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.     

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.     

Predatory asteroids and the decline of the articulate brachiopods

Various causes, such as increased predation pressure, the lack of planktotrophic larvae, a 'resetting' of diversity, increased competition from benthic molluscs and the decline of the Palaeozoic fauna, have been suggested to explain the failure of the brachiopods to reradiate following the Permo-Triassic mass extinction. Increased predation pressure has hitherto appeared improbable, because typical predators of brachiopods, such as teleostean fish, brachyuran crabs and predatory gastropods, did not undergo major radiation until the late Mesozoic and early Cenozoic. However, new evidence strongly suggests that one important group of predators of shelly benthic organisms, the asteroids, underwent a major radiation at the beginning of the Mesozoic. Although asteroids appeared in the early Ordovician, they remained a minor element of the marine benthos during the Palaeozoic acme of the brachiopods. However, these early asteroids lacked four important requirements for active predation on a bivalved epifauna: muscular arms (evolved in the early Carboniferous); suckered tube feet, a flexible mouth frame and an eversible stomach (all evolved in the early Triassic). Thus radiation of the Subclass Neoasteroidea coincided with both their improved feeding capability and the decline of the articulates. The asteroids were the only group of predators of brachiopods that underwent a major adaptive radiation in the earliest Mesozoic. The asteroids may therefore have contributed to inhibiting a Mesozoic reradiation of the brachiopods. Epifaunal species lacking a muscular pedicle may have been particularly vulnerable. Unlike bivalve molluscs, modern brachiopods show only a limited range of adaptations to discourage asteroid predation. □ Asteroidea, Brachiopoda, evolution, predation, functional morphology.     

Intense drilling in the Carboniferous brachiopod Cardiarina cordata Cooper, 1956

The brachiopod Cardiarina cordata, collected from a Late Pennsylvanian (Virgilian) limestone unit in Grapevine Canyon (Sacramento Mts., New Mexico), reveals frequent drillings: 32.7% (n = 400) of these small, invariably articulated specimens (<2 mm size) display small (<0.2 mm), round often beveled holes that are typically single and penetrate one valve of an articulated shell. The observed drilling frequency is comparable with frequencies observed in the Late Mesozoic and Cenozoic. The drilling organism displayed high valve and site selectivity, although the exact nature of the biotic interaction recorded by drill holes (parasitism vs. predation) cannot be established. In addition, prey/host size may have been an important factor in the selection of prey/host taxa by the predator/parasite. These results suggest that drilling interactions occasionally occurred at high (Cenozoic-like) frequencies in the Paleozoic. However, such anomalously high frequencies may have been restricted to small prey/host with small drill holes. Small drillings in C. cordata, and other Paleozoic brachiopods, may record a different guild of predators/parasites than the larger, but less common, drill holes previously documented for Paleozoic brachiopods, echinoderms, and mollusks.     

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.     

Traces of predation/parasitism recorded in Eocene brachiopods from the Castle Hayne Limestone,North Carolina,USA

Schimmel, M., Kowalewski, M. & Coffey, BP. 2011: Traces of predation/parasitism recorded in Eocene brachiopods from the Castle Hayne Limestone, North Carolina, USA. Lethaia, Vol. 45, pp. 274–289. The Castle Hayne Limestone (Middle Eocene, North Carolina), noted for its diverse macro‐invertebrate fossils, was sampled to assess if Early Cenozoic brachiopods from eastern North America record any traces of biotic interactions. Systematic surveys of two North Carolina quarries yielded 494 brachiopods dominated by one species: Plicatoria wilmingtonensis (Lyell and Sowerby, 1845). Despite subtle variations in taphonomy, taxonomy and drilling patterns, the two sampled quarries are remarkably similar in terms of quantitative and qualitative palaeoecological and taphonomic patterns. In both quarries, brachiopods contain frequent drillholes (24.5% specimens drilled). The majority of drillholes were singular, perpendicular to shell surface and drilled from the outside. Ventral valves were drilled slightly more frequently than dorsal ones, but site‐selectivity in drilhole location was not evident. Larger brachiopods were drilled significantly more frequently than smaller ones. However, drillhole diameter did not correlate with brachiopod size. The drillholes are interpreted as records of ‘live‐live’ biotic interactions, representing either predatory attacks or parasitic infestations or a combination of those two types of interactions. A notable fraction of specimens bear multiple drillholes, which is consistent with either parasitic nature of interactions or frequent failed predatory events. The high drilling frequency reported here reinforces other reports (from other continents and other epochs of the Cenozoic), which suggest that brachiopods may be an important prey or host of drilling organisms in some settings. The number of case studies reporting high frequencies of drilling in brachiopods is still limited and thus insufficient to draw reliable generalizations regarding the causes and consequences of these occasionally intense ecological interactions. □Brachiopods, drilling parasitism, drilling predation, Eocene, North Carolina, taphonomy.     

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.     

Avian brain evolution: new data from Palaeogene birds (Lower Eocene) from England

Investigation of how the avian brain evolved to its present state is informative for studies of the theropod–bird transition, and as a parallel to mammalian brain evolution. Neurological anatomy in fossil bird species can be inferred from endocranial casts, but such endocasts are rare. Here, we use computed tomographic analysis to determine the state of brain anatomy in two marine birds from the Lower Eocene London Clay Formation of England. The brains of Odontopteryx (Odontopterygiformes) and Prophaethon (Pelecaniformes) are remarkably similar to those of extant seabirds, and probably possessed similar somatosensory and motor capabilities. Each virtual endocast exhibits a degree of telencephalic expansion comparable to living avian species. However, the eminentia sagittalis (wulst), a feature characteristic of all living birds, is poorly developed. Our findings support the conclusion that much of the telencephalic expansion of modern birds was complete by the end of the Mesozoic, but that overall telencephalic volume has increased throughout the Cenozoic through dorsal expansion of the eminentia sagittalis. We suggest that improvements in cognition relating to telencephalic expansion may have provided neornithine avian clades with an advantage over archaic lineages at the Cretaceous–Tertiary boundary, explaining their survival and rapid diversification in the Cenozoic. © 2009 The Natural History Museum. Journal compilation © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 198–219.     

Mosaics in the mangroves: allopatric diversification of tree‐climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo‐West Pacific

The Indo‐Australian Archipelago (IAA) is the richest area of biodiversity in the marine realm, yet the processes that generate and maintain this diversity are poorly understood and have hardly been studied in the mangrove biotope. Cerithidea is a genus of marine and brackish‐water snails restricted to mangrove habitats in the Indo‐West Pacific, and its species are believed to have a short pelagic larval life. Using molecular and morphological techniques, we demonstrate the existence of 15 species, reconstruct their phylogeny and plot their geographical ranges. Sister species show a pattern of narrowly allopatric ranges across the IAA, with overlap only between clades that show evidence of ecological differentiation. These allopatric mosaic distributions suggest that speciation may have been driven by isolation during low sea‐level stands, during episodes preceding the Plio‐Pleistocene glaciations. The Makassar Strait forms a biogeographical barrier hindering eastward dispersal, corresponding to part of Wallace's Line in the terrestrial realm. Areas of maximum diversity of mangrove plants and their associated molluscs do not coincide closely. © 2013 The Natural History Museum. Biological Journal of the Linnean Society © 2013 The Linnean Society of London, 2013, 110 , 564–580.     

Latitudinal and depth gradients in marine predation pressure

Aim There is a general paradigm that marine predation pressure increases towards the tropics and decreases with depth. However, data demonstrating global trends are generally lacking. Rhynchonelliform brachiopods inhabit all the oceans and often survive shell‐crushing predator attacks. We investigate shell repair in brachiopods across a range of Southern Hemisphere and tropical Northern Hemisphere latitudes and depths. Location The Southern Hemisphere and tropical Northern Hemisphere. Methods We analysed the frequency of shell repair in 112 bulk samples, over 70% of which showed traces of shell damage and repair. Results The pattern of shell repair frequency (RF) was more complicated than the anticipated increase with decreasing latitude, with low levels at both polar and tropical sites but high levels at temperate latitudes. This pattern is only evident, however, in shallow water assemblages; and there is no latitudinal trend in water depths greater than 200 m, where shell RF is systematically low. There was a significant logarithmic relationship between RF and depth. Low polar repair rates reflect reduced predation pressure, directly supporting the global paradigm. Low rates in the tropics appears counter to the paradigm. However, tropical brachiopods are generally very small (micromorphic) in shallow water and below the minimum size at which damage is recorded anywhere. Main conclusions Predation pressure decreased logarithmically with depth. At shallow depths (< 200 m) RF showed its highest levels in the mid temperate latitudes with decreasing frequency towards both the tropics and the poles. Low levels of shell repair at high latitudes are likely to be due to a lack of crushing predators, but in the tropics it is suggested that the low frequency is a result of the small size of tropical brachiopods. We hypothesize that micromorphy in this region may be an outcome of high predation pressure.     

FILM REVIEWS

Book reviewed in this article:
Lucy in Disguise . 1980. Written, produced and codirected by David Smeltzer in cooperation with the Cleveland Museum of Natural History.
The First Family . 1980. Produced by WVIZ in cooperation with the Cleveland Museum of Natural History.     

The type material of Mantodea (praying mantises) deposited in the National Museum of Natural History,Smithsonian Institution,USA

The collection of Mantodea of the National Museum of Natural History, Smithsonian Institution, includes 26 holotypes, 7 allotypes, 4 lectotypes, 23 paratypes, and 1 paralectotype. Four type specimens were designated as lectotypes within this work. Highly accurate measurement data, high resolution images of specimens and labels, verbatim label data, georeferenced coordinates, original and newly assigned database codes, and bibliographic data are presented for all primary types. Label data for all paratype specimens in the collection are provide in tabular form. The location of the USNM collection has been moved to the Cleveland Museum of Natural History as a loan under the Off-site Enhancement Program.     

Variations in the middle ear of the Mammalia

The comparative morphology of mammalian middle ear structures has been studied by examining the histological sections of the ears of 144 specimens of mammals which had been collected, with a few exceptions, in two expeditions to the Caribbean area. Fifty-four species of six orders of mammals were represented. Sixteen structural features or relationships were specifically studied and the bearing of taxonomic groupings on the variability was analysed.
A taxonometric analysis using these middle ear features indicated that Microdipodops may lie nearer to Dipodomys than to Perognathus.
A table presenting the detailed results has been deposited with the British Museum (Natural History).     

Drilling predation on scaphopods and other molluscs from the Upper Cretaceous of Manitoba, Canada

Abundant information on drilling predation upon fossil bivalves, gastropods, and brachiopods has been accumulated during the past several decades, but very little is known about the predation on marine, infaunal carnivorous scaphopods. A survey of over 440 specimens of the scaphopod Dentalium gracile collected from the Upper Cretaceous Millwood Member (Campanian) of the Pierre Shale at a site near Russell, Manitoba displays a drilling percentage of approximately 35%. This figure is higher than previously reported for the scaphopods of similar age elsewhere, but is comparable to or lower than that of the sub-Holocene (34–61%). The difference in drilling percentage among different collections may be taxon-related or affected by the composition and structure of the fossil community. Morphologically, the drill-holes, produced by predatory drilling, are beveled with a countersunk profile of clean sharp edges. The drill-hole inner margin is ovate whereas the outer margin is nearly circular. Among the Manitoba specimens, drill-holes seem to be more common on the lateral side. Presumably due to the lack of effective defense strategy, the prey effectiveness is low (∼3.2%). There is no correlation between drill-hole dimension and scaphopod prey size, indicating that predator size does not correlate with the prey size. Nearly 480 specimens of cephalopods, pelecypods, and gastropods were also collected from the same community. There were only a total of 16 drill-holes on this additional material. Over 400 specimens of the only naticid Euspira obliquata were recovered from the same site and are believed to be the predators of the scaphopods and other molluscs. The low percentage (∼2.5%) of drilling on the abundant gastropod E. obliquata may indicate mistaken or opportunistic attacks by the predator, or that the cannibalism is not common in this community.     

Ainu: Spirit of a Northern People

Ainu: Spirit of. Northern People. National Museum of Natural History, Smithsonian Institution, Washington, DC, April 30,1999-January 2, 2000.
Ainu: Spirit of. Northern People. William W. Fitzhugh and Chisato O. Dubreuil. eds. Washington, DC: Arctic Studies Center, National Museum of Natural History, in collaboration with University of Washington Press, 1999. 415 pp.     

Has the kouprey (Bos sauveli Urbain, 1937) been domesticated in Cambodia?

The kouprey (Bos sauveli Urbain, 1937) is a very rare bovid species of Cambodia, which may be extinct in the wild, as no living specimen has been observed for a long time. Here, we describe a complete taxidermy mount, which presents astonishing morphological similarities with the kouprey. The animal was mounted in 1871 at the National Museum of Natural History in Paris, where it was referenced as No. 1871-576. It was deposited at the Natural History Museum of Bourges, France, in 1931, where it is still conserved today. To clarify the taxonomic status of the specimen of Bourges, DNA was extracted from a piece of bone taken on the mandible, and two different fragments of the mitochondrial cytochrome b gene were independently amplified and sequenced. The phylogenetic analyses show that the specimen of Bourges is robustly associated with the holotype of the kouprey, and that both are related to other wild species of Bos found in Indochina, i.e., banteng (B. javanicus) and gaur (B. frontalis). Because of doubts for sexing the animal, we applied a molecular test based on the PCR amplification of a DNA fragment specific to the Y chromosome. The results indicate that the specimen of Bourges is a male. The comparisons with male kouprey previously described in the literature reveal important differences concerning the body size, general coloration and horns. As these differences involve phenotypic traits that are strongly selected in case of domestication, we suggest that the specimen of Bourges was a domestic ox. This implies therefore that the kouprey may have been domesticated in Cambodia, and that several extant local races may be directly related to the kouprey.     

Coevolution of post-Palaeozoic arthropod basibiont diversity and encrusting bryozoan epibiont diversity?

We hypothesize that the diversification of motile marine arthropods with hard carapaces resulted in a concurrent increase in the diversity of encrusting marine bryozoans whose larvae exploited those substrates through the Mesozoic and Cenozoic. To test this, family-level data were tabulated from the literature on the post-Palaeozoic diversity of motile marine arthropod basibionts and sessile marine bryozoan epibionts. We found strong temporal correlation from general to more specific basibiont-epibiont relationships (i.e. arthropods and bryozoans in general to decapods and encrusting gymnolaemates to robust decapods and encrusting cheilostomes in particular). We compared the diversification of bryozoans to another common group of basibionts (i.e. molluscs) and found weaker correlations. This suggests that the diversification of motile arthropods with hard carapaces (e.g. brachyuran crabs) may have played a role in the diversification of sessile bryozoans (e.g. encrusting cheilostomes) in the post-Palaeozoic.