The post‐Palaeozoic fossil record of drilling predation on lingulide brachiopods

Research on drilling predation, one of the most studied biological interactions in the fossil record, has been concentrated on prey with calcareous skeletons (e.g. molluscs, echinoids, rhynchonelliform brachiopods). Based on a compilation of literature sources and surveys of paleontological collections of the Florida Museum of Natural History and the National Museum of Natural History, we provide a tentative evaluation of the post‐Palaeozoic history of drilling predation on the organophosphatic brachiopods of the family Lingulidae. Despite temporal, geographical and methodological limitations of the data assembled here, the results indicate that lingulide brachiopods have been subject to drilling predation since at least the Eocene. Variation in drilling frequencies at the locality level suggests that lingulides may occasionally experience somewhat elevated predation pressures from drilling organisms. Overall, however, drilling predation on lingulide brachiopods has been infrequent in the Cenozoic and may have been absent in the Mesozoic. The Mesozoic‐to‐Cenozoic increase in drilling frequencies on lingulides is similar to the trends observed in other marine benthic invertebrates and consistent with the hypothesis that predation pressures increased through time in marine ecosystems.     

Drill holes in Australian Cainozoic brachiopods

The fossil record of drill holes in marine invertebrates has received a considerable amount of interest from paleontologists, primarily due to its importance for reconstructing the history of interactions between drilling predators and their prey. Such drill holes have been described in numerous studies of Paleozoic brachiopods but rarely in those focusing on brachiopods of the post-Paleozoic, a striking pattern given that in the late Mesozoic and Cainozoic drilling gastropods diversified and frequencies of drilled molluscs increased dramatically. During the past several years, however, drilled brachiopods were reported in several studies of the Mesozoic and Cainozoic, suggesting that this phenomenon may be more common than has been previously assumed. Here we report on 10 genera of brachiopods from four Cainozoic basins in Australia of which 7 shows evidence of having been drilled by predators. Of 298 specimens examined, 38 contain a single complete hole. Drilled specimens were identified in all 4 basins and in all stratigraphic units. When considered in the context of recent reports of drilled Cainozoic brachiopods, these Australian brachiopods further imply that drilling predation on these invertebrates was geographically, taxonomically and temporally widespread.     

Traces of predation in the Cambrian

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.     

Grazing bioerosion in Jurassic seas: a neglected factor in the Mesozoic marine revolution?

Grazing bioerosion, notably by chitons, gastropods and regular echinoids, is a powerful destructive force in many recent shallow-marine environments and impacts significantly on sessile epibionts through grazing predation and/or unselective dislodgement. Grazing bioerosion was an important component of a major phase of biotic escalation; the Mesozoic marine revolution. Recent investigations of hard substrates in southern British Jurassic marine formations have identified widespread ichnofossils attributable to grazing activity by gastropods and/or chitons, and regular echinoids. The co-occurring benthic macrofaunas include groups that would have been vulnerable to grazing disturbance and dislodgement; notably articulate brachiopods. The emerging ichnological evidence strengthens the argument for grazing bioerosion as a significant contributor to the Mesozoic–Cenozoic decline of the articulate brachiopods, and their retreat to deep-water and/or cryptic refugia.     

Evidence of predation damage in Pliocene Apletosia maxima (Brachiopoda)

Abstract:  Little is known about predation of Mesozoic and Cenozoic articulated brachiopods, but it is far from clear whether this is because they suffered very little predation pressure or because there have been few attempts to search for evidence of it. A study of 248 museum specimens of the large Pliocene terebratulid Apletosia maxima from the Coralline Crag (UK) has revealed that more than 16 per cent of them show evidence of having been attacked by predators. The styles of damage can be attributed to drilling muricid gastropods (most of which were successful) and failed crushing attacks probably by decapods. Brachiopods are usually thought to offer a poor tissue yield to potential predators, but in this instance it appears that A. maxima was attractive to predators even though they were living with a rich molluscan fauna. It is suggested that the mass of adductor and diductor muscles (likely to be spicule-free) of these particularly large brachiopods may have made them profitable. Further studies of post-Palaeozoic brachiopod faunas are required, particularly those from mixed shallow-water communities, before it can be established whether articulated brachiopods have or have not been driven into refugia by increasing predation pressure.     

High frequency of drill holes in brachiopods from the Pliocene of Algeria and its ecological implications

The fossil record holds a wealth of ecological data, including data on biotic interactions. For example, holes in the skeletons of invertebrates produced by drilling activities of their enemies are widely used for exploring the intensity of such interactions through time because they are common and easily distinguished from non-biotic holes or holes produced by other types of interactions. Such drill holes have been described in numerous studies of Palaeozoic brachiopods but rarely in those focusing on brachiopods of the post-Palaeozoic, a striking pattern given that in the late Mesozoic and Cenozoic drilling gastropods diversified and frequencies of drilled molluscs increased dramatically. During the past several years, however, drilled brachiopods were reported in several studies of the Mesozoic and Cenozoic, suggesting that this phenomenon may be more common than has been previously assumed. Here we report on drilled brachiopods from a Pliocene locality in Algeria where 90 of 261 (34.5%) specimens of Megerlia truncata show evidence of predatory drilling. These data confirm that Cenozoic drilling frequencies of brachiopods may be locally high and, when taken together with other published data, that drilling frequencies are highly heterogeneous in space and time.     

Cambrian to Cretaceous changes in hardground communities

The changing nature of the communities of boring and encrusting taxa found on upward-facing hard-grounds has been studied from the standpoints of (a) diversity, (b) faunal composition, and (c) nature of the niches occupied. After a rapid initial increase in the early Palaeozoic, diversity remained at much the same level from the Middle Ordovician until the late Cretaceous. However, there is a considerable turnover in the identity of the individual taxa between successive sample intervals. The incoming and outgoing of the major groups parallel their fortunes in the marine realm as a whole. Niche analysis suggests that the same feeding levels are occupied for most of the history of hardground communities, but Mesozoic faunas contain a much higher proportion of species with true exoskcletons, or which lived infaunally. The evolution of these forms was probably influenced by the Mesozoic radiation of marine predators and duriphages, but it also resulted in Mesozoic hardground faunas being more resistant than their Palaeozoic counterparts to episodic corrasion. Resulting higher population densities in the Mesozoic were probably one reason why cavity faunas beneath some of these hardground surfaces are more diverse than those beneath Palaeozoic examples. □ Hardground, community, evolution.     

Origin of planktotrophy--evidence from early molluscs

The size of early ontogenetic shells (protoconchs) of ancient benthic molluscs suggests that feeding larvae occurred at about 490 myr (approximately, transition from Cambrian to Ordovician). Most studied Ordovician protoconchs were smaller than Cambrian ones, indicating smaller Ordovician eggs and hatchlings. This suggests substitution of nutritious reserve matter such as yolk by plankton as an energy source for larvae. The observed size change represents the first direct empiric evidence for a late Cambrian to Ordovician switch to planktotrophy in invertebrate larvae. It corroborates previous hypotheses about a possible polyphyly of planktotrophy. These hypotheses were primarily based on molecular clock data of extant clades with different types of larva, change in the overall body size, as well as increasing predation pressure on Early Paleozoic sea floors. The Early Ordovician is characterized by an explosive radiation of benthic suspension feeders and it was suggested that planktotrophy would prolongate escape from benthic predation on hatchlings. This biological escalation hypothesis does not fully explain why planktotrophy and suspension feeding became important at the same time, during a major biodiversification. An additional factor that probably included availability of nutrients must have played a role. We speculate that an increasing nutrient supply and availability of photoautotrophic plankton in world oceans have facilitated both planktotrophy and suspension feeding, which does not exclude a contemporaneous predation-driven escalation. It is very likely that the evolution of planktotrophy as well as increasing predation contributed to the Ordovician radiation.     

Ecological succession among late palaeozoic and mesozoic tetrapods

Natural selection and the development of new taxa are associated with ecological replacement and the increase in number of niches with time. Continental faunal interchange was possible globally because of the existence of the super-continent Pangaea during much of the Upper Palaeozoic and Mesozoic. Figures of tetrapod niches vs. time and discussion of this concept for that period are presented for the first time. Four habitat divisions are used, namely marine, fresh-water, lowland and upland.The marine habitat was colonised rather late by tetrapods and these may have been the first predators on the early bony fishes which had diversified in the Permian. The radiation of bony fishes in the Jurassic was followed by a further increase in variety of their reptilian predators. Predators seem to develop some time after the radiation of a new potential prey group.Most early amphibians occupied fresh-water habitats in “crocodile” or “frog” niches, but from the Triassic tetrapods moved from fresh-waters and lowlands into the uplands also.In terrestrial habitats, the replacement of mammal-like reptiles by dinosaurs is tentatively explained in terms of palaeoclimatology and thermoregulatory physiology. Ornithischians capable of dealing with tough vegetation evolved to occupy the new niches produced by the radiation of conifers in the Jurassic. The extinction of dinosaurs appears to have been connected with temperature and habitat changes.Conclusions are supported by a summary of published opinions on the palaeoecological roles of early tetrapods.     

Larval ecology and morphology in fossil gastropods

The shell of marine gastropods conserves and reflects early ontogeny, including embryonic and larval stages, to a high degree when compared with other marine invertebrates. Planktotrophic larval development is indicated by a small embryonic shell (size is also related to systematic placement) with little yolk followed by a multiwhorled shell formed by a free‐swimming veliger larva. Basal gastropod clades (e.g. Vetigastropoda) lack planktotrophic larval development. The great majority of Late Palaeozoic and Mesozoic ‘derived’ marine gastropods (Neritimorpha, Caenogastropoda and Heterobranchia) with known protoconch had planktotrophic larval development. Dimensions of internal moulds of protoconchs suggest that planktotrophic larval development was largely absent in the Cambrian and evolved at the Cambrian–Ordovician transition, mainly due to increasing benthic predation. The evolution of planktotrophic larval development offered advantages and opportunities such as more effective dispersal, enhanced gene flow between populations and prevention of inbreeding. Early gastropod larval shells were openly coiled and weakly sculptured. During the Mid‐ and Late Palaeozoic, modern tightly coiled larval shells (commonly with strong sculpture) evolved due to increasing predation pressure in the plankton. The presence of numerous Late Palaeozoic and Triassic gastropod species with planktotrophic larval development suggests sufficient primary production although direct evidence for phytoplankton is scarce in this period. Contrary to previous suggestions, it seems unlikely that the end‐Permian mass extinction selected against species with planktotrophic larval development. The molluscan classes with highest species diversity (Gastropoda and Bivalvia) are those which may have planktotrophic larval development. Extremely high diversity in such groups as Caenogastropoda or eulamellibranch bivalves is the result of high phylogenetic activity and is associated with the presence of planktotrophic veliger larvae in many members of these groups, although causality has not been shown yet. A new gastropod species and genus, Anachronistella peterwagneri, is described from the Late Triassic Cassian Formation; it is the first known Triassic gastropod with an openly coiled larval shell.     

Co-evolution Between Fishes and Crustaceans

In aquatic ecosystems fishes are, in general, the most important predators of crustaceans. This must not have been so at the beginning of the Mesozoic. Evolution of aquatic vertebrates led from heavy, benthic, microphagous animals to buoyant, agile, omnivorous ones. Key events of this evolution took place in fresh water, while most crustacean groups probably evolved in marine habitats. It is pointed out that until early Jurassic times the caridoid escape reaction obviously ensured the survival of eucarid species, while the later radiation suggests that selection favoured well-calcified and short-tailed species. It is suspected that the radiation of the marine Teleostei is a main cause of this evolution. This hypothesis is in accordance with the occurrence of the most archaic crustacean relict species in fish-free habitats.     

Phylogeny and classification of the Asteroidea (Echinodermata)

Post-Palaeozoic asteroids share a large number of derived characters of the ambulacral column and the mouth frame, and constitute the crown group of the monophyletic group Asteroidea. This crown group is here called the Neoasteroidea (new subclass). The stem species of the crown group lived in the Permian or early Triassic and so the evolution of the asteroids parallels that of the echinoids. Character distribution within the Neoasteroidea, especially morphology of the skeleton, digestive system, larvae and tube feet, allows subdivision into four orders (Paxillosida, Notomyotida, Valvatida, Forcipulatida). The latter three orders possess the synapomorphy of suckered tube feet and are united as the Surculifera (new superorder); the Paxillosida are their primitive sister group. Palaeozoic asteroids represent the stem group of the class, and may be divided into plesions according to the order of appearance of synapomorphies with the crown group. Classification of Palaeozoic asteroids requires much further study. The appearance of new characters within the crown group asteroids, such as suckered tube feet, implies that these were absent in the stem group. The range of life-habits possible in Palaeozoic asteroids can thus be partly deduced from evidence provided by living asteroids. Palaeozoic asteroids are deduced to have lacked suckered tube feet and were presumably unable to evert the stomach; hence they were precluded from life on hard substrates and extraoral feeding on epifaunal organisms. It is suggested that they lived on soft substrates by deposit feeding, scavenging and predation on small benthos.     

Facies and fauna of the Pennsylvanian Buckhorn Asphalt Quarry deposit: a review and new data on an important Palaeozoic fossil <Emphasis Type="Italic">Lagerstätte</Emphasis> with aragonite preservation

The Pennsylvanian Buckhorn Asphalt Quarry contains the best-preserved Palaeozoic mollusc fauna in the world. Early impregnation of mixed siliciclastic–carbonate rocks (mudstones, pack to grainstones, shell beds, and conglomerates) with hydrocarbons prevented aragonite destruction (“Impregnation Fossil Lagerstätte”). The exceptional preservation comprises shell microstructures, microornaments and early ontogenetic shells. Most gastropods had planktotrophic larval development indicating a high primary production although the remains of phytoplankton are very rare in this and other Late Palaeozoic deposits. Deposition occurred close to a shallow-water coastal area. Mass flow processes (density currents) triggered by storms were involved in the transport mechanisms of some units. Shells of benthic molluscs yield the most diverse known Palaeozoic microboring assemblage, indicating at least partly euphotic conditions. The invertebrate fauna comprises about 160 species and is dominated by molluscs, which is unusual for a Palaeozoic deposit, suggesting that aragonite dissolution produces a major bias in the fossil record. However, most mollusc genera in the Buckhorn deposit are also known from other Pennsylvanian occurrences as recrystallised shells. This shows that preservation bias via preferential aragonite dissolution may be overestimated.     

Signs of predation in the Middle Jurassic of south-central Poland: evidence from echinoderm taphonomy

Distinct faunal aggregates are described from the Middle Jurassic (uppermost Bajocian/lowermost Bathonian and Middle Bathonian) clay deposits of Częstochowa area, south-central Poland. These aggregates are composed of molluscs (scaphopods, gastropods, bivalves, ammonites and belemnites), articulate brachiopods and echinoderms (asteroids, crinoids and echinoids). A large percentage of the fossils, especially bivalves, are fragmented, but some fossils are complete. Although most of the fossils are crushed and fragmented, they are still identifiable to at least the genus level. Thorough statistical analysis of taphonomic features indicates that the preservation of asteroid marginal plates is distinct from the ossicles derived from the host clays. The high frequency of bite marks and the good state of preservation suggest that the accumulations are the products of predation activities and most probably are the effect of regurgitation. Taking into account the rich and diverse fauna, the predator was a bottom-feeding generalist. Possible predators include palaeospinacid sharks, a tooth of which was collected from the same bedding surface, but not associated with regurgitated remains. Although the bite marks on the asteroid ossicles point to sharks as potential producers of regurgitates, other vertebrates, like durophagous pycnodontiform fish, cannot be excluded.     

A tale of two eras: Phytoplankton composition influenced by oceanic paleochemistry

We report the results of simple experiments which support the hypothesis that changes in ocean chemistry beginning in the Mesozoic Era resulted in an increase in the nutritional quality per mole of C and per cell of planktonic algal biomass compared to earlier phytoplankton. We cultured a cyanobacterium, a diatom, a dinoflagellate, and a green alga in media mimicking aspects of the chemistry of Palaeozoic and Mesozoic‐Cenozoic oceans. Substantial differences emerged in the quality of algal biomass between the Palaeozoic and Mesozoic‐Cenozoic growth regimes; these differences were strongly affected by interspecific interactions (i.e., the co‐existence of different species alters responses to the chemistry of the medium). The change was in the direction of a Mesozoic‐Cenozoic biomass enriched in protein per mole C, although cells contained less carbon overall. This would lead to a lower C:N ratio. On the assumption that Mesozoic‐Cenozoic grazers’ assimilation of total C was similar to that of their earlier counterparts, their diet would be stoichiometrically closer to their C:N requirement. This, along with an increase in mean cell size among continental shelf phytoplankton, could have helped to facilitate observed evolutionary changes in the Mesozoic marine fauna. In turn, increased grazing pressure would have operated as a selective force for the radiation of phytoplankton clades better equipped with antigrazing capabilities (sensu lato), as found widely in phytoplankton with biomineralization. Our results emphasize potential links between changing seawater chemistry, increased predation pressure and the rise to ecological dominance of chlorophyll a+c algae in Mesozoic oceans. The experiments also suggest a potential role for ocean chemistry in changes of marine trophic structure from the Palaeozoic to the later Mesozoic Era.     

Predation in the Ordovician and Silurian of Baltica

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.     

Disturbance and contrasting patterns of population structure in the brachiopod Terebratulina septentrionalis (Couthouy) from two subtidal habitats

The population structures of Terebratulina septentrionalis (Couthouy) from exposed upper rock surface and semi-cryptic rock wall habitats at 33 m depth in the Gulf of Maine differ. Over a 3-yr period, population densities were consistently higher in rock wall habitats. Although both populations were dominated by juveniles (1–4 mm shell length), size-frequency distributions constructed from upper rock surface and rock wall populations were significantly different, as a result of a greater frequency of large brachiopods (> 20 mm shell length) in rock wall populations. Prominent modes occurred at 14–15 mm shell length in upper surface populations and at 19–20 mm length in rock wall populations. Recruitment was higher in rock wall habitats where ambient light intensities were significantly lower than on upper rock surfaces. Differences in recruitment are either the result of larval selection for shaded rock walls or differential juvenile mortality between habitats. The larvae of Terebratulina settle on a diverse array of substrata. These include bedrock, sandy polychaete tubes and algae in upper surface habitats and bedrock, calcareous polychaete tubes, and ascidians in rock wall habitats. Individuals attached to polychaete tubes and algae in upper surface habitats do not attain large body size (> 13 mm shell length). It is suggested that these differences in population structure reflect the greater intensity of disturbance in upper surface habitats. For example, the cod, Gadus morhua (Linnaeus), ingests brachiopods attached to algae and polychaete tubes in this habitat. Gastropod predation affects brachiopods in upper surface habitats but not in rock wall habitats. Predation by gastropods and asteroids is not size-specific. These results are consistent with the hypothesis that predation contributed to the decline in the abundance and diversity of articulate brachiopods since the Mesozoic, and suggest that the restriction of recent populations to semi-cryptic rock wall and crevice habitats is, in part, controlled by disturbance.     

SOME ORDOVICIAN AND SILURIAN BRACHIOPOD ASSEMBLAGES

Finds of brachiopod assemblages in graptolitic shales give support to Ruedemann's theory of an epiplanktic life of certain Palaeozoic brachiopods. The possibly epiplanktic brachiopods observed in the present study belong to the families Obolidae ('Obolus') and Sowerbyellidae (Sericoidea). Other assemblages might have been benthic, and a representative of Chonetidae (Shagamella) is thought to yield an example of this kind of biocoenosis.     

Palaeoecological and phylogenetic implications of a new scleractiniamorph genus from Permian sponge reefs,south China

Scleractinian corals are the most important constituents of modern coralgal reefs. For many years, it was thought that they first appeared in the Middle Triassic and subsequently underwent explosive radiation. However, abundant scleractinian-like corals with ancestral morphological traits have recently been recovered from Middle Permian sponge reefs in China, which not only confirms a role in Permian reef ecology but also suggests a possible Palaeozoic origin for the group. Two species of a new Permian scleractiniamorph genus from China are described herein as Houchangocyathus wangi gen. et sp. nov. and Houchangocyathus yaoi gen. et sp. nov. Putative Palaeozoic Scleractinia may have evolved over a substantial time interval and diverged into stem lineages by the end of the Permian. These forms evolved within both the rigid framework of their basic body plan and the morphological constraints characteristic of each lineage. The Middle Permian development of calcisponge reefs was closely related to habitat expansion, which would have provided an ideal dwelling for scleractinian-like corals and enhanced their chances of fossilization. Such scleractiniamorphs disappeared at the end-Permian extinction, but may have survived as progenitors of Triassic Scleractinia.     

Evidence of predation in Early Cretaceous unionoid bivalves from freshwater sediments in the Cameros Basin,Spain

Here, we present evidence of possible vertebrate predation on freshwater bivalves from the Lower Cretaceous strata of the Cameros Basin (Spain). The described collection contains the largest number of vertebrate‐inflicted shell injuries in freshwater bivalve shells yet reported in the Mesozoic continental record. Several types of shell damage on fossil shells of Protopleurobema numantina (Bivalvia: Unionoida) are described and their respective modes of formation interpreted in the context of morphological attributes of the shell injuries and the inferred tooth morphology of predators that could have inflicted such injuries. Detailed study of these bite marks shows similarities with the well‐documented injuries in the shells of marine molluscs, namely ammonoids, that have likewise been attributed to reptilian predators. The most parsimonious interpretation suggests crocodiles as the vertebrates interacting with the bivalves in the Cameros Basin. □Barremian–Aptian; bite marks; freshwater bivalves; predation; reptile; Unionoida.