Global bio-events: a critical approach : proceedings of the first international meeting of the IGCP Project 216, ''Global biological events in earth history''

Lecture Notes in Earth Sciences Edited by Somdev Bhattacharji, Gerald M. Friedman, Horst J. Neugebauer and Adolf Seilacher

8 Global Bio-Events A Critical Approach Proceedings of the First International Meeting of the IGCP Project 216: "Global Biological Events in Earth History"

Edited by Otto H. Walliser

Springer-Verlag Berlin Heidelberg NewYork London Paris Tokyo

Editor Prof. Dr. Otto H. Walllser University of GSthngen Institute and Museum for Geology and Palaeontology Goldschmidt-Str. 3, D-3400 Gottingen, FRG

ISBN 3-540-17180-0 Springer-Verlag Berlin Heidelberg New York ISBN 0-38?- 17180-0 Springer-Verlag New York Berlin Heidelberg

This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translatzon,reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under § 54 of the German Copyright Law where copies are made for other than private use, a fee is payable to "Verwertungsgesellschaft Wort", Munich © Springer-Verlag Berhn Heidelberg 1986 Printed in Germany Printing and binding' Druckhaus Beltz, Hemsbach/Bergstr 2132/3140-543210

PREFACE This 5.

volume

contains

ALFRED

Republic

the c o n t r i b u t i o n s

WEGENER-Conference

of Germany,

international biological

21 - 24 May

meeting

events

of

(a) the s t a t e - o f - t h e - a r t

the i n t e r n a t i o n a l intended

During

according

Deutsche

of

of P r o j e c t

carried

out in

216.

a more c r i t i c a l

have b e e n

University,

100

registered.

and 34 f u r t h e r

to the i n t e n t i o n

about

contributions

have b e e n

that the d i s c u s s i o n

s h o u l d prevail.

a few D e v o n i a n events have been d e m o n s t r a t e d

through

(International

Paliontologische

programme

from G S t t i n g e n

at out-

Schiefergebirge.

The c o n f e r e n c e was s p o n s o r e d by

IPA

first

of g l o b a l b i o - e v e n t s .

, 7 key-notes

in the R h e i n i s c h e

UNESCO/IUGS

to the r e c o g n i t i o n

are n e e d e d for f u r t h e r research,

24 c o u n t r i e s

At the f o u r t h day, crops

the

21 6 : " g l o b a l

of their c a u s e s

to a c h i e v e w i t h these d i s c u s s i o n s

from

3 days

discussed,

in r e s p e c t

to the m e m b e r s

participants

Pro j ect

to d i s c u s s

cooperation

to the p r o b l e m s

In a d d i t i o n

I GCP

of new d a t a

(c) the s t r a t e g i e s w h i c h

approach

This c o n f e r e n c e was

the

and to the a n a l y s i s

(b) the p r e s e n t a t i o n

It was

1986.

at the

Federal

in e a r t h h i s t o r y " .

The aim of the c o n f e r e n c e was,

bio-events

w h i c h have b e e n p r e s e n t e d , held in GSttingen,

the f o l l o w i n g

institutions:

IGCP Palaeontological

Association)

Gesellschaft

Forschungsgemeinschaft

IGCP N a t i o n a l

C o m m i t t e e of the F e d e r a l

R e p u b l i c of G e r m a n y

State of N i e d e r s a c h s e n U n i v e r s i t y of G S t t i n g e n The c o n f e r e n c e w o u l d not have b e e n p o s s i b l e w i t h o u t these

institutions.

members during a short

But great

thanks

and s t u d e n t s of my institute, the c o n f e r e n c e . time w i t h o u t

the s u p p o r t

of

is also due to all those staff who s u p p o r t e d me b e f o r e

and

This volume w o u l d not have b e e n p r e p a r e d

the e x t r a o r d i n a r y

e f f o r t of Mrs.

in such

Martina Noltk~mper.

Otto H. W a l l i s e r

CONTENTS

Ill

PREFACE

INTRODUCTION WALLISER, O,H.: The IGCP in earth history" WALLISER,

O.H. : Towards

Project

216

"Global

biological

events

1 a more

critical

approach

to b i o - e v e n t s

LOTTMANN, J., SANDBERG, Ch.A., SCHINDLER, E., WALLISER, O.H. ZIEGLER, W.: Devonian events at the Ense area (Excursion to the R h e i n i s c h e s Schiefergebirge) GENERAL

& 17

ASPECTS

* BOUCOT, A.J.: magnitude,

E c o s t r a t i g r a p h i c c r i t e r i a for e v a l u a t i n g c h a r a c t e r and d u r a t i o n of b i o e v e n t s

* SEPKOSKI, J.J., periodicity * HOLSER, W.T., variations

Jr.:

Global

bioevents

and the q u e s t i o n

PRECAMBRIAN

25 of 47

TO

LOWER

PFLUG, H.D. & REITZ, Proterozoic

E.:

factors 75

changes

in the 95

M.A. : Global

THE

biological

events

in 105

M. : P r e c a m b r i a n - C a m b r i a n TO

boundary

BOUNDARY

biotas

and events

& CHEN

Xu:

A big

109

ORDOVICIAN/SILURIAN

* BARNES, C.R.: The faunal e x t i n c t i o n event n,ear the O r d o v i c i a n Silurian boundary: a c l i m a t i c a l l y induced crisis RONG J i a - y u China

63

CAMBRIAN

Evolutionary

* SOKOLOV, B.S. & FEDONKIN, the late P r e c a m b r i a n

ORDOVICIAN

the

MAGARITZ, M. & WRIGHT, J.: Chemical and isotopic in the world ocean d u r i n g P h a n e r o z o i c time

* WILDE, P. & BERRY, W.B.N.: The role of o c e a n o g r a p h i c in the g e n e r a t i o n of g l o b a l b i o - e v e n t s

BRASIER,

5

event

of latest

Ordovician

121

in

127

~TORCH, P. : O r d o v i c i a n - S i l u r i a n b o u n d a r y (Prague Basin - B a r r a n d i a n area)

event

in Bohemia 133

ERDTMANN, B.-D.: Early O r d o v i c i a n eustatic cycles and their b e a r i n g on p u n c t u a t i o n s in early n e m a t o p h o r i d (planktic) graptolite evolution

139

LINDSTROM,

153

SILURIAN

M.: TO

Global

bio-events

in the O r d o v i c i a n ?

PERMIAN

SCHONLAUB, H.P.: S i g n i f i c a n t g e o l o g i c a l events in the zoic r e c o r d of the Southern Alps (Austrian part)

Paleo-

163

CHLUP~, I. & KUKAL, Z.: R e f l e c t i o n of possible global D e v o n i a n events in the B a r r a n d i a n area, C.S.S.R.

169

BECKER, R. Th.: Ammonoid e v o l u t i o n the " K e l l w a s s e r - e v e n t " - r e v i e w results

181

* C o n t r i b u t i o n which Event C o n f e r e n c e

has been

before, during and after and p r e l i m i n a r y new

presented

as k e y - n o t e

during

the Bio-

Vl

FARSAN, N.M.: Frasnian mass event or c u m u l a t i v e ?

extinction

- a single

catastrophic 189

FEIST, R. & CLARKSON, E.N.K.: E v o l u t i o n of the last c o r y p h i n a e (Trilobita) d u r i n g the Frasnian

Tropfdo199

RACKI, G.: Middle to Upper Devonian b o u n d a r y beds of the Cross Mts: b r a c h i o p o d r e s p o n s e s to eustatic events HLADIL, J., KESSLEROVA, event in Moravia

Z. & FRI~KOVA,

O.:

Holy 203

The K e l l w a s s e r 213

MC GHEE,G.R., Jr., ORTH, Ch.J., QUINTANA, L.R., GILMORE, J.S. & OLSEN, E.J.: Geochemical analyses of the Late Devonian "Kellwasser Event" s t r a t i g r a p h i c horizon at S t e i n b r u c h Schmidt (F.R.G.)

219

KALVODA, J.: Upper Frasnian and Lower T o u r n a i s i a n events e v o l u t i o n of c a l c a r e o u s f o r a m i n i f e r a - close links to climatic changes

225

and

STREEL, M.: Miospore c o r r e l a t i o n b e t w e e n North American, German and Uralian (Udmurtia) deep facies through Appalachian, Irish and Belgian platform and c o n t i n e n t a l facies near the Devonian/Carboniferous boundary

237

INGAVAT-HELMCKE, R. & HELMCKE, D.: Permian f u s u l i n a c e a n of Thailand - event c o n t r o l l e d e v o l u t i o n

241

TRIASSIC

TO

JURASSIC

SCHAFER, P. & FOIS-ERICKSON, E.: Triassic Bryozoa e v o l u t i o n a r y crisis of Paleozoic S t e n o l a e m a t a WHATLEY, R.: ostracoda

faunas

Biological

events

in the e v o l u t i o n

and the 251

of Mesozoic 257

RIEGEL, W. , LOH, H., MAUL, B. & PRAUSS, M. : Effects in a black shale event -- the T o a r c i a n P o s i d o n i a NW Germany

and causes Shale of 267

CRETACEOUS

* KAUFFMAN, E.G.: High r e s o l u t i o n event and global C r e t a c e o u s Bio-events

stratigraphy:

BRUMSACK, H.-J. : Trace metal a c c u m u l a t i o n the C e n o m a n i a n / T u r o n i a n Boundary Event

regional 279

in black

shales

from 337

HILBRECHT, H. , A R T H U R , M . A . & S C H L A N G E R , S.O. : The C e n o m a n i a n Turonian b o u n d a r y event: sedimentary, faunal and g e o c h e m ical c r i t e r i a d e v e l o p e d from s t r a t i g r a p h i c studies in NWGermany

345

DAHMER, D.-D. in Europe

353

~ ERNST,

CRETACEOUS/TERTIARY

G.:

Upper

Cretaceous

Event-Stratigraphy

BOUNDARY

BESSE, J., BUFFETAUT, E., CAPPETTA, H., COURTILLOT, V., JAEGER, J.-J., MONTIGNY, R., RANA, R., SAHNI, A., VANDAMME, D. & VIANEY-LIAUD, M.: The Deccan Trapps (India) and C r e t a c e o u s T e r t i a r y b o u n d a r y events

365

KNOBLOCH, E.: P a l a e o f l o r i s t i c and p a l a e o c l i m a t i c changes in the C r e t a c e o u s and Tertiary periods (Facts, problems and tasks)

371

MART~N-CLOSAS, C. & SERRA-KIEL, J.: Two examples of e v o l u t i o n c o n t r o l l e d by large scale abiotic processes: Eocene nummulitids of the S o u t h - P y r e n e a n Basin and Cretaceous C h a r o p h y t a

VII 375

of Western Europe HANSEN, H.J., GWOZDZ, R., HANSEN, J.M., BROMLEY, R.G. & RASMUSSEN, K.L.: The diachronous C/T plankton extinction in the Danish Basin

381

SIREL, E., DA~ER, Z. & SOZERI, B.: Some biostratigraphic and paleogeographic observations on the Cretaceous/Tertiary boundary in the Haymana Polatlz Region (Central Turkey)

385

WIEDMANN, J.: Macro-invertebrates boundary

397

and the Cretaceous-Tertiary

FEIST, M. (Coordinator): Bio-events in the continental realm during the Cretaceous-Tertiary transition: a multidisciplinary approach

411

TERTIARY

PANTIC, N.: Global Tertiary climatic changes, graphy and phytostratigraphy

paleophytogeo-

SCHLE!CH, H.H.: Reflections upon the changes of local Tertiary herpetofaunas to global events

419 429

THE IGCP PROJECT EVENTS IN EARTH

216 "GLOBAL HISTORY"

~T A

contribution to Project

BIOLOGICAL

GLOBAL BIO EVENTS

u WALLISER,

Otto

H.

*)

A b s t r a c t : The IGCP P r o j e c t 216 is c o n c e r n e d w i t h w o r l d w i d e t r a c e a b l e e x c e p t i o n a l c h a n g e s ( " e v e n t s " ) w i t h i n the b i o s p h e r e . The p r i n c i p l e o b j e c t i v e s of t h i s p r o j e c t , w h i c h n e e d s i n t e r d i s c i p l i n a r y cooperation, are the f o l l o w i n g : (I) S t u d y of t h o s e a b i o t i c ( g e o l o g i c ) p r o c e s s e s and e v e n t s w h i c h c a u s e global biological events (geological level); (2) R e c o n s t r u c t i o n of the o v e r a l l e f f e c t of g l o b a l g e o l o g i c e v e n t s on the b i o s p h e r e or p a r t s of it ( e c o l o g i c a l l e v e l ) ; (3) E v a l u a t i o n of the i n f l u e n c e of g l o b a l e v e n t s on e v o l u t i o n and e v o lutionary mechanisms (evolutionary level); (4) R e f i n i n g of s t r a t i g r a p h i c a l s c a l e s and of c o r r e l a t i o n m e t h o d s by c o m b i n a t i o n of b i o s t r a t i g r a p h y and e v e n t - s t r a t i g r a p h y (chronological level).

The

establishment

global

events

events"

cations

But

possible

Different may

have

which

In that

216

such

available

for

IGCP

-- s u c h

boundary

Project

216

as a b o u t

is the e l a b -

as d e t a i l e d

as e v e r

methods.

In so far,

and less

on hypothetical

be u s e d

a renewed

In the m e a n t i m e

theoretical

to d i s p r o v e in

for

their

own working

the f o l l o w i n g

this

project assump-

discussion

the m e m b e r s

possibilities.

data.

-- has b e e n

modern

should

publi-

of e x t r a - t e r r e s -

and l a b - w o r k

all

are d i s c u s s e d

of the

be as p r e c i s e

biological

the g e o s c i e n c e

of n e w g e o c h e m i c a l

new data

goal

with

the r e l e v a n t

as i m p a c t s

a main

should

"Global

time,

on a c c o u n t a few

are c o n c e r n e d

grew within

Cretaceous/Tertiary

then

or

only

and

which

events,

assumed

and e f f e c t s .

be o p e n

They

about

of the

Project

should

not

hypothesis.

A f e w of

paragraphs.

level global

abiotic

a direct

finally

or e v e n *)

all

to p r o v e

Geological

years.

both

at the

on f i e l d -

processes

latter

been

which

events'*

interest

few

which

The n e w d a t a

hesitate the

Therefore,

based

should

"Rare

to the b i o s p h e r e

by using

is m a i n l y

216

have

of n e w d a t a

causes,

last

microfossils

available. oration

199

on s p e c t a c u l a r

These

in r e s p e c t

projects

the b r o a d

the

stressed

planctic

IGCP

Project

during

bodies.

tions.

two

-- r e f l e c t s

community

tric

--

of

more

effect

then events

lead

events

may

have

different

on the b i o s p h e r e to an e v e n t .

and/or

processes

I n s t i t u t u n d M u s e u m fur G e o l o g i e D-3400 G~ttingen, F.R.G.

or t h e y

We a l s o may und

causes. may

should

overlap

These

trigger

consider

or a m p l i f y

Paliontologie

der

causes

processes, that

each

two other.

Universitit,

Lecture Notes in Earth Sciences, Vol. 8 Global Bio-Events. Edited by O. Waltiser © Springer-Verlag Berlin Heidelberg 1986

The

processes

may

interactions. periodic

exist

very

Furtheron,

or b o t h

logically

be

long

kinds

the may

lasting.

events

of

extremely

of

this

of

events

with

but

and/or

they

not

different

possible

sea-level

manifold

processes

overlap;

Last

In s p i t e cases

complex

may

least

events be

we

reactions

may

very

be

short

should

be

and

episodic

or

or

even

geo-

aware

that

there

effectiveness.

multicausality,

changes

actions,

or/and

we

black

observe

shales

that

play

in m o s t

an

important

role. In o r d e r and

chemistry, with

have

biosphere, whole

i.e.

range

event

chain

other

sedimentology,

or n e t w o r k

of

geosciences,

tectonics,

causes

such

geophysics,

as g e o and

even

of

that

a global

only

one

biotopes

happened,

or or

may

event

affects

a few

special

facies.

Those

then

influence

only

certain

biotopes

or

biotopes

parts

facies

in w h i c h

neighbouring,

of

within

a bio-

primarily

un-

biotopes.

Certainly, changes

complex with

level

the

logical

whole

is n e c e s s a r y .

to c o n s i d e r

affected

the

cooperation

oceanography,

the

the

evaluate a close

astrophysics,

Ecological We

to

processes,

an

within

food-chain.

attention

to

important

ecosystems, Therefore,

the

and

often

i.e.

deciding

role

in

to b i o - e v e n t s ,

play

the

relations

we

shall

pay

in

the

investigation

of

marine

phytoplancton.

realm,

we

observe

respect

to m a j o r within

special

Evolutionary level With

all

quence: mass This the

hitherto in

the

differences one

hand

events,

bio-events,

of

changes sudden

the in

be

strong

those

additional

the

other

especially

hand.

after

a main

"normal"

Chronological Many

the

It w i l l

an e v e n t --

between

and on

of

followed

indicates

evolution.

events

time-span

extinctions, clearly

known

a comparable

there

occur

a certain

influence task

of

evolutionary ones

Already

extinctions,

now

have

it

is

and

been

events

mechanisms by

obvious

the

radiations. have

elaborate

caused

quite

accelerate

to

se-

or e v e n

-- b y

global

project

steps

which

extinctions

interval

which

the

biological

on

the

on

the

global

that

global

evolution.

level

hitherto lithology,

occurrence

discovered

biological

sedimentology

of b l a c k

shales.

and

events

facies,

Often,

these

are

and

connected

for

example

litho-events

with with

occur

a

only

in

a certain

zone

and

part

can

thermore,

of

a biozone,

be u s e d

these

for

events

i.e.

this

a refinement

can

often

be

of

event the

traced

is

shorter

as

chronological

globally

a bio-

scale.

Fur-

within

the

relevant

there

are

world-

facies. Another

aspect

wide

developed

from

those

be u s e d

in

for

If

in

the

the

other

in

the

times.

are

for of

climatic

times

which

on

the

time-specific

the

they

may

be

in

ecological

the

cycles

of

the

may

these

well

differ

can

also

chronological

such

used

Often

side

facies

features,

units.

other

refinement.

refinement

periodic

as

certain

chronological

short-termed,

as w e l l

in

facies,

for

biostratigraphical

long-term

that of

These

and

possibility

lithology

that, of

fact

types

decipherment

latter

smallest

the

same

correlation

A further lays

is

the

as c l i m a t i c

for

the

subdivision

microcycles pattern.

contribute

scale cycles.

are

In c o n t r a s t

to

the

of

documented to

occurrence

bio-events. If we

cal

combine

scales,

graphy, least

event-stratigraphy

as b i o s t r a t i g r a p h y ,

volcanostratigraphy,

. Such

reached

accuracy

correlation scale

a combined

of

role

causes

but

scale

respect

smallest

contains

ordinary

in

and

processes

and

with and

In a d d i t i o n , data,

to

stratigraphi-

lithostrati-

etc.,

us

that

deciffering

led

other

we r e c e i v e

at

h olostratigraphical

resolution

manifold

which

a

provide

time

units.

in r e c o g n i z i n g

and

finally

will

to

time

so m u c h

with

chemostratigraphy,

magnetostratigraphy,

a multistratigraphical,

scale

the

this

such

the they

the

un-

worldwide

holostratigraphical will

global

these

an h i t h e r t o

to

play

events

an e x t r a -

as w e l l

as

events.

Subprojects All

the

above

concentrating to c e r t a i n

mentioned to

questions

certain

problems.

time

From

are

to

effets

on

--

Late

Precambrian

events

logical

innovations

play

--

are

The

been

events

much

Event

which

within

the

the

can

Some

those

of

have

been

event

investigated of

chosen,

are,

by

fossils

to c a u s e s

key-studies

of b i o l o g i c a l

important

be

groups

in r e s p e c t

the

(as c a s e s

will

to c e r t a i n

other

Devonian

and

analysis

be c o m p a r e d

Cretaceous Middle

problems

and

which

as w e l l

for

events,

as

example:

in w h i c h

bio-

role); (as

a case

of b i o l o g i c a l

its

boundaries.

events

a glaziation);

an o v e r a l l

then

younger

an

events

each

boundary

by

within

for

of

biosphere.

triggered

chosen

System, the

the

Ordovician/Silurian

which --

different

the

assumedly

and

levels,

System.

Devonian

as

of with

all

events

corresponding

Of s p e c i a l a case

of

The

within

has

investigations

interest a black

Devonian

a Palaeozoic

are:

shale

in

the

otomari

event

which

caused

only

Famennian the

minor

biotic

boundary

biosphere,

as

such

as

the

Devonian/Carboniferous sea

changes[

a black

the

shale

worldwide

boundary

Kellwasser

event

with

Event

extinction

event

as

at

enormous

the

of b i o h e r m a l

a case

of

Frasnian/

consequences

to

reefs[

short-time

the

change

in

level:

-- C l i m a t i c

changes

taxa

late

in

-- T h e

the

and

the

cesses

for and

an e v e n t changes

-- R e l a t i o n

acme

and

extinction

of

plant

Palaeozoic;

Palaeozoic/Mesozoic

example

appearance,

(Permian/Triassic)

which

has

in c l i m a t e

between

abiotic

eventually and

sea

events

boundary

been

event

caused

by

(as

an

long-term

pro-

level);

and

renewal

and

evolution

of

Jurassic

ammonoids: -- E s t a b l i s h i n g , graphy

in

the

comparison --

The

with

and

Relation

tion

of

extratelluric

these

case

as

cooperation

we the

of

biosphere Even

to if

by

(as

event

met

of

the

within

an e v e n t - s t r a t i -

Mesozoic,

the

and

for

Devonian):

a case

in w h i c h

probably

together);

glaziation

about

a grave

events

and

into

given

investigation

are

the

speciation

and

extinc-

also

of

the

indication

as m a n y

based

of g r e a t

partly

processes of

past

out

with

comparable

Only

with

this

about

details

of

methods

international

the

global

the

reaction

on palaeontological importance

stratigraphical

Furthermore

ecosystems

world.

synof

the

as p o s s i b l e .

to c o n s i d e r

of

carried

the

clear

about

correlation,

disturbance

value

to b e of

is m a i n l y

possible.

long-term

of g r e a t

and

refining

world-wide now

receive

project

results

have

regions

events

these

The

until

the

within

investigations

events

studies

shall

the

expected

sciences.

facts

correlation

example

boundary

quaternary

possible

chronism

than

worldwide an

mammals.

All

itate

equivalent

between

in as m a n y

the

and (as

Cretaceous/Tertiary

telluric --

refining

Cretaceous

the the

with the

in t h e

scales

biosphere, These

long-term

effects

existing

is

also

the

human

will

aspects

much

are

to o u r

facil-

smaller us

with

be

also

interference

In so own

geo-

triggered

might

recent

population.

a contribution

will

provide

which

of

applied

methods

time-intervals project

ecosystems.

by

for and

methods,

far

this

future.

TOWARDS BIO-EVENTS

A

MORE

CRITICAL

APPROACH

A contribution

TO

to Project

GLOBAL UIVES~

WALLISER,

Otto

H.

B I0 EVENTS

W-

*)

A b s t r a c t : G l o b a l b i o - e v e n t s are m a n i f o l d in r e s p e c t to c a u s e s and e x t e n t . T h e r e can be r e c o g n i z e d the f o l l o w i n g p a t t e r n s of g l o b a l b i o - e v e n t s : (]) i n n o v a t i o n - e v e n t s ; (2) r a d i a t i o n - e v e n t s ; (3) s p r e a d i n g - e v e n t s ; (4) e x t i n c t i o n - e v e n t s with stepwise or/and contemporaneous, rapid extinctions. P r o b a b l e c a u s e s for g l o b a l b i o - e v e n t s are (I) c o s m i c c a u s e s , s u c h as (la) c h a n g e s c a u s e d b y the r e v o l u t i o n of the s o l a r ' s s y s t e m w i t h i n the G a l a x y and (Ib) i m p a c t of c o s m i c b o d i e s ; (2) e a r t h - b o r n c a u s e s , n a m e l y (2a) b i o l o g i c a l c a u s e s ( m o s t l y b i o l o g i c a l i n n o v a t i o n s ) and (2b) a b i o t i c = g e o l o g i c a l c a u s e s . The l a t t e r imply, a m o n g o t h e r s , s e a - l e v e l c h a n g e s , c h a n g e s of the p h y s i c a l and c h e m i c a l c o m p o s i t i o n of the o c e a n and the a t m o s p h e r e , c h a n g e s of c l i m a t e , c h a n g e s of o c e a n o g r a p h i c p a r a m e t e r s . I m p a c t s of c o s m i c b o d i e s m a y then h a v e a c a t a s t r o p h i c e f f e c t , if o n e or s e v e r a l of the a f f e c t e d s y s t e m s are a l r e a d y n e a r to i n s t a b l e c o n d i t i o n s , the l a t t e r c a u s e d i n d e p e n d e n t l y b y o t h e r g e o l o g i c p r o c e s s e s . Holostratigraphy, that is the c o m b i n a t i o n of all a v a i l a b l e s t r a t i g r a p h i e s , s u c h as b i o - s t r a t i g r a p h y , event-stratigraphy, chemo-strat i g r a p h y a n d o t h e r s , p r o v i d e us w i t h an e x t r e m e h i g h t i m e - r e s o l u t i o n . In n e x t f u t u r e , the m a i n t a s k in o r d e r to a n s w e r o p e n q u e s t i o n s in r e s p e c t to c a u s e s a n d p r o c e s s e s in c o n n e c t i o n w i t h g l o b a l b i o - e v e n t s , w i l l b e to e l a b o r a t e m o r e and m o r e d e t a i l e d and p r e c i s e d a t a . T h e r e f o r e we need interdisciplinary cooperation.

Introduction Why

are g l o b a l

years,

do

Science; as for

why

short

for

historical

biotic

that

catastrophes, course, tions

of

with

a "higher",

all

why,

such

important

since

might

several

as N a t u r e in s e v e r a l

of e v o l u t i o n a r y

questions

or at l e a s t of e a r t h

science,

i.e.

time,

find

important

history, since

or respect,

and of g e o -

an a n s w e r

Deluge

t h a t the c a t a s t r o p h e s

more

"progressive"

extinctions,

especially

the

by

a

(e.g.G.

were

in the

known

late

have been

and m o r e mass

changes

are w e l l

about

the e v e n t s

to the s c r i p t u r a l

a macro-evolutionary

dominated *)

drastic

pre-Darwin

according

As a r e a c t i o n

journals

enormously

of t h e s e

of o u r

in the

terms:

happenings;

in

understanding

documentation

it w a s b e l i e v e d

In m o d e r n

space

reminiscence. means

or a b i o t i c

There,

fascinating

events

Parts

very beginning

tury.

such

permanent

a better

processes?

Events,

the

have

are g l o b a l

example

logical

events

they

since

18th c e n -

interpreted de C u v i e r ) .

followed "perfect"

extinctions,

as Of

by new creastandard. are c o n n e c t e d

pattern.

to D a r w i n ' s

considerations,

theory, thus

I n s t i t u t u n d M u s e u m fur G e o l o g i e D-3400 GSttingen, F.R.G.

gradualism

denying und

sudden

(or m i c r o - e v o l u t i o n ) changes.

Paliontologie

der

The

latter

Universitit,

Lecture Notes in Earth Sciences, Vol, 8 Global Bio-Events. Edited by O. Walliser © Springer-Verlag Berlin Heidelberg 1986

have been

explained

(e.g.

Lyell).

Ch.

In the e a r l y mental e.g.,

also

cussed

and

Since

the

versus

a certain

such

as t h a t

enormous

by

happenings and m o r e

of n e w

5th

have

available

review

data,

paper of

ontogenetic

changes.

have been

But

at the

have been

same

dis-

again but

versus

such

a few years hypothesis

ideas

this

of m a j o r

of s u c h

in o r d e r

became

assumption

of

bio-events.

hypotheses,

hypotheses

as the m o d e l l i n g

investigations

macro-evo-

discussion

to s e v e r a l

The

only

as p u n c t u a t e d

and the

lead

Nemesis.

k i n d of

phylogenetic,

the o c c u r r e n c e

latter

to s o m e

at the b e g i n n i n g

have

been

spectacular

to r e c e i v e

new

data.

international

But

meeting

to h y p o t h e s e s

then,

especially

at G S t t i n g e n

new

of

the

about

these

hypotheses

ones,

presented

to c o m b i n e

the

authors

]986,

IGCP

which

Project

the c a u s e s became

has

216,

tested

by

the

at the c o n f e r e n c e . standpoint

the

of b i o - e v e n t s

with

The

a short

the c o n f e r e n c e .

the

Each happening term,

we s h o u l d an e v e n t

intermediate

terms

restrict has

times

developments, is i n d e p e n d e n t

initial

processes. event

important

or u l t i m a t e

whether

chemistry,

which

of r e l a t i v e l y

means,

event,

bio-event

in o r d e r

it to e x t r a o r d i n a r y

This

of the q u e s t i o n

several

global

Therefore,

a time-span

respectively.

another,

Global

event,

is an e v e n t .

cases

ocean

to

back

revived,

WEGENER-Conference

attempts

H o w to h a n d l e

not

Since

as w e l l

in r e s p e c t

been discussed.

present

two

new basic

the f i r s t

funda-

Schindewolf,

events,

Simpson)

definitions,

impact

the d a r k d w a r f

basic

state-of-the-art

G.G.

swung

was

phyletic

in r e s p e c t

ALFRED

O.H.

in an e a r l y

biological

record

the f o u r t i e s ,

of g e o l o g i c a l

(e.g.

or/and

of the

stimulating

precise

also been

views

terms

of t h e s e

as also

At the

for r a p i d

micro-evolution.

about

into

of g e o l o g i c a l

again.

transformations besides

gradualism,

periodicity

and

discussed

the p e n d u l u m

on a c c o u n t

The c o m b i n a t i o n

and/or

preferred.

years,

versus

intensified

century,

The o l d d i s c u s s i o n

labels

equilibrium lution

genetic

processes,

subsequently

information

this

gradualistic

a few

of

have been

explanations

more

catastrophism. under

major

Orthogenetic

as c a u s a l

time,

of

changes

emphasized

used

a lack

decades

biological

phase.

by

that

stable

but

this

it is d e t e c t a b l e

etc.)

under

in m o s t

than

weak

event

it is the

the

affected

result

by

of o n e or

Thereby

(such

and

definition

is c a u s e d

worldwide.

consideration

has b e e n

this

or s l o w c h a n g e s

intended

final

event or whether

Then,

shorter

conditions

necessary

to w i s h y - w a s h y

happenings.

is r e m a r k a b l y

whether

the o b j e c t

biosphere,

not

it is

as l i t h o l o g y ,

in a g r e a t

extent

or o n l y

in a s m a l l

The

sector.

term biological

ordinary

event,

shortened:

change

within

the w o r l d

change,

Faunen-

and/or

Floren-Schnitt).

Pattern

of global

It c a n be

taken

a change

of

events

or

as a r u l e

appearance

meters,

such

tology)

or c h e m i c a l

After

it c a u s e

different In-the

emigration another,

some

extinction

of o n l y

If

a higher

rank,

Another is that

aspect

where

only

one

of

such

a species

taxon

categories

or

analysis

or

only

out

bio-

para-

as s e d i m e n -

a region,

one

imply

it n e e d s

or g l o b a l ; or

a few

already

of g l o b a l

important

taxa

the

event-

affects

burst,

are

regional

by

families,

member

extinction

we s h o u l d etc.,

one

be aware consist

that

have been

an e n t i r e

to be p r o v e n

of that

A.J.

eliminated. (a)

aspects,

(this

caused

we c o m e

species.

t a x a of

bio-events, In o r d e r

to

a careful

of the r e l e v a n t

the c r e a t i o n

and e x t r a o r d i n a r y ,

mentioned

of the

of s p e c i e s .

of g l o b a l

Boucot

the

of s p e c i e s .

we need

analysis

that

considerations,

and b a c k g r o u n d - c r e a t i o n cases

total

of o t h e r

in the f o l l o w i n g

in t h o s e

the

possible

is an i m p o r t a n t

subsequent

species

bio-events

a regional

it is w e l l

event-extinctions

unusual

listed. extent

of e c o s y s t e m s

always

Thereby

in the k e y - n o t e

these

for

are

regional

population

replacement

which

important

(b)

bio-events

or r e s t r i c t e d

or r e g i o n a l

level,

a few

it has

all

of g l o b a l

is i n c l u d e d

from

and

is r e a l l y

Considering

abiotic

or b a s i n - w i d e

questions

bio-event

the

as g e n e r a , which

background-

words:

the d i s -

In so far,

as w e l l

to

contemporaneously).

species,

the s p e c i e s

as p o i n t e d

other

lead

a change

of the

it a f f e c t

local

species.

triggers

is e s p e c i a l l y

cause-effect system,

one

aspect,

distinguish

did

of taxa,

of b a c k g r o u n d - e x t i n c t i o n

This

With

here

local

are v e r y

a global

or m o r e

food-chain,

I mention

such

a change

These

of

already

In our d e f i n i t i o n ,

general

parameters

connected.

in a s e c t i o n

categories

if t h e y

immigration

of o n e

an e v e n t

mass-mortality

existed

population

with

petrography

is it o n l y

categories

even

and

an e x t r a -

floral

of b i o - e v e n t s .

(e.g.

which

that

by

(including

extinction?

following,

stratigraphy

of a b i o t i c

is a l w a y s

in all r e g i o n s ;

mass

bio-event

not d i s c u s s e d ,

for

and/or

composition.

recognized

pattern

Thereby,

means

indirectly

its d i m e n s i o n :

is it s y n c h r o n o u s

a change

This

lithology

having

to e x a m i n e

or d i d

that

of t a x a

can b e r e c o g n i z e d as

stands

(faunal

bio-events

the e c o s y s t e m .

appearance

bio-event,

of o r g a n i s m s

eco-

volume).

or e x t i n c t i o n by

to the

the e v e n t . following

(I)

Innovation-event. in the w o r l d

Biological

of b i o t a .

As an e x a m p l e

tain cepha!opodes , leading ammonoids.

It has

an i m p o r t a n t times & M.A.

vation

and

of DNA,

existing

Radiation-event. certain short

taxa

Pflug

M. B r a s i e r ~

this

volume).

tion-event.

example

for

also

occur

Nevertheless,

occurs

innovation

and

of c e r -

innovations

B.S.

The f i r s t

Sokolov

foundation

structures, totally

affected

played

Phanerozoic

& E. Reitz,

the

inno-

new worlds

also

the p r e -

system. in Fig.

I, r a d i a t i o n s

to e x t i n c t i o n - e v e n t s , implies

of the

an u n u s u a l

mostly high

of

after

percentage

taxa.

Radiation-events

radiation

out b e l o w

subsequently

coiling

the

opened

and

feedback

A radiation-event

of

etc.,

changes

domination

of m e t a z o a n

influenced

to m a j o r

and e a r l y

H.D.

As p o i n t e d

interval.

of c r e a t i o n s

in P r e c a m b r i a n

in a d y n a m i c

appear

serve

of

surely

ecosystems

may

lead

that biological

mineralization,

This

may

long-termed

the c r e a t i o n

of s k e l e t a l

of e c o s y s t e m s .

(2)

especially

contributions

Fedonkin

of p r o t e i n ,

to the

to b e s u g g e s t e d

role

(compare

innovations

this

subsequently

in some

cases

only

after

kind

of d e p e n d e n c i e s

of m a m m a l s

and

such

to a b i o l o g i c a l

an i n n o v a t i o n - c a u s e d

an e x t i n c t i o n - e v e n t .

their

main

exists

innova-

The most

in the

radiation

evident

late

Triassic

the

K/T mass

after

extinction-event. (3)

Spreading-event. out

of s p e c i e s

very

difficult

spreading

out

one

from

hand,

example wide

may

Under

to c l e a r l y of n e w

unusual

serve

lites

of

(compare

also

latest

Silurian, Devonian

this

life

but

The m o s t

especially

when

example

at the

Event,

thus

a certain

event,

shows

well

the

groups several

of o r g a n i s m s .

extinctions

steps,

even

In o t h e r if t h e s e

to m a s s

may

within

cases, happen

Event occur

As an world-

due

among

to t h e

grapto-

this

Event

volume)

in the

at the b e g i n n i n g

occurrence.

are the

extinction-

extinctions, or

sudden

at the b e g i n n i n g

burst

bio-events

Kellwasser

hand.

Erdtmann~

its f i r s t

it is

on the

occurs

example

of B.-D.

a sudden

also

is o b v i o u s l y

transgrediens

increase

producing a real catastrophe

out

Another

after

evident

they

which

transgression

habit.

the

but

spreading

that

bio-events

on the o t h e r

spreading

After

species

events,

O/S

between

flabelliforme,

the g l o b a l

of s u d d e n

the n o r m a l ,

in t i m e s

the c o n t r i b u t i o n

System,

(4) E x t i n c t i o n - e v e n t s .

During

distinguish

Its u n u s u a l

uniformis.

of

the

with

kinds

to be m e n t i o n e d ,

spreading-events

a floating

is M o n o q r a p t u s

several

it has

"Dictyonema"

Ordovician.

innovation

term But

species

at and t o g e t h e r

of the

this

are u n i t e d .

the

in a v e r y

the a f f e c t e d

the e x t i n c t i o n s in a r e l a t i v e l y

as for

K/T Event. short

time,

ecosystems may short

occur time

or in

a

(compare the contributions Fois-Erickson; wise the

ones same

may

this occur

event.

Cretaceous

regressions,

by

several

change

of a l b e d o , for

the

has

been

Another The

above

al e v e n t s events

At the

change

of J.

mentioned

radiation. ent groups,

the The

may be explained

might

within

the f o l l o w i n g

step-

one

and

and e n d have been

of b i o t o p e s ,

parameters,etc.). are g i v e n

et al.,

the

in

in this

extinction

i.e.

& E.

and

during

loss

Hansen mass

event,

Event

then

iridium-event.

(see b e l o w ) . reflect

extinction

be c o m p o s e d

ones

K/T E v e n t

H.J.

stepwise

for e x a m p l e ,

sequence:

may happen by

at the

and of

of b i o - e v e n t s Thus,

Schifer

extinctions (e.g.

and of o c e a n i c

extinctions

this

of P.

of o r g a n i s m s

stepwise

of c l i m a t e ,

Kellwasser

patterns

latter

which

which

and

catastrophic

the e n d - P a l a e o z o i c

b y an a d d i t i o n a l

is the

following

groups serve

Wiedmann

to the e v o l u t i o n .

show

Becker

types,

of the r e g r e s s i o n s

K/T boundary,

superimposed example

may

during

effects

stepwise

the c o n t r i b u t i o n s volume).

Th.

Both

in d i f f e r e n t

As e x a m p l e s

caused

Examples

of R.

volume).

of

hypothesis

nomismogenesis l

influence

-- s h o r t

several

a certain

the

all k n o w n

major

interval

radiations

time-span. (compare

~

regularity

1):

< evolut, rate > select, stress
>

niches

-I-

s.L

nomismogenesis

> increase
' ' ' ' n to pmoles

these e l e m e n t s

are b o u n d

in the living o r g a n i s m

in

proportion.

In oxic waters, into

w a t e r column:

or not s u f f i c i e n t -

of d i s s o l v e d m a t e r i a l

of m a r i n e biomass,

and n u t r i e n t

[(CH20)I06(NH3)I6H3PO4] ~Ni

ments b a c k

acti-

to a t m o s p h e r i c c o n t e n t

removal

and the m a r i n e b i o l o g i c a l

use of an ideal m o l e of a p h y t o p l a n k t o n ,

In the p h o t i c

to

interactions within marine

introduction

as pH or redox p o t e n t i a l .

of sea w a t e r

their p r o p e r

The d i s s o l v e d c o n s t i t u e n t s

(4) n o n - c o n s e r v a t i v e

include

b a s e d on the b u l k c o m p o s i t i o n

productivity

interaction

(composition related

(composition related and

volcanoes,

cipitation,

toplankton.

and sea floor.

the n u t r i e n t

influence biological

content

to the c o m p o -

assigned.

Processes that

food webs.

that r e l a t e

and the c h e m i c a l

( c o m p o s i t i o n b u f f e r e d c h i e f l y by b i o l o g i c a l

in sea water);

to be

are those

of the ocean

(I) c o n s e r v a t i v e

(3) n o n - n u t r i e n t gases

and s o l u b i l i t y

would

processes

content

upon death,

sea w a t e r

oxidative decay matter

as the o r g a n i c

particle

sinks

releases through

the elethe

78

2

~

{

~ °s

~

z

~

z

d

0 O,.20

~dSd

-

~

V

~s~

Z

V

7

Z

o~oN

~ ~

F i g u r e 3. Periodic table from Quinby-Hunt & Turekian

of the d i s s o l v e d (1983).

elements

in

o

o

sea-water.

Data

79

M o l e P h y t o p l a n k t o n + ]38 02 --> 106 C O 2 + 132 H 2 0 + H N O 3 + H 3 P O 4 + T r a c e M e t a l s Thus,

dissolved

in the

photic

remain

constant

zone~

In a n o x i c nitrite

nutrients

rapidly

increase

in d e e p w a t e r

waters,

as the

and n u t r i e n t - r e l a t e d

(Fig.

initially,

oxidant

with

metals

to the b a s e

are

of the

at a m i n i m u m

pycnocline,

then

4).

oxygen

ammonia

is r e p l a c e d

being

the

by nitrate

reduced

nitrogen

and end

pro-

duct: M o l e P h y t o p l a n k t o n + 53 N O _ - --~ 106 C O 2 + 16 NH 3 + 53 NH 4 + H 3 P O 4 + T r a c e M e t a l s Trace

metals

would

a m m o n i u m ion w a s + NH 4 + Trace

build

Metals

As the c o n c e n t r a t i o n tions

of

up

available

in s u c h

--~

Metal

of a m m o n i a

the d i s s o l v e d

waters

to p r o d u c e

trace

until

soluble

sufficient

metal

ammonia

as

amine:

Amines

in this

metals,

zone

most

far

of

exceeds

the m e t a l s

the c o n c e n t r a would

be c o m -

plexed. As a n o x i c i t y fate becomes

increases

upon

consumption

However,

dissociated

H2S would

complexes

react

amine column

of d i s s o l v e d

Trace

metals

( N i , Z n , C u , C d , H g , etc.)

The d e c a y

of p h y t o p l a n k t o n

reduction

of n i t r o g e n

to s u l f i d e

with

to the

constant

Black

Sea

by

state,

transfer

50 to

100

ductivity given

of

the r e l e a s e d

sulfides]

thus

metals

depleting

+ S 2- --~ anoxic

to a m m o n i a , of t r a c e

waters

waters

Sulfides

would

then reduction metals

metals

rich

Metal

proceed

of s u l f a t e

as s u l f i d e s .

would

decrease

in s u l f i d e s

by

Thus

con-

or r e m a i n

as s e e n

in the

5).

and c h e m i c a l

from depth

changes

occur.

locations,

stratification

significant

of w a t e r

meters, could

into

dissolved

in a n o x i c

any

the g e n e r a l

various

sul-

interactions

the d e n s i t y

a steady

compounds

(Fig.

many

particulate

sinking

situation,

depth

Physical-chemical With

nitrite,

metals:

the p r e c i p i t a t i o n

oxic

with

to f o r m

with

and m e t a l

modern

and

thermodynamically

the water

trary

of n i t r a t e

favored oxidant: 2_ M o l e P h y t o p l a n k t o n + 53 SO 4 --~ HCO 3 + 53 H2S + 16 NH 3 + H 3 P O 4 + T r a c e M e t a l s

106

the

name

effect into

in s p e c i e s

The

process

of u p w e l l i n g

and f r o m

various

of

the o c e a n

on m a r i n e

the p h o t i c

zone.

composition

of v e r t i c a l and

occurs

depths.

biota

generally

must Here,

in

the u p p e r

and p h y t o p l a n k t o n

advection at v a r i o u s

in

be c a u s e d

of w a t e r rates,

prois at

80

F i g u r e 4a. Oxic C o n d i t i o n s in m o d e r n open ocean: N u t r i e n t s after B a i n b r i d g e (1979) and Q u i n b y - H u n t et al. (1981).

I

I

I

I

Pacific Ocean Near Hawaiian Islands 200

400

600

800

100(

120( 0

10

20

30

50

40

~;:~:i:::::~:i.1 Nitrate Concentration (pmol/kg)

0 ~

1.0 2.0 3.0 4.0 Phosphate Concentration (pmol/kg)

5.0

8604-002

F i g u r e 4b. Oxic C o n d i t i o n s in m o d e r n open ocean: Trace m e t a l s d a t a after B r u l a n d (1980).

0,~-~----]----.i & ~ , 200

I

t

I • ......

o--,--.-,

\~ '\\

400

A

%. \.

Cadmium Copper Nickel

Zinc

\ \!

600

•

\

!l !l ~J

O

I

800

"\ \

"\

x

71 :!

?1

40 &

1000

h'

: to.,, .A A

2000

3000 I

0

~

I

l

100

200

....

I

300

,,

400

t \

i

i

500

600

700

Concentration (ngtkg) 8604-005

81

10 -s Moles Phosphorus 0

2

4

6

Figure 5. Anoxic C o n d i t i o n s in the Black Sea. Phosphorous after Fonselius (1974). Trace Metals after Brewer and Spenser (1974

8

+200 - ~ " ~ " 0 -f~

t

-soo -I

'\

\ a

\

I -1000 ]

t

\

I

L

-I~oo J~

{

~ ;z

1

-2000

'

0

I

L

I

i

2

4

6

8

10

t 0 -8 Moles

Types

of u p w e l l i n g

Three

main

time

scales:

t y p e s of u p w e l l i n g

may be i d e n t i f i e d

functioning

with

various

I. P l a n e t a r y A. d i v e r g e n c e depth

of water

relatively

shallow

10 to 80 m e t e r s / m o n t h B. d i s p l a c e m e n t

Peng

1982,

p.

known but II.

& Reed

renewal

resident

or one mixing

of deep water

presumably

much

of the wind

stress.

Effective Vertical

rise:

1963). of water

time

masses

is about

at source.

1000 years

Slow:

(Broecker

&

cycle.

to the surface less

caused by winds.

(Wilde

than one modern

& Berry mixing

1984).

Rate un-

cycle.

Regional

A. seasonal lar

Present

20)

transport

as a f u n c t i o n

(Wooster

by c o n t i n u a l

millimeters/day.

C. o v e r t u r n

by Ekman

Ekman

to p l a n e t a r y

B. o f f - s h o r e

(high pressure Ekman

advection

near

the equator).

III.

Local

A. o b s t r u c t i o n & Hogg

1980)

B. c l o s e d

moving

by a s e a m o u n t

through

eddy c i r c u l a t i o n

West

(Wooster

by current

of current

and uplift

off

upwelling

Coast). & Reid off

Characteristics

shore

producing

simi-

1963). (Peru-Chile

Taylor

Current

columns

(Owens

continuity.

producing

low p r e s s u r e

(Robinson

1983).

82

C. B e r n o u l i

uplift

D. b r e a k i n g

internal

Duration The

through

constricted

waves

straits

(Stommel

in s h a l l o w

water

(Hall

different

water

into

et

& Pao

al.

1983).

1971).

of u p w e l l i n g

injection

of c h e m i c a l l y

have

affects

tion

of the u p w e l l i n g

for modern

related

coastal

be s i g n i f i c a n t l y

to v o l u m e event.

upwelling.

greater

of w a t e r ,

Fig.

depth

6 indicates

The v o l u m e s

than

those

the

of

the

the

layer

source,

volumes

generated

produced

surface

by

by

and d u r a latitude

overturn

in c o a s t a l

would

would

upwelling.

Changes in density stratification Destabilization

of

into

zone

ges

the p h o t i c

the

in the c h e m i s t r y

tion

of c h e m i s t r y

the o x i d i z e d the s u r f a c e through

of

surface layer

(2) o p e n i n g

of o v e r t u r n

I shows

thinnest water tion

the s t a b i l i t y

glacial

the b a s e

from warm

displacement bility

climates

pycnocline.

from

sources

Fig. of

some

by normal

for

geophysical

the c h e m i s t r y

life

of f o r m e r l y

or w h e n

processes

or b y

for

and

a change

are

the

of

a transi-

initiated

oceans

(3)

demonstra-

for displacement

can b e the

taxa,

niches,

conditions

zone

in

change

taxa.

climates

stabile

of

of w a t e r s

restricted

affected

to the p h o t i c

Overturn

event

chan-

a func-

of e x i s t i n g

for v a r i o u s

the m o s t

be

evolutionary

range

the e n e r g e t i c s

climates.

physical

the

would

deep-water rapid

to the c h e m i s t r y

instituting

the o c e a n

7 gives

of

bring

inducing

Such changes

to n e u t r a l l y

the p y c n o c l i n e

to c o o l e r

by

of

produces

would

for

in r e l a t i o n

Modification

or e x p a n d i n g

food

which

potential

ocean.

or r e s t r i c t i n g

of n e w n i c h e s

Generation

column

water

the p o t e n t i a l

restricting

that

the u p p e r

layers.

has

or

ting

water

the g r e a t e s t

of the u p w e l l e d

(I) d e s t r o y i n g

opening

Table

oceanic

has

by massive

at l o w

sta-

in the m a j o r

source

of w a t e r . As s h o w n by climatic

in Fig. change

graphically. latitudes the c o l d

edge

relatively der b u t

of

less

the

saline

the

(HLWb)

becomes

the

high

high

latitude denser

of

maximum

water

forms

latitude water

where

(SMW)

the

deep

can be caused

water

water

water

At the o n s e t fall

below

salinity

shifts

temperature

Thus,

overlain of

a cold

5 ° C,

maximum,

geo-

at h i g h

is f o r m e d

in m i d - l a t i t u d e s .

the b o t t o m

water.

from

the

the d e n s e s t

temperatures

than water

of o v e r t u r n

the d e n s e s t

climates

5 ° C (HLWa),

salinity salty

occurrence

source

non-glacial

n o t go b e l o w

warm but

where

an o b v i o u s

where

During

does

8,

at this

by colperiod

this

water

displacing

83 F i g u r e 6. Coastal Upwelling Volumes-Eastern Atlantic Ocean - a f t e r W o o s t e r & R e i d (1963). •

.- C~ Q::, 60

, , , ,

0. . C.lCO

, ,

.

.

,"~ :~

OCO

, ,

_ North 40

~

Pacific

I 20

0

20

40 ~. South Pacific 60

% .

J I , I t 30 20 t0

0

/110 -

Transport

30

20

10

0 -10

(10 ~kg s~1 m-1) B604-006

12

I

11

J

3: "6

7+ foe

I

-

F i g u r e 7. Potential E n e r g y of D i s p l a c e m e n t for Overturn Conditions preceding cold climates. From Wilde & B e r r y (1984).

0.929

10

0.755

9

0.052

I :E

0.44

% II

p0.299

6

5

_ t

~ 200

400

600

800

1000

1200

I

0.592

1400

Rise in Meters Contours in E R G S t g r a m assuming I cm 3 weighs 1 gram

84

UNIT

Z

(thickmess

I08E

(m -I )

I04N/2~ 2

m)

(Hz)

/N X 60

I04U

(min.)

(sec. -I )

Non-Glacial

CLIMATE P r e - or Postglacial

Glacial (Modern)

2900

1400

900

43

84

523

3

4.5

51

36.8

6

8.9

11 14.8 21

T a b l e I. S t a b i l i t y in m a i n p y c n o c l i n e for v a r i o u s C l i m a t e s - a f t e r W i l d e & B e r r y (184, p. 148). Z = m o d e l t h i c k n e s s of p y c n o c l i n e - after W i l d e & B e r r y (1982); E = s t a t i c s t a b i l i t y ( H e s s e l b e r g & S v e r d r u p 1915); N = buoyancy frequency (Gill 1982, p. 5 1 - 5 2 ) ; b = v e r t i c a l s h e a r (Munk 1966, p. 710). S t a b i l i t y i n c r e a s e s w i t h h i g h e r v a l u e s .

F i g u r e 8. Global Oceanog r a p h i c M o d e l as a f u n c t i o n of c l i m a t e - m o d i f i e d a f t e r W i l d e & B e r r y (1984). D a s h e d lines encompass major water mass sources. EQW STW SMW SSW MW

Equatorial Water Sub Tropical Water Salinity Maximum Water Shelf Sea Water Metiderranean Water (model f o r h i g h s a l i n i t y s h e l f sea) HLW = High Latitude Water

Density as Sigma-T 3O

2s/~?,

= = = = =

Climates: Warm Climate (light pattern) - high latitude (HLWa) s e a t e m peratures greater than 5 ° C and SMW densest and d e e p w a t e r . Pre or P o s t Glacial Climate (light p a t t e r n and i n t e r m e d i a t e pattern) - high latitude (HLWb) s e a t e m p e r a t u r e l e s s t h a n 5°C b u t no s i g n i f i c a n t s e a ice f o r m a t i o n .

2O

P :3 ,¢,,.,

$

15

c~

E }-~D

i -2 32

33

HLWc 34 35

36

37

38

Salinity %o 8604-004

85

that ice

water will

(HLWc) the

in

an o v e r t u r n .

form

as

in

seen

mates

the

tudes

warms

high

ditions

potential

For

above

the

of

near

of

the

the

tropical

the

deep

tudes.

For

low

pics as

due

the

to

of

less

of

a particular

of

the

dense

at

the

occurs

The

deep

likely

water due

to w a r m e r

oceans

at h i g h

becomes

denser

major

is

of

latithan

prior

forming, by

water

as

can

the the

latitude

be

overturn During

only

slightly

climates,

would

be

of

case

the

the

high

in

tro-

areas

wind

overturn

lati-

the

case,

significant

effects

the

at m i d - l a t i -

to b o t h

In e i t h e r

have

high

the

the

and

volume

masses. the

such

produced

first

due

simple

rate

of

climate. At

global

model

water

Thus,

deep

divergence

be

found

Such

where

areas

or d i v e r g e n c e

ceased

or

other the

could

decimated

water

has

areas

biota be

the

will

where

or

of

locations,

water.

has

dis-

formation

principally

formation

extent

locations

However,

such

overturn

At c o n v e r g e n c e s

overturn.

displacement

The

geographic

the

and

of

rising

downsinking

areas

For

volume

elsewhere.

maximum,

would

usual

density.

a function

mass

of

of g r e a t e r

occur

surface

refuges the

water.

the

the

cold

effect

which the

on

thus

water

repopulate

of

con-

change.

formed

mid-latitude

a special

the

salinity with

would

the

sup-

from

an

already

become where

would

"homes"

during

as be

the

water

be

for

relataxa

overturn.

overturn

effect of

to

cli-

climatic

equator.

was

at

overturn,

of

the

cessation

deep

water

climates,

surface

thermal

and

varies

into

of d e e p

Accordingly,

toward

from

by w a t e r

insignificant.

unaffected

function

the be

depends

density,

mix

tively

of

the

an o c e a n i c

climates

experience

upward

sources,

where

later

is

sea

advection.

overturn

sources

are

Types

would

overturn

masses

that

The

vertical an

volumetrically

by

would

or d o w n s i n k i n g

will

or

caused

warm

oceans,

or d i l u t e d

overturn

denser

return

direction

cold

subtropical

cold

mid-latitude

pressed

of

latitudes

during

for

the

source.

of

the

latitude

convergence

onset

of

water

various

on

upwelling

enhanced

here,

even

again

source

required

boundaries

In g e n e r a l ,

high

and

water

upwellings,

the

the

overturn

the

to

cussed

as

water

eventually,

overturn Upon

occur

geographic

high

latitudes.

eastern

coastal

no

column.

would

depend

by

an o v e r t u r n the

the

from

as

and

continues,

to p r o d u c e

However

mid-latitude

mid-latitude

colder

tudes

trend

water

stability

overturn,

of

the

would

caused

generated

direction

ocean.

overturn

in

zero

overturn

displaces be

of

5 ° C and

change

an o v e r t u r n

would

cold

quantities

modern

for

the

water.

to

sequence

source

the

stability

latitude Due

sufficient

in

increasing

If

on b i o t a the

in

the

chemistry

photic of

the

zone deep

of

such

water

overturns

displaced

to

would the

be

a

surface.

88

Oxic

overturn:

metals ents

causes

upwelling problem toxic

area,

effects.

followed

to the

water

for

1982)

showed

the

fact

by

Experimental

increased

on o t h e r

potent

and

in o t h e r

occur

turn, warm due

higher

effects during

potentially

lack

of

a toxic

available

chelation red-tide

may be some

taxa.

thermal

effect

on o x y g e n - b r e a t h i n g

Anoxic

overturn:

organisms

with

the

effect

potential

(1984)

and W i l d e

waters

based

Benthic

on t h e

organisms

of v a r i o u s

their

shifts

upward

the c h e m i c a l

habitat.

species

ocean.

rate

could

& Caperon

to i n c r e a s e d to u p w e l l e d 1963).

found

phowater

Secondary

with

"red

algae

mortality

pro-

in f i s h

mortalities

level

of n u t r i e n t

of

an o x i c

over-

climate

to a

initially

of o v e r t u r n

available have

is c o l d

by s u r f a c e

of the s o u r c e However,

was

slow

nutrients

catastrophic and well

not b e

could effects

oxygenated,

any s i g n i f i c a n t

would

would

change

during

differ

as the

a chance

of

oxidants

invasion

However,

water

of

in

anoxic of h o w

et al.

anoxic (Fig. anoxic

of the w a t e r

relatively

to c o n v e r t

the r i s e .

Wilde

or d i s p l a c e m e n t ,

During

dwelling

as a f u n c t i o n

zonation

the c o m p o s i t i o n

upwelling

became

equilibria.

dissolved

most

of u p w e l l i n g .

may have

of the u p w e l l e d

a chemical

of d o m i n a n t

be effected

to n e a r - s u r f a c e

if the w a t e r

the c h e m i c a l

proposed

In s i m p l e

composition

(Terry

for

of d e e p

a glacial

there would

organisms

change

have

at the r a t e

This

responses

sequence

would

compositions

lying

equilibrium

(1986)

If the

conditioned

productivity

additional

thought,

viability

effect

from

primary

but

is n o t

they

Such mass

a threshold

have

in the

and o t h e r

mass

1963).

may

a

organisms.

on s u c h

would

cause

As this w a t e r

the d e p t h

well-oxygenated

depth

the r e d o x

with

the

effects,

Inhibitory

increase

the m o d e r n

metals.

been

if the less

that

programs

The major

reduce

type bloom.

on n o n - p h o t o s y n t h e s i z i n g there

occur.

then

on the

have

trace nutri-

waters

lag,

chelators

a transition

would

The

as G o n y a u l a x

suggesting

toxic

noted

in P r o v a s o l i

upwelling

(Provasoli

the b l o o m

occur

to p e r m i t

produce

organisms

of

metals

inhibitory

of m e t a l

such

is

not properly

Conversion

if i n g e s t e d

would

climate~ to the

which

before

which

enough

during

This

trace

studies

initially

Dinoflagellates

neurotoxins

at the p e a k

enrichment

was

photosynthesis

(1971)

was

and

additional

in s u r f a c e

(see d i s c u s s i o n

organisms

conditions.

duce

of

al.

water

addition

productivity

et

Also,

Energy

nutrients of

is a v a i l a b l e .

productivity.

the d e e p

organisms. Thermal

increased

of n u t r i e n t s

increased

deep water

water, upwelling

upwelling

Barber

rise

that

Ocean that

tosynthesis.

tide"

for light

However,

initial

use b y p h o t o s y n t h e t i c

effects

In g e n e r a l ,

conditions

tropics.

inhibitory

immediately

has

well-oxygenated

depth.

favorable

in the

or

due

with

occurs when sufficient

Peru-Chile

was

In p r e s e n t

increase

each

9). waters over-

zone

slow upwelling,

to the

local

during

overturn

redox the

87

Brochert (1965)

Wilde (1986)

Oceanic Concentration

i

OXIC

Rhodes and Morse (1971)

Demaison et aL (1983)

AEROBIC

OXICAEROBIC

02 Zone DYSAEROBIC