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Journal of the Geological Society, London, Vol. 147, 1990, pp. 663-674, 3 figs 2 tables. Printed in Northern Ireland

An oceanic model for lithological and faunal changes tested on the Silurian
record

Abstrack A model for oceanic cycles is presented. The cycles are expressed as changes between more
humid low latitude and cooler high latitude climates (P episodes), and dryer low latitude and warmer
high latitude climates (S episodes). The cyclicity may be self-regulating through changes in the CO,
storage capacity of the deep ocean due to temperature changes and through changes in the oceanic
input and output of dissolved carbonate. The model permits over 30 predictions regarding observable
changes in both deepwaterand shelflithologies and variations in the abundance, diversity and
extinction rate of planktic and benthic faunas. Sedimentary changes involve fluctuations in the rate of
clay deposition and carbonate production, the advance and retreat of carbonate deposition on the
shelf edge, the timing of oolite, reef, and black shale formation, and changes between oxic and anoxic
deepwater. Fauna1effects include both iterative evolution and intermittently widespread taxa
('Lazarus taxa'). These predictions agree wellwith changes observed inwellknown sequence of
Wenlock and Ludlow age, as well as changes seen in selected older and younger sequences.

Profound cyclic lithological andcorrelatedfaunalchanges model makes a number of predictions (Table 2) which are
took place during the Silurian Period. These can be compared with changes observed in successions of Silurian
exemplified
by changes in the conodontfaunas
and age.
associated sediments (see Jeppsson 1987a). Some intervals
are characterized by high conodontdiversity,often with Effects in the deeper parts of the ocean
platform-equipped taxa.
Associated lithologies show a Global oceanographyresponds to variationsin climate
strong decrease in carbonate deposition in outer carbonate (Wilde & Berry 1984, 1986), one feature being a difference
platform settings, an increase in clay deposition in the in theformation of thedensestdeep-water.When high
carbonateareas,
and less anoxic tomore oxygenated latitude water temperature remains above 5 "C, the densest
deep shelf and/or oceanic sediments(Jeppsson 1975, water is formed at the cold edge of the salinity maximum at
1985, 1987a,b). The two
best
examples are
the mid-latitudes(Schopf1980). This deep water is poor in
Pterospathognathodus amorphognathoides Chron which oxygen, and anoxic or euxinic conditions may develop
spans the Llandovery-Wenlock boundary,
and
the (Bralower & Thierstein 1984). Duringcolderintervals,
Polygnathoides siluricus Chron (the Cardiola event of when high latitude waters are below 5 "C, these waters are
Schonlaub1986) in late earlyLudfordiantimes ( = late the densest and they form the oceanic bottom water which is
Leintwardinian of previous usage). Such intervals are rich in oxygen (Wilde & Berry 1986).
designated P episodes (Fig. 1, Table 1). Anintermediate
state
seems possible. If the high
Other intervals are characterized by less diverse latitiude cool waterdoesnotbecomedenseenough to
conodontfaunas,andincludetaxaabsent or rare in P replace the most salinity-dense water, a three-layered Ocean
episodes. Associated lithologies indicateastrong increase might form. In this situation the temperature-dense water
and seaward spread of shallow water carbonate production, would form the middle layer, as for example in the present
widespread reef formation and perhaps minor development day Mediterranean out-flow which is not sufficiently dense
of oolites. In the oceans, euxinic conditions are indicated by to form thebottomwaterbutdoes forman intermediate
such sediments as alum shales (Jaeger 1976, 1977; Jeppsson layer (Armi et al. 1988). The resulting sediments would thus
1987b). The best examples of these conditions are probably remain anoxic in thedeepestareas,butbecome well
in the early Sheinwoodian, the late Homerian and the late oxygenated in more shallow areas,forexampleonthe
Ludfordian ( = Whitcliffian of previous usage). Such deeperparts of the shelf andperhaps in evendeeper
intervals are identified as S episodes (Fig. 2 , Table 1). environments.
Lithological and faunal changes in the Silurian have been Many of the deepersequences
remained euxinic
interpreted previously as being caused by transgressions or throughout Silurian times into the Early Devonian (Jaeger
regressions (Schonlaub 1986), butamodel whichinvolves 1976), except during one P episode (Fig. 1; Table 2 ) which
only habitat displacement is inadequate. We need to explain started during late early Ludfordian times. Such a tripartite
faunal changes thatare widespread endugh to preclude them division of this timeinterval is represented in alarge
being due merely to latitudinal shifts inresponse tothe number of sequences in central,westernandsouthern
movement of climatic belts, or to oceanward or shoreward Europe,northernAfricaandMiddleEast(Jaeger 1976),
migrations in response to regressions and transgressions. and in the Polar Urals, Pai-Khoi, and Central Asia (Koren
The model introduced here is based on changes between 1987). Some of these sequences were deposited below the
intervals when the climate is either wetter at low latitudes carbonate compensation depth
(compareJaeger 1977).
and colder at high latitudes, and intervals when the climate Other P episodes are less well represented in deep water
is dryer at low latitudes and warmer at high latitudes. The sequences. A P episode can be recognized in the Gorstian of

663
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664 L .J E P P S S O N

P-STATE
LATITUDES
HIGH Initially: absorption of CO2 by the
ocean from the atmosphere
< + 5OC I l l / /
High planktic production
supports I LUz I
Diverse, abundant faunas l LOW 1
Freshening, nutrients, clay.etc.

Intense

(Black shales)

Temperature stratified ocean

carbonate
deposition
Thermal contraction

Cold oxic water, rich in CO2

Fig. 1. Some characteristics of an oceanic
e. P shale, grey limestone etc.

southeastern Sardinia (compare Jaeger 1976), and the late water.Eitherathree-layeredoceandeveloped, or the
Llandovery-earliest Wenlock P episode is recorded as a spread of theoxygenatedtemperature-densewater was
limestoneunit in thegraptoliteshalesequence of the restricted by a topographical barrier from reaching all the
northern Yukon (Jackson & Lenz 1962). oceans.
The Silurian stratigraphical record agrees with a model The warm (S) episodesarerecorded, as expected, by
in which, during the earliest episodes discussed above, the deep-watereuxinicsedimentsexcept
that
oxygenated
waters
were
not cold enough to cause complete
a conditionscontinuedthroughparts of thePridoliEpoch
replacement of all the oxygen-poor, salinity-dense bottom after the late early Ludfordian P episode.
Table 1. Some characteristics typical of P and S episodes. Not all episodes will conform rigidly to all these criteria because of the influence of
local conditions

P episode S episode

more Low latitudes: humid dryer

er High latitudes: warmer
Rate of weathering: high low
Carbonates: pure
platform
Carbonate margin: retreat expansion
Reef growth: scarce extensive
Black shale formation preferentially
in: shallow water deep water
Deep oxic anoxic
Typical deep water
sediments:
limestone,
grey-green
shale, alum shale with graptolites
bioturbated sediments
Characteristic low latitude, shallow
condensation;
limestone;
argillaceous
sediments:
water reefs, pure limestone; initially
ore minerals oolites
Sea level: higher somewhat somewhat lower
Sea clear
Water depth preference of benthic
communities inphotic
the
zone:
shallower deeper
Most importantsource of nutrients:
terrestrial
weathering; upwelling benthic recycling
ction: Benthic low high
ction: Planktic high low
Faunas depending on the planktic
; abundant; production: specialists sparse; few species; specialists in
with narrow niches low productivity environments
Evolution: iterative low rate of evolution
leneck taxa:Characteristic Lazarus’ ‘Lazarus’
severe End: taxa
for characteristic smooth transition
extinction events
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OCEANIC MODEL TESTED ON THE SILURIAN 665

S-STATE \ /
HIGH
Initially : release of CO2 from the
‘n--
/W\
LATiTUDES
>+5OC
ocean to
atmosphere
the / \
planktic Low D R Y LOW
production LATITUDES
Expansion Reef Ooiites
Faunaswith
of carbonates growth initially
few, rare taxa

limestones
Pure of weathering

‘1
I
Benthicrecycling
1.2.10e&s of nutrients

Salinity stratified ocean

Thermal expansion

Warm, saline
with L
Canoxic
O2 little water
Fig.2. Some characteristicsof an oceanic
S state.
Blackshales

Thedeep oceansedimentsarethus in accord with a Organic carbon productivity changes

model in which there was a long-term trend towards lower Bralower & Thierstein (1984) calculatedthatthereplace-
oceanictemperaturesat high latitudes.Thefluctuations
ment rate of anoxic,salinity-densedeepwaterduring
between warmer and colder climates were strong enough to
various mid-Cretaceous episodeswas less than a tenthof the
produce distinct, short-term episodes; that is, the effect of
rate of temperature-densedeepwaterlikethat of the
the long-term trend during a single cycle was low compared
present. Thus the rate of vertical advection would be less
to the influence of the cycle itself.
than a tenth of that of the present. The content of nutrients
transported to the surfacewould be lowered still more, since
the trace metals would be bound effectively as sulfides and
Effects on the clay and metal content of shelf the upwelling water would be depleted in them. They also
sediments calculatedthatoceanicplankticproductivity in asalinity
Duringwetterclimaticintervals, therate of chemical stratified ocean would one order of magnitude lower than
weathering, erosion and transport of clay to the sea would that of the present. Applied to the Silurian, this means that
be increased ( K a u b a n 1986). There is ample evidence of planktic productivity duringthe S episodes would have been
this effect during the Wenlock and Ludlow P episodes (see a tenth of that of the P episodes. The increased amount of
references in Jeppsson 1987~).However, only sections from terrestrial
weatheringduring P episodes would have
low latitudes or those of uncertain latitudinal position are constituted
another
source of minerals
for
organic
well known (compare maps in Ziegler et al. 1974; Van der production.
three-layered
aIn Ocean, the planktic
Voo 1988), and thus itis only possible to infer alow latitude productivity could havebeensimilar tothat duringan
humidity change. episode cold enough to have formed a two-layered ocean,
During dryer climatic intervals, limestones would be less because the terrestrial weatheringwould be that typical of a
argillaceous. In the Silurian record, the contrast of these, to wetter episode. Tappan (1986) emphasized the importance
those in thewetterepisodes is distinct(seereferences in of terrestrialweathering as nutrient
a sourceforthe
Jeppsson 1983, 1987~). plankton. Furthermore, the increasedupwelling would bring
Deposition of variousironand otherore minerals in up increased amounts of nutrients, although the concentra-
epeiric seas and on the shelf areas seems to increase during tion would dependpartlyonthearea of seabottom in
P episodes, for
example in theLlandovery-Wenlock contact with the cold, dense, oxic water.
boundary episode (Jeppsson 1987~).This may be due to the A ten-fold increase in primary productivity would permit
fact thatmetalsare
deposited as sulfidesin oceanic acorrespondingincrease in numbers of individualsat all
sediments during periods with anoxic ocean bottom water, levelsin thefood web andenablethedevelopment of a
and the upwelling water is consequently depleted (Wilde & different community structure. Thus the P episodes would
Berry 1986). Increasedweatheringduringthewetter (P) becharacterized by moreabundantandmorediverse
episodes would also increase the flowof metal ions to the planktic and nektic faunas than the S episodes.
oceans.In
addition, the reduced plankticproductivity K a u b a n (1986), in his review of Cretaceous sequences
duringdryer (S) episodes would decreasetherate of in western North America, registered a sedimentologically
transport to the bottom; the removal of metals by sinking significant increase in organic carbon
production by
particles may be significant in depleting metal ions from the organic-walled algae(dinoflagellates,etc.)duringthe wet
photic zone (Bostrom et al. 1974). All three processeswould episodes. Such studies are not available for the Silurian, but
contribute to a higher concentration of metal ions in the a strong increase in planktic production is indicated by a
water-sediment interface on the shelf during P episodes. strong increase in conodont abundance duringP episodes. A
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666 L. JEPPSSON

Table 2. Some causes and effects of climatic lithological and faunal changes

Humid low latitudes result in:

1. high rate of weathering, erosion, transport and deposition of clay etc; the increased run off may result in
2. salinity stratified epeiric seas and coastal waters (with a ‘fresh water lid’) and
3. blooms of organic walled plankton which may result in
4. shallow water black shale events and
5 . a carbon isotope anomaly; 1, 2, 3 and 13 (below) combine to cause
6. inhibition of reef formation and
7. decreased benthic carbonate production resulting in
8. retreat of the margin of the carbonate belt; the increase in oceanic turbidity due to 1 and 3 causes
9. light-influenced benthic communities to move to shallower waters; 1, 6, 7 and 8 result in a gradual increasein CaCO, concentration in sea
water, increasing the likelihood of
10. oolite formation during the early part of the succeeding dry period.
High latitude temperatures below 5 “C result in temperature stratified oceans which have:
11. oxic deep water sediments and a
12. high rate of upwelling which together with 1 results in
13. high planktic production, supporting
14.high abundance and
15. high diversity conodont faunas including taxa specialized in high productivity environments. The thermal contraction of the deep oceanic
water due to the lowered temperature results in a
16. minor regression, which may be enhanced if the lowered temperatures at high latitudes result in glaciation. T h e upwelling oxic bottom
water is not depleted in metals; this together with 1 and 11 results in
17. increase in deposition of ore minerals on the shelf areas, which is especially evident in areas affected by 6, 7 and 8.
A change to warmer high latitudes would initially result in a cessationof the formation of new oceanic deep water and thereby a cessationof deep
upwelling, which together with 31 below would lead to an:
18. extremely low planktic production during a
19. severe period of about 0.1 Ma, resulting in an
20. extinction episode. Modification by Milankovitch cycles may cause interruptions of the severe period and both extention of it and
21. stepwise extinctions. When the deep ocean water lost enough density, a salinity stratified ocean would develop characterized by a
22. low rate of upwelling (10% of that of 12) which together with 31 (below) causes
23. low planktic production (less than 10% of 13) resulting in clear water which permits
24. increased benthic production, especially in marginal areas; the increase in skeletal carbonate causes a
25. seaward expansion of limestone formation. Often the salinity stratified ocean has anoxic bottom waters resulting in
26. black shale formation in deep oceans. Point 23 further results in
27. lowered abundance at all trophic levels relying on planktic production and a
28. lowered conodont faunal diversity; the fauna consists of
29. taxa specialized in low planktic productivity environs or those more dependent on benthic production. Point 23 further results in
30. lower 613cvalues.
Dry low latitudes result in a
31. decreased rate of weathering, erosion and deposition of clay etc. resulting in
32. more pure carbonates and decreased freshing of coastal waters permitting
33. widespread reef growth, further promoted by a
34. transgression due to thermal expansion of the deep oceanic water and melt off from any glaciation.
repetition of most or all of points 1-34 results in
35. lazarus taxa if enough refuges exist and
36. iterative evolution, if refuges are lacking, due to the repetition of the changes in selection pressure.

changeinconodontfrequency is observed in mostplaces area that correlations cannot be made, whereas the two P
where those from late Llandovery-earliest Wenlock and the episodefaunas havemany taxa in common. Evidently
lateearlyLudfordian P episodescan becompared with plankticproductivity differences increased inan offshore
those from succeeding strata. In the Silurian of Gotland, the direction as may be expected from the model. The better
Llandovery-Wenlock transition can be studiedinsurface oceaniccirculation duringthe P episodes would transport
exposures,
and
in the Narborrningen 1 core which moreland-derivednutrients
farther offshore, and the
palaeogeographicallywas situated 45 km fartheroffshore. nutrients derived from upwelling would be distributed more
The Lower Visby Beds (Llandovery-Wenlock boundary equally. Thus the Nar area was so similar to the area closer
beds) in the surface exposures belong to Benthic to the shoreduring the P episode that most of the taxa were
Assemblage 4 (Gray et al. 1974). The decrease in conodont the same, whereas during thesucceeding S episode only rare
frequency up section is about two orders of magnitude in the individuals of a few taxa, specialized for lowproductivity
core, where the lithologies seem to differ little, but only one environments, could survive so far away from the terrestrial
order of magnitude in the shallower area represented by the nutrient source.
surface exposures.Furthermore,the incoming S episode The deep water response during the P episode spanning
fauna in the Nar area is so different from that in the outcrop the Llandovery-Wenlock was small, indicating that vertical
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O C E A N I C M O D E L TESTED ON THE S I L U R I A N
advection was low. Thus the overall strong faunal response
is another indication that terrestrially derived nutrients were
the most important source for planktic production.
Effects on calcium carbonate deposition
During most of the Silurian Period, planktic calcium
carbonateproduction wasnegligible, so it isonly benthic
carbonateproductionanddeposition thatneedsto be
considered.
The wetter climateduring the Pepisodes affected
benthiccarbonateproductionanddepositiononthe shelf
areas negatively inseveraldifferent ways. The relative
importance of these factors varied with the environmental
setting. The increase in availability of nutrient in itself may
have been detrimental, as Hallock(1986) found evidence for
‘the hypothesis of excess nutrient availability as a factor in
reef drowning . . . because major carbonate-producing reef
organisms
have life history strategies specialized to
nutrient-deficient conditions,they are unable tocompete
when nutrients become plentiful’.
Both the increased planktic productionand the increased
amount of clay reaching the shelf waters would decrease the
penetration of light; thus, benthic photosynthetic production
would decrease, since both free-living algaeandzooxan-
thellae would be affected. This would affect not only calcium
carbonateproducingbenthic algae andthehosts of the
zooxanthellae but also the whole food web dependent on
them or on other benthic algae. The result would be most
severe in the marginal parts of the carbonate platforms and
in similar settings. Duringthelate Llandovery-earliest
Wenlock P episode,carbonateformationretreatedabout
500 km in parts of eastern North America (Jeppsson1 9 8 7 ~ ) .
In a study of Cretaceous climatic effects, Kauffman
(1986)concluded that brackish water may formasurface
layer during wetter episodes in some epicontinental settings.
The flow of oxygen to the bottom may then have been so
low that black shale was formed. Such Pepisodeepeiric
black shales have not yet been identified from the Silurian,
but some of those reported from Devonian cycles (House
1985) seem to have formed under such conditions.
Alternatively, the shelf waters may havebeen well
mixed,allowingbrackish water to come into contact with
the bottom. Brackish water has a strongly negative effect,
especially on reef formation. The cyclicity in the formation
of reefsandreef-relatedlimestone onGotland has long
been known (Murchison 1847;van Hoepen 1910; Hede
1921, 1960; Martinsson 1967; Laufeld 1974). The details of
this cyclicity correlate wellwith the cycles notedherein.
Similarly, the MuchWenlock Limestone Formation in the
Welsh Borderland was formed during one of the strongest
Silurian S episodes, that of the late Homerian.
Changes in temperature would also directly affect both
carbonate production and deposition, producing a latitudi-
nal shift of climatic belts. Another setting where
temperature effects might be registered is on the deepest
shelf and slope areas, where different kinds of deep water
were in direct contact with the bottom. These effects cannot
yet beseparatedfromthose discussed above. Since the
change in temperature may have been rather small (a few
degrees in the high latitude waters would suffice to cause the
oceanic effects) it is unlikely that a shift in the climatic belts
would be detected and separated from the other effects.
667
Eustatic effects
If consideredseparately,some of the lithological changes
discussed above fitwellwith a eustatic explanation. Thus,
the change betweenlimestoneandcarbonate-poor sedi-
ments in marginal carbonateareas is thatexpected from
rapidsea-level changes.In such settings the colderand
wetterintervals would be likely to be described as
transgressive.Such aninterpretation fails to explain the
contemporaneous change more to oxic deep water
conditions registered in the sediments. Furthermore, during
the succeeding regression, when limestone was formed in
the area again, we would expect to find at least parts of the
diverse earlier fauna further offshore. As noted above, that
is not the case.
Theincrease in turbidity of the seawaterduringP
episodeswould decrease the penetration of light. One
consequencewould that
be
light-dependent
benthic
communities would live in shallower waters duringP
episodes and in deeper waters during S episodes. Thus, even
this tool for identification of sea-level changes is affected by
the cycles.
However, some eustatic effects are to be expected as a
consequence of the modelsuggested here. If the high
latitude temperature was low enough during the P episodes,
and land existed in suitable places, ice sheets would form.
Glacially controlled eustatic
changes would result.
Moreover, Milankovitchcyclesmighthaveinfluenced ice
volume to cause a series of geologically very rapid eustatic
changes. Therefore, in addition tothe possiblesea-level
changecaused by the changesbetweencolder(P)and
warmer ( S ) episodes, any regression might be modifiedby
rapid oscillations. Such eustatic changes would enhance the
general effects outlined above. Thus, the lowered sea level
would openfurtherareasforweatheringanderosion,
increasing the transport to the oceans of nutrients, dissolved
carbonates, and clay.
Anothereustatic effect wouldalso beexpected. The
increase in volume of sea water with temperature is such
that a rise in temperature from 0°C to 12°C for a water
column of 3.44 km (see below)would cause a rise in sea
level of 4.6m. Compensationforthe shelf areas would
lower the value to about 3 m. Such small transgressions have
recently been discussed in the Silurian (Cotter 1988), but
they are not dated well enough to permit correlation with
the cycles noted herein.
These eustatic changes would combine and be opposite
in effect to those deduced sometimes from the sediments:
the lowered sea level would coincide with episodes of retreat
of the edge of carbonate deposition on the shelf areas, and
themajor sea-level rise wouldcoincidewith the seaward
expansion of carbonatedeposition. The growth of large
reefs during these episodes fits well, however, since a slowly
rising sea level is important for the formation of large reefs.
Thus some eustatic changes may be interpreted only as
an effect of global changes in humidity and temperature and
they do not provide a driving force for sedimentological and
faunal changes. This agrees well with the fact that the large
eustatic changes in theQuaternaryhavenotresulted in
extinctions in the marinebiota(Eckert 1988). Further,
before the presumedspecies-area effect is invoked, the
critical comments by Stanley
(1984a, b ) should be
considered. Many eustatic changes may have other causes,
such as variations in the rate of sea-floor spreading.
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668 L. JEPPSSON

Some transition effects periods would be more complicated, including a number of

When the production of dense, deep water ceased in high Milankovitch-induced brief returnsto P stateconditions,
latitudes it would take some time before the bottom water thereby prolongingthetransition
period.Further,
the
lost so much of its densitythatit could be replaced by resultingsedimentary andtor faunal sequences would
salinity-dense water (Schopf 1980). One effect causing a loss include a number of repetitions. The detailed effects would
of density would beheatingfrom below (Schopf 1980). vary between different transition periods due to the timing
Worthington (1968) calculated that the present oceanic heat of the perturbating forces.
Thestratigraphicalrecord
flow suffices to raise the temperature of a 1km column of provides someexamples. The faunalresponseduringthe
water 7 "Cin 15,000 years. Bralower & Thierstein (1984) two best-studiedSilurianchangesfromwettertodryer
counted
temperature
on
a difference
between mid- climate did take some time (Jeppsson 1983, 1987~).I have
Cretaceous and Recent bottom water of 12 "C and provided earliercalculated(Jeppsson1987u), based on sediment
data on the area and volume of the deep water reservoir thickness, that the first faunal change occurred 90,000 years
which give an average of 3.44 km for the thickness of the before the change in clay lithology at the contact between
deepwaterreservoir. 88,000 years of currentheat flow the Lower Visby Beds (Llandovery-Wenlock boundary
would raisethetemperature of that column of water by interval) and the Upper Visby Beds (Wenlock) on Gotland.
some 12 "C. A further interval of time would be needed to Thelatterformationrepresentsthelaterpart of the
heat the sea bottom, since after a long interval of contact transitionperiod and is somewhatthickerthanthestrata
with 0°C bottom water the temperature gradient through from the early part. The length of the transition period was
the rock would be in equilibrium with this water. evidently in the order of a few hundreds of thousands of
Agradual warming of thetemperature-densebottom years, or less.
water would decrease its density. The stability of the Furthermore, several of the best known extinction events
stratification would then decreasegradually
and
the seem now torepresentthe sum of several minor events,
threshold for a brief return to a modified P-state would be separated by shortintervals of time.Forexample,the
lowered.Thus, slighthigh latitudeclimaticdeteriorations earliest Wenlock conodont extinction episode consists of at
that would not have affectedthe production of bottom water least two pulses followed by a period with extinctions of rare
at the beginning of a transition period (Fig. 3) could do so survivors (Jeppsson 1987~).Additionally, especially after
somewhat later. Similarly, it is likely that the first inhibition the second pulse, several taxa that did not become extinct
of deep water production athigh latitudes could be triggered are absent from the lower part of the 'inter-pulse' strata.
by any slight climatic amelioration. Theirreappearance reflects
a brief return
to
P-state
These calculations predict a number of effects. First, for conditions.Consideringthe likely rate of sedimentation,
example, a geologically detectable interval of time is thickness of strata, and amount of time available, it may be
expected during a change from P to S episodes before the theobliquitytermthat was behindtheextinctions.This
new oceanic state is in balance with the heat flux. Secondly, gains further support from the observation that deviations in
deep upwelling would beextremely low duringthetime insolationat high latitudesare mostly afunction of the
when no new bottom water was produced. obliquity term, whichis ,currently41,000years(Berger
The calculated length of the transition period is longer 1977).
than the shortest Milankovitch period. The strong effect of The changefromsalinity-densetotemperature-dense
Milankovitch periods on high latitude temperature iswell bottomwater,onthe other hand, would be smooth. The
established(Saltzman 1987). Thus,instead of asteadily production of salinity-dense bottom water would continue
progressing change duringthe
transition
period
these until forced to ceaseby the denser temperature-dense water.

P+ S TRANSITION
PRODUCTION:
PLANKTIC
LATITUDES
HIGH
MILANKOVITCH
CYCLES: extremely
low
temperoturo8
hlgher higher
lower temperatures

FAUNAS:
extfnctfon8, entry o f h8rdy S-rtote toxo
Lazerus taxa, hardy P-state taxa

Fig.3. The transition period. The ocean

TEMPERATURESTRATIFIEDOCEAN alternates between two differentstates.
The first state (bold italics) is charac-
teristic of intervals with higher tempera-
tures at high latitudes; thesecond state
(thin, Roman-style lettersand thin
circulation arrows) is associated with
HEAT FLOW lower temperatures at highlatitudes.
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O C E A N I CTESTED
MODEL O N THE SILURIAN 669

Inaddition, the effect of theheat flowwould increase post-extinction fauna. If this process wererepeated,each
temporarily due to the fact that the temperature gradient rebuilding of a high diversity fauna would start from those
was in balance with theformer, warmerbottomwater. forms which had remained in the low productivity environs
Therefore upwelling would increase. during the high diversity, episode.These would probably
have evolved but little. Furthermore, evolution would take
place under closely
a similarselection pressure.Thus,
Effects on conodont faunas iterative evolution would be expected.
As yet, there is no agreement on some of the fundamental Iterative evolution has not yet been identified in Silurian
questionsregardingconodont ecology,including whether conodonts, reflecting the fact that refuges were available for
they were pelagic or benthic (Seddon & Sweet 1971; Barnes many of the high diversity taxa.
However,iterative
& Fghraeus 1975; Klapper & Barrick 1978; Ethington et al. evolution is well known in the Cambrian-early Ordovician
1987). However there is a range of potential habitats, from trilobitebiomeres(Stitt 1971). It has also been identified
thosedependententirelyon pelagic production to those recently in Devonian to mid-Carboniferous conodonts
dependent completely on benthic resources. Considering the (Ziegler & Lane 1987; Sweet 1988).
large variation in morphology and the variability of The model of oceanic cyclicityprovides an explanation
distributional patterns, conodonts presumably exploited not for long periods in which some lineagesmaylacka fossil
just a limited part of this range buta significant portion of it. record, and it seems likely that more cases of intermittent
Therefore, changes in the relative importance of benthic and records of lineages will be identified. Iterative evolution may
planktic production would affect differentconodonttaxa yet be found in Silurian conodonts. The difference between
differently. Similarly, changes in all theother conditions the Silurian and the Devonian to mid-Carboniferous record
discussed above would also influence which taxa thrived. may be onlyadifference in the relativeimportance of
TypicalP episodetaxaappear to havedepended to a iterativeevolutionandreappearancesfrom refugia for
larger extentonplankticproductionthan S statetaxa. A re-establishing a high diversity fauna.
tendency to form local populations is likely tobemore
pronounced in the latter, and certainly seemsto characterize Average length of B cycle
the Ozarkodina steinhornensis group (Jeppsson 1975, 1989),
typical S state taxa. The time scale of the average length of the episodes cannot
Many Silurian conodont lineages have turned out to be yet be defined precisely. It can be estimated by taking the
intermittently widespread and confined to unknown refuges duration of the Silurian to beabout 35 to40Ma(data
between these widespread episodes (Jeppsson 1984, 1987a; combined from Gale 1985; Odin et al. 1986). The relative
and Wilde & Berry 1986, have discussed where such refuges length of the Wenlock and Ludlow epochshasbeen
may be found). The large number of such ‘Lazarus’ taxa in estimated to be 21% and23%, respectively, of the total
the Silurian indicates that suitable refuges for different kinds Silurian Period (Boucot 1975). As noted above, there are at
of specialists did exist through this interval. least four cycles duringthese two epochs, if the late
Asnotedabove,thetransitionfrom aP episodestate Llandovery-earliestWenlock episodeand the dryerlate
with temperature-densebottomwater (or a three-layered Ludfordian episode (the end of whichispresentlypoorly
ocean) toan S episode withsalinity-dense bottomwater known) are included. Thus, the average length of acycle
would be much more severe than the reverse. Furthermore, would be about 4 Ma.
thechange would hit ahigh-diversity fauna with more The average time
between the eight ammonoid
specialized taxa. Theconodontfaunal response tothe extinction events in the Devonian discussed by House (1985)
lowering of oceanic productivity was massive and worldwide is about 4.3 Ma. Only three conodont cycles were identified
(Jeppsson 1987~).Largeconodontextinctioneventsare by Ziegler & Lane (1987) during this interval, with
found during the two most pronounced changes of this kind boundaries that were not alwayscoevalwith those of the
in the oceanic regime during Wenlock to Piidoli times. For ammonoids, but different taxonomic groups might be
expected to react differently. Also different criteriafor
example,morethan half of the known conodont species
establishinga significant extinctioneventwere usedby
disappeared attheend of thelate Llandovery-earliest
Wenlock P episode (Aldridge 1976, 1988; Jeppsson 1987~). House and by Ziegler & Lane. The characters of the four
Silurian cycles counted herein are very different, and only
two of them are easily detected by use of conodont
Effects on evolution extinction evidence. Moredetailed
studies are needed
The cyclic changes outlined above would result in acyclic before the number of events recognized in the Silurian and
change in selection pressure. As noted previously, decreases Devonian is complete and correct.
in planktic productivity resulted in extinctions within highly
diverse conodont faunas. On the other hand, it seems likely
that specialists in low productivity environments would have Reservoir effects
a good chance to find a refuge when necessary since ‘oceanic Both the strength and the length of the Silurian cycles seem
deserts’ wouldalways exist somewhere. The faunal to differ greatly. The differences in character are probably
succession of conodonts in the Silurian is in agreement with related to both thelength of the cycle and to the state of the
this. Thoselineagesthatcharacterize the early Wenlock oceanic and other reservoirs at the beginning of the cycle.
fauna after the extinctions have an intermittent record well The characterandlength of the previous cycleswould
into the Devonian and show little change. therefore affect the character of the succeeding cycles.
If all members of a high diversity fauna became extinct, It is obvious that faunal characters would be the most
the nexthighdiversity fauna would have to evolvefrom sensitive to such historical effects, since the number of
those few taxa that constituted the low diversity lineages present and their degree of specialization constitute
 Downloaded from http://jgs.lyellcollection.org/ at New York University on May 24, 2015
670 L. JEPPSSON
the reservoir with both the longest build-up time and the
shortest emptying time.
However, other types of reservoir need to be considered.
A large number of ionshaveresidencetimes in the
present-day oceans of such a length that their concentrations
would be affected significantly by changes in input, andJor
in deposition, over the length of time of an average cycle.
Here I discuss one example.
Calcium presentlyhasaresidencetime of0.85 Ma
(Schopf 1980). As noted above, for mostof the Silurian only
benthic carbonate production was of budgetary importance.
Thus, during the P episodes, the decreased output due to
the decrease in benthic carbonatedeposition
and
the
increased inputdueto increasedweatheringanderosion
would bothlead to agradualincreasein the oceanic
reservoir of calcium. If the concentration became sufficiently
high, it might be registered in the sediments formed.
The formation of oolites requires water oversaturated in
calcium carbonateanda high temperature(Fisher 1984).
Oolites are rare in the Silurian, but during an early phase of
at least two of the warmeranddryerepisodes,oceanic
calcium carbonate concentrations in Gotland reached a level
where oolites were formed. The Bara Oolite Member (Hede
1928)was deposited during a very early phase of the late
Wenlock S episode. The Burgsvik Oolite (Hede 1921) was
formedduringa similar phase of thelate Ludfordian S
episode.
Other Palaeozoic examples for which this model
may be valid
The early Ordovician
Fauna1 and lithological changeshavebeendocumented
widelyfrom theOrdovician (Miller1984; Leggett et al.
1981; Lindstrom 1986; Erdtmann 1986; Thickpenny &
Leggett 1987). A good example was provided by Ethington
et al. (1987), who studied one of the biomere boundaries in
the early Ordovician of North America. In several ways this
boundary coincides with a change from a high productivity
to low productivity ocean (a P to S change). In their Ibex
section the faunal change took place over more than 8 m of
strata, with the conodonts showing a decrease in abundance
of over 60%. A lowdiversity and low abundance interval
followed the main extinction, which appears to have been
worldwide; the extinction may have been stepwise but the
collections aretoo small to becertain.The extinction is
followed, locally, by siliceous oolites,andadecreasing
amount of mud occurs stratigraphically upwards across the
interval. The interpretation of the faunal changes
as
consequences of a change from a P episode to an S episode
deviates from earlier explanations, in which the lithological
changes were ascribed to shallowing (Stitt 1983) and brief
coolingwassuggested asthe causefor the extinction
(Lochmann & Duncan 1944; Stitt 1975). The reliability of
the suggested eustaticchanges was ranked verylow by
Steckler (1984),who noteda lack of agreementonthe
number, time and magnitude of minor sea-level changes.
The Devonian
Devonian and Carboniferous cycles andeventshavebeen
documented in many publications (e.g. House 1985;
Schonlaub 1986;Chluptic & Kukal 1986;Ziegler & Lane
1987). Here I discuss only two episodes.
One is theend of the episode when the Silurian-
Devonian ‘Oberen
Graptolithenschiefer‘, a deep water
black shale indicative of an anoxic environment, was
deposited in Thiiringia; it is succeeded there by the
TentakulitenKnollenkalk (Jaeger 1976), the boundary
between the two units representing a change to oxic deep
waters. The Knollenkalk includes conodonts with platform
elements of the ‘ancyro’ group referred to Ancyrodelloides
by Bischoff & Sannemann (1958). Similar elements seem to
be characteristic of other P episodes during the late Silurian
(Ancoradella) and Devonian. The Knollenkalk and its fauna
form thelateLmhkovian(Chlupac et al. 1985)high-
diversity episode of Ziegler & Lane (1987). The end of this
episode was distinguished as the Lochkovian-Pragian
boundaryevent by Schonlaub (1986). Inthe Barrandian
area this event is expressed lithologically as a change from
darker, shale-rich stratato markedly lighter carbonates
(Chlupac & Kukal 1986). Profound changes in other faunal
components were alsodescribed(Chlup6c & Kukal 1986).
There is thus good evidence for concluding that this high
diversity episode represents a P episode, during which high
latitude parts of the ocean became so cool that the oceanic
bottom water was formed there. When the episode ended, it
seems as if thedeep oceanremainedbetteroxygenated,
although the faunal responseand the responsein
clay-limestone deposition were typical.
The thirdappearance of conodonts with ‘ancyro’
platforms was in the latestGivetianandFrasnian, when
both Ancyrodella and Ancyrognathus flourished (Feist &
Klapper1985;Ziegler & Lane 1987). Theend of this
episode is the famous Kellwasser event (House 1985), at the
end of the deposition of the Upper Kellwasser Limestone
(Ziegler 1984; McGhee et al. 1986;Ziegler & Lane 1987;
Walliser1986;Walliser et al. 1988). If this sequence is
interpreted according to the model suggested here, then a P
episode would begin near the beginning of the Frasnian, and
end at the top of the Upper Kellwasser Limestone. There is
some evidence for such an interpretation. In the Montagne
Noire, a marginal carbonate area with a pelagic sequence,
theearlypart of theepisode includes achange tomore
argillaceous sediments (Feist & Klapper 1985); a similar
sequence is foundintheHiihnertal section in theHarz
Mountains(Lottmann et al. 1988). 13C dataindicatesthat
the euxinic character of the Kellwasser limestones resulted
from
blooms of phytoplankton (McGhee et al. 1986)
analogous to those described by Kauffman (1986,) from the
Cretaceous.In the Cretaceous, also the lack of benthic
oxygen dueto salinity stratifaction of the watercolumn
restricted macrofaunas to, at best,
sparse low-oxygen
tolerant inoceramidbivalves(Kauffman1986), and large
bivalveshave beenreportedfromthe euxinic Devonian
sequences (Lottmann et al. 1986).
The sequence of events shown by the Devonian faunas is
closelysimilar to that registered during the severe period
after the Llandovery-Wenlock boundary episode,
P
although the Silurian strata lack evidence of a euxinic phase.
A sharp decrease in conodont abundance is recorded across
the Frasnian-Fammenian boundary. Sandberg et al. (1988),
for example, noted that in the Steinbruch Schmidt section
there is a drop from over 11,OOO elementsper kg in the
lowest sample from the linguiformis Zone, the last zone in
the Frasnian, to less than loo0 elements per kg in the richest
 Downloaded from http://jgs.lyellcollection.org/ at New York University on May 24, 2015
OCEANIC
MODEL
TESTED ON
SlLURIAN
THE 67 1

sample,from the oldestFammenian Zone; diversity also
drops from at least 25 species to 9 or 10. There is an
accompanying change in the composition of the conodont
fauna, with Icriodus increasing strongly in frequency. This
Icriodus bloom hasbeenreportedfrom many areas,for
exampleSouthChina(JiQiang 1988), Poland(Matyja
1988), Germany (Walliser et al. 1988; Sandberg et al. 1988),
Belgium, Nevada, and Utah (Sandberg et al. 1988).
Furthermore,theextinction of differentgroups was not
contemporaneous but stepwise (Walliser et al. 1988). Becker
(1986)calculated aduration of severalhundredthousand
years for this Devonianfaunal crisis, and Walliser et al.
(1988) concluded that it lasted at least several lOKa, which
is the same order of magnitude calculated for that beginning
attheend of the P. amorphognathoides Zone (Jeppsson
1987a).
The above interpretation of the Kellwasser event differs
considerably from that presented by Sandberg et al. (1988 p
296)which embodied 12 independent (including bolide
impact) events.They claimed thatthe first of these,a
transgression, drowned the reefs whereas the extinction was
connected with the eighth event, a second bolide encounter.
No explanation was given for
the first of their
transgressions, butprobably the most rapidtransgressions
are caused by the melting of a major ice sheet, and the end
of reef growth is often ascribed to drowning. However the
repeated Quaternary advance and retreat of ice sheets has
not caused widespread reef drowning; on the contrary, most
reef tops are close to present sea level, and major marine
extinctions were not connected with the eustatic changes of
this period.
Walliser et al. (1988) did not invoke any eustatic causes
for the Kellwasser eventbutwhat brief suggestionsthey
madeaboutits origin agree with the model of oceanic
overturn outlined herein. The Kellwasser event is thought
by the present author to be the consequence of change in a
fewclimatic factors, and not to be unique with regard to
cause; even the effects were not unique although they were
more severe than usual.
Post-Triassic examples
The possible usefulness of the model to assess post-Triassic
changes
is unknown; the majorchange in
carbonate
deposition pattern and CO, storage pattern causedby the
appearance at this time of sedimentologically significant
planktic carbonate production (Fisher1984) may or may not
be important. However, some of the discrepancies between
theobservationsand the modelsinvoked to explain the
Cenomanian-Turonianoceanicanoxicevent(seeJarvis et
al. 1988)would be dispelledby application of the model
presented here.
Forcing mechanism
Only a few observationspertinent to thisquestion are
available currently. They include the following.
(1) The cycles seem to be of varyinglengthbut the
average length of each is several Ma, which rules out an
explanation based on Milankovitch cycles. Moreover, they
are much shorterthansomeotherproposed cycles, for
example those discussed by Fisher & Arthur (1977).
(2) The cyclicity was not limited to the Silurian but can
be found at least from the Cambrian into the Triassic.
(3) The magnitude of the faunal changes varies strongly.
(4) The evidence points to acontrol byhigh latitude
temperature changes and correlated low (-middle?) latitude
humidity changes. The temperature changes themselves may
be of three different types.
(a)Changes in the global averagetemperature but
without a changed latitudinal temperature gradient. Among
possible causes would be changes in the solar constant and
in the albedo of the earth.
(b) The global average temperature remained more or
less constant but the latitudinal gradient varied.
(c) Only the high latitude temperature fluctuated.
The firsttwo scenarios involve temperature changes in
low latitudes, which maysomehow produce associated
humidity fluctuations; ameans of modifyinglow-latitude
humidity is not apparent in the third scenario.
There is a strong difference in the severity of successive
episodes. One cause of an effect that is stronger than the
average is found in the stepwise decrease in marine primary
productivity due to increases in the retention of nutrients in
terrestrial ecosystems (Tappan 1982; compare alsoFisher
1984). The transitional period subsequent to such a decrease
would be especially severe. Another cause of the variation
in severity would be whether or not the most severe part of
a cycle coincided with a similar part of a Milankovitch cycle.
The forcing mechanisms that are possible may be divided
into three groups.
(1) Astronomical, such as variation in the solar constant.
The models for energy production in the sun do not indicate
any fluctuation or cyclicity, although the detected neutrino
flux is presently only a third of that expected.
(2) Terrestrial,butexternal tothe ocean-atmosphere
system. This group includes changesin the rateof volcanixn
(for climatic effects see Lamb 1970; Toon 1984) and in the
rate of sea floor spreading. In such cases, the ultimate origin
of the cyclicity remains to be identified.
(3) Terrestrial, within the ocean-atmosphere system. If
one oceanic state leads to a change in a reservoir crucial for
the maintenance of either of the oceanic states, then the
chain of causes would include a closed loop. A model of this
kind
discussedby
Saltzman(1987),
involving the
late
Quaternary climatic changes with a 0.1 Ma length of cycle,
stresses the importance of atmospheric CO, concentration.
COz fluctuationswouldalso beexpectedfrom the model
presented here and a model could be developed to account
for this.
The ‘metronome and engine’
Only thatfraction of the CO,found in theatmosphere
regulatesglobal temperature. A much largerfraction is
stored in the ocean as dissolved CO2, carbonic acid and its
ions. All these reservoirs are in equilibrium.
At the onset of a P episode, the strong decrease in the
temperature of the deep oceanic waters would increase the
oceanic storage capacity considerably,extractingcarbon
dioxide fromtheatmosphereandthereby lowering the
average global temperature. The result would stabilize the P
state conditions far beyond the duration of the perturbation
which caused the beginning of the P episode.
During the P episode, the increased input and decreased
output(seeabove) would fill the storage capacity of the
oceanandincrease the atmosphericcontent of CO,. The
 Downloaded from http://jgs.lyellcollection.org/ at New York University on May 24, 2015
672 JEPPSSON L.

resulting temperature increase would decrease the stability 91-104.

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Received 16 June 1989; revised typescript accepted 1 December 1989.
Note added in proof
The effects of changes in oceanic water with regard to turbidity of
the
water, benthic carbonate production and net deposition,
community displacements, ‘drowning’ of reefs, etc. have been
analysedby Hallock andher collaborators (Hallock & Schlager
1986; Hallock 1987, 1988; Hallock et al. 1988). Their data underline
verystrongly the importance of the presence of, or the lack of,
nutrients for allbiologicaleffects, both with regard to which
organisms were thriving and what sediments such organisms
produced or destroyed.
Climatic effects due to variation in the burial of organic carbon
have been well studied. Evidence for such variation during the
Silurian is lacking, whereas there is ample evidence for variation in
the removal of carbon in the form of carbonate.Therefore my
discussion above was limited in the latter.
Several papers published afterthe manuscript was submitted
are also pertinent. Berger et al. (1989) have recently calculated the
length of Milankovitch cycles back to 440 Ma, when the present-day
41 000 years obliquity term is estimated to have been only about
30 500 years. The base of the Wenlock has been dated as 430 Ma
(Odin et al. 1986). Based on sediment thickness I previously
calculated that the time between the first and the fourth extinction
datum was 90000 years (Jeppsson 1987~).Above, I noted that the
obliquity term was the mostlikely cause of the timing of these
extinctions; that is, the length of time between the first and fourth
extinctions should bethree times the obliquity term.The new
values for three times the obliquity-term is not about 93 OOO years,
instead of the previous estimated 123 000
years. This fits
unexpectedly wellwith theduration calculated fromsediment
thickness.
Another link between oceanic upwelling, nutrient concentration,
ocean productivity, atmospheric CO2and climate has been
discussed by Wilson (1989). These effects would strengthen those I
have described.
Also, McKinney (1989) has discussed ‘the rebound effect’. This
is the rebound time that is needed after a major extinction before
there are enough extinction-sensitive taxa present again to permit a
second major extinction.
Regarding stepwise extinctions, some similarities which may or
may not be accidental can be noted. The number of steps identified
is usually between five and ten; the strongest extinction effects are
found at oneof the earlier datums, although not at the first; and the
R. J. (ed.) Paleobiology of conodonts
Mid-Carboniferous.In: ALDRIDGE,
EUis H o w d Ltd Chichester, for the BritishMicropalaeontology
Society, 147-164.
distance in terms of metres of sediment between the steps in the
latter part of the sequence is larger than between steps at the
beginning. The patterns of extinction and the replacement datums
across the Cenomanian-Turonian boundary (Edler 1989), for
example agree in all these characters with those across the
Llandovery-Wenlock boundary interval.
Opinions between different authorsaboutthe length of time
involved with an extinction event vary strongly, both with regard to
individual events (e.g. see McGhee 1989) and with regard to
individual events. An extinction event based on my model would
not be expected to deviate much from a few hundred thousand
years. However, a series of S and P cycles each ending with an
event (each one probably with stepwise extinctions) could result in
a 10Ma long extinction period of the kind described by Prothero
(1989) from the later Eocene and Oligocene, although the
extinction steps he described may well have other causes.
Additional references
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