well-developed wood , o nly the cen tral part is lignified. Th is is a feature of many modern halophyte (Le. salt- resistant) plants, so it is assumed that Brachyphyllllm livell on salty soi ls and was a weak-stemmed, shrubby plan t (see also pa laeoecology, p.72). Palaeocyparis is figured in 7.8 (sec also fig. 7.7b). It d iffers from Brachyphy/.

•

,

Fig. 7.7 Gymno~pcrm.'> of the Solnhofen 1'1:l lIcn].,al]., (ll) /lrlllllr/l/n-lllllll t",ig. (hI P(I{(lI!oc)'{Jtlri.f twig; (c) II "hroltl.II/{':' conc: (d) Arillicari,lII n lllc ~(IIIl'

Plants: vascular - gymllosperms

II ~ 7.8 Gymnosperm: Palaeocyparis prin ceps Saporta, Mornsheim beds at Daitingj 1.lIlIl h 490 mm: BSPHGM 1964 XX II 149.

the arrangement o f its leaves on the sma ller branches and twigs. The • llle ·llke Icuvcs are arranged in the shape of a letter X, with a central scale IIIl n undcd by four o thers , alt hough on the larger branches the leaves are Illunltcd in a spiral. as in Brac"ypllyllum. Both BrachYP"yllwlI and Palaeocy1,,"'1 lire relatively comnlon at quarries at Daiting in the west and Kelheim in 1111 ~'['~t. Cont!l! :Ire known only from onc gen us in the Solnhofen Platte nkalk, ""'mJlll.l."ilt.~ (fig. 7.7c), alt hough isolatcd cone sca les assigned to Araucaria (l1~' 7.7d) have 011so heen found . 1111/1 III

III

Thefossils

ANIMALS Invertebrates SPONGES Sponges are amongst the simplest of the metazoans with a bag-l ike body . variously folded in on itself and permeated by numerous canals . The body is supported by small need le-like struts, ca ll ed spicul es , made eithe r of silica or calcium carbon ate, which on occasion may grow into one another to mak e an in terconnected structure. The form of the spicule is very useful in classifying the sponges. Sponge thickets helped trap sediment and build the mounds which both underlie and subdivide the plattenkalk basins (see pp. 26-33). At the time of deposition of the Sol nhofen Plattenkalk most of the lagoona l sponges were dead and so they are rare ly found as fo ssils in the plattenkalk sedime nt. TremadiclyQn , a sponge with an intergrown network of si liceous spicules (fig . 7.9), is found in the reefal localities of the Kelheim area. It almost certain ly grew locally on the reefs, and avalanched into the basin when the sediment pile gave way. The sponge Ammollella is well known both from its body fossils, and from its spicules, which have three mutu ally pe rpendicu lar axes. CNlDARJANS

C nidarians are perhaps best known from the soft-bodied creatures such as the sea-anemone or je ll yfish which demonstrate the radial symmetry. The body has a simple design of either a tubular polyp , or disc-like medusoid. One end has all opening surrounded by a ring of tentacles. T his aperture serves as both mouth and anus and opens into a gut. Duri ng the life cycle there is oft en an alternation of body form. In the polyp stage the closed end of th e elongate body is attached to the substrate and the tentacles dangle freely in the water. In the frccswimming, medusoid stage, the body takes on a much ftatter, umbrella-like shape . The medusoid stage is the sexua l stage o f the life cycle. Jellyfish (scyphozoans)

Scyphozoans are best known in their medusoid stage , and on ly fossil m ed u ~ n have been preserved in the Solnhofen Plattenkalk . In the Scyphozoa . IIw mouth is on the underside of the bell-shaped body, is cross-shaped and

Fig.7.9 Sponge; Trellllldiclyoll .\'1)., Kapfclhc rg hei Ke lhcim ; nw'(imu(Il di amClf;'1 144 mm ; I3SPI1 G M 1975 1 1(1).

11 2

Allimals: inl1erlebrales - cllidarilll1S

, ,

• \

, ,

.,.

(& ~~

J, "

"

..... ,,

•

~ ,{f ,

'l

I1J

Th efossils

surrounded by small protuberances. Internal part itions divide the body into four lobes and there are four o r eight gonads present. Both radial and ring canals occur. The edge of the bell has tentacles, either four or else a mulliple of four in number. The Solnhofen Plattenkalk is renowned for its jellyfish, most of which have come from the quarry region of Ou ngoldi ng- Pfalzpaint where their presence in

four different horizons is recorded. Isolated occu rrences are also known from the Eichstalt area where they are not as well preserved. The impression of the

lower surface of Rhizostom;tes (fig. 7. 10) is illustrated. Often over 50 em in diameter, this is onc of the largest , as well as the commonest of the plattenkll lk

,}

.. • .

l'

,

,

r'

/.,. ,

:~f(

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' r "~

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I

.,

r rUt

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,

/

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if.

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,

:ms. Culcareous colon ial corals were important fra mework 1 "u l the Mluthc rn conl l reef,> , ~ u c h as those now exposed at Neuburg an 1) 11111111 J he polyp.. sc.llkd on a firm 1>UbstraIC, constructing a star-shaped I

Thefossils

basnl plate, with internal divisions numbering usually six (or a multiple thereof) and enclosed by an outer wall. As new polyps arise by budding the coral colony increases in size, its shape being determined by the species that built it. Finr fragm ents of these corals were probably important contributors to the plattenkalk sedi ment, although little un equivocal evidence of this remains . There are also some examples of the preservation of soft-bodied cora ls in tho Solnhofen Platt e nkalk . The species in fig. 7. 11 is tentat ively attributed to tho gorgonians , a group of corals with an eight-rayed symmetry. Gorgonians have a central axis made of proteinaceous material (keratin). which is on ly very rarel),

Fig. 7.1 1 ParI of n pos,ihle gorgoni;.n ; Solnhofen ; f\ISI\ V

Animals: in vertebrates - annelid worms

11hiliLCd , or else ca lcium carbona te . From this ax is arise the polyp-bearing t" ,1I1l:hcs (see fig. 7.12 , drawing of a modern gorgonian). T he branches are Ill-Ilk of small spicules of calcite which normally disintegrate on foss ilization. 111\· ligured specimen resembles the living sea-fan Iridogorgonia in its construc!lun and constitutio n , but is co nsiderabl y larger.

ANNELID WORMS

um: lid worms are typified by the soft , squashable terrestrial earthworm. The t'lllla l characteristics of the group are the segmentation of the body and the lul.ll cra l symmetry. Worms are not exclusively terrest rial ; there are a lso I1 hUIil!.! worms , some of which build themse lves a tube-shaped casing o f calcium I (I hOllate, and these are quite commonly fossilized. These worms, mostly 11,·n the generic name of SerplIla , were common encrusters in reefal environW' ·1l1\ In the Jurassic Period (as indeed they are today). Solnhofen serpulids 11 \ I II (In small pieces of driftwood and ammonite shells wh ich floated into the II uun. t here are also rare examples of free-l ivi ng, soft-bodied mari ne worms, 1111'11 lived either on or in t he plattenkalk sediment. Two genera are recog111 ,11 and the smaller of the two , Clelloscolex , is illust rated (fi g. 7.13). An 1111 pression of the segmented body can be seen together with the bunches of Io d ~ ll cs, or chaetae. It is these structures that give their name to this group of III l1d id worms, the polyehaetes. Chaetae are more prominent on the other \0 11111 , Eill/icites , and there are also remnants of the jaw apparatus. AhOlit 15 different genera and species of so-called Soln hofe n 'worms' have I. I Il named. but most of them are too poorly preserved to provide certain 1.1, 1I11hcations and , in fact , many may be fossil faeces (e.g. Lumbricaria , see Iii ~. I & 5.2 under pa laeoeco logy . pp. 75-6). URVOZOANS

I h, "ufl parts of modern bryozoans show that these animals are closely related thf,.' brachiopods, as both show the same basic construc\tion, of a ring of

401

Fig. 7. 12

Sh,

,

,,

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.

.,,,

~

,

,

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Fig.7.27

, "

DCCllpod crUSI:ICClm: Cycferyoll {Jr0fJlm/1U11 (Schlolhl:llll) , r id,sltll 1: "11111

Icnglh 157 mill : BSI'IIGM AS V 'I.

., '

The fossils

Fig.7.29 Decapod crustacean ; Magil« l(IIim(ma Munster, Schern reld bei Eichstiitt : total length 28 mm ; BSPHGM 1964 XX III 163.

Fig.7.30 Drawing of Magi/a.

Xiphosurulls T he xiphosunms include o nc of the Plattenkalk 's most fam o us fos~ il s. Mesoli· muius (fig. 7.35 and figs. 5.5 & 5.7. p, 78 & 80). a clO'ic n:l;.tll\lC of thc modern horse-shoe crab Umll/us (see fi g. 5.6, p . 80). Tod:l)' //1/1111 /1\ li ve!'. ill .. hallow cOilstal WiHc r). and ;s fou nd in great IlUmbe r.. \111 IIll.' l'll'!l (,'nll't l)f Nnrth

Animals: invertebrates - arthropods

f,

J •

,

'f

.,• •• .

.

!-ig. 7.31 S(omatopod crustacean; SClIlda spinosa Kunth,localilY given as Solnhofen hut more likely to be Kelheimj total1cngth 32 mm; BSPt-l GM AS 1 8 12.

Fig. 7.32

Drawing of SCIf/tit,

The/Qssi/s

Fig.7.33 Barnacle ; Archaeolepas refllenbac/.ui (Oppel). Kelheim: x2: BSPJ-I GM AS 1806.

Ame rica, on some Pacific islands and in Japan. Limu lids bu rrow in the mud to shallow depths in search of sma ll invertebrates for food. They may spend short periods out of the water, and they are part icularly tolerant to " uctuations in the sa li nity or temperat ure of the water. Conseq uently. it is not surprising that they were one of the few an imals sti ll alive when Ihey reached the poisonous lagoon floor , being responsible for some of the famou s spiral tracks or 'deat h marches' which lermi nate with Ihe body at the centre. Some of the fossils are the rem•• ins of moulted exoskeletons r:llher than body fossils. In constrast to some of the crustacea ns, limulid moults are almost impossible to d isti nguish from the body fossils. This is because the shell splits only along its anlerior margi n and the animal slips oul leaving behind the intact, rigid exoskeleton with all ils appendages. More about the trace fossils and activity of M esolimlllwi in the lagoon is given in the chapter on palaeoecology (pp. 77- 9). Almost all fossi l lilllu lids from the western area of Solnhofen and Eichstiitt arc thought to be juveni les. a conclusion reached by comparison with much large r individuals from the reefal areas of Kelhcim. M esolimllllls has a large ca rapace covering the head and front of the thorax, on the top of which lire two kidney-shaped eyes. There arc six pairs of limbs under the front o f Ihe body. the first beari ng large pincers. the next five uscd for wall-into:. th e 1:. ,1 of which has a brush·like cnding and is u,ed to lever the hody 1m WllHh

A llimals: iI/ vertebrates - arth ropods

Jlg.7.34

Barnaclc; A rchaeolC/Jas redrellbachcri (Oppel). Kclhcim ; x2; BSPI-I GM

AS [807. Arach"ids

A rathe r dubious palpigrade is recorded from the Solnhofen Platte nkalk under the name o f Stertlarlhrotl . Palpigrades are relat ives of the spiders, but have legs wh ich li re fair ly long and thin and a body wh ich ta pers to a flagell um. T he body i ... a curious spi ndle sha pe and sho ws neit her the swolle n poste rior nor the division beJween the anterio r and posterior typical of mode rn spiders. Mode rn pli lpigradcs live under sto nes and in crevices and arc neve r greate r than 3 mm III lengt h. The Solnho fen example is nearer20 mm in length , so if it is generally cOlll para bl e to modern palpigmdcs it must have had a very different mode of I1fe. Insects In ...ects arc terrestrial arthropods. some of which have become adapted for a hfc in o r lI round freshwate r. They li re the most diverse gro up of all living .ulIllla ls and they also account for the greatest proportion of species in the Solnhofen Plattenk alk . T he body is divided into head, tho rax and a bdomen lind thcse are stro ngly differentiated from each other. On th e thoracic segme nt.. insects have th ree pa irs o f legs and o ften two sets o f wi ngs, wh ilst the head i.. adorned with a high ly developed set of mouthparts and o ne set of ~e ll ..ory an ten nae. All the types o r insects fossi lis..."d in the Solnhoren beds ure those which possessed ex ternal jaws covered by upper and lower naps o r nltlcle. O nly winged inS(.'Cts arc known. presumably beca use they were hl()wn out in to the Sol nhoren lagoon (!«!C also p. 86). 14 1

Tilt fossils

Fig. 7.35 Xiphosuran chcliccr:tte; Mesolimu/llS IWllchi DCSIll:lrc~t, Max bcrg bci Soln· hofen: width of carap:lce 92 1111ll ; MSA V.

The insects o f Late Jurassic limes were .. Iready fairly advanced and around two-fifth s of the modern o rders .. rc rcpresented in the Solnhofen Plaltcn kalk Characteristics al the genus and particularly at the species level are ofte n difficu lt to determine because the preservation. which is mainly as impression .... is not sufficienllyclea r. Most insects from the Soln hofen Platten kalk have c.:omc from the Eichstall quarries and a few from the Solnhofen area. Mayflies (Ephemeroptera) Mayfli es have nllrrow bod ies, gene rally aro und 3 em III le ngt h , wit h two long tail hairs (fig. 7.36). The front pairofwingl> j, l ~ .ri:c und 1111..' hind pUlr\mu ll In

142

Animals: iI/vertebrates - arthropods

Fig. 7.36 Drawing of modern mayfly .

modern mayflies practically the entire life cycle is spenl in the larval stage when the insects swim and feed on plants. The adult , when hatched . has only days, o r ,'ve n hours, to breed, lay eggs and die. The illustrated HexagetJiles (fig. 7.37) is n typica l example. Dragonflies (OdOllOta)

Dragonflies are also dependent on freshwater for the larval stage. Dragonflies h,lve a head with large, prominent eyes and short antennae. The thorax is , ho rt , wh ilst the abdomen is long and narrow, terminating in two short tail h'lIrs. Two long, but narrow pairs of membranous wi ngs with a vine venat ion oII" l' attached to the thorax. To accommodate the wings, the legs are shifted far Ili the front of the thorax. As the dragonflies are amongst the most aesthetica lly pkasing of the Solnhofen fossils, here two different genera are figured.

1·' 1t 717 Mhyny: I!t:rfl[;t'flili'. \· cdlulo.HH 11"I'11 0 r\'1 A S I ,'IOI(

Hagen. EichsliHI: IOlal length 52.5 mm ;

143

The[ossifs

Libellulium (fig. 7.38) has large, powerful wings which in the illustrated specimen are spread out, a common type of preservation. O n this specimen the delicate venation is enhanced by the precipitat ion of dark iron oxide. In Allisophlebia (fig. 7.39) the wings are folded back over the body. Cockroaches (Blattoidea) The body is dorso·vent rally fl attened and a shield covers the thorax and back of the head. The front wings are large and powerful with a characterist ic venation , whilst the hind wings are small and delicate. In Lithoblatta (fi g. 7.40) the wings are extended. Waler skulers (Pltasmida) The light body and long, splayed·out legs enable the creature to walk o n the water su rface. Chresmoda (fig. 7.41) may have lived in st reams oron the surface of the lagoon o r, as some modern forms , it might have been full y marine.

Fig. 7.38 Dragonfly ; Libell!IJillll1 /ongilal!lll1 wingspan 138 mm ; USPIIGM AS VI 36.

144

(Gcrmar) ,

Fichsllilt ; maximum

AI/imals: iI/vertebrates - arthrOfJo{/.1

Fig.7.39 Dragonfly; Allisophlebia heffe (Hagen). Solnhofen; body lenglh 92 mm ; BSPH GM AS J 804.

I ;i~ .

7.40 Cockroach; Ul/rob{fl/wlil/lllf,hilf/ (Gcnnar) . nSl'llGM AS V 13.

Eieh~'iill; wing~pun ~5 . 5

111m ;

I,


lt feeder.

159

The fossil$

Sea-cucumbers (holothurians) I-Io lothurians are shaped like stretched-out ech ino ids, the mouth at o ne end . usua lly surrounded by tentacles. and the an us at the o ther. Moreover. calcll rcous plates are reduced to small microscopic ossicles, isolated fro m each other lind embedded in the skin . On dea th the ossicles disperse. with the large numbe r making them an important cont ribution to the sed iment. The forms which live on the seafloor arc protected by a leathery ski n. whi lst those wh ich burrowed are more delicately built. As expected, given their effectively soft -bodied nature, the fossil record of ho lothurian body fossils is extreme ly meagre. Thus the Solnhofe n specimens . although of very poor quality, arc of great palaeontologicul importance. PrQloilolotllllria (unillustrated) serves as an exam ple although it is extremely rare . It has a tube-shaped body, around 6 cm in length , with a coarsely granulate outer surface, and one e nd beari ng fi ve to six indistinct. shrivelled tentacles. In some silicified portions o f Soln hofen Plal1e nkalk , the cnicile ossicles o f holot hurians may be replaced by silica nnd so may be dissolved out of the sediment by acid preparation in the laboratory. The small (0.1-0.3 mOl) plates differ in shape and are assigned to various form-genera and form-spec ies. It is recognized that these names are unlikely to correspond to original taxa of holothurians in as much as seve ral ossicles of diffe rent sizes and shapes we re present in the same body.

Vertebrates nSI-I

The fi sh arc by far the most nume rous of the Solnhofen vertebrates, primari ly because a few genera are found in grea t numbers at certain loclliities and strat igraphic leve ls. Most genera arc. however, quite rare and o nly known from a few specimens. The Solnhofe n fish may be conveniently classified into wellknown living groups.

Shark-like cartilaginous fish (Chondrichthyes) This includes th ree separate groups, the sharks and rays and the more di stantly related ratfish . Instead of the heavy bony skeletons of their anccstors, th eir skeletons li re made of the lighter lind more flexibl e mal crial , cartilage. A primitive characteristic present in both the Solnhofen and modern-day cartilaginous fi sh is a body covcring of dcnticles o r skin teeth. Each denticle con tain'i an inner pulp cavity surrounded by dcnline with nn ou teT coa ling o f enamel. The similarity be tween this arrangement and ou r Hwn Il'l' lh IS IllOTe than coincidental .... they 1lI0'i1 probahly cvolVl'd from dtrHrlk, I1l1lhe ~klll coveTIng

Animals: I'ertebrares - fish

the edges of the jaws. In cartilaginous fi sh the water necessa ry for res pira tio n is drawn in through the mouth on the underside , as well as through a pair of circular structures known as spiracles (actually mod ified gi ll slits), and passed out through the remaini ng gill sli ts. which a re usuall y located o n the side o f the animal. Sharks (selachialls) Sha rks move by s inuous curves of the tai l, particularly o f the upper fl a nge, wh ich provides t he forwa rd thrust. The body, being torpedo-sha ped , is ... treaml ined a nd the backbone may be weakl y calcified fo r stre ngth . T he large lateral fin s, o r pectora l fi ns, act like t he wings of a n aeropla ne providing a n upwa rds thrust which counteracts the weight of the head and the te nde ncy o f the a ni mal to nose-di ve. As the o ve rall body de nsity is greate r than the wate r, ... hnrks m ust sti ll keep moving in order to re main nfl oat. The continual "wimming makes the m either act ive hunte rs searching fo r and chnsing prey, o r else suspensio n feeders filte ring the seawnte r. Sha rks' teeth a re usually shar ply pointed but in some forms the re mny be fla ller teeth adapted fo r crushing. The jtlW is not a ttached to the brain case, but is supported behind a mod il1ed gill arch . Fi ve types of sharks are recognized fro m the Soln ho fe n Platte nkalk. T he Galeoidea cont ains the majorit y o f the living sha rks, most o f which a re harm lcss c no ugh , a round a met re in le ngth, nnd inhabiting a rcas o f the wo rld's ocea ns. Ma ny are to be found in the Nort h Sea , whe re t hey fe ed o n fi sh. A mongst the Solnhofe n assemblage Palaeoscyllium (fi g. 7.56), has the typically long and narrow body o f the galeoid sha rks a nd promi nent unpaircd fin s, two dorsal a nd one a nal, without supporting spines. The paired fi ns a re visible, .l lthough o nly in oblique orie ntation , a nd the tip o f one o f the pectora l fi ns pT{llrudes slightly a bove the head. The vertebral colum n is very promine nt , as the ve rtebrae are heavil y calcifi ed . In the fossil, the gill slits a re no lo nge r recogn iza bl e and th e spiracle is known 10 be absent in this genus. T he le ft eye uppears slightly e levated from the surface, whilst the right o ne, wh ich is onl y Just visib le, is pushed to thc top of the skull. T he teeth (no t rea ll y visible in th is photograph, but relat ively frequen tly found de tached from the body) have ,ha rp cutt ing edges wilh o ne mai n po inl , or de nt icle , and som et imes smalle r u\\(}ci.. ted denticles, I' ,\'el/(Iorllill" ( fi g. 7.57) is placed in the Squaloidea . This type of sha rk fl'"ernb lcs a ray in ils dorsa-ventrall y flatte ned body, well suited fo r gliding over the sCHfloo r. Un like a ray, the gill sl its (accompani ed by a la rge spiracle) open on the side of the a nima l a nd the mouth, with sma ll ras ping tee th , is , .tunted al the front o f the head . In addit io n , t he pecto ral fin s a re very large a nd dearly ~c para t ed from the head , the pelvic fins bei ng somewha t sma ll er. As in 11 11 "'lua loid ~ h urk." the re i~ no anal lin and although Ihe spines supporti ng the tlor"ul fin., do occur in rciat cd ~('nera, suc h as Prolospillux (un figurcd), in I lu, tlllar/1I1/1I the)' Ufe nh'l,.l1 1111' vertehrae afe weak ly o~~ ifie d ,

Tile fossils

,• .,..:~.

'J ,

, Fig.7.56 Shark; Pu/ueoscyl/illm formosum Wagner, Solnhofen; length 313 mill; BSPHGM AS I 589.

1111

Animals: vertebmtes - fish

Th e fossils

(

1

.

•

\

"

,/ " "

•

11\.1

Animals: verlebrates -fish

Rays (batoideatls) Rays have aba ndoned the life of -(:ontinual motion needed by most sharks to keep them in mid-water. Instead they Jive on the boltom, camouflaged or sl ightly covered by the sedi ment . T he tail is now no longer o f prime importance in swimming and the muscles have di min ished leaving a whip-like o rgan useful only in navigation. Th e ray moves by passing ripples along the pectoral fins , which arc very large and fused to the head . The mouth is o n the underside of the animal as rays feed on moll uscs lind crustaceans fro m the seafloor which they chew with a battery of sma ll teeth, united in thick rows, or in a chewing palate. A current of water must be supplied to the gills, but to avoid also inhaling sediment, water is drawn in through the spiracles on the top of the head and then ejected through the gill sl its which are always on the lowersideof the body. Fig. 7.58 shows Aellopos, probably a relation of the modern 'guitar fis hes', such as RhinobalUS. It does not show the extreme flatn ess o f some of th e rays and is still torpedo-shaped, like the sharks . although it is widened laterally by its pectoral and pelvic fins. However, the pectoral fin s pass int o the side of the head and the mouth and gills are on the underside making A ellopos a true ray. Dorsal and tail fins are also well developed , the dorsa l fin s each supported by a spi ne. This example is a ma le and is recogn ized by the supporting strut s of the pectora l fin s. Chimaenformes, ratfislr (holoceplraliatls) This third group o f cartilagi nous fish is o nl y distantly related to the sharks and the rays. The upper jaw is fu sed to the skull for extra strength . and hence gives the name to the group, Holocephali . The vertebrae are not additionally "deified . The few teeth are strong and flat ratherthan pointed and are used for l'fushing shellfish. Unlike the sharks and rays the gills are protected by a lobe of .,ki n. which is not dissimilar in construction to that of the bony fish. The first dorsal fi n is suppo rted by a stro ng spine. Ischyodus (fig. 7.59) is II typical Chimaera , o ne of the two types o f Solnhofen ratfish. The head is particularly large and slants forward , whilst the body tapers backwards produci ng a ~tream lined shape. The spine of the first dorsal fin is clearly visible be hind the head whi lst the second dorsal fin is e longate and of small size. Both paired pectoral and pelvic fi ns are large.

Hony fish (Osteichthyes) Hoy-ji'wed fish (act; lIopterygiolls) Of the bony fi sh , the main type found in the Solnhofen Plattenkalk are the rayhnned fi sh, so called bcC;:IUse the fins arc supported by stro ng bo ny rays or hI! . 7.~7 Shark ; 1'\'j'/11lorll/ll/l 1IIIfail HSPIIOM AS Vll 4

(MlIn~lcr), Eich~tatt:

tot:ll length 463 mm ;

,

..

Thefossils

166

Animals: vertebrates -fish

spi nes . There are three groups, two of which , the Chond rostei and the Holostei, may be intermediate in the evol ut ionary road to the modern bony fish , the Teleostei. Cart ilaginous ganoid fis h, or chond rosteans, may be conside red the most primi tive. They are covered by a restrictive coat of thick, enamel-covered scales (ga noid scales). The scales are finnly held together in rigid circles around the body , but these may articulate between one another. T his allows the body 10 bend side to side in snake-like movements, but not to change in shape. The vertebrae remain unossified. The tail and pectoral fins are still requi red to produce an upward thrust for the heavy body (even though the fis h may possess a swim-bladder). T he tail shape remains asymmetrical with the upper flap "upported by the vertebral column and the lower flap fles hy and not covered with sca les. A living relat ive of the chondrostea ns may be the caviar-producing "t urgeon. From the Solnhofen Plattenkalk only one species of the gen us, Coccolepis (unfigu red), is recorded. Bony ganoid fis h, or holosteans, make up most of the Sol nhofen fish. A bony Inner skeleto n takes over the function o f moveme nt regu lation ma king the "trong and heavy scale coating of lesser importance , especially for those fo rms which are rounder in cross-section and less suscepti ble to distortion. Sca les are IIlte rmediate between rhomboid ganoid scales and overlapping round cycloid 'icilles. T he tail is genera lly smaller than in the chond rosteans and in some forms may be almost symmetrical. A swim-bladder enables the fis h to be neutrally buoyant. The mou th region has become shortened and the upper jaw more manoeuvrable. The specimen o f Lepid otes (fig. 7.60) is the most noteworthy o f the Solnhofen holosteans because of its extremely large size , which may reach 2 m in length . Il owever, en tire specimens o f Lepidotes are generally very rare. The t hick, l' l1 amel-covered scales are very prom inent and arc more often found d isarticulatcd from the skeleton, as are the hem ispherical teeth. Gyro fi c/IfIs (fig. 7.61) , belongs to a different grou p within the holosteans, and rese mbles the modern parrot fish. T he body is very th in and disc-shaped , ~ upported in ternally by a delicate network of bones , and the front part is ~' ()vc rcd externa lly by large scales. The fis h manoeuvred by the dorsal , ventral +lI1d symmetrica l caudal fi n as well as the rela tively sma ll paired fins , e nab ling UvrO I1c/IIIS to weave between coral bra nches. Gyrol1chus, as well as its larger relative Gyrot/us , has small round teeth (fig. 7.62), which in the parrot fish are udapted for chewing co ral.

II ~_ 7 . 5~

Ray: Al'lIop Ol IJIIRC,I;(/CIIJ (ThiolliCrc ), Langcnalthc im: width of PCCIOTllt 1\20111111 , BS PIIOM AS I 'i( )(l Copyri ght of Ihal collection. Photograph M. I lrc,~ l cr ,

IlIt~

The fossils

Fig.7.59 R:ltfish; /schyodlls qllensledli Wagner, Eichstiitt ; lengt h 1550 mill : B$PHG M 1954 I 366. Copyright of that collection. Photograph M . Dressler.

, Fig. 7.60 Sl!lllionotifoTIll. h()I\)~tl!a n fhh; tl'pilJOII'1 mUll/lUll WIIJ.tllcr. I Jlllgclllll Ihci m; lenglh 2tl.'i1l 111111. NMSI' P 2~~l!. CUI)) Tij.:hl nf lhul mlN·um

Animals: vcrtebrllfes - fish

1(0

Thefossils

f

I ...•

';4; .

,

Fig. 7.61 Pycnodontiform, holostc.tn fish; GyrollchliS mllcroplerus (Wagner). Blumenberg bei Eichstiin; total1cngth 162 mm : BSPHGM 1939 1 19.

Calurus (fig. 7.63), with a streamlined body and al most sym metrical deeply forked tail, must have been an agile and fa st-moving swi mmer. It 100 possesset\ a thick cove ring of scales. which conceals the vertebral col umn . Juven iles (such as the fi gu red exa mple) sti ll had unossified vertebrae whi ch do not preserve well. On the ventral side the intestinal contents are preserved in phosphate. The related Urocles (unfigured) has a body shape which indicates that it too was a good swim mer and an active predator. It is easily recognizable from ils brushli ke tail. The remains o f smaller fish on which il preyed have been foun d in the gut o f some specimens. Urocles specimens arc peculiar ililhal they have never been found on the beddi ng planes, bu t OCCU I" in .. idc llilll UI1II~ olllllinOI" planes

170

Allimals: vertebrates - fish

Ilg.7.62 Teeth of Gyrodlls cirClllaris Agassiz, Solnhofen?, approximately natural \ II.C:

BSPHGM 1972 XX 137.

of weak ness. From another related fami ly comes Histionotu$ (fig. 7.64), which

hils very large and disti nctly shaped do rsal and caudal fi ns which could have enabled precise naviga tion amongst the reefs. A very different shape is demonstrated by the e longate . pike- like Aspidorhynchlls (fig. 7.65) and related BeJollostomus (fig. 7.66). The scales are large on the flanks but small on the underside and back. When the fish died and started 10 decay the scales tended to stay together and become detached from the underlying tissue as a scale envelope (sec p. 91). The best-preserved specimens arc of moderate size and have been obtained fro m Zandt and Kelheim . Lastly PllOlidophorlls (unfigured) is a holostean which possesses some of the Iwits of the modern bony fish , the Teleostei. In o utli ne Pholidophorus looks ra ther like the modern he rri ng and it may have had a sim ilar life style. The body " spindle-shaped and the tail symmetrical. The sk ull shows the 'adva nced characteristics' of a movable upper jaw. and in a re lated genus t he lower jaw protrudes over the upper. The vertebrae are fully ossified. The armour o f sca les 1\ li ghter , the scales being thin ner. Modern bony fis h , or (eleosts, have a skeleton which is completely ossified li nd supports the body. The scales, which are no lo nger needed for support , are ~ lI1atl and round and constitute the body covering. The body is a simple spindle ... hapc with a deeply forked symmetrical tail. The uppe r jaw is movable. A ll of the Solnhofen fi sh be long to the same group as the modern herring alt hough ~O I1l C reached a size of around a metre. The aesthetically pleasing PachYlliris\ /JpS (fig. 7.67) is a typical exampl e . The sprat-like fis h assigned to the ge nus 1A!IJ/olepides ( fi g. 6.4, pp. 94-5) are undoubted tcleosteans but a re probably luvc nilcs belonging to a variety of separate species. These sma ll , superficially ~ il1lita r. fi sh occurred in swarms such as are preserved in the so-ca lled Solnh orl!tl " Fi,fciiflill z' (sec p. 92), 17 1

The fossils

Animals: L1erlebrates - reptiles

Lobe-finfled fish (crossopterygia lls) T hese are also bon y fish but only distantly related to the Acti nopterygii . The lobe-fi nned fish are very important in vertebra te evol utionary history as one group of these is thought to have given rise to the amphibians by the development o f li mbs and an internal nostril. The coelacanths lack an internal nostril, and have modified the pri mitive lung into an air-filled swim-bladde r with ossified walls. In the Solnhofen coelacanths we call see the characterist ically pecu liar fin structure with the stout base and brush-like end , which , in Palaeozoic forms. evolved into the limbs of modern land animals. Coccoderma (fig. 7.68) is a small coelacanth with a narrow body and pelvic fin s in an ante ri o r position near to the pectora l fins. It has a promi nent scale cove ring and the vertebra l column is unossified. The weakly ossified swi m-bladde r is seen nea r the fron t of the body, visi bly outli ned by a brown st ain .

REPTLLES

The So lnhofen Platten kalk has yielded a varied selection o f repti les whose ha bit ats were eithe r marine or te rrestrial. The primary classification o f the rept il es is based on t he number and position of the skull openings which allow fo r a bulging of muscles when they are relaxed (see fi g. 7.69).

Turtles (chelonians) rhe turtles are anapsid reptiles (fig. 7.69a). In this group the only skull openings are those o f the eyes, nostrils and brain stem. The most obvious Icature of the turtles is their body armour , consisting o f upper and lowershe lls. rhe uppe r shell is made of an intergrowth of ribs, backbone plates and o ther ma rginal plates, and the lower shell of the lower ri b cage and pa rts o f the , houlder girdle. In modern turt les the she ll is overgrown by a horny substance, which is not preserved in these fossils. In some turtles, includi ng some Solnhofen genera , the armour is lightened by leaving ' windows' between the plates presumably to facilitate swimming. However, the Soln hofen turtles l"illlnot have been true open-sea swimmers (as is the example in fig. 7.70) hCl.:ause instead of fin s they had stumpy toed limbs which were evide ntly ilI .tdapted to swim ming. T hey probably lived partly in freshwate r and partly ulong Ihe sea coast. A ll specimens found so far are , at half a metre in le ngth, ,mall in comparison to today's turtles and may be juveniles. Of the fo ur easily Iccognizable genera from the Solnhofen Plattenkalk Ellrystermun is figured (1Igs. 7 .71 & 7.72). T his genus is cha racterized by a shallow shell wi th narrow,

lilt 7 ~III II ;

(I)

Amiiform , h (l l o' I ~'H n lI'h : Ca ll1l"r/l [ /lrm/tll' Ag.lssiz, Schcrnfcld bci EichfIIlll , IIStJll(,M 1%·1 XX III 1:'i 9

10lat Icnglh 1(-):'1

1

Allimals: vertebrates - reptiles

Fig. 7.65

Drawing of AspidorhYllclms.

,., .'

Fig. 7.66 A~pid\lrhvll..:hitnnn . holm,lea n fish ; Be/OIlOS/OIllIl.... IIlII ells/cri Ag,.ssiz. htllt1t.lcngth 345 111111, B:-i !'1 I(;M 11)~7 I J:W

,"

J

/

Thefossils

Fig. 7.67 Teleost fish; PlIchythrissops propufl/s (Wagner). Pain tcn bci Kclhcim ; to tal Icngth 385 mm; BSPHGM 1964 XII I 154.

Fig.7.68 Crossopterygian fish; Coccotiefllw 1IIIlIIIIII Rei' . Kclhclln: towl length 188 OlIO : IlS I'IiGM 1887 VI ~Ol.

,.,,.

Animals: verteb rates - reptiles

" a

c d Fig. 7.69 Side views of reptilian skulls to show various types of temporal opening. (a) No opening: 'anapsid' condition . (b) A lower opening with postorbital and sq uamosal meering above: 'synapsid' condition . (e) An upper opening with postorbital and -.quamosal meeting below: 'parapsirl' condition. (d) Both openings prescnt : 'diapsid' condition. Abbreviations: j , juglll; pa. parictal ; po, postorbital ; sq, squamosal. From Romer (1962).

Fig. 7.70

Drawing of modcrn sea turtle.

marginal 'windows', The lower shell too shows one central and two lateral

openings.

Ichthyosa urs Ichthyosaurs. as most swimming reptiles (apart from thc turtlcs), are parapsid . They have onc additional skull open ing, high on the head and bordcred on the lowcr

·1

,

I·lg,7.76

.

Lizard ; Eid,\'f(lelllfllllrtIHcllrocflcri (Broili) , Wintcrshofbci Eichstiitt; skull

Ic nglh 19 111m ; !lSI'] 10M 11}17 1 1

IRI

The fossils

Rhynchocephalians The beaked lizards fro m the Solnhofen Plattenkalk are of si mila r small size to the true lizards an d the nearest living relative was thought to be Spltenodon, from New Zealand (fig. 1.77). Rhynchocephalia means 'beaked head', which j.., a reference to the shape of the snout. Apart from a few problematic forms the species can be accommodated in two genera; Kallimodotl (fig. 7.78) has a longer and narrower skull than its relative Hom eosaurus (unfig ured), bolh having elonga te skull openings. A small chambe r is often preserved in the apex o f the skull . Kallimodoll was about a quarte r the size of Spizellodoll, with shorter and stouter appendages. The swimmi ng lizards, such as Pleurosaurlls (fig. 7.79) and the smaller Acrosaurus (possibly ajuve nile of Plellrosollrlls), are probably also members of the Rhynchocephalia. Since they all have a beaked skull, elongate body and short legs, the lack of t he lower skull ope ning is probably secondary. The litt lc legs would probably have functio ned o n land but were more o ft en used in the water as stabilizers. With its snake·like form and laterally compressed tai l PleurOS(lllrllS was probably a highly effi cient swimmer in the manner of an eel. T he nasal openings at the top of the head are fairly far back so that the animal needed only to bring its head to just above the water surface in o rde r to

Fig.7.77 Drawing of purported rhynchocep halian from New Zealand Sphello(/oll . approx. length 600 mm.

Fig.7.78 Rhynchoccphnlilln: Kallimmloll Iwlehe/llu (ZlllCI). 1" ll l1tcn !lei Kclhcim, length 175 mm; BSP II GM IRS7 V12.

182

Animols: vertebrates - reptiles

J

I

.

'. '111

I R1

Th e fossils

Fig. 7.79

Rh ynchocephalian; Pleurosallrus gold/llss; Meyer , Sappenfeld bel Eich·

statt ; length 1520 mm ; BSPHGM 1925 11 8. breathe. It also bore six-sided sca les. Plellrosallrlls is found most frequently where there is an accumu lat ion of driftwood which suggests that it lived in a stream and its dead body was washed into the lagoon.

Crocodiles Crocodiles are also diapsid repti les. All the Sol nhofen specimens have an inner nasa l open ing which is not shifted as far back as it is in the modern crocod il es. There are three types of Solnhofen crocodile . The fi rst , exemplified by Alligalorellus (fig. 7.80), is a sma ll crocodile by modern standards at on ly 50 Clll long, and much of the body is covered by an armour of bony phllcs. T he long limbs enabled the animal to hunl on land as well as in the watcr. As is the case for all land animals, the Solnhofen Plattenka lk has yiclded few specimens and they have all come from the Eichstiitt and Kelheilll regions. The second type are coastal crocodiles, but Ihes!; nrc ex tremely rarc. A reconstructed specimen of SUmCOS(lflrtl$ . some 4 In In lenglh . i~ on diosil C !,lIge) Arr hm'OI}/l'ryr litlrogrllphica Meyer; Be rlin speci men , BluI1fl'nhcrg hci Eiclhtil tl ; ICIIj.\lh of ' kill! ~2 111m ; PAMN II UB MB 1880181.4598, cou n tc rJlilll " 'it}!)

,00

The fossils

Fig. 7.95 Rcconstructionor Arclweoplt'ryx based on the model in the British Museum or Natural History.

?flO

Allimals: vertebrates - birds

Fig. 7.96 Archaeopteryx /itllOgraphica Meyer; the isolated feat her. BSPHGM .

2111

8

Conclusions: The Solnhofen Plattenkalk and comparisons to other plattenkalk lagerstatten Summary of the characteristics of the Solnhofen Plattenkalk

The Solnhofen Pl attenkalk contains the bodies of both marine and terrestria l

organisms and must have bee n deposited in a shallow sea. This water was protected from ocean turbulence by an outer barrier of coral reef, and water movement was further restricted by the dead sponge- algal mounds which subdivided the Platten kalk lagoon into sma ller basins. Pulses of fine marine sediment rained down through the quiet waters to the bottom o rlhe basins and these sed iments now form the Hat sheets of Plattenkalk rock. There are tWO lithologies: pure homogeneous limestone beds, varying in thickness from (typically) 1 to 30 em , separa ted from each other by thinner beds of argill aceous limestone wh ich range in thickness from th in laminae to beds 1-3 Clll thick. Chemically, the limestone beds are al most pure calcium carbonate , wi th a low concen tration of organic matter which gives the th ickest beds a bluc-grey colour when freshly broken. The limestone beds may be fin ely laminated on a millimetric scale, but do not usually cleave along these planes. When a limestone slab is removcd from the section it shows distinct upper and lower surface textures, and the fossi ls occur almost exclusively on the underside of a slab. Fossils are hard to find in the Sol nhofen Platten kalk and a day's coll ect ing may produce nothing. Only from the established collections is it possible to compare the relative abundances of different sections o f the fauna and flora although we must bear in mind that only the best preserved and most showy of specimens may be on display; poorly preserved specimens are too oftcl! disca rded, even though they may be a valuable source of info rmation. In Ihe assemblage of marine fossils there are main ly planktonic and small nektonic organisms. Benthos is under-represented . Presumably this is because. in contrast to the inhabitants of the surface waters. they were less likely to he swept away from their normal ha bitats into the Plattenkalk basins. Most of the an im als died as soon as they were plu nged into the Plattenkalk waters so they had no opportunity to make t racc fossil s. The only trace fossils whi ch :Irl' presen t can be att ributed to species whose livi ng relati ves urc pa rticularly resistant to extremes in si,l inity or temperaturc. The he~ t known o f Ihese is the

OIlier plrlllellkulks

horse-shoe crab , Mesolimll/IIS. which crawled disorientatedly in a spira l over the sediment before it succumbed to the hostile cond itions. The bod ies of tcrrestrial organisms also came to be buried in the Plattenkalk sediment. These mcl ude a large and d iverse collection offtying insects as well as reptiles, such as ptcrosaurs and lizards, and that most celebrated bird, Arcliaeolneryx.

Other plattenkalks with exceptionally preserved fossils Of the other plattenkalks which show special preservation, the nearest deposit In Solnhofen , both in ti me and space, is at Nusplingen in the adjacent Swabian Alb (e .g. Temmler 1966). Th is plattenka lk is slight ly younge r, being KimmerIdgian in age . and in a very simila r geological setting. The Nusplingen platte nkalk was also laid down in a hollow between sponge- algal mounds on a hroad shelf, but the sides of the Nusplingen basin were probably slightly , teeper than is gene rall y the case for the Solnhofen area. Piles of sedime nt regularly collapsed downslope depositing slump beds and turbidites. T he re are illso coarse breccia beds in the sequence , which represent mate rial eroded from the sides o f the sponge mou nds. Lithologically , the Nusplingen plattenka lk is lh ffcrent from that o f Sol nhofen. There is no eq uivalent of the thick, pure homogenous limestone beds nor of the intervening marls. The beds are equally Impure (about 10% insoluble residue) a nd cleave on a millimetric scale along mternal laminations. The Nuspli ngen plattenkalk also lacks the regular fine~ra in cd constitu tion of the Solnho fen limestone slabs which make them ,> wluble for lithography; there is more coarse de tritus, sponge spi cules and a cll lour banding due to varying concentrations of organic matter. The distinct ive '>urfa ce textures of the Solnhofen slabs a re nOI present (perhaps, as suggested hy I Hickel 1970, because marly part ings do not separate the limestone beds), und the fossils are not restricted to the underside of limesto ne slabs. However . thc collection of fossil s is similar in conte nt to those from Solnho fen in so far as thcre ; Ire the bodies of both marine and terrestrial organisms, and in the marine cullcction nekton and plankton predominate . Spiral death marches are , howevc r, not known from Nusplingen . Many of these fossi ls are also superbly prc~e rved and provide fu rther insigh ts into L.:"l te Jurassic life . A lt hough frequent ly compared to the Solnhofen Plattenkalk , when exa mIIIC(l closely , the plattenkalk of Sierra de Montsech (e.g. Barale et at. 1984) lIum the Mesozoic sequences o n the sout hern flanks of the Pyrenees is Itt hulogicall y qui te (l issimilllr. Its ilge is Ea rly Cretaceous (Be rriasianV" IHngini an) , M) it is younge r than the Soln hofe n deposit. It must have been la td down in a broadly similar , lagoon -type setting. However, th is lagoon "rollithly lay ve ry clo'>e to the C(l w,1 and , although the plattenkalk itself shows 110 ,t ruClurc, IIldl CitllV I' o f ~· t1t(· r~C Il CC. other units of this sequence are intcr- or ? f)1

ConclusiOlls

supra-tidal. There are stumps in the sequence denoting a slight slope and implying that the plattenkalk was deposited in some kind of protected basinal area. The Montsech plattenkalk is similar to the Soln hofen in that it is a finely laminated limestone but here the lithological comparison ends. Instead of a homogeneous fine-grained texture , the beds are rhythmically layered wit h horizons of ostracods and other bioclastic debris and pieces of torn alga l mat. The plattenkalk may be quite impure, generally reddish-purple in colou r and somet imes colour ba nded due to differences in clay, iron oxide and organic conten t. As at Solnhofen , the beds themselves are a few centimetres in thickness and do not cleave internally . The fossi ls are no t located exclusively on the underside of limestone beds , but split evenly between the under- and overlying slabs. In most respects the collection of fossils preserved in the Montsech plattenkalk is quite different from the Solnhofen assemblage. Although the fish are most numerous (includ ing some sha rk eggs), in Sierra de Montsech many genera are endemic and were probably brackish water in habitants. The rc are also amphibians, bird remains, many insect and some spider fossils, a nd large, well-preserved pieces of plants. Unlike Solnhofen most of these animals lived in the depositional basin in conditions which were probably quite equitable, and there are trace fossils including coprolites. Another important plattenkalk outcrops in the south of France, in the southernmost parts of the French Jura mountains. near to the small village of Ceri n (e.g. Enay & Hess 1970. Gall & Blot 1980). The Ceri n plattenkalk , being Kimmeridgian , is rough ly comparable in age to the Soln hofen and is also in a back-reef position . The Cerin beds are not dissim ilar in colour or text ure to those of Solnhofen. In thickness, entire beds do not reach the 30 cm achieved at Solnhofen , but the bedding on a scale of 0.5-10 cm is roughly com parabl e. The limestone beds are separated by fine marly partings and certainly in the lower part of the sequence there is an internal lamination. The surface of the beds is slightly undulose and some are covered by a pattern suggestive of the erosion of a thin algal mat (G all etal. 1985). This feature , together with the celebrated reptile tracks which adorn some bedding planes and the predominance of small burrows in the upper part of the thicker slabs, suggest episodic sedimentation interrupted by periods of elllersion (or at any rate very shallow water). Fossils are usuall y found on bedding planes although they do not preferentially adhere to the upper surface as is the case at Solnhofen. The assemblage is representative of a littoral environment and is do minated by fi sh, reefal and benthic form s being much commone r than trul y pelagic variet ies. Invertebrates are less common , although theircoprclites and faecal pellets may be quite abundant. The terrestrial reptile fossils (rhynchocephalians, tortoises , crocodiles and pterosaurs) are also an important part of the Cerin collections . The four examples given above - Solnhofen , Nusplingen. Sie rra de Mont sech and Ccrin - arc all plattenkalks deposited in a brondly IHgooni.II se tting. However, plattenkalk wi ll form in other gcographicll i ~c llin g .. gIven the

Other plauenkalks

prerequisite of a protected basin and periodic supply of lime mud. For example , there are also shallow~water deposits wh ich are not associated with reefs, such as depressions in shelves of tectonic origin (e.g. Bear Gulch plattenkalk of Central Montana, USA ; Williams 1983) , or t he plattenkalks of Ilaquel and Hjoula in Lebanon (Huckel 1970). The latter is also an example of ,I plattenkalk deposited under water too deep for benthic microbial/algal mats and the sediment is mostly pelagic. There are also plattenkalks which have fo rmed in lakes , such as the Green River plattenkalks of Colorado, Wyoming and Utah (e.g. Bradley 1931 , 1964). All the examples given here are p l atten ~ kalks wh ich con tain an exceptionally preserved fa una and flora. For an expanded discussion of plattenkalk deposition the reader is referred to the review article by Hemleben & Swinburne (i n press). T hus, to conclude, the Soln hofen Plattenkalk is one of the most celebrated examples of exceptional preservatio n in the foss il record. It is a la ndmark in the history of life and its classic stat us confers an importance and re levance to all who seek to understa nd the gene ral problems of evolution. A nd yet it is only one of a number of plattenkalks, most of wh ich yield some well~preserved fossils, although none approach the diversity and richness of Solnhofen. These lIcposits show significant featu res , but each also has uniq ue characters and owes its overall nat ure to special combinat ions of geology and palaeoenviro nment. T he Solnhofen Plattenkalk owes much o f its foss il fa me to the conti nuing quarryi ng and the sharp eyes of generations of Bavarian workmen. Without A rchaeopteryx, the celebrated death marches of doomed limul ids, and the exquisi tely preserved fl ying reptiles and fish , palaeontology and evolutionary hiology would be much impove rished . T he Solnhofen Platten kalk is a unique deposit , but it is evident t hat a greater investment in to systematic and scie ntific cxcllvations of plattenkalks (and othe r typesof lagerstiitte) wi ll open new vistas mlO the va nished worlds of the past.

•

Appendix Faunal and floral list

The macrofossil list is derived directly from Barthel 's book with only minor corrections and addit ions, and so many names may already be out of date. The microfossils come from two additional sources: the entire list of foraminifera from Groiss (1967) and the calcareous nan nopla nkton from Keupp (1977).

Monerans Cyanobacteria

Protists Coccolithophor ids Bidiscus bellis (Noel) Bidiscus ignofus (Gorka)

Biseulwn ellipticllm (Gorka) Cyc/age/osphaera margereli Noel ElfipsagelosplJaem britannica (Strandne r) EllipsageiosplJaera keftalrempti Grli n Ellipsagelosphaera OIlOW (Bukry) Microstaurus a/ell/one"s;s Keu pp Podorhabdus cylil/dratus Noel Podorhabdus dietzmanlli (Reinhard t) Slaurorhabdus quadriarcullus (Noet) Stephanolithion bigoti Deflandre Watznaueria bamesae (B lack) Watzllulleria deflandrei (Noel) Zellgrhabdot/ls noeli Rood , Hay & Barnard Zellgrhabdotlls snlillllm (Noel) ?Loxolitlllls sp. ?Tetmlitl1l.lS py ramidlls Garde t ?

,

Faunal and floral list

Calcareous nannoplankton of uncertain systematic position Pselldolithl'llphidites multibacillmus Keu pp Pseudolithraphidites quattl/orbacil/atus Keupp Calcispheres Pithonellll gustafson; Bolli Pithonella cf. thayeri Bolli Pithonella piriformis Keupp f'ora miniferans Gaudryina bukowiensis C ushman & O lazewski Marginu/imj distorta Kusnetzowa Nodosaria euglypha Schwager Patellina feifeli ( Paalzow) Quinque/oculillo egmonfetlsis Lloyd R1'I dioiaria indet,

Plants Non-vascular plants I'haeophyles, brown algae Phyllotllllllus

Vascul ar plants (;ym nosper ms I)teridospe rms Cycadopreris Bennettitales Sphellolamites Zmnites Ginkgos Baiera (someti mes known as Furci!o/ium) Gi"kgo Conifers ArtlllCllriO

ArrhrOUIt/rl!l' (prevo Edlillostrolms) IJr{/chVfI" yll /I'" POf(Il'Of 1'//(11/ \

Appendix

Animals Invertebrates Sponges

Ammonella Tremadictyon Cnidarians

Scyphozoans, jellyfish Cannostomites Epiphyllina Eulithota Leptobrachites Quadrimedusina Rhizostomites (includes Myogramma , Hexarhizites & Ephyropsites) Semaeostomites Hydrozoans Acalepha Acraspedites Hydrocraspedota Anthozoans, corals 'Iridogorgo nia' Annelids

Ctenoscolex Eunicites Serpula Bryozoans Brachiopods

Lacuflosella Loboidothyris Septa/iphoria Molluscs

Bivalves Arcomytilus H/lchia (prev. Aucellf/) EOI)eCleli

208

FauI/al and floral list

Inoceramus Liostrea Pinna Solemya

Gastropods Ditremaria Globularia ' Patella' Rissoa Spinigera Cephalopods Squids & cUlllefish Acal1lhoteuthis Celaenoteuthis Geopeltis Kelaeno Leptoteuthis Palaeololigo Plesioteuthis Trachyteutlris ...cc also Engeser & Reitner (1981) & Engeser (1986) Belem nites Duvalia Hibolites Raphibelus (possibly a juvenile of Duvalia) Nauti loids Pseudaganides Ammonites Aspidoceras Glochiceros lithographicum (Oppel) • Glochiceras solenoides (Quenstedt) Gravesia Hybonoticeros Irybonotum (Oppel) Lithacoceras Neochetoceras steraspis (Oppel) Subplanites Sumeria Taramellict!rQS prolithographicum (Fontannes)

Appelldix

Arthropods

Crustaceans Malacostracans Mysidaceaos Elder Frallcocaris Isopods Palaega Urda Decapods Natantians Acanllzochirana Aeger Amrimpos B1aculla Bombllr Bylgia Drobna Dusa He/riga Raul/a Udora Udorella Replantians Cancrinos CycleryolJ Eryma Eryoll Ell/llonia Glyphaea Knebe{ja Magila Mecochirus NodoprosOpOfl Palaeopeillacheles PalaeolJOlycheles Pa/inurina (and juveni le prevo Phyllosoll/a) PselldastaClis Sfelloclliru.f Slomalopods SClllda

210

Faunal alld floral list

Cruslaceans of unknown affinity Allihollema Uuvenile crustacean) Pfllpipes Uuvenile crustacean) Ostracods Cirri pedes, barnacles A rellaeolepas Brachyzap/es (trace fossil)

Chelicerates Xiphosurans Mesolimllius

Arachnids Siernarlitron

Insects Ephemopterans, mayflies Hexageniles

Odonatans , dragonflies

•

Aeschnidillm Aescilnogomphlls Anisophlebia £Ilphaeopsis Isophlebia Ube/llllillln (prev. Cymarophlebia) NamlOgomphus PrOlolindenia Pselldoellphaea Sieleopteron Stellophlebia Tarsophlebill UrogomplllIs

Blattoideans, cockroaches Uthoblatta Megalocerca Progeotrupes

Phasmidans, water skaters Chresmoda En~ifcrans,

locusts & crickets

. £IClllIll' JI/muo/mlca

PYlfIOP" It'I'/11 21 1

Appendix

Heteropterans, bugs & water scorpions Mesobelostomum Mesocorixa Mesonepa NOlonectiles Sphaerodemopsis 'Auchenorryhnchans', cicadas Archepsyche Eocicada Limacodites Prolopsyche Neuropterans, lacewings Archegeles Kalligramma Mesochrysopa Osmylites Coleopterans, beetles Actaea Amarodes Apiaria Cerambycinus Curculionites Eurthyreites Hydrophilus Malmelater No tocupes Omma O ryctites Procalosoma Pseudohydrophillls Pseudothyrea Hymenopterans, bees & wasps Pselldosirex Trichopterans, caddis fli es A rchotalllius MesolalllillS Dipterans, fli es Empjdia ProhirmOfleura see updated review in Ponomarenko (1985)

212

Fauna/ alld (lora/list

Echinoderms Crinoids, sea·lilics Miller;crinus Pterocoma (prev. Amedon) Saccocoma Sofaflocrinites Asteroids, starfish UthaSler Pentaster;a sec also Hess ( 1986) Ophiuroids , brittle slars Geocoma Oplliopsammlls (prev. Ophioctell) Ophillreifa Echinoids, sea-urchins Colfyropsis Hemicidaris Peditw Plrymoped;fla Plegiocidaris Pselldodiadema Rlwbdocidaris Tetragramma Holothurians, sea-cucumbers Acltistrum EocQudina Hemisplraerallllros Priscopedmus Proto/lOlotllll ria Pselldocaudillll Th ee/ia Trace rossils Lllmbricaria illlestiflllm Lumbricaria recta

Appendix

Vertebrates Fish Chondrich thyes Seiach ians , sharks Go/ells (prev. Prisrillrus) Hererodolltus (prev. Paracesrracion) Hexanc!ws (prev. Noridll/lus) Hybodus Orectolobus (prev. Palaeocrossorhinus & Crossorhillus) Pa/aeocarcharias Pa/aeoscyllium PhorcYllis Protospillax (= Belenlllobatis) Pseudorhilla (prev. Squatina) Batoideans, rays Aellopos (prev. Spath obatis) A sterodermus Ch imaeri forms, rattish Chimaeropsis Ischyodus Osteichthyes, bony fish

Acti nopterygii , ray-fi nned fish Chondrosteans, cartilaginous ganoid fis h Coccolepis Holostea ns, bony ganoid fi sh Sem ionotiformes Heterostrophus Lepidotes Pycnodontiformes Eomesodoll Gyrodus Gyronchlls (prcv. Mesodoll ) MeslIIrus Proscinetes (prev. Microdoll ) Am iiformes Asthenocormus Callopterus CmuYlls Clilums (S,robilOlllls) 214

Fa/mal ami floral list

Eurycormus Eusemius Furo (prev. Eugnathlls & Isopholis) Histionows f1ypsocormus I OIlOSCOP"s LiOllesmus Macrosemius Notagogus Ophiopsis Orthocormus Propter/IS Sauropsis Urocles (prev. Megalurus)

Aspidorhynchi fo rmes AspitlorhYllchus 8 elol/ osrom us

Phol idophoriformes O/igopleflr/IS Pholidophorus Pleuropholis

Teleosts, modern bony fi sh Allothrissops A lI(lethalioll Ascalabos Leptolep ides Orthogonikleithrus Pachythrissops Tllflrsis Tlrrissops -,ca also Nybelin (1974) & Arralia (1988)

C rossopterygii, lobe-finned fis h Coccoderma J-/oIOI)haglls (prev. UlIdina) Libys

Kcptilcs Chclonians. turt les Elirystemlwl Itliochely.f P/(l/ycht!ly\ PfI'\;of"lll'l\'\"

215

Appendix

Ichylhyosaurs Leptopterygills Macroprerygius

Plesiosaurs SlretosaurllS (one tooth)

Lacertilians , lizards ArdeosaurIIS BavarisaurllS Eic!lslaettisau rlts Palaeo/acer/a Proaigia/osalt rIIS

Rhynchocephalians AcrOSllUrlIS f!omeosaurus Kallimodon Piocormus Plellrosallrlls

Crocodiles Aeolodon Alligmorel/us AlligalOrium Aloposallrus Dacosallrlls Geosaurus Stelleosaurus

Sa urischian dinosaurs Compsogllathlls

Plerosaurs Aflllrogllalhlls Ctenoclrasma GermanodaclY!us G"alhosaurus PlerodacrylIlS Rlwmphorhy"chus SCa f)hognathus

Uirds Archaeopteryx

216

Bibliography

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51-74. Feduccia , A. & Tordoff, H . B. (1979). Feathers of Archaeopteryx: Asymmetric vanes indicate aerodynamic function. Science , 203 , 1021- 2. Fesefe ldt , K. ( 1962). Schichtenfolge und Lagerung des obere n Weissjura zwischen Solnhofen und der Donau (Siidliche Frankenalb). Erlanger geologisclle Abhandlullg , 46 , 80 pp. Fisher, D. C. (1975a). Evolution and fu nctional morp ho logy of the Xiphosurida . Un publ ished PhD Dissertation , Harvard University. Fisher, D. C. (I975b). Swimming and burrowing in Limllius and Mesolimlllu.f. Fossils and Strata, 4, 28 1- 90. Fliigel, E. & Franz, H . E. (1967). Elektronenmikroskopischer Nachweis von Coccol ithen im Solnhofener Plattenkalk (Oberer Jura) . Neues lllhrbllch fiil" Geologie ulld PaliiolllOlogie. A blzalldlullgell , 127 (3), 245-63 . Forste r, R. ( 1967). Zur Kenntnis natanter jura-Dekapoden. M itreiltmgell der Bayerische Staatssammhmgen fur Paliioflfologie und Izistorische Geologie, 7,

157- 74. Freyberg , B., von ( 1958). Johann Jacob Baiers Oryktographia Norica ncb:;t Supplementen . B. von Freybcrg, H. Hermann & F. Hel ler (eds.). Erlal/ger geologische Ahhlllldlllllg , 29 , 133 pp. Freyberg, 13 ., von ( 1964). Geologie des Weisse n J ura zwh..clt cn Eichsliiu und

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Ne uburg/Do nau (S lidliche Franke nalb) . Erl(lIIger geologische Ablumdlung , 54 , 97 pp. Freyberg, B., von (1968). Ubersicht iiber den Maim de r Altmuhl ·Alb . Erlanger geologische AbhalldlulIg, 70, 37 pp . Freyberg, B. , von (1972). Die erste erdgesc;hichtliche Erforschungsphase Mittelfrankens (1840--1847). Eine Bricfsam mlung lOr Geschichte der Geologie, erliiuterl von B. Freyberg, Erlallger geologische Abhundflmg, 92 , 33 pp. Freyberg, B. , von (1974a). Das geologischc Schrifttum iiber Nordost-Bayern ( 1476--1965). Teill , Bibliographie. Geologica Bavarica, 70 , 476 pp . Frcyberg, B. von (1974b) . Oas geologische Schrifllu m iibe r Nordost-Bayern (1476--1965). Tcil II , Bibliographisches Autoren-Register. Geologica Bavarica, 71 , 177 pp. Frisc;hmann. L. ( 1853). Versllch einer Zusammellstellullg der bisjetzt bekallntell Thier- will Pjlaflzell-Oberreste des lilhographischefl Kalkschie/ers ill Bayem, Programm des bisch6f1. Lyceums Eichstatt. Fritz, P. (1 965). OIl,!/0 16- lsotopenanalysen und Paleotemperaturbestimmun gen an Bele miniten aus de m Schwab. Jura . Geologische RtlfldsclwlI , 54 , 261- 9. Gall , J .-c. , Be rnie r, P., Gai llard , C., Barale, G ., Bourscau, J .- P., Buffe taut , E. & Wenz, S. (1985). Palaeoecologie-in flue nce du developpement d' un voile alga ire sur la sedimentatio n et la taphonomic des calcaires lithographi ques. Exemple du giseme nt de Cerin (Kim meridgian Superieur , Jura meridional fram;a is). Comptes rel1dus de l'academie des sciellces, 302 (series 11) , 8,547'- 52. Ga ll , J .-c. & Blot , J . (1980). Rcmarquables gisements fossi lifcres d'Europe occidentale. [Fine fossilife rous localities in Western Europe. ) Geobios. Memoire speciale, 4, 11 3-75. Gocht. H . ( 1973). Einbettungslage und Erhaltung von Ostracoden-Gehausen im Solnhofener Pl attenk alk (Un ter-Tithon, SW-Deu tschland). [Burial position and preservation of ostracod carapaces in the Soln hofen lithograhic limestone. ] Nelles lahrbllch /iir Geologie lind Puliiol1lologie . MOlllluhe/re, 1973 (4) , 189-206. Goldring, R. & Seilacher, A . (1971). Limulid undertracks and their sedime ntological implications. Neues lahrbllch [iir Ge%gie Imd Pa/iiolliologie. Abhalldlllllgell , 137 (3), 422-42. Go lu bic, S. (1973). The rela tionsh ip between blue-gree n algae and ca rbonate deposit io n. In : N. G. Carr & B. A. Whitton (eds .) , Th e Biology of BllleGreen Algae, Blackwell. London , 434--72 . Guuld, S. J . (1987). The fossil fraud that never was. New ScielllisI , 113,32-6. (;miss, J . T . (1967). Mikrop:llaontologische Untersuchunge n der Solnhofe ner Schil.:hten im Gebict urn Eichstiitt (Slid liche Frankenalb) . Erlanger geolo/-:i\C/u! Ablwl/(lIl11l/-:. 66 . 75 96.

Bibliography Groiss, J. T. (1975). Eine Spurenplatte mit Kouphidm;um (Mesolinwlu,f) walch; (Desmarest, 1822) aud Solnhofen. Geologische Bliilfer fiir Nordost· Bayern , 25 , 80-95. Gumbel, C. W. von (1889). Kurze Erliiulemng ZII dem Blaue /ngo/stadt (No. XV) tier geognostisciJell Karte des Konigreichs Bayem, T. Fischer Verlag, Cassel. Gumbel, C. W. von (1891). Geogllostische BeschreibuIIg des KOlligreichs Bayern , Vierte Abtheilllllg. Geogllostisch Beschreibuflg der Friinkischen Alb (Frankenjura) mit dem anslOpendell friinkischen Keupergebiete, T. Fischer Verlag, Cassel. Gumbel, C. w. von (1894). Geologie von Bayern . II . Geologisclte Beschreibung von Bayern. VIII , T. Fischer Verlag , Cassel. Hadding, A. (1958). Origin ofthe lithographic limestones. KlllIglige FysiogmflSka Siillskapels I Lund Fdrhalldlingar, 28 , 21-32. Hecht , M. K., Ostrom , J . H. , Viohl. G. & Wellnhofer, P. (eds.) (1985). Till' Beginnings of Bire/s, Proceedings of the Imernatiollal Archaeopteryx Conference , 1984. Freunde des Jura-Museums. Eichstiitt. Heller, F. (1959). E in dri tter Ardtaeopteryx-Fund aus den Solnhofener Plattenkalken von Langena ltheim/Mfr. Erlanger geologisclte Abhandhmg, 31 . 25 pp. Hemleben , C. (1977). Autochthone und aullochthone Sedimentanteile in den Solnhofener Plattenkalken. [Autochthonous and allochthonous components in the Solnhofen lithographic limestones.] Neues lahrbuch fiir Geologie und Paliiolltologie. Monmshe/te, 1977 (4), 257-71. Hemleben, C. & Freels, D. (1977). Algen-laminierte und gradiert e Plattenkalke in der Oberkreide Dalmatiens (Jugoslawien). [Algal-lam inated and graded lithographic limestones from the Upper Cretaceous of Dalmatia (Yugoslavia).] Neues laltrbuchfiir Geologie lind Paliiolllologie. Abhandlt/llgell, 154 (J), 61-93. Hem leben, C. and Swi nburne, N. H. M. (in press). Cyclic deposi tion of the plattenkalk facies. In: Cycles alld Events in Siraligraphy, G. Ein sele. W. Ricken & A. Seilacher (eds.), Springer Verlag. Hess, H. (1977). Neubearbeitung des Seesterns Pelllaceras jurassiclls aus den Solnhofener Plallenkalken. [Redescription of the starfish Pemoceras jflrassicus from the Solnhofen lithographic limestone (Lower Tithonian, Bavaria).] Neues lllhrblich fur Geologie und Paliiontologie. Mona/slte/tl!, 1977 (6), 321-30. Hess , H. (1986). Ein Fund des Seesterns Termil1(fstercall cnformis (Quenstedt) aus den Solnhofener Plal1enkalken . Archaeopteryx. ialtreneitschri/l cler Freunde des lura-Museums EicllSliilt, 4. 47- 50. Hirmer. M. (1924). Zur Kenntnis von Cyclll/opteris Zigno . Palaeolllogmphica, 66, 127-62. I-Ioffstctler. R. (1964). Lcs Sauria du J ura~sique superie ur ('\ ~p~cia Ie /llenl Ie'"

Bibliography Gc kkota de Baviere et de Mandchourie. Senckenbergimw Bi%gia, 4.5 , 28 1-324 . Howga te, M. E. ( 1984a). The teeth of Archaeopteryx and a reinterpretation of the E ichstiitt speci men. Zoological Journal of the Linllean Society, 82, 15975. IIowgate, M . E. ( 1984b). On the supposed difference betwee n the teeth of the London and Berlin speci mens o f Archaeopteryx lithographica. Ne lt.es lahrbuch fur Geologie Iwd Palaomologie. MOl/atshefte , 1984 (I I ), 654--B. 71. 89. 203 Regen sbu rg 19 Regell sbu rg embaymen t 18 Ren aissance. usc of plattcnkalk in 4 reptili a n skull openings 173. 177 Rles 19 Ries metcorile eraler 22. 25 ROgli("1g Beds 27.35 Romans. usc of plattenkalk by

Painten 4.33.58.61.68.79. 96 palaeoclimate 71 palaeocurrent Indicators 91-3 palaeoenviroll("1en I rcstricled basin model S-8 subaerial expostlrC hypothesis 56 palaeogeography 24-37 palaeotemperature calculati ons 71 PalaeOZOIc rocks 17. 23 l'UIJil"rsdur!er ('paper shale ') 37 1';Ippen hCl m 13 I'CI'Illian \Cdl!l1 C1l1~ 17 l'ful lP3tnl 59.79 .90.1 14. I :!I 1'11I/lI"'rl",,~

,

ROle "'1ers ..1 Lag .. (red marl byer) 34 sahnity of water (su IInder lagoonal water) Schernfeld 38. 43 .48 Schlo.heim. Baron Friedrich von 10 scul ptures 4 sediment deposit ion ll1eorics 65-70 scdi ment transport by ~at et 72 by ..... ind 72 sedi ments allochthonous 67-8 aut ochthonous 67-8 Scilacher. A. 67-8 ScncEelder, Alois 4-6 SIf,fr o-Erlwlllllrg

2

24

Soht r l ;tke. SIn;1!

37

55

General index

Solnhofcn 2-4,14, 16,20, 21,31 ,33,35,37,38-40, 46,48, 6 1,63,67,68, 70,74,79,91, 140, 142 Solnhofcn Plancnkalk 2,4, 19,27,33,38,49,55,67 and many instanccs in Chaptcr7 characteristics of 202-3 day chemistry of Lower 35,61 Upper 35-7,61, 127 Southern Franconian Alb 1,

sa,

n

2,4

geological hiSlOry of 17-24 palaeoclimate of 71-3 palaeoecology of 71-88 palaeoenvironment of 56 palaeogeography of 24--37 sponge-algal mounds 3, 26-32, 58, 67, 73, 79, 112,

202,203, 204 Spurmschie[er 35 Stone Age, use of plattenkalk in 4,9

strontium in c3rbonat es and pore water 52-4 styloli tes 52 Swabian Alb 203 terrestrial ecosyslems 84-8 Tertiary deposils 17 Tertiary uplift 23 Tethys Ocean 18- 19,25,33, 37.57.61 tiles floor and wall 4 roof 4.7-8 Tithonian 2 palaeogcography 34--7 Toulouse-Lalltrec lithographs 6 trace fossils 35,56.74--5. 202,204 Trenllfmde Kmmme Luge 35,

36 Treuchtlingcn marble 27,28. 124 Triassic sediments 17, 23

turbidites

34,67-8, 203

uses of plancnkalk 4-8 Veiler, J , 62 vencbrate evolution 173 Vinddieisch Land 17.18-19, 25

Viohl. Gun te r

14,63

Wagner. Andreas 10 Walther, Johannes Die Fall/In dt'r Solnhofem!r I'IOlltmkulke 13 WcJlnhofcr. Peter 14 Wchcnburg ]S5 Wickramasinghe. Cha nd ra 13 Winte rshof 23, 49 bndt 153. ]S5, 171 zone fossils 127 Z ...OI/-Aposte/-Ftlstn 3 1

The celebrated Solnhofen Limestone is among the most important foss il deposits because of its astonishing diversity of organisms , many exquisitely preserved. Marine and terrestrial creatures and plants, buried 150 million years ago in soft lagoonal muds, provide a unique glimpse into the true diversity of Jurassic life. Articulated skeletons are preserved, as well as some soft-bodied animals that otherwise would be too delicate to survive fossilisation. Among the highlights are superbly preserved jellyfish , crustaceans, sq uid , fish and flyi ng reptiles. Perhaps most important of all is Archaeopteryx - the celebrated ' missing li nk' which has the skeleton of a dinosaur but is covered in feathers , revealing a crucial evolutionary transition between the reptiles and birds. Solnhofen opens a window into a vanished world , and reveals the unexpected richness of a land and sea teeming with life. This book is a revised and updated translation of Werner Barthel's classic work Solnhofen: Ein Blick in die Erdgeschich/e. In revising the text , Nicola Swinburne and Simon Conway Morris have added a considerable amount of new material whilst preserving the spirit of tbe original book. This is an authoritative account of the geological history , palaeoecology , palaeoenvironment and fossil taxonomy of th is classic location. Not only will it be of great interest to palaeontologists and evolutionary biologists, but it will also be of value to amateur collectors , natural historians and also those with an interest in the history of life. 'This book is an excellent testament to one of the most fa mous limestones in the geologic record. ' American Scientist 'Geologists are now indebted to Nicola Swi nburne and Simon Conway Morris for an excellent general survey .. . of the considerable body of research by German sedimentologists and taphonomists during the last 25 years'. Nature

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