Dissecting latitudinal diversity gradients: functional groups and clades of marine bivalves

The latitudinal diversity gradient, with maximum taxonomic richness in the tropics, is widely accepted as being pervasive on land, but the existence of this pattern in the sea has been surprisingly controversial. This is partly due to Thorson's influential claim that the normal latitudinal diversity gradient occurs in marine epifauna (taxa living on the surface of the substratum) but not in infauna (burrowing or boring into the substratum), a contrast he attributed to the greater spatial and temporal environmental homogeneity of infaunal habitats. In an analysis of 930 species of north-eastern Pacific marine shelf bivalves, we found that bivalves as a whole, and both infauna and epifauna separately, show a strong latitudinal diversity gradient (measured as number of species per degree latitude) that is closely related to mean sea surface temperature (SST), even in analyses of residuals and first differences. This agrees with results for marine gastropods, but contradicts Thorson's environmental homogeneity hypothesis. The relationship between SST and diversity is consistent with a species-energy hypothesis, but the linkages from SST to diversity remain unclear. Most bivalve clades within broad functional groups conform to the general latitudinal trend, except for the deposit-feeding protobranchs. This group's non-directional pattern may be related to its mode of development, because a similar effect is seen in several other groups locked into this low-fecundity, non-feeding larval mode.     

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.     

Epibiont habitation patterns and their implications for life habits and orientation among trigoniid bivalves

The bivalve superfamily Trigoniacea has persisted from the Late Paleozoic to the Recent. Late Jurassic and terminal Cretaceous mass extinctions decimated this once-dominant group in shallow marine facies; only a single genus with seven species survives today in the Austral Province. Trigoniacea retain a vestigial byssus and primitive but efficient schizodont dentition. They have been widely considered as infaunal bivalves, burrowing with a very large foot to shallow depths, with inhalant and exhalant apertures at or slightly below the sediment-water interface (SWI). Yet the Trigoniacea are poorly adapted for this life habit. The mantle in living species is unfused and non-siphonate, and some fossil Trigoniacea have permanent shell gapes over these apertures, enhancing the probability of sediment fouling of feeding and respiratory structures. Some living Neotrigonia , e.g., N. margaritacea , solve this problem by having a semi-infaunal life habit, with the inhalant and exhalant apertures elevated above the SWI and the zone of active sediment transport. Semi-infaunal species commonly have epibionts cohabiting the exposed posterior-posteroventral portion of the shell. Numerous well-preserved species of South American Mesozoic Trigoniacea have phototropically and geotropically oriented epibionts on co-attached valves, strongly suggesting a semi-infaunal life mode for at least some members of these taxa. These shell symbionts allow orientation of extinct trigoniid shells relative to the SWI during life, as well as analysis of their depth of burial. Careful analyses of the kinds, size classes, orientation, and dispersion of various epibionts on fossil Trigoniacea thus yield important new information on their life habits, and demonstrate that semi-infaunal life modes were far more common than previously supposed.     

Autecology of Labrocuspis, a Middle Devonian omphalotrochid gastropod

Labrocuspis kobayashii , a Middle Devonian euomphalacean omphalotrochid gastropod from Japan has a wide callus pad on its base similar to that seen in the living trochid Umbonium. L. kobayashii could support the shell over the cephalopedal mass in a similar fashion to that seen in the Umbonium species. This reconstruction indicates that the labral projection in the outer lip was above the head, and the opisthocyrt basal lip and the sinus in the outer lip were loci of inhalant and exhalant currents, respectively. Therefore, the animal is assumed to have had only one ctenidium (the left). The presence of a wide callus pad is indicative of an infaunal life habit for the Umbonium species, Labrocuspis and some Paleozoic gastropods; clamping of the shell against the foot is also suggested for some omphalotrochid gastropods. These features indicate an additional adaptive strategy employed by euomphalaceans.     

Marine extinction risk shaped by trait–environment interactions over 500 million years

Perhaps the most pressing issue in predicting biotic responses to present and future global change is understanding how environmental factors shape the relationship between ecological traits and extinction risk. The fossil record provides millions of years of insight into how extinction selectivity (i.e., differential extinction risk) is shaped by interactions between ecological traits and environmental conditions. Numerous paleontological studies have examined trait‐based extinction selectivity; however, the extent to which these patterns are shaped by environmental conditions is poorly understood due to a lack of quantitative synthesis across studies. We conducted a meta‐analysis of published studies on fossil marine bivalves and gastropods that span 458 million years to uncover how global environmental and geochemical changes covary with trait‐based extinction selectivity. We focused on geographic range size and life habit (i.e., infaunal vs. epifaunal), two of the most important and commonly examined predictors of extinction selectivity. We used geochemical proxies related to global climate, as well as indicators of ocean acidification, to infer average global environmental conditions. Life‐habit selectivity is weakly dependent on environmental conditions, with infaunal species relatively buffered from extinction during warmer climate states. In contrast, the odds of taxa with broad geographic ranges surviving an extinction (>2500 km for genera, >500 km for species) are on average three times greater than narrow‐ranging taxa (estimate of odds ratio: 2.8, 95% confidence interval = 2.3–3.5), regardless of the prevailing global environmental conditions. The environmental independence of geographic range size extinction selectivity emphasizes the critical role of geographic range size in setting conservation priorities.     

The geographic, taxonomic and temporal distribution of determinate growth in marine gastropods

Many marine gastropods are characterized by determinate growth, as inferred from the presence of unique terminal elaborations of the shell's aperture. Although determinate growth has evolved repeatedly in most major gastropod clades, it is especially frequent among siphonate caenogastropods. Analyses of shallow-water assemblages show that the incidence of species with determinate growth is far higher in the tropics (especially the tropical Pacific and Indian Oceans) than at higher latitudes. Compilations of fossil assemblages from warm-water environments indicate that, although determinate growth occurred in some Palaeozoic gastropods, it became widespread only in the Neogene. In some groups, terminal apertural elaborations arose in lineages whose growth was more or less continuous and indeterminate, but in others it was derived either from or was ancestral to episodic growth. The hypothesis that periodic or terminal apertural elaborations evolved as a means to dispose of calcium carbonate once growth in the spiral direction ceased is rejected in favour of functional interpretations. Among the latter, the roles of modified apertures in defence and in mate recognition are explored, but no firm conclusion regarding the latter possibility can be drawn owing to our ignorance of mate recognition in gastropods.     

The Tommotian phase of the Early Cambrian Agronomic Revolution in the carbonate mud environment of central Siberia

The profound ecological change of the marine benthos that eventually led to the almost complete destruction of the Precambrian matgrounds by benthic grazers and bioturbators (the agronomic revolution) was largely completed in the Tommotian. At that time, burrows produced by bottom‐dwelling animals as shelters against predators were supplemented by burrowing for food by predators and sediment feeders. The limy mud ichnofauna of that age in Siberia was very different from the roughly coeval sand bottom faunas of Baltica. Although the exact zoological identity of the animals forming the infaunal Tommotian traces remains unknown, they probably mostly represent various kinds of early nemathelminthes. No apparent locomotion traces of mollusc origin have been encountered in the Early Cambrian, despite the abundant occurrence of skeletal fossils attributed to molluscs. Possibly the standard muscular foot, typical of modern molluscs, had not yet developed. Ichnotaxa represented are Teichichnus isp., Rhizocorallium isp., Chondrites isp., possibly the Buren ichnocomplex and others.     

Larval and juvenile gastropods from a Carboniferous black shale: palaeoecology and implications for the evolution of the Gastropoda

A horizon in the late Visean Ruddle Shale from Arkansas contains the oldest well-preserved gastropod protoconchs known from the Americas. The gastropod fauna consists of a diverse larval shell assemblage and a low diversity assemblage of juvenile gastropods that probably had a benthic life habit. Gastropod larval shells are always isolated, i.e. the gastropods did not complete their life cycle (no metamorphosis) and were unable to become benthic. This was caused by unfavorable environmental conditions on the soft muddy bottom that was probably due to anaerobic to exaerobic conditions. The absence or scarcity of bioturbation caused by invertebrate detritus or sediment feeders in both shale and concretions (formed before compaction) favored preservation of the delicate larval shells. The lack or scarcity of infauna and bioturbation as well as the low diversity of the presumed benthos supports an interpretation of a quasi-anaerobic to exaerobic benthic environment. The superbly preserved larval shells demonstrate that there are more caenogastropod clades present in the late Palaeozoic than suggested previously. Some larval shell types have an openly coiled first whorl followed by a planktotrophic larval shell; openly coiled initial whorls are unknown from modern caenogastropods. The vetigastropods have a smooth protoconch of two whorls clearly demarked from the following whorls - a pattern unknown in modern vetigastropods which have a protoconch of less than one whorl and build no larval shell during their planktonic stage. This could indicate a link between Palaeozoic vetigastropods and the caenogastropods.     

Evolutionary Dynamics in the Southwest Indian Ocean Marine Biodiversity Hotspot: A Perspective from the Rocky Shore Gastropod Genus Nerita

The Southwest Indian Ocean (SWIO) is a striking marine biodiversity hotspot. Coral reefs in this region host a high proportion of endemics compared to total species richness and they are particularly threatened by human activities. The island archipelagos with their diverse marine habitats constitute a natural laboratory for studying diversification processes. Rocky shores in the SWIO region have remained understudied. This habitat presents a high diversity of molluscs, in particular gastropods. To explore the role of climatic and geological factors in lineage diversification within the genus Nerita, we constructed a new phylogeny with an associated chronogram from two mitochondrial genes [cytochrome oxidase sub-unit 1 and 16S rRNA], combining previously published and new data from eight species sampled throughout the region. All species from the SWIO originated less than 20 Ma ago, their closest extant relatives living in the Indo-Australian Archipelago (IAA). Furthermore, the SWIO clades within species with Indo-Pacific distribution ranges are quite recent, less than 5 Ma. These results suggest that the regional diversification of Nerita is closely linked to tectonic events in the SWIO region. The Reunion mantle plume head reached Earth’s surface 67 Ma and has been stable and active since then, generating island archipelagos, some of which are partly below sea level today. Since the Miocene, sea-level fluctuations have intermittently created new rocky shore habitats. These represent ephemeral stepping-stones, which have likely facilitated repeated colonization by intertidal gastropods, like Nerita populations from the IAA, leading to allopatric speciation. This highlights the importance of taking into account past climatic and geological factors when studying diversification of highly dispersive tropical marine species. It also underlines the unique history of the marine biodiversity of the SWIO region.     

Constructional morphology of strombid gastropods

The extreme diversity in shell shape of strombid gastropods is interpreted as the result of three independent factors: (1) The terminal growth pattern of the Strombidae allows the circumvention of geometric constraints on shell morphology found in gastropods with continuous or periodic growth patterns. (2) Shell morphology in the Strombidac is adaptive in epifaunal locomotion, burrowing. infaunal or semi-infaunal habits, and passive protection from predators. Specialization for one of these functions often conflicted with the others. thus bringing about a forced 'choice' among mutually exclusive morphological characters. (3) Conservatism in life habits and anatomy of the soft parts has allowed the multiple evolution of extreme shell morphologies, as well as the secondary return to relativcly unspecialized morphologies. □ Constructional morphology, functional morphology. growth. behaviour. evolution, locomotion, burrowing, predation, exoskeleton. shell. Mollusca. Gastropoda. Strombacea. Strombidae.     

Host-associated speciation in the coral barnacle Wanella milleporae (Cirripedia: Pyrgomatidae) inhabiting the Millepora coral

Speciation by host shift is a common phenomenon observed in many symbiotic animals. The symbiont–host interaction is highly dynamic, but it is poorly documented in the marine realm. In the present study, we examined the genetic and morphological differentiation of the coral barnacle Wanella milleporae (obligate to fire corals) collected from four different Millepora host species in Taiwan to investigate the host specificity of this barnacle. Phylogenetic analysis of mitochondrial COI gene for 241 individuals of Wanella revealed five distinct clades, whose sequence divergences are comparable to values between other cogeneric barnacle species. The five clades also differ in shell and opercular plate morphology and colour. Genetic and morphological differentiations together strongly suggest the presence of cryptic species. Although the five clades do not display species-level host specificity, they showed a significant difference in preference on host growth form. Clades 1 and 2 were predominantly found on encrusting Millepora exaesa and Millepora platyphylla , while clades 3, 4 and 5 live exclusively on branching-form fire corals Millepora dichotoma and Millepora tenella . Phylogeny inferred from the combined mitochondrial COI, 16S and 12S (2182 bp) analysis suggests the division of the five clades into two major lineages congruent with the morphology of the host coral. Multiple independent invasions to the same form of host and subsequent speciation are evident in the Red Sea and Taiwan. Our results indicate that ecological/sympatric speciation could occur in marine symbiotic invertebrates through host shift and specialization. It appears that, as in their terrestrial counterparts, host–symbiont radiations in the marine realm are more prevalent than we expected and thus warrant further investigation.     

An evolutionary framework for the polar regions

Abstract: Our knowledge of the nature, generation and maintenance of largescale biodiversity patterns is still far from complete. This is particularly so in the Southern Hemisphere and in the marine realm, where recent taxonomic investigations of Mollusca and other invertebrate groups has cast doubt upon the existence of a simple cline in species richness between the tropics and the pole. Comparatively high regional diversity values for the shelled gastropods and other epifaunal taxa implies a considerable evolutionary legacy; this is supported, at least in part, by available evidence from the fossil record. Certain families within the living gastropod fauna maintain their prominence when traced back 40 m.y., and perhaps even longer; in addition, several Southern Ocean gastropod and bivalve genera can now be traced back to at least the late Eocene. Use of a variety of refugia may have enabled many taxa to survive repeated glacial advances. As we begin to revise our concept of the nature of latitudinal diversity gradients, so we also need to examine regional variations in evolutionary rates. Clearly this is a complex issue. but recourse to a pilot study based on the molluscan fossil record suggests that there may be no significant difference between the rates of radiation of tropical and cold-temperatdpolar taxa. The most diverse clades, which are all tropical, are simply the oldest. What data are available from the fossil record indicate that there is no appreciable latitudinal variation in rates of extinction either. Time, but not necessarily environmental stability, would appear to be crucial to the development of pockets of high taxonomic diversity. Recent improvement in our understanding of the biology of many polar marine invertebrates suggests that life in cold water is not an insuperable evolutionary problem. Of qual importance to any intrinsic properties of organisms which may have governed the differentiation of large-scale biodiversity patterns is the role of extrinsic processes. Foremost among these has almost certainly been repeated range shifts in response to Cenozoic climatic cycles.     

Major issues in the origins of ray‐finned fish (Actinopterygii) biodiversity

Ray‐finned fishes (Actinopterygii) dominate modern aquatic ecosystems and are represented by over 32000 extant species. The vast majority of living actinopterygians are teleosts; their success is often attributed to a genome duplication event or morphological novelties. The remainder are ‘living fossils’ belonging to a few depauperate lineages with long‐retained ecomorphologies: Polypteriformes (bichirs), Holostei (bowfin and gar) and Chondrostei (paddlefish and sturgeon). Despite over a century of systematic work, the circumstances surrounding the origins of these clades, as well as their basic interrelationships and diagnoses, have been largely mired in uncertainty. Here, I review the systematics and characteristics of these major ray‐finned fish clades, and the early fossil record of Actinopterygii, in order to gauge the sources of doubt. Recent relaxed molecular clock studies have pushed the origins of actinopterygian crown clades to the mid‐late Palaeozoic [Silurian–Carboniferous; 420 to 298 million years ago (Ma)], despite a diagnostic body fossil record extending only to the later Mesozoic (251 to 66 Ma). This disjunct, recently termed the ‘Teleost Gap’ (although it affects all crown lineages), is based partly on calibrations from potential Palaeozoic stem‐taxa and thus has been attributed to poor fossil sampling. Actinopterygian fossils of appropriate ages are usually abundant and well preserved, yet long‐term neglect of this record in both taxonomic and systematic studies has exacerbated the gaps and obscured potential synapomorphies. At the moment, it is possible that later Palaeozoic‐age teleost, holostean, chondrostean and/or polypteriform crown taxa sit unrecognized in museum drawers. However, it is equally likely that the ‘Teleost Gap’ is an artifact of incorrect attributions to extant lineages, overwriting both a post‐Palaeozoic crown actinopterygian radiation and the ecomorphological diversity of stem‐taxa.     

Sinistral snail shells in the sea: developmental causes and consequences

The paucity of sinistral (left-coiling) relative to dextral (right-coiling) species of gastropods in the marine realm is an enigma. In Conus , one of the most diverse marine animal genera, sinistral shell coiling has evolved as a species-wide character only once. Fossils of this species, Conus adversarius , are found in Upper Pliocene and lowermost Pleistocene deposits in the southeastern USA. Conus adversarius had nonplanktonic larval development; this may have been a critical factor for the early establishment of the species, as well as sinistral marine species in other clades. Notably, most specimens of aberrantly sinistral modern Conus are derived from typically dextral species that have nonplanktonic development. If C. adversarius was reproductively isolated from dextral conspecifics, then this species may provide an example of nearly instantaneous sympatric speciation in the fossil record. Furthermore, the common and widespread – while geologically short-lived – fossil shells of C. adversarius show large amounts of variability in form and this variation may be related, at least in part, to a pleiotropic effect associated with the reversed coiling direction of this species.     

End‐Triassic bivalve extinction: Lombardian Alps,Italy

A major biotic crisis affecting virtually all major marine invertebrate clades occurred at the close of the Triassic. Species‐level data on bivalves from the Lombardian Alps of Italy record the extinction and suggest a possible causal mechanism. A significant decline in species richness is observed during the lower Rhaetian, where 51% of bivalve species, equally distributed among infaunal and epifaunal filter‐feeders, went extinct. The taxonomic loss at the middle Rhaetian was more severe, where 71% of the bivalve species were eliminated, including all infaunal and 50% of the epifaunal species. The data indicate that the extinction selectively eliminated infaunal bivalves.

An initial loss of bivalve species richness during the middle and upper Rhaetian correlates with changes in sedimentary facies related to a fall in relative sea level. This sea level fall is marked by the onset of peritidal micrites and shifting ooid shoals which may have rendered substrates unsuitable for both epifaunal and infaunal bivalves. The possible influences of temperature and salinity fluctuations are difficult to assess, but they may also have had a deleterious effect on the local bivalve fauna. The loss due to peritidal conditions is not consistent with the selective survivorship of epifaunal taxa recurring in overlying Jurassic rocks.

We propose that physiologic differences and selective resistance to physical stress are consistent with the pattern of selective extinction. Facies shifts associated with the marine regression are not sufficient to account for the extremely high magnitude of infaunal extinction. This selection against infaunal bivalves is probably caused by their decreased capacity to filter feed relative to their metabolic demands. A decrease in primary productivity could have selectively eliminated the infauna. Oceanographic processes or atmospheric darkening, perhaps caused by an extraterrestrial impact, could drastically limit food resources (primary productivity) and is consistent with the selective extinction at the end of the Triassic.     

Diversity,palaeoecology and systematics of a marine fossil assemblage from the Late Triassic Cassian Formation at Settsass Scharte,N Italy

A marine fossil assemblage from the Late Triassic (Early Carnian) Cassian Formation is reported. It was retrieved by bulk sampling, including wet sieving and quantitative picking, and by quantitative surface collection. The collection consists of c. 460 specimens (foraminiferans not included) representing 54 species. In terms of abundance and species richness, it is strongly dominated by molluscs, especially gastropods. 97 % of the individuals are molluscs. The most abundant species are the gastropods Goniospira armata, Schartia carinata and Helenostylina convexa, followed by the scaphopod Plagioglypta undulata and the bivalve Palaeonucula strigilata. Disarticulated echinoderm ossicles (mostly echinoids, crinoids, few holothurians) comprise almost all of the rest of the assemblage. The studied assemblage shows moderate diversity, similar to those of previously reported assemblages or associations from basinal settings. However, it differs considerably in taxonomic composition from previously described associations of the Cassian Formation. The abundance of small gastropods is a result of their primary abundance in these ancient living communities and of the sampling method (sieving at 0.5 mm), because most of the previous associations were obtained by performing surface collections, in which small gastropods are easily overlooked. The fauna is interpreted as an autochthonous soft bottom assemblage dominated by species that lived in low epifaunal to shallow infaunal habitats. Detritivory, deposit feeding and microcarnivory represent the main feeding types. Most of the species were fully motile but slow, and either infaunal (scaphopods, nuculids, the gastropod Domerionina) or epifaunal (most other gastropods, echinoids). The present assemblage underlines the pronounced heterogeneity of the Cassian biota. The low grade of lithification and diagenetic alteration facilitates bulk sampling and the investigation of small species. This minimizes possible sampling and preservation biases, so the studied assemblage reflects the alpha diversity of this ancient living community to an unusually high degree. The following gastropod taxa are new: Helenostylina convexa n. sp., Schartia carinata n. gen. n. sp., Schartiinae n. subfam.; Cassianastraea Bandel non Volz is replaced with Bandelastraea nom. nov.     

Ratchet riposte: more on gastropod burrowing sculpture

Recent warnings concerning paleobiological inferences based upon gastropod shell morphology (Houbrick 1991) merit serious consideration, although the dangers have been overstated. Ratchet sculpture, an asymmetrical sculpture that assists marine invertebrates in burrowing, is not qualitatively different from sculptures that apparently do not aid in burrowing. Therefore, the interpretation of such sculpture might be problematical. Nevertheless, the large body of empirical evidence demonstrating the function of ratchet sculpture in burrowing by bivalves, gastropods, carpoid echinoderms, brachiopods, and arthropods and the lack of evidence supporting alternative functions in the Gastropoda warrant the continued, although cautious, association of ratchet sculpture with burrowing in marine gastropods. □ Functional morphology, Gastropoda, ratchet sculpture, burrowing.     

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.     

Correlation of shape and habit with sediment grain size for selected species of the bivalve Anadara

Burrowing ability was experimentally determined for a range (3–150 g) of sizes of the blood ark, Anadara ovalis , in sieved sediments ranging from very coarse sand (-1 φ) to silt (>4 φ). Small individuals (<5 g) could penetrate the complete range of sieved sediments, but the ability to reburrow apparently decreases with size (age) so that adults larger than 50 g can only reburrow in fine sands (2.5–3.0φ). This semi-infaunal species has a length/height ratio of 1.16 and is most commonly found in sand. The infaunal A. cheninitzi and A. braziliana , with L/H values of 1.10 and 1.18, respectively, are also most common in sands. The ovate (L/H ratio = 1.07–1.25) forms of the Pliocene A. trilineata from the Kettleman Hills of California are inferred to have been infaunal, based on the absence of muricid boreholes and the infrequent occurrence of epizoans. Ovate specimens are restricted to fine sands (2–3φ) in the Etchegoin and San Joaquin Formations, sediment textures into which the adults could probably reburrow. Elongated shell morphotypes (L/H = 1.5–1.67) have muricid boreholes and epizoans concentrated posterodorsally on the inferred exposed portion of the shells in life position. Populations of elongated shells were associated with gravelly, sandy and silty sediments, as is the living epibyssate A. lienosa (L/H = 1.72) and A. transversa (L/H = 1.48).□ Anadarids, burrowing, epifounal, infaunal, Pliocene, sediment influence .     

The chondrichthyan fauna from the Middle-Late Triassic of Guanling (Guizhou province,SW China)

The Recent volutids Cymbiola and Melo burrow in a forward direction. This agrees well with the spiral terrace pattern observed in Eocene Volutidae, thus strengthening the idea that they were forward burrowers as well. The presence of collabral terraces near the aperture in the Cassidae suggests an oblique burrowing direction, by convergence with the Nassariidae. This is confirmed by observations on living cassids. The sutural canals and associated structures of the Olividae and Seraphidae are functional in detecting whether the posterior region of the shell is buried. This confirms the idea that fossil Strombidae possessing similar features were burrowers.

In contrast with these “predictable”; observations, burrowing was observed in a few gastropods the shell morphology of which would seem to exclude such behaviour, such as Architectonica, Cymatium, Volema and Bolinus. Thus, a few morphologic criteria appear to be reliable and of general applicability in inferring burrowing habits in gastropods. At the same time, the broad variety of burrowing mechanisms and life habits of gastropods makes it unlikely that general criteria will ever be found to exclude burrowing habits on the basis of shell morphology.