ArticlePDF Available

Predation in the Ordovician and Silurian of Baltica

Authors:

Abstract and Figures

Signs of predation appear in the Middle Ordovician of Baltica. Shell repair dominates over the predatory borings in the Ordovician and Silurian. Predators attacked molluscs, brachiopods and tentaculitoids in the Ordovician and molluscs, tentaculitoids, brachiopods and ostracods in the Silurian. There is an increase in the number of prey species in the Late Ordovician, which could be related to the Great Ordovician Biodiversification Event. Molluscs are the favourite prey taxon in the Ordovician, but in the Silurian, molluscs became less dominant as the prey. This is probably not an artefact of preservation as Ordovician and Silurian molluscs are equally well preserved.
Content may be subject to copyright.
Full Terms & Conditions of access and use can be found at
https://www.tandfonline.com/action/journalInformation?journalCode=ghbi20
Historical Biology
An International Journal of Paleobiology
ISSN: 0891-2963 (Print) 1029-2381 (Online) Journal homepage: https://www.tandfonline.com/loi/ghbi20
Predation in the Ordovician and Silurian of Baltica
Olev Vinn
To cite this article: Olev Vinn (2017) Predation in the Ordovician and Silurian of Baltica, Historical
Biology, 29:1, 11-16, DOI: 10.1080/08912963.2015.1092964
To link to this article: https://doi.org/10.1080/08912963.2015.1092964
Published online: 01 Oct 2015.
Submit your article to this journal
Article views: 147
View Crossmark data
Citing articles: 2 View citing articles
HISTORICAL BIOLOGY, 2017
VOL. 29, NO. 1, 11–16
http://dx.doi.org/10.1080/08912963.2015.1092964
Predation in the Ordovician and Silurian of Baltica
Olev Vinn
Department of Geology, University of Tartu, Tartu, Estonia
ABSTRACT
Signs of predation appear in the Middle Ordovician of Baltica. Shell repair dominates over the predatory
borings in the Ordovician and Silurian. Predators attacked molluscs, brachiopods and tentaculitoids in the
Ordovician and molluscs, tentaculitoids, brachiopods and ostracods in the Silurian. There is an increase
in the number of prey species in the Late Ordovician, which could be related to the Great Ordovician
Biodiversication Event. Molluscs are the favourite prey taxon in the Ordovician, but in the Silurian
molluscs became less dominant as the prey. This is probably not an artefact of preservation as Ordovician
and Silurian molluscs are equally well preserved.
© 2015 Informa UK Limited, trading as Taylor & Francis Group
KEYWORDS
Predators; prey; molluscs;
brachiopods; biological
interactions; early Palaeozoic
ARTICLE HISTORY
Received 8 August 2015
Accepted 8 September 2015
CONTACT Olev Vinn olev.vinn@ut.ee
1. Introduction
Vermeij (1977) demonstrated that predation has been an
important evolutionary factor throughout the Phanerozoic. It
is known that predation intensities increased with time from
the Paleozoic to the Recent (Vermeij 1977, 1987; Vermeij et al.
1981). is increase is reected by the morphological evolution
of the gastropod shell (Vermeij 1977, 1987; Vermeij et al. 1981).
ere have been two episodes of a general increase of predation
intensities during the Phanerozoic. e Silurian–Devonian was
described as the rst interval of general increase in predation
intensities (Signor and Brett 1984; Brett and Walker 2002; Brett
2003; Nagel-Myers et al. 2013). is event is commonly termed
as the Mid-Paleozoic revolution of predation (Signor and Brett
1984; Brett and Walker 2002; Nagel-Myers et al. 2013). More
recent quantitative analysis, which were based on combined
data on predatory borings and shell repair, suggests that there
was also a general increase in predation intensities in the
Ordovician (Huntley and Kowalewski 2007). In the Mesozoic
second major increase in predation intensities took place (Stanley
1974, 1977) and it is known as Mesozoic revolution of predation
(Vermeij 1977).
e Phanerozoic record of predation is mostly based on shell
repair and predatory borings in various shelly fossils. e earliest
predatory drillings occur in Cloudina from the Ediacaran. e
Paleozoic record of predation is mostly based on borings known
from the shells of brachiopods and a lesser number from mollusks
(Brett and Walker 2002; Brett 2003; Harper 2003; Huntley and
Kowalewski 2007). e major predator groups of marine inverte-
brates appeared and diversied in the Early Paleozoic. A consid-
erable number of new marine predators appeared by the Middle
Ordovician. ese predators include asteroids, various arthropods,
larger cephalopods, and probably also drilling gastropods (Brett
and Walker 2002). e priapulids, cephalopods, phyllocarids and
eurypterids were the most important durophagous predators (hard
shell feeding) in the Silurian (Brett and Walker 2002).
ere is an excellent record of shell repair and drilling pre-
dation for the Ordovician and Silurian of Baltica (Peel 1984;
Holmer 1989; Ebbestad and Peel 1997; Lindström and Peel 1997;
Ebbestad and Högström 2000; Kröger 2004; Vinn 2009, 2012).
However, overview papers on the early Palaeozoic predation of
Baltica are lacking. It is possible that some of the bivalve and
gastropod record of predation traces could be biased by the mode
of preservation of the shells; if they are recrystallized calcite or
silicied they show the traces much better than in moulds. Poor
preservation of the aragonitic shells constitutes a potential major
bias in fossil record of predation.
e aims of this paper are: (1) to analyse the distribution
of predation in the Ordovician and Silurian of Baltica, (2) to
analyse taxonomic composition of prey taxa, and (3) to discuss
the predation methods (i.e. drilling vs. durophagous predation).
2. Geological background
A shallow epicontinental sea covered Scandinavia, the modern
Baltic states and NW Russia in the Ordovician. e Ordovician
succession is relatively complete and represented mostly by
carbonate rocks except for the terrigenous Lower Ordovician
sequence. e climate in Baltica changed drastically during the
Ordovician, as the result of the dri of the continent from the
southern high latitudes to the tropical realm (Torsvik et al. 1992).
An increased sedimentation rate of carbonates was induced by
the warming of the climate. During the Late Ordovician deposits
that are characteristic of an arid and tropical climate appeared in
the sequence (Mõtus and Hints 2007). In the Early and Middle
Published online 01 Oct 2015
12 O. VINN
next most numerous prey taxa are the brachiopods (N=5) and
ostracods (N=3) in the Silurian.
3.4. Predation frequency
e highest shell repair frequencies occur in gastropods in the
Ordovician (Ebbestad and Peel 1997). In the Silurian the highest
shell repair frequencies occur in tentaculitoid tubeworms (Vinn
2012).
4. Discussion
4.1. Drilling vs. durophagous predation
e dominance of durophagous predation over drilling predation
in the Ordovician and Silurian of Baltica likely does not represent
a study bias. It is possible that shell breaking was the dominant
predation method in Ordovician and Silurian of Baltica. Drill-
ing predators were likely less numerous than the durophagous
predators in the Ordovician and Silurian. Drilling predation is
usually associated with predatory gastropods. ese gastropods
may have been relatively rare in the Orodovician and Silurian
marine communities. Durophagous predation is associated with
nautiloids, arthropods and sh. Nautiloids were abundant in the
Ordovician of Baltica. In the Silurian in addition to nautiloids
also large arthropods were common predators (Brett 2003).
4.2. Predation frequency in the Ordovician and Silurian
e highest shell repair frequencies occur in gastropods in the
Ordovician because gastropods presumably were favourite prey
taxon for the Ordovician predators in Baltica. Shell repair occurs
in 7 percent of gastropod individuals from the late Ordovician
Kullsberg and Boda limestones (Ebbestad and Peel 1997), which
is relatively low as compared to the maximal Silurian predation
frequences in Baltica. e highest Silurian shell repair frequences
(20–29% of individuals) in tentaculitoid tubeworms (Vinn
2009; Vinn 2012) possibly indicate their important role as prey.
However, molluscs are more abundant in the Silurian of Baltica
than the tentaculitoid tubeworms (Larsson 1979; Raukas and
Teedumäe 1997; Vinn 2009) and most likely they were more
important as the prey taxon for Silurian predators. e shell
repair frequencies of Late Ordovician (Ebbestad and Peel 1997)
and Silurian gastropods (Lindström and Peel 1997) were similar
indicating similar predation intensity.
4.3. Stratigraphic distribution of predation
e relatively steady number of prey species in the Darriwilian
to Katian indicates that there was not rapid increase in number
of prey species in the Ordovician. However, there are remarkably
more prey species in the Late Ordovician than in the Middle
Ordovician. Both Middle and Late Ordovician of Baltica are
equally well studied and this change in the number of prey spe-
cies is presumably not an artefact of sampling. us, most likely
predators aected more species in the Late Ordovician than in
the Middle Ordovician. e lack of predation records from the
Early Ordovician and Dapingian presumably does not represent
a study bias as the Middle Ordovician to Late Ordovician and
Ordovician Baltica was situated in a temperate climate zone and
these types of deposits were lacking (Jaanusson 1973). e rst
signs of warm climate in the early Katian include the appearance
of tabulate corals, stromatoporoids and reefs. However, it was
not until the Hirnantian that they became prevalent (Mõtus and
Hints 2007).
During the Silurian the Baltica was located in equatorial
latitudes (Melchin et al. 2004). An epicontinental basin cov-
ered the area of Scandinavia and the Baltic states. The Silurian
basin was characterized by diverse biotas and a wide range
of tropical environments including reefs and lagoons (Hints
2008). Nestor and Einasto (1977) described five main envi-
ronments in that basin: tidal flat/lagoonal, shoal, open shelf,
transitional (basin slope), and a basin depression. The first
three environments formed a carbonate platform (Raukas and
Teedumäe 1997).
3. Results
3.1. Drilling predation vs. durophagous predation
Predatory borings appear in the Darriwilian of Baltica
(Table 1). Predatory borings occur in the lingulate brachiopods
in the Ordovician (Figure 1) and in the ostracods and rhyn-
chonelliform brachiopods in the Silurian (Table 1). Earliest signs
of durophagous predation are known from the Darriwilian (Table
1). Shell repair occurs in molluscs, brachiopods and tentaculitoids
in the Ordovician and Silurian of Baltica (Table 1) (Figures 2–4).
e Ordovician and Silurian record of predation is dominated by
shell repair (Table 1).
3.2. Stratigraphic distribution of predation
Number of prey species is relatively evenly distributed in the
Ordovician (Darriwilian to Katian) and Silurian (Wenlock to
Ludlow). ere is increase in the number of known prey species
from Middle Ordovician (nine species) to Late Ordovician (18
species). e entire Early Ordovician and Dapingian lacks the
record of predation. ere is a gap in the predation record in
latest Ordovician (Hirnantian) and early Silurian (Llandovery).
e number of prey species is lower in the Pridoli (N=2) than
in the Wenlock (N=10) and Ludlow (N=8).
3.3. Taxonomic composition of prey
Ordovician and Silurian prey are equally diverse (Table 2). e
taxonomic composition of prey in the Ordovician (i.e. gastro-
pods, nautiloids, tergomyans, inarticulate brachiopods, articulate
brachiopods, cornulitids) is somewhat dierent from that of the
Silurian (i.e. gastropods, bivalves, tentaculitids, cornulitids, Anti-
calyptraea, articulate brachiopods, ostracods). e Ordovician
record of predation is dominated by the molluscs (20 species;
incl. gastropods, nautiloids and tergomyans). Gastropods are
favourite prey among the Ordovician molluscs in number of
species. Brachiopods (six species) and tentaculitoids (one spe-
cies) form a minor part in the Ordovician predation record. In
the Silurian molluscs became less dominant than in the Ordo-
vician. Molluscs (N=6) and tentaculitoids (N=6) are the most
numerous among the prey species of the Silurian of Baltica. e
HISTORICAL BIOLOGY 13
Table 1.Distribution of predatory borings and shell repair in the Ordovician and Silurian of Baltica.
Prey
Type (drilling
vs. shell repair)
% of
speci-
mens
with
traces Age Locality References
Shaleria ornatella (Davidson)
(brachiopod)
Shell repair Pridoli Kaugatuma, Estonia Rubel (2011)
Anticalyptraea calyptrata (Eichwald
1860) (tentaculitoid tubeworm)
Shell repair 29% Pridoli Kaugatuma, Estonia Vinn (2012)
Lepidoleptaena poulseni (brachiopod) Shell repair Ludlow Gotland, Sweden Hoel (2014)
Odessites majstriensis (tentaculitid) Shell repair Ludlow Kättelviken, Sweden Larsson (1979)
Lonchidium extensum (tentaculitid) Shell repair Hamra beds, Ludlow Bankvät, Sweden Larsson (1979)
Oehlertia gradate (gastropod)
(Lindström 1884) (gastropod)
Shell repair Ludlow Gotland, Sweden Lindström and Peel (2005)
Crenilunula limata (Lindström 1884)
gastropod)
Shell repair Ludlow Gotland, Sweden Lindström and Peel (2005)
Hammariella pulchrivelata (ostracod) Boring Ludlow Hammarudden, Sweden Martinson (1962)
Neobeyrichia nutans (ostracod) Boring Ludlow Hammarudden, Sweden Mar tinson (1962)
Beyrichia globifera (ostracod) Boring Ludlow Hammarudden, Sweden Martinson (1962)
Cornulites cellulosus (Herringshaw et al.
2007) (cornulitid)
Shell repair 20.7% Sheinwoodian (Wenlock) Saaremaa, Estonia Vinn (2009)
Leptaena purpurea Cocks (brachiopod) Shell repair Sheinwoodian (Wenlock) Lätiküla, Estonia Rubel (2011)
Leptaena depressa visbyensis
(brachiopod)
Shell repair Visby Formation, Wenlock Gotland, Sweden Hoel (2014)
Leptaena depressa depressa
(brachiopod)
Boring Halla Formation, Wenlock Gotland, Sweden Hoel (2014)
Volonites muldiensis (tentaculitid) Shell repair Wenlock Mulde Tegelbruk, Sweden Larsson (1979)
Seretites? versabilis (tentaculitid) Shell repair Wenlock Follingbo, Sweden Larsson (1979)
Euomphalopterus (gastropod) Shell repair Wenlock Gotland, Sweden Peel (1984)
Poleumita sp. (gastropod) Shell repair 10% Wenlock Gotland, Sweden Lindström and Peel (1997)
Lophospira gothlandica (Ulrich in Ulrich
and Scofield 1897) (gastropod)
Shell repair Wenlock Gotland, Sweden Lindström and Peel (2005)
Nuculodonta gotlandica (Bivalve) Shell repair Wenlock Gotland, Sweden Liljedahl (1985)
Harjumena schmidti Gagel
(brachiopod)
Shell repair Late Ordovician? Estonia Rõõmusoks (2004)
Leptaena rugosa Dalman (brachiopod) Shell repair Dalmanitina beds (Late
Ordovician)
Sweden Rõõmusoks (2004)
Salpingostoma Megalostoma
(gastropod)
Shell repair Erratic, Ordovician Bromberg, Germany Ebbestad et al. (2009)
Strophomena? arachnoidea (Lindström
1880) (brachiopod)
Shell repair Fjäkka Formation (Katian) Dalarna, Sweden Ebbestad and Högström
(2000)
Ruedemannia sp. (gastropod) Shell repair Boda Limestone (Katian) Kallholn, Sweden Ebbestad and Peel (1997)
Holopea harpa (Lindström 1880)
(gastropod)
Shell repair Boda Limestone (Katian) Kallholn, Sweden Ebbestad and Peel (1997)
Holopea sp. 1 (gastropod) Shell repair Boda Limestone (Katian) Osmundsberget, Sweden Ebbestad and Peel (1997)
Holopea sp.2 (gastropod) Shell repair Boda Limestone (Katian) Kallholn, Sweden Ebbestad and Peel (1997)
Holopea? sp. (gastropod) Shell repair Boda Limestone (Katian) Osmundsberget, Sweden Ebbestad and Peel (1997)
Gyronema rupestre? (Eichwald 1842)
(gastropod)
Shell repair Boda Limestone (Katian) Kallholn, Sweden Ebbestad and Peel (1997)
Platyceras canaliculatum (Lindström
1880) (gastropod)
Shell repair Boda Limestone (Katian) Kallholn, Sweden Ebbestad and Peel (1997)
Megalomphala crassiuscula
(gastropod)
Shell repair Herøya Formation (Katian) Skien, Nor way Ebbestad et al. (2009)
Cornulites sp. (cornulitid) Shell repair Katian Northern Estonia Vinn (2009)
Cyrtonotreta striata (lingulate
brachiopod)
Boring Dalby Limestone (Sand-
bian)
Sweden Holmer (1989)
Cyrtonotreta sp. (lingulate brachiopod) Boring Dalby Limestone (Sand-
bian)
Sweden Holmer (1989)
Bucania gracillima (Koken 1896)
(gastropod)
Shell repair Kullesberg Limestone
(Sandbian?)
Amtjern, Sweden Ebbestad and Peel (1997)
and Ebbestad (1998)
Straparollus (Euomphalus) obtusangu-
lus (Lindström 1880) (gastropod)
Shell repair Kullesberg Limestone
(Sandbian?)
Kullesberg, Sweden Ebbestad and Peel (1997)
Bucania czekanowskii (Schmidt 1858)
(gastropod)
Shell repair Viivikonna Formation
(Sandbian)
Vanamõisa, Estonia Isakar and Ebbestad (2000)
Bucania erratica (Frisk and Ebbestad
2007) (gastropod)
Shell repair Dalby Limestone (Sand-
bian)
Tvären, Sweden Frisk and Ebbestad (2007)
Conotreta? siljaensis (lingulate
brachiopod)
Boring 5% Ryd Limestone (Darriwilian) Sweden Holmer (1989)
(Continued)
14 O. VINN
best be explained by a sampling bias. e Hirnantian and Lla-
ndovery faunas of Baltica are best studied in Estonia (Raukas
and Teedumäe 1997), but most previous research was devoted
to stratigraphy and taxonomy and not to paleoecology. However,
Early Ordovician invertebrates are equally well studied (Raukas
and Teedumäe 1997). It is possible that the predation intensi-
ties increased in the Middle Ordovician of Baltica. e lack of
predation records from the Hirnantian and Llandovery could
Figure 1.Possible predatory boring in dorsal valve of Conotreta? siljanensis from Ryd Limestone, Ordovician, Sweden after Holmer (1989).
Figure 2.Line drawings of multiple shell repairs in Bucania czekanowskii (Schmidt 1858) based on specimen TUG 666–37 modified after Isakar and Ebbestad (2000).
Prey
Type (drilling
vs. shell repair)
% of
speci-
mens
with
traces Age Locality References
Scaphelasma mica (lingulate brachi-
opod)
Boring 2% Ryd Limestone (Darriwilin) Sweden Holmer (1989)
Joleaudella sphenonotus (gastropod) Shell repair Elnes Formation (Darri-
wilian)
Asker, Norway Ebbestad et al. (2009)
Baltiscanella christianiae (Tergomya) Shell repair Elnes Formation (Darri-
wilian)
Helgøya, Norway Ebbestad et al. (2009)
Orthoceras regulare (Schlotheim 1820)
(nautiloid)
Shell repair Grey Orthoceratite Lime-
stone (Darriwilian)
Vilm, Germany Kröger (2004)
Orthoceras sp. (nautiloid) Shell repair Upper Grey Orthoceratite
Limestone (Darriwilian)
Rügen, Germany Kröger (2004)
Nilssonoceras nilssoni (Boll 1857)
(nautiloid)
Shell repair Upper Red Orthoceratite
Limestone (Darriwilian)
Gransee, Germany Kröger (2004)
Plagiostomoceras laevigatum (Boll
1857) (nautiloid)
Shell repair Upper Red Orthoceratite
Limestone (Darriwilian)
Öland, Sweden Kröger (2004)
Orthoceras scabridum (Angelin 1880)
(nautiloid)
Shell repair Upper Grey Orthoceratite
Limestone (Darriwilian)
Öland, Sweden Kröger (2004)
Table 1. (Continued)
HISTORICAL BIOLOGY 15
of prey in the Ordovician as compared to the Silurian of Baltica
(Table 2). Predators attacked selectively more mollusc species
than barchiopods in the Ordovician and Silurian of Baltica
(Table 2). Tentaculitoid species were also selectively more
attacked by predators than the brachiopods in the Silurian of
Baltica (Table 2). It is possible that predators selected more oen
mollusc and tentaculitoid prey than brachiopods because of their
voluminous so body.
4.5. GOBE and O/S mass extinction
e increase in prey species number in the Late Ordovician of
Baltica could be explained by the Great Ordovician Biodiver-
sication Event (GOBE). During the GOBE probably also the
predators diversied. Increased number of predators presumably
it is possible that the predation intensities in the Hirnantian and
Llandovery may have been lower than in the Katian and Wen-
lock. e low number of prey species in the Pridoli presumably
represents also a study bias as there is only a small number Pridoli
fossil localities in Baltica. It is possible that variable preservation
of aragonitic shells may alter perceptions of relative frequency
of predation traces.
4.4. Taxonomic composition and selection of prey
Predators aected similar number of invertebrate higher taxa in
the Ordovician (N=6) and Silurian (N=5) of Baltica (Table 1).
However, in the Ordovician predators preferred molluscs over
the other prey species (Tables 1 and 2). It is possible that pre-
dation aected molluscs relatively more than the other groups
Figure 3.Leptaena depressa visbyensis,Gnisvärd, Upper Visby Formation (Sheinwoodian); repaired crescentic wound after Hoel (2014).
Figure 4.Shell repair in Anticalyptraea calyptrata from Kaugatuma Formation (Pridoli), Saaremaa, Estonia, after Vinn (2012).
Table 2.Total numbers of species of gastropods, clams, nautiloids etc. at various levels and numbers of taxa that actually display traces of predation.
Note: Numbers of species derived from Paleobiology Database (2015), Raukas and Teedumäe, Rõõmusoks (1970), Kaljo (1970), and Larsson (1979).
Stratigraphy Gastropods Clams Nautiloids Tentaculitoids Brachiopods
Pridoli 9 3 5 6 (N=1, 16.6%) 15 (N=1, 6.6%)
Ludlow 57 (N=2, 3.5%) 10 15 18 (N=2, 11.1%) 70 (N=1, 1.4%)
Wenlock 83 (N=3, 3.6%) 11 (N=1, 9.1%) 10 21 (N=3, 14.3%) 80 (N=3, 3.8%)
Llandovery 30 2 2 5 40
Hirnantian 2 7 9 1 35
Katian 45 (N=8, 17.7%) 130 50 6 (N=1, 16.6%) 65 (N=1, 1.5%)
Sandbian 30 (N=4, 13.3%) 26 10 2 90 (N=2, 2.2%)
Darriwilian 40 (N=2, 5.0%) 10 30 (N=5, 16.6%) 1 50 (N=2, 4.0%)
16 O. VINN
Kröger B. 2004. Large shell injuries in Middle Ordovician Orthocerida
(Nautiloidea, Cephalopoda). GFF. 126:311–316.
Larsson K. 1979. Silurian tentaculitids from Gotland and Scania. Fossil
Strata. 11:1–180.
Lindström A, Peel JS. 1997. Failed predation and shell repair in the
gastropod Poleumita from the Silurian of Gotland. Sweden Bull Czech
Geol Surv. 72:115–126.
Lindström A, Peel JS. 2005. Repaired injuries and shell form in some
Palaeozoic pleurotomarioid gastropods. Acta Palaeont Pol. 50:697–704.
Liljedahl L. 1985. Ecological aspects of a silicied bivalve fauna from the
Silurian of Gotland. Lethaia. 18:53–66.
Martinson A. 1962. Ostracods of the family Beyrichiidae from the Silurian
of Gotland. Bull Geol Inst Univ Uppsala. 41:1–369.
Melchin MJ, Cooper RA, Sandler PM. 2004. e Silurian period. In:
Gradstein FM, Ogg JG, Smith AG, editors. A geologic time scale 2004.
Cambridge: Cambridge University Press; p. 188–201.
Mõtus MA, Hints O, editors. 2007. Excursion Guidebook. In: 10th
International Symposium on Fossil Cnidaria and Porifera. Excur-sion
B2: Lower Paleozoic geology and corals of Estonia. August 18–22, 2007.
Institute of Geology at Tallinn University of Technology; 66 p.
Nagel-Myers J, Dietl G, Handley J, Brett CE. 2013. Abundance is not
enough: the need for multiple lines of evidence in testing for ecological
stability in the fossil record. PLoS One. 8(5):e63071.
Nestor H, Einasto R. 1977. Model of facies and sedimentology for
Paleobaltic epicontinental basin. In: Kaljo DL, editor. Facies and fauna
Silurian of Baltica. Tallinn: Institute of Geology AN ESSR; p. 89–121
[in Russian].
Peel JS. 1984. Attempted predation and shell repair in Euomphalopterus
(Gastropoda) from the Silurian of Gotland. Bull Geol Soc Den. 32:
163–168.
Raukas A, Teedumäe A. 1997. Geology and mineral resources of Estonia.
436 pp. Tallinn: Estonian Academy Publishers.
Rõõmusoks A. 2004. Ordovician strophomenoid brachiopods of northern
Estonia. Fossilia Baltica. 3:1–151.
Rubel M. 2011. Silurian brachiopods Dictyonellida, Strophomenida,
Productida, Orthotetida, Protorthida and Orthida from Estonia.
Fossilia Baltica. 4:1–65.
Signor PW, Brett CE. 1984. e mid-Paleozoic precursor to the Mesozoic
marine revolution. Paleobiology. 10:229–245.
Stanley SM. 1974. What has happened to the articulate brachiopods. Abstr
Programs Geol Soc Am. 6:966–967.
Stanley SM. 1977. Trends, rates, and patterns of evolution in the Bivalvia.
In: Hallam A, editor. Patterns of evolution, as illustrated by the fossil
record. Amsterdam: Elsevier; p. 209–250.
Schmidt F. 1858. Untersuchungen ber die Silurische Formation von
Ehstland [Investigations ber the Silurian formation of Ehstland],
Nord-Livland und Oesel. Archiv für die Naturkunde Liv-, Ehst- und
Kurlands, 2: 1–248, Dorpat.
Torsvik TH, Smethurst MA, Van der Voo R, Trench A, Abrahamsen
N, Halvorsen E. 1992. Baltica. A synopsis of vendian-permian
palaeomagnetic data and their palaeotectonic implications.Earth Sci
Rev. 33:133–152.
Vermeij GJ. 1977. e Mesozoic marine revolution: evidence from snails,
predators and grazers. Paleobiology. 3:245–258.
Vermeij GJ. 1987. Evoluton and escalation: an ecological history of life.
Princeton (NJ): Princeton University Press. 527 pp.
Vermeij GJ, Schindel DE, Zipser E. 1981. Predation through geological
time: evidence from gastropod shell repair. Science. 214:1024–1026.
Vinn O. 2009. Attempted predation on Early Paleozoic cornulitids.
Palaeogeogr Palaeoclimatol Palaeoecol. 273:87–91.
Vinn O. 2012. Shell repair in Anticalyptraea (Tentaculita) in the Late
Silurian (Pridoli) of Baltica. Carnets de Géologie CG2012_L01.
p. 31–37.
aected the larger number of prey species. e lack of predation
records from the Hirnantian and Llandovery of Baltica presum-
ably represents a study bias (Table 2). However, it is possible that
the number of predators decreased during the O/S mass extinc-
tion and fewer predators aected smaller number of prey species.
Acknowledgements
is paper is a contribution to IGCP 591 ‘e Early to Middle Palaeozoic
Revolution. I am grateful to C. E. Brett and an anonymous reviewer for the
constructive reviews.
Disclosure statement
No potential conict of interest was reported by the author.
Funding
Financial support to O.V. was provided by the Palaeontological Associa-
tion Research Grant, Sepkoski Grant (Paleontological Society); Estonian
Research Council projects ETF9064 and IUT20-34.
References
Brett CE. 2003. Durophagous predation in Paleozoic marine benthic
assemblages. In: Kelley P, Kowalewski M, Hansen T, editors. Predator-
prey interactions in the fossil record. Lawrence (KS): Kluwer Academic-
Plenum Publishers; p. 401–432.
Brett CE, Walker SE. 2002. Predators and predation in Paleozoic marine
environments. In: Kowalewski M, Kelley PH, editors. e fossil record
of predation. Vol. 8. Paleontol Soc Pap; p. 93–118.
Ebbestad JOR. 1998. Multiple attempted predation in the Middle
Ordovician gastropod Bucania gracillima. GFF. 120:27–33.
Ebbestad JOR, Högström AES. 2000. Shell repair following failed predation
in two Upper Ordovician brachiopods from central Sweden. GFF.
122:307–312.
Ebbestad JOR, Lindström A, Peel JS. 2009. Predation on bellerophontiform
molluscs in the Palaeozoic. Lethaia. 42:469–485.
Ebbestad JOR, Peel JS. 1997. Attempted predation and shell repair in
Middle and Upper Ordovician gastropods from Sweden. J Paleont.
71:1007–1019.
Frisk Å, Ebbestad JOR. 2007. Paragastropoda, tergomya and gastropoda
from the late Ordovician dalby limestone of Sweden. GFF. 129:81–99.
Harper EM. 2003. Assessing the importance of drilling predation over
the Palaeozoic and Mesozoic. Palaeogeogr Palaeoclimatol Palaeoecol.
201:185–198.
Hints O. 2008. e Silurian system in Estonia. In: Hints O, Ainsaar
L, Männik P, Meidla T, editors. e seventh Baltic stratigraphical
conference. Abstracts and eld guide: Geological Society of Estonia,
Tallinn; p. 46.
Hoel OA. 2014. Palaeobiology of Silurian Leptaeninae (Brachiopoda) from
Gotland, Sweden. Paleont J. 2014(1–2):1–14.
Holmer LE. 1989. Middle Ordovician phosphatic inarticulate brachiopods
from Våstergotland and Dalarna, Sweden. Fossil Strata. 26:1–172.
Huntley JW, Kowalewski M. 2007. Strong coupling of predation intensity
and diversity in the Phanerozoic fossil record. Proc Natl Acad Sci.
104:15006–15010.
Isakar M, Ebbestad JOR. 2000. Bucania (Gastropoda) from the Ordovician
of Estonia. Paläont Z. 74:51–68.
Jaanusson V. 1973. Aspects of carbonate sedimentation in the Ordovician
of Baltoscandia. Lethaia. 6:11–34.
... Durophagous predators first appeared at low latitudes in the early Cambrian (Skovsted et al. 2007). Shell repair dominates over the predatory borings in the Ordovician and Silurian of Baltica (Vinn 2017), which is different from the Cambrian where borings are slightly more common than scar, presumably because durophagous taxa were more diverse later on than in the Cambrian. Predators attacked molluscs, brachiopods and tentaculitoids in the Ordovician of Baltica and molluscs, tentaculitoids, brachiopods and ostracods in the Silurian of Baltica (Vinn 2017). ...
... Shell repair dominates over the predatory borings in the Ordovician and Silurian of Baltica (Vinn 2017), which is different from the Cambrian where borings are slightly more common than scar, presumably because durophagous taxa were more diverse later on than in the Cambrian. Predators attacked molluscs, brachiopods and tentaculitoids in the Ordovician of Baltica and molluscs, tentaculitoids, brachiopods and ostracods in the Silurian of Baltica (Vinn 2017). Trilobites were also important as prey in the Ordovician (Owen 1985). ...
... This is somewhat different from the situation in the Cambrian. Molluscs are the favorite prey taxon in the Ordovician of Baltica (Vinn 2017). Thus, Cambrian predation record is more brachiopod and trilobite dominated than the Ordovician record. ...
Article
Full-text available
Series two marks a revolution in Cambrian predation when new predators and new predation methods appeared, which led to general increase in predation intensities and in the diversity of prey groups. The number of bored taxa and taxa with the predation scars is similar in the Cambrian. Most of the borings are associated with brachiopods and most of the scars with trilobites. Brachiopods, arthropods, molluscs, cnidarians and echinoderms were the most common prey in the Cambrian. The Cambrian record of predation is dominated by damage inflicted on brachiopods and trilobites. The fossils with predation signs are known from a majority of paleocontinents and all the Cambrian series.
... However, malformations are known from the fossil record too, and have been reported from individuals of different fossil groups, including foraminifera (Ballent & Carignano, 2008), trilobites (Owen, 1985;Babcock, 1993), brachiopods (Copper, 1967;He et al., 2017), bivalves (Savazzi, 1995), gastropods (Lindström & Peel, 2010), cephalopods (De Baets, Keupp & Klug, 2015;Hoffmann & Keupp, 2015;Mironenko, 2016;De Baets, Hoffmann & Mironenko, 2021), echinoderms (Thomka, Malgieri & Brett, 2014), graptolites (Han & Chen, 1994), insects (Vršanský, Liang & Ren, 2012), conodonts (Weddige, 1990), shark teeth (Itano, 2013), amphibians (Witzmann et al., 2013), reptiles (Buffetaut et al., 2007), primate teeth (Tougard & Ducrocq, 1999), and plankton (Vandenbroucke et al., 2015;Bralower & Self-Trail, 2016). In addition to gene mutations or embryonic developmental disorders, malformed fossils may also have resulted from healed injuries and pathology (Owen, 1985;Babcock, 1993;Kelley, Kowalewski & Hansen, 2003;Vinn, 2017Vinn, , 2018. Malformed fossils provide important evidence of both organisms-organisms and organisms-environment relationships during geological history. ...
... For the specific identity of endoparasites, shell-less organisms and microorganisms, it is difficult to confirm because of their poor preservation or they are indiscernible in the fossil record. Some brachiopod malformations were caused by predators, which presented the fractures, indentations, and scars on their shells, and were often accompanied by signs of repair (e.g., Alexander, 1986;Kowalewski, Flessa & Marcot, 1997;Happer, 2005;Vinn, 2017). In these malformed athyrid shells from western Junggar, except for a pair of indentations on the opposite valves of specimen BGEG-CLJ01 (Fig. 1H), no wounds or scars were found on other specimens. ...
Article
Full-text available
Although malformations are found in both extant organisms and the fossil record, they are more rarely reported in the fossil record than in living organisms, and the environmental factors causing the malformations are much more difficult to identify for the fossil record. Two athyrid brachiopod taxa from the Upper Devonian Hongguleleng Formation in western Junggar (Xinjiang, NW China) show distinctive shell malformation. Of 198 Cleiothyridina and 405 Crinisarina specimens, 18 and 39 individuals were malformed, respectively; an abnormality ratio of nearly 10%. Considering the preservation status and buried environment of the abnormal specimens, and analysis of trace elements and rare earth elements from whole-rock and brachiopod shells, we conclude that the appearance of malformed athyrids is likely related to epi/endoparasites, or less likely the slightly higher content of heavy metal in the sea.
... The pre-Cretaceous record of drilling predation has long been considered as scarce, discontinuous and questionable, but recent works document a much richer fossil record in the Paleozoic and early Mesozoic. Drill holes have since then been reported from the Tonian (Porter, 2016), Ediacaran (Bengtson and Zhao, 1992;Brain, 2001;Hua et al., 2003;Becker-Kerber et al., 2017), Cambrian (e.g., Miller and Sundberg, 1984;Vinn et al., 2021), Ordovician (e.g., Rohr, 1991;Vinn, 2017) and Silurian (e.g., Rohr, 1976;Liljedahl, 1985;Vinn, 2017). Drill holes illustrating preyspecies preferences and site stereotypy tentatively attributable to platyceratid gastropods have been reported from the Cambrian onwards in brachiopods and from the Ordovician onwards in molluscs . ...
... The pre-Cretaceous record of drilling predation has long been considered as scarce, discontinuous and questionable, but recent works document a much richer fossil record in the Paleozoic and early Mesozoic. Drill holes have since then been reported from the Tonian (Porter, 2016), Ediacaran (Bengtson and Zhao, 1992;Brain, 2001;Hua et al., 2003;Becker-Kerber et al., 2017), Cambrian (e.g., Miller and Sundberg, 1984;Vinn et al., 2021), Ordovician (e.g., Rohr, 1991;Vinn, 2017) and Silurian (e.g., Rohr, 1976;Liljedahl, 1985;Vinn, 2017). Drill holes illustrating preyspecies preferences and site stereotypy tentatively attributable to platyceratid gastropods have been reported from the Cambrian onwards in brachiopods and from the Ordovician onwards in molluscs . ...
Article
Drill holes are trace fossils relatively common on post-Paleozoic ostracods (especially from the Cretaceous onwards) and usually ascribed to predation by Muricidae and Naticidae gastropods. In the last few decades, increasing reports of these marks on Late Paleozoic and Early Mesozoic ostracods appeared in the literature. In this paper, we review the drill holes on marine and mixohaline ostracods with emphasis on the Permian and Triassic intervals, based on a detailed analysis of their published records, as well as new specimens and data. Based on ichnological principles, analyzes of several types of marks observed on ostracods are carried out to distinguish holes caused by predation on live individuals from the effects of dissolution and post-mortem bioerosion. The twenty-two marks identified as drill holes in this work are classified into nine types, ranging from Kungurian-Roadian to the Rhaetian in age. The diversity of drill hole patterns supplies new evidence that the end-Permian biotic crisis influenced not only the global ostracod diversity, but also the trophic interactions. In the Late Triassic, the drilling predators were already deterred by increased thickness or complex ornament of bairdiid shells. Although the data do not permit precise identification of drilling predators, it is assumed that different animal groups might have performed this action throughout geologic time, due to the diversity of paleoenvironments and broad chronostratigraphic occurrence of drill holes analyzed. Potential drilling predators both in marine and mixohaline (paleo)environments are briefly discussed.
... A detailed compilation and analysis of evidence for Phanerozoic predation by Huntley andKowalewski (2007, but see López-Villalta, 2016) showed that predation intensity increased during the Ordovician, tracking increasing diversity levels. Intriguingly, a regional study examining predation patterns in the Ordovician of Baltica (Vinn, 2017) could not document Ordovician predatory trace fossils from strata lower than the Darriwilian, in which evidence is preserved of both borings and durophagy. ...
... A detailed compilation and analysis of evidence for Phanerozoic predation by Huntley andKowalewski (2007, but see López-Villalta, 2016) showed that predation intensity increased during the Ordovician, tracking increasing diversity levels. Intriguingly, a regional study examining predation patterns in the Ordovician of Baltica (Vinn, 2017) could not document Ordovician predatory trace fossils from strata lower than the Darriwilian, in which evidence is preserved of both borings and durophagy. ...
Article
The Ordovician Period records an extraordinary biodiversity increase known as the Great Ordovician Biodiver- sification Event (GOBE), which coincided with a series of environmental changes to the Earth System, notably a cooling global ocean, increased oxygenation, and increased nutrient supply from volcanism and continental weathering. The co-evolution of Earth and its biota during this interval has been studied in various contexts on multiple paleocontinents. Emerging patterns depend on the lens of investigation. Here we summarize the current state of understanding by reviewing and synthesizing the fossil and sedimentary records. Recent paleontological studies, mainly focused on rhynchonelliform (articulated) brachiopods, bryozoa, cephalopods, trilobites, graptolites, echinoderms, and reef organisms, have documented details of diversification, body size increase, development of ecosystem complexity, and intensification of inter-continental dispersal from the late Cambrian through Late Ordovician. Biomass increased markedly between the Early and Middle Ordovi- cian. Furthermore, diversification rates increase statistically during the mid-Darriwilian Age both globally and regionally. Coincident with these biotic changes, geochemical proxies record significant changes to Earth's physical sys- tem. Oceanic temperatures decreased, and atmospheric oxygen levels increased to near modern levels in con- cert with the Middle Ordovician diversification of shelly fauna. Anoxic pulses ceased and evidence for deep ocean ventilation prevails in Middle Ordovician strata. Furthermore, a major Middle–Late Ordovician change in oceanic strontium isotopic composition indicates increased weathering of juvenile volcanic rocks and delivery of nutrients to marine settings. This multi-proxy dataset records near-simultaneous changes in fossil-rich shallow marine environments during exactly the interval of greatest diversification. By integrating biotic and geochemical datasets, a clear picture of the co-evolution of Earth and its biota emerges indicating that the Darriwilian was the critical interval of the GOBE. We recommend restricting the term “GOBE” to indicate this short interval of rapid diversification and ecosystem change, and using “Ordovician Radiation” when referencing the sum of diversifications that occurred throughout the Ordovician Period.
Article
While it is well established that the shapes and sizes of shells are strongly phylogenetically controlled, little is known about the phylogenetic constraints on shell thickness. Yet, shell thickness is likely to be sensitive to environmental fluctuations and has the potential to illuminate environmental perturbations through deep time. Here we systematically quantify the thickness of the anterior brachiopod shell which protects the filtration chamber and is thus considered functionally homologous across higher taxa of brachiopods. Our data come from 66 genera and 10 different orders and shows well‐defined upper and lower boundaries of anterior shell thickness. For Ordovician and Silurian brachiopods we find significant order‐level differences and a trend of increasing shell thickness with water depth. Modern (Cenozoic) brachiopods, by comparison, fall into the lower half of observed shell thicknesses. Among Ordovician–Silurian brachiopods, older stocks commonly have thicker shells, and thick‐shelled taxa contributed more prominently to the Great Ordovician Biodiversification but suffered more severely during the Late Ordovician Mass Extinction. Our data highlight a significant reduction in maximum and minimum shell thickness following the Late Ordovician mass extinction. This points towards stronger selection pressure for energy‐efficient shell secretion during times of crisis.
Article
Predation on ancient shelled prey is an often-studied topic in paleoecology, but the early Paleozoic and the brachiopods that dominated the seafloor at that time are relatively underrepresented in the predation literature. We assessed predatory repair scar frequencies among the brachiopod genera from the Early Richmondian (Late Ordovician) Oregonia Member (Arnheim Formation) near Flemingsburg, Kentucky. We found higher repair frequencies on the concavo-convex Rafinesquina and Leptaena relative to the bi-convex genera. There were no trends in repair frequency through the stratigraphic section and no relationships between repair frequency and community diversity metrics. It is possible that concavo-convex brachiopods’ flat shape, thin shell profile, and free-lying (no pedicle attachment) lifestyle made them more likely or appealing targets of Ordovician crushing predators. It is also possible that concavo-convex brachiopods were better suited to survive crushing attacks than biconvex taxa. We also found differences in shell ornament that may influence the visibility of repair scars.
Article
Full-text available
Tertiary and Recent marine gastropods include in their ranks a complement of mechanically sturdy forms unknown in earlier epochs. Open coiling, planispiral coiling, and umbilici detract from shell sturdiness, and were commoner among Paleozoic and Early Mesozoic gastropods than among younger forms. Strong external sculpture, narrow elongate apertures, and apertural dentition promote resistance to crushing predation and are primarily associated with post-Jurassic mesogastropods, neogastropods, and neritaceans. The ability to remodel the interior of the shell, developed primarily in gastropods with a non-nacreous shell structure, has contributed greatly to the acquisition of these antipredatory features. The substantial increase of snail-shell sturdiness beginning in the Early Cretaceous has accompanied, and was perhaps in response to, the evolution of powerful, relatively small, shell-destroying predators such as teleosts, stomatopods, and decapod crustaceans. A simultaneous intensification of grazing, also involving skeletal destruction, brought with it other fundamental changes in benthic community structure in the Late Mesozoic, including a trend toward infaunalization and the disappearance or environmental restriction of sessile animals which cannot reattach once they are dislodged. The rise and diversification of angiosperms and the animals dependent on them for food coincides with these and other Mesozoic events in the marine benthos and plankton. The new predators and prey which evolved in conjunction with the Mesozoic reorganization persisted through episodes of extinction and biological crisis. Possibly, continental breakup and the wide extent of climatic belts during the Late Mesozoic contributed to the conditions favorable to the evolution of skeleton-destroying consumers. This tendency may have been exaggerated by an increase in shelled food supply resulting from the occupation of new adaptive zones by infaunal bivalves and by shell-inhabiting hermit crabs. Marine communities have not remained in equilibrium over their entire geological history. Biotic revolutions made certain modes of life obsolete and resulted in other adaptive zones becoming newly occupied.
Article
Pleurotomarioid gastropods typically develop a spiral band called the selenizone in the outer whorl face of the shell that is formed by the closure of an open slit in the apertural margin. The slit and selenizone may be important in controlling the extent to which fractures induced by predatory attacks propagate across the whorl surface. A prominent selenizone can prevent fractures from traversing the entire whorl. Study of six Palaeozoic pleurotomarioid gastropod species with repaired shell injuries shows that repaired injuries are dependent on both the nature of the selenizone and shell form. The species can be divided into three morphological groups (turbiniform, trochiform and planispiral) and show a variety of selenizones with different degrees of prominence Turbiniform shells show more repaired injuries than planispiral forms, indicating that species in the former group more often survive predatory attacks. The studied material is too sparse for meaningful statistical analysis, but individual case studies suggest that the combined influence of shell form and the nature of the selenizone can make the interpretation complex.
Article
Shell breakage and subsequent repair are desribed in Poleumita Clarke and Ruedemann, 1903, a widely distributed Silurian gastropod, from the Hogklint Beds and Slite Beds of Gotland, Sweden. The specimens display a variety of shell injuries interpreted as non-lethal attacks by an unknown predator, but the identity of the attacker remains obscure. About 10% of the examined 430 specimens preserve repaired significant injuries, mainly in growth stages larger than about 20 mm diameter; larger specimens often show repeated damage and repair. Injuries are typically arcuate breaks in the outer and lower whorl surfaces penetrating a short distance adapically from the apertural margin at the time of the attack.
Article
Most minerals are of sedimentary origin. Bauxite is an important resource and oil, natural gas, copper, manganese and iron ores also occur, as do other radio-active and non-metallic ores. Much of the bauxite is found conveniently close to coal reserves and the Danube, providing power and transport. Well developed karst causes problems in mining and the characteristics and location of the resources are described. The country has an ambitious multi-mineral exploration 'Eocene Program' aimed at raw mineral self-sufficiency and which includes construction of cement, alumina and electric power plants.-M.S.OliverWATER English
Article
Repaired shell injuries are reported in 11 specimens including six genera and eight species of gastropods from the upper Middle Ordovician (Caradoc) Kullsberg Limestone and the Upper Ordovician (Ashgill) Boda Limestone, Siljan district, Sweden. The specimens are of different sizes and morphologies, including one isostrophic, three low-spired, and six moderately high-spired turbinate forms. Single and repeated episodes of shell damage and subsequent repair are preserved, the breaks ranging from simple arcuate or scalloped fractures to removal of large sections of the apertural margin. Both early and late growth stages show damage, but the injuries are usually restricted to only one whorl. No shell repairs were discovered on the 65 more or less complete specimens of the subulitids, although the smooth shell makes observation difficult. Injuries in the Siljan samples are found in both microgastropods and large specimens, but the sample is too small for meaningful quantification. The shell repair frequency is about 7 percent, based on examination of 404 specimens (54 from Kullsberg Limestone and 350 from Boda Limestone). Most of the repaired injuries are attributed to failed predation, adding significantly to the Lower Paleozoic documentation of predation on gastropods. The gastropod shells are morphologically weak by modern standards, but do show some architectural strengthening features such as narrow apertures, collabral or spiral threads, and other ornamentation. The identity of the predator(s) is unknown.
Article
The mid-Paleozoic was punctuated by a rapid radiation of durophagous (shell-crushing) predators. These new predators were primarily placoderm and chondrichthyan fishes but probably also included phyllocarid and eumalacostracan arthropods. Coincident with the radiation of these durophages, beginning in the mid-Devonian, there was an increase in the frequency of predation-resistant morphologies in a variety of marine invertebrate taxa. Among bellerophontid molluscs, disjunct coiling disappeared and umbilici became less common while the frequency of genera with sculpture increased. The abundance of brachiopod genera with spines on one or both valves increased dramatically. Sculpture became more pronounced and common among genera of coiled nautiloids. Inadunate and camerate crinoids showed a marked increase in spinosity, and all three crinoid subclasses tended to develop thicker thecal plates. Trends toward increasing relative frequencies of predation-resistant features were formed in different ways. Bellerophontid genera lacking predation-resistant features tended to go extinct, leaving the sculptured, tighdy coiled forms as the predominant forms. Among Brachiopoda, the radiation of productids provided the tremendous increase in numbers of spinose genera. Among crinoids, predation-resistant features were acquired through evolution within established clades. These observations suggest that predation by shell-crushing predators has been an important control on the morphology and composition of the marine invertebrate fauna since at least the Middle Devonian. The mid-Paleozoic radiation of durophages and response of the marine fauna was in many respects similar to events of the Mesozoic Marine Revolution, in effect, the Paleozoic precursor to that event.
Chapter
This chapter presents insight into large-scale evolution through the study of a class of animals that is well represented both in the modern world and in the fossil record. Among the most distinctive features of the Heteroconchia are crossed-lamellar shell structure and heterodont dentition. This subclass includes the majority of post-Paleozoic burrowing clams. Another specialized group, the Trigoniacea, expanded dramatically in the Mesozoic through the evolution of somewhat unusual adaptations for efficient burrowing (Stanley, in preparation). Because of their unusual morphology, they too became an evolutionary cul-de-sac, after relatively unspecialized early representatives gave rise to the freshwater Unionacea. The rudists (Hippuritacea), which originated in the Jurassic, represent another divergent group of heteroconchs. The evolution of modern predators has probably been responsible for the decline of endobyssate taxa and the survival of only those epifaunal taxa with mechanisms for avoiding predation.