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The diversification of macroscopic pelagic arthropods such as caryocaridid archaeostracans was a crucial aspect of the Great Ordovician Biodiversification Event, and the plankton revolution. A pelagic mode of life has been inferred for caryocaridids from their common presence in black graptolitic shales alongside carapace morphologies that appear streamlined. However, the hydrodynamic performance within the group and comparisons with other archaeostracans were lacking. Here we use a computational fluid dynamics approach to quantify the hydrodynamic performance of caryocaridids, and other early Palaeozoic archaeostracans including Arenosicaris inflata and Ordovician ceratiocaridids. We show that streamlining of the carapace was an important factor facilitating a pelagic mode of life in caryocaridids, in reducing the drag coefficient and facilitating a broader range of lift coefficients at different angles of attack. However, comparable hydrodynamic performance is also recovered for some ceratiocaridids. This suggests that alongside carapace streamlining, adaptations to appendages and thinning of the carapace were also important for a pelagic mode of life in Ordovician caryocaridids., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Pates, Xue. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

The exchange of information and social interactions on broad spatial scales between human groups in the past can be studied through the provenance of key indicators of distant origin recorded at archaeological sites. The remains of shells of mollusk species, especially when crafted as elements of personal ornaments, express aspects of the behaviors and valuations for the populations that selected, transformed, and exchanged such items. In the southern cone of South America, past hunter-gatherer groups traveled long distances and interacted with communities distributed throughout the territory to acquire goods for technological use, visual display or considered highly valued materials. When recorded at distant locations, these goods of extra local origin are very informative regarding the differences between commonly used home ranges and the occasional access to remote spaces. We present the results of the analysis of the archaeomalacological assemblage of the Baño Nuevo 1 site, a cave with exceptional preservation conditions in Central West Patagonia. This site has yielded a diverse group of artifacts made of shells with origins from multiple distances, as well as evidence of the use of marine, freshwater, and terrestrial species. Its deposits, which extend over the last 11,000 years, reveal an antiquity of at least the middle Holocene for the acquisition, manufacture, use and transport of goods as personal ornaments from shells in the macroregion., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Hammond et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

A fossil oyster bed (Ostrea edulis) was recently encountered offshore Helgoland (German Bight). Oysters are important filter feeders in marine environments and their habitat structure supports a large associated biodiversity. The European flat oyster Ostrea edulis has historically occurred in vast populations in the North Sea, but declined massively in the early 20th century. The ecological restoration of Ostrea habitats is a current focal point in the North Sea. To better understand the mechanisms that caused the local collapse of the oyster population, this study investigated the size structure, weight, and age of the shells, along with the spatial dimensions, seafloor properties, and environmental context of the oyster bed. The results show that the demise of the population occurred around 700 CE, ruling out excessive harvest as a driver of decline. Synchronicity of increased geomorphological activity of rivers and concurrent major land use changes in early medieval Europe suggest that increased sedimentation was a viable stressor that reduced the performance of the oysters. The shells provided no indication of a demographically poor state of the oyster bed prior to its demise, but manifested evidence of the wide-spread occurrence of the boring sponge Cliona sp. Our study challenges the assumption of a stable preindustrial state of the European flat oyster in the North Sea, and we conclude that the long-term variability of environmental conditions needs to be addressed to benchmark success criteria for the restoration of O. edulis., Competing Interests: The authors have declared that no competing interests exist.

Nipponites is a heteromorph ammonoid with a complex and unique morphology that obscures its mode of life and ethology. The seemingly aberrant shell of this Late Cretaceous nostoceratid seems deleterious. However, hydrostatic simulations suggest that this morphology confers several advantages for exploiting a quasi-planktic mode of life. Virtual, 3D models of Nipponites mirabilis were used to compute various hydrostatic properties through 14 ontogenetic stages. At each stage, Nipponites had the capacity for neutral buoyancy and was not restricted to the seafloor. Throughout ontogeny, horizontally facing to upwardly facing soft body orientations were preferred at rest. These orientations were aided by the obliquity of the shell's ribs, which denote former positions of the aperture that were tilted from the growth direction of the shell. Static orientations were somewhat fixed, inferred by stability values that are slightly higher than extant Nautilus. The initial open-whorled, planispiral phase is well suited to horizontal backwards movement with little rocking. Nipponites then deviated from this bilaterally symmetric coiling pattern with a series of alternating U-shaped bends in the shell. This modification allows for proficient rotation about the vertical axis, while possibly maintaining the option for horizontal backwards movement by redirecting its hyponome. These particular hydrostatic properties likely result in a tradeoff between hydrodynamic streamlining, suggesting that Nipponites assumed a low energy lifestyle of slowly pirouetting in search for planktic prey. Each computed hydrostatic property influences the others in some way, suggesting that Nipponites maintained a delicate hydrostatic balancing act throughout its ontogeny in order to facilitate this mode of life., Competing Interests: The authors have declared that no competing interests exist.

We here report on fossil remains of the earliest known crown-Testudo, an extant clade of Mediterranean testudinid tortoises from the late Miocene (Vallesian, MN 10) from the hominoid locality Ravin de la Pluie (RPl) in Greece. The material studied is a small, nearly complete carapace with a clearly distinct hypo-xiphiplastral hinge. This supports the sensu stricto generic assignment. This new terrestrial testudinid specimen is characterized by a possible tectiform, narrow, elongated shell with a pentagonal pygal and a long, posteriorly elevated, lenticular and rounded dorsal epiplastral lip. These unique features differ from those of other known Mediterranean hinged forms and allow the erection of the new species Testudo hellenica sp. nov. This taxon is phylogenetically close to two Greek species, the extant T. marginata and the fossil T. marmorum (Turolian, around 7.3 Ma). This record provides evidence for the first appearance of the genus Testudo sensu stricto at a minimum age of 9 Ma., Competing Interests: The authors have declared that no competing interests exist.

Species of terebratulide brachiopods have been largely characterized qualitatively on the basis of morphology. Furthermore, species-level morphological variability has rarely been analyzed within a quantitative framework. The objective of our research is to quantify morphological variation to test the validity of extant named species of terebratulide brachiopods, focusing on the lophophore-supporting structures-the "long loops." Long loops are the most distinctive and complex morphological feature in terebratellidine brachiopods and are considered to be phylogenetically and taxonomically informative. We studied eight species with problematic species identities in three genera distributed in the North Pacific: Laqueus, Terebratalia, and Dallinella. Given how geometrically complex long loops are, we generated 3D models from computed tomography (CT) scans of specimens of these eight species and analyzed them using 3D geometric morphometrics. Our goal was to determine ranges of variation and to test whether species are clearly distinguishable from one another in morphospace and statistically. Previous studies have suggested that some species might be overly split and are indistinguishable. Our results show that these extant species of terebratellidines can be reliably distinguished on the basis of quantitative loop morphometrics. Using 3D geometric morphometric methods, we demonstrate the utility of CT beyond purely descriptive imaging purposes in testing the morphometric validity of named species. It is crucial to treat species described and named from qualitative morphology as working hypotheses to be tested; many macroevolutionary studies depend upon the accurate assessment of species in order to identify and seek to explain macroevolutionary patterns. Our results provide quantitative documentation of the distinction of these species and thus engender greater confidence in their use to characterize macroevolutionary patterns among extant terebratellidine brachiopods. These methods, however, require further testing in extinct terebratellidines, which only rarely preserve the delicate long loop in three dimensions. In addition, molecular analyses of extant terebratellidines will test the species delimitations supported by the morphometric analyses presented in this study. [Species determination; morphological variability; 3D geometric morphometrics; terebratulide brachiopods; long loops.]., Competing Interests: The authors have declared that no competing interests exist.

The function of Late Archaic period (5000-3000 B.P.) shell rings has been a focus of debate among archaeologists for decades. These rings have been variously interpreted as a product of seasonal feasting/ceremonial gatherings, quotidian food refuse generated by permanent dwellers, or a combination of seasonal and perennial activities. Seasonality of shell rings can be assessed by reconstructing the harvest time of oysters (Crassostrea virginica), the primary faunal component of shell rings. We estimated the timing of oyster harvest at St. Catherines Shell Ring (Georgia, USA) by statistical modeling of size frequency distributions of the impressed odostome (Boonea impressa), a parasitic snail inadvertently gathered by Archaic peoples with its oyster host. The odostome samples from three archaeological excavation units were evaluated against resampling models based on monthly demographic data obtained for present-day populations of Boonea impressa. For all samples, the harvest was unlikely to start earlier than late fall and end later than late spring, indicating that shell deposits at St. Catherines Shell Ring formed seasonally with substantial harvesting activities restricted to non-summer months. For all samples, the resampling models indicated that harvesting activities likely occurred during multiple months. However, these analytical outcomes would also be expected in the case of extensively time-averaged records of short-term, non-summer harvest events. Regardless of the exact harvest duration, the results point to seasonal harvesting and suggest that Archaic populations may have opted out of consuming summer oysters to focus on other resources, avoid unpalatable food, decrease pathogen risks, or ensure sustainable harvesting., Competing Interests: The authors have declared that no competing interests exist.

During the past decade, over 3000 shell middens or shell matrix deposits have been discovered on the Farasan Islands in the southern Red Sea, dating to the period c. 7,360 to 4,700 years ago. Many of the sites are distributed along a palaeoshoreline which is now 2-3 m above present sea level. Others form clusters with some sites on the shoreline and others located inland over distances of c. 30 m to 1 km. We refer to these inland sites as 'post-shore' sites. Following Meehan, who observed a similar spatial separation in shell deposition in her ethnographic study of Anbarra shellgathering in the Northern Territory of Australia, we hypothesise that the shoreline sites are specialised sites for the processing or immediate consumption of shell food, and the post-shore sites are habitation sites used for a variety of activities. We test this proposition through a systematic analysis of 55 radiocarbon dates and measurement of shell quantities from the excavation of 15 shell matrix sites in a variety of locations including shoreline and post-shore sites. Our results demonstrate large differences in rates of shell accumulation between these two types of sites and selective removal of shoreline sites by changes in sea level. We also discuss the wider implications for understanding the differential preservation and visibility of shell-matrix deposits in coastal settings in other parts of the world extending back into the later Pleistocene in association with periods of lowersea level. Our results highlight the importance of taphonomic factors of post-depositional degradation and destruction, rates of shell accumulation, the influence on site location of factors other than shell food supply, and the relative distance of deposits from their nearest palaeoshorelines as key variables in the interpretation of shell quantities. Failure to take these variables into account when investigating shells and shell-matrix deposits in late Pleistocene and early Holocene contexts is likely to compromise interpretations of the role and significance of shell food in human evolutionary and socio-cultural development., Competing Interests: The authors have declared that no competing interests exist.

Plesiochelyidae were relatively large coastal marine turtles, which inhabited the epicontinental seas of Western Europe during the Late Jurassic. Their fossil record can be tracked in Germany, Switzerland, the United Kingdom, France, Spain and Portugal. The Jura Mountains, in northwestern Switzerland, have been the main source for the study of this group, mostly thanks to the rich and famous historical locality of Solothurn. In the last two decades, numerous plesiochelyid remains have been collected from Kimmeridgian deposits (Lower Virgula Marls and Banné Marls) in the area of Porrentruy (Canton of Jura, Switzerland). This material was revealed by construction works of the A16 Transjurane highway between 2000 and 2011, and led to the recent description of the new species Plesiochelys bigleri. In the years 2014 and 2016, new fragmentary turtle material was collected from the Banné Marls (Reuchenette Formation, lower Kimmeridgian) near the village of Glovelier, Canton of Jura, Switzerland. The new material consists of a complete shell, additional shell elements, a few bones from the appendicular and vertebral skeleton, and a fragmentary basicranium. This material can be confidently assigned to the species P. bigleri. It supports the presence of this species in the Banné Marls, slightly extends its spatial distribution and confirms the differences with the closely related species P. etalloni. The new material reveals that the split between the cerebral and palatine branches of the internal carotid artery occurs in a vertical plane in P. bigleri. This condition could not be observed in the type material due to poor preservation. This new character clearly distinguishes P. bigleri from P. etalloni and seems to be unique among thalassochelydians., Competing Interests: The authors have declared that no competing interests exist.

Marine protists play an important role in oceanic ecosystems and biogeochemical cycles. However, the difficulties in culturing pelagic protists indicate that their ecology and behavior remain poorly understood; phylogeographic studies based on single-cell genetic analyses have often shown that they are highly divergent at the biological species level, with variable geographic distributions. This indicates that their ecology could be complex. On the other hand, the biomineral (calcareous) shells of planktic foraminifers are widely used in geochemical analyses to estimate marine paleoenvironmental characteristics (i.e., temperature), because the shell chemical composition reflects ambient seawater conditions. Among the pelagic protists, planktic foraminifers are ideal study candidates to develop a combined approach of genetic, morphological, and geochemical methods, thus reflecting environmental and ecological characteristics. The present study precisely tested whether the DNA extraction process physically and chemically affects the shells of the planktic foraminifer Globigerinoides ruber. We used a nondestructive method for analyzing physical changes (micro-focus X-ray computed tomography (MXCT) scanning) to compare specimens at the pre- and post-DNA extraction stages. Our results demonstrate that DNA extraction has no significant effect on shell density and thickness. We measured stable carbon and oxygen isotopes on the shell of each individual in a negative control or one of two DNA-extracted groups and detected no significant differences in isotopic values among the three groups. Moreover, we evaluated isotopic variations at the biological species level with regard to their ecological characteristics such as depth habitat, life stages, and symbionts. Thus, our examination of the physiochemical effects on biomineral shells through DNA extraction shows that morphological and isotopic analyses of foraminifers can be combined with genetic analysis. These analytical methods are applicable to other shell-forming protists and microorganisms. In this study, we developed a powerful analytical tool for use in ecological and environmental studies of modern and past oceans., Competing Interests: The authors have declared that no competing interests exist.

The geographical spreading of new fishing activities and the increasingly deeper locations of these activities have shown the worldwide distribution of gerionid crabs and new descriptions of Chaceon taxa. However, incomplete penetrance, variable expressivity, and phenotypic overlap make the morphometric identification of these species difficult. In this study, partial sequences of the cytochrome c oxidase subunit 1 (COI) and 16S mitochondrial ribosomal RNA (16S rRNA) genes have been analyzed in Chaceon species from the Eastern Central and South Atlantic and compared with sequences of species from Western Atlantic. Our results corroborate the proposed morphological species and highlight the significant separation of the Eastern Atlantic species and those from Atlantic coasts of South America for both markers (97% Bayesian posterior probability, BPP / 83% Bootstrap replicates, BT). Interestingly, Chaceon sanctaehelenae shows a closer relationship with the species of the American coast than with those from the Eastern Atlantic. On the other hand, while COI marker clearly separates Chaceon atopus and Chaceon erytheiae species (99 BPP / 91% BT), these species share haplotypes for the 16S rRNA marker, pointing to a recent speciation process. Moreover, a close relationship was observed between Chaceon maritae and Chaceon affinis (94% BPP / 77% BT). The topologies of the trees obtained indicate that the ancestor of this genus was closer related to those species from South America than to those from the Eastern Atlantic., Competing Interests: The authors have declared that no competing interests exist.

The slipper limpet Crepipatella dilatata, native to Chile and Argentina, was introduced in Spain in 2005. The species was thought to inhabit the region of Rias Bajas, yet recently, putative C. dilatata populations have been documented on the coast of north-central Spain and in the Ebro Delta of the Spanish Mediterranean. Here we undertook a multidisciplinary approach to study the invasion biology of this species. Specifically, two geographically distant populations, one being a successfully established population from O Grove and the other a declining population from Gijon, were studied over the course of four years. Analyses of morphological and developmental traits as well as genetic information confirmed the presence of C. dilatata in these sites. The results revealed polymorphism in anatomical traits and shell shape. Shell shape polymorphism was unevenly distributed among sites and among sexes. Males were monomorphic, while females were polymorphic. Of the female morphotypes encountered, one was absent in the declining population from Gijón. Size at first female maturation and female size were greater in the declining population than in the established population. Reproductive success varied seasonally but not spatially among populations. In the established population, gregariousness was significantly greater; the size when sex changes was found to be plastic and socially controlled. The sex ratio of the declining population was female biased while in the established population the sex ratio changed during the study period from being balanced to being female biased. This change in sex ratio was probably due to higher male mortality. Molecular analyses pointed to the localities of Corral Bay in southern Chile and Puerto Madryn in southern Argentina as potential population sources. The intercontinental import of fresh mussels cultivated in Chilean farms is a likely source of this mussel in Spain. Comparison with available data of native populations of C. dilatata strongly indicate that ecophenotypic plasticity, socially controlled sex change, high gregariousness, increased nurse egg supply to viable larvae during the encapsulated developmental period, later maturation and larger female sizes altogether enhance establishment success of this non-indigenous species. Human-mediated factors like the intraregional mussel trade and transplantation are also likely secondary dispersal mechanisms favouring the spread of this organism., Competing Interests: The authors have declared that no competing interests exist.

Determining the magnitude and causes of intrinsic variability is a main issue in the analysis of bivalve growth. Inter-individual variability in bivalve growth has been attributed to differences in the physiological performance. This hypothesis has been commonly tested comparing the physiological rates of fast and slow growers after size differentiation has occurred. This experimental design may detect a link between growth and physiological performance, but we cannot interpret the posterior physiological performance as a driver for the prior growth variability. Considering these limitations, this work introduces a new methodological framework for the analysis of bivalve growth variability. We have conducted sequential measurements of size and physiological performance (feeding, digestion and metabolic rates) in even-sized mussels growing under homogeneous environmental conditions. This experimental design allows us to distinguish between changes over time within individuals, i.e. growth and trends in the physiological rates, from differences between individuals with respect to a baseline level. In addition, Functional Data Analysis provides powerful tools to summarize all the information obtained in the exhaustive sampling scheme and to test whether differences in the physiological performance enhance growth dispersion. Our results report an increasing dispersion in both size and physiological performance over time. Although mussels grew during the experiment, it is difficult to detect any increasing or decreasing temporal pattern in their feeding, digestion and metabolic rates due to the large inter-individual variability. Comparison between the growth and physiological patterns of mussels with final size above (fast growers) and below (slow growers) the median found that fast growers had larger feeding and digestion rates and lower metabolic expenditures during the experimental culture than mussels with slow growth, which agrees with the hypothesis of a physiological basis for bivalve growth variability., Competing Interests: The authors have declared that no competing interests exist.

Flattening of coiled shells has occurred in several gastropod lineages, while the evolutionary process of shell flattening is little known. The subfamily Fossarininae of the top shell family (Trochidae) is unique, because it includes four genera at various stages of shell flattening. Broderipia and Roya, have zygomorphic shells that has lost coiling, while the sister genera, Fossarina and Synaptocochlea, have respectively turbiniform and auriform shells. Therefore, comparisons of biology, habitats and detailed morphology among these four genera may help us to detect selection pressure driving shell flattening and loss of coiling. Although Broderipia has recently been identified as living symbiotically in the pits of sea urchins, the habitats and biology of the other three Fossarininae species, especially Roya are poorly known. After an extensive search on rocky shores of the Japanese Archipelago, we found live Roya eximia snails on intertidal/subtidal rock surfaces exposed to strong waves. Roya snails crept on the bare rock surface to graze periphyton at low tide, and fled into vacant barnacle shells at high tide. Comparison of the morphology of soft bodies in Fossarininae revealed that the columellar muscle of flattened species has been drastically elongated and arranged in posterior semi-outer edge of the flattened shell as observed in true limpets. The flattering and loss of coiling of the shell in Roya caused acquisition of a zygomorphic flat body, retraction of coiled visceral mass, and expansion of the foot sole. All of these changes improved tolerance against strong waves and the ability to cling to rock surfaces, and thus enabled a lifestyle utilizing both wave-swept rock surfaces and the inside of vacant barnacle shells., Competing Interests: The authors have declared that no competing interests exist.

Invasive snails in the genus Pomacea have spread across Southeast Asia including Peninsular Malaysia. Their effects on natural and agricultural wetlands are appreciable, but species-specific effects are less clear because of morphological similarity among the species. Our objective was to establish diagnostic characteristics of Pomacea species in Malaysia using genetic and morphological criteria. The mitochondrial COI gene of 52 adult snails from eight localities in Peninsular Malaysia was amplified, sequenced, and analysed to verify species and phylogenetic relationships. Shells were compared using geometric morphometric and covariance analyses. Two monophyletic taxa, P. canaliculata and P. maculata, occurred in our samples. The mean ratio of shell height: aperture height (P = 0.042) and shell height: shell width (P = 0.007) was smaller in P. maculata. P. maculata co-occurred with P. canaliculata in five localities, but samples from three localities contained only P. canaliculata. This study is the first to confirm the presence of two of the most invasive species of Pomacea in Peninsular Malaysia using a molecular technique. P. canaliculata appears to be the more widespread species. Despite statistical differences, both quantitative and qualitative morphological characteristics demonstrated much interspecific overlap and intraspecific variability; thus, shell morphology alone cannot reliably verify species identity. Molecular techniques for distinguishing between these two highly invasive Pomacea species are needed to understand their specific ecological niches and to develop effective protocols for their management.

Late Cretaceous members of Peritresius belong to a diverse clade of marine adapted turtles currently thought to be some of the earliest representatives of the lineage leading to modern hard-shelled sea turtles (Pan-Cheloniidae). Prior studies have suggested that Peritresius was monospecific, with a distribution restricted to Maastrichtian deposits in North America. However, new Peritresius specimens identified from Alabama and Mississippi, USA, show that the genus contains two taxa, Peritresius ornatus, and a new species Peritresius martini sp. nov. These two taxa are characterized by the presence of a generally cordiform carapace with moderately serrated peripherals, well-developed ventral flanges beginning at the third peripheral, squarish umbilical and lateral plastral fontanelles, and a narrow bridge formed by the contact between the hyoplastron and hypoplastron. Peritresius martini sp. nov. can be distinguished by its lack of dermal ornamentation and the presence of a 'rib-free' 10th peripheral. These new specimens represent the first occurrences of Peritresius from the Late Cretaceous Mississippi Embayment and extend the temporal range of this genus back to the early Campanian. When tested within a global phylogenetic context, Peritresius is placed on the stem of Cheloniidae (Pan-Cheloniidae) along with Ctenochelys and Allopleuron hofmanni. The heavily vascularized and uniquely sculptured dermal elements of P. ornatus are interpreted here as potentially relating to thermoregulation and therefore may have been one of the key factors contributing to the survival of Peritresius into the Maastrichtian, a period of cooling when other lineages of Campanian marine turtles (e.g., Protostegids, Toxochelys, and Ctenochelys) went extinct.

Characterization of morphological variation in the shells of extant Eastern Box Turtles, Terrapene carolina, provides a baseline for comparison to fossil populations. It also provides an example of the difficulties inherent to recognizing intraspecific diversity in the fossil record. The degree to which variation in fossils of T. carolina can be accommodated by extant variation in the species has been disagreed upon for over eighty years. Using morphometric analyses of the carapace, I address the relationship between modern and fossil T. carolina in terms of sexual dimorphism, geographic and subspecific variation, and allometric variation. Modern T. carolina display weak male-biased sexual size dimorphism. Sexual shape dimorphism cannot be reliably detected in the fossil record. Rather than a four-part subspecific division, patterns of geographic variation are more consistent with clinal variation between various regions in the species distribution. Allometric patterns are qualitatively similar to those documented in other emydid turtles and explain a significant amount of shape variation. When allometric patterns are accounted for, Holocene specimens are not significantly different from modern specimens. In contrast, several geologically older specimens have significantly different carapace shape with no modern analogue. Those large, fossilized specimens represent extinct variation occupying novel portions of morphospace. This study highlights the need for additional documentation of modern osteological variation that can be used to test hypotheses of intraspecific evolution in the fossil record.

Empty mollusk shells may act as colonization surfaces for sclerobionts depending on the physical, chemical, and biological attributes of the shells. However, the main factors that can affect the establishment of an organism on hard substrates and the colonization patterns on modern and time-averaged shells remain unclear. Using experimental and field approaches, we compared sclerobiont (i.e., bacteria and invertebrate) colonization patterns on the exposed shells (internal and external sides) of three bivalve species (Anadara brasiliana, Mactra isabelleana, and Amarilladesma mactroides) with different external shell textures. In addition, we evaluated the influence of the host characteristics (mode of life, body size, color alteration, external and internal ornamentation and mineralogy) of sclerobionts on dead mollusk shells (bivalve and gastropod) collected from the Southern Brazilian coast. Finally, we compared field observations with experiments to evaluate how the biological signs of the present-day invertebrate settlements are preserved in molluscan death assemblages (incipient fossil record) in a subtropical shallow coastal setting. The results enhance our understanding of sclerobiont colonization over modern and paleoecology perspectives. The data suggest that sclerobiont settlement is enhanced by (i) high(er) biofilm bacteria density, which is more attracted to surfaces with high ornamentation; (ii) heterogeneous internal and external shell surface; (iii) shallow infaunal or attached epifaunal life modes; (iv) colorful or post-mortem oxidized shell surfaces; (v) shell size (<50 mm2 or >1,351 mm2); and (vi) calcitic mineralogy. Although the biofilm bacteria density, shell size, and texture are considered the most important factors, the effects of other covarying attributes should also be considered. We observed a similar pattern of sclerobiont colonization frequency over modern and paleoecology perspectives, with an increase of invertebrates occurring on textured bivalve shells. This study demonstrates how bacterial biofilms may influence sclerobiont colonization on biological hosts (mollusks), and shows how ecological relationships in marine organisms may be relevant for interpreting the fossil record of sclerobionts.

We report the results of 212 radiocarbon determinations from the archaeological excavation of 70 shell mound deposits in the Wathayn region of Albatross Bay, Australia. This is an intensive study of a closely co-located group of mounds within a geographically restricted area in a wider region where many more shell mounds have been reported. Valves from the bivalve Tegillarca granosa (Linnaeus, 1758) were dated. The dates obtained are used to calculate rates of accumulation for the shell mound deposits. These demonstrate highly variable rates of accumulation both within and between mounds. We assess these results in relation to likely mechanisms of shell deposition and show that rates of deposition are affected by time-dependent processes both during the accumulation of shell deposits and during their subsequent deformation. This complicates the interpretation of the rates at which shell mound deposits appear to have accumulated. At Wathayn, there is little temporal or spatial consistency in the rates at which mounds accumulated. Comparisons between the Wathayn results and those obtained from shell deposits elsewhere, both in the wider Albatross Bay region and worldwide, suggest the need for caution when deriving behavioural inferences from shell mound deposition rates, and the need for more comprehensive sampling of individual mounds and groups of mounds.

Identifying impacts of exotic species on native populations is central to ecology and conservation. Although the effects of exotic predators on native prey have received much attention, the role of exotic prey on native predators is poorly understood. Determining if native predators actively prefer invasive prey over native prey has implications for interpreting invasion impacts, identifying the presence of evolutionary traps, and predator persistence. One of the world's most invasive species, Pomacea maculata, has recently established in portions of the endangered Everglade snail kite's (Rostrhamus sociabilis plumbeus) geographic range. Although these exotic snails could provide additional prey resources, they are typically much larger than the native snail, which can lead to lower foraging success and the potential for diminished energetic benefits in comparison to native snails. Nonetheless, snail kites frequently forage on exotic snails. We used choice experiments to evaluate snail kite foraging preference in relation to exotic species and snail size. We found that snail kites do not show a preference for native or exotic snails. Rather, snail kites generally showed a preference for medium-sized snails, the sizes reflective of large native snails. These results suggest that while snail kites frequently forage on exotic snails in the wild, this behavior is likely driven simply by the abundance of exotic snails rather than snail kites preferring exotics. This lack of preference offers insights to hypotheses regarding effects of exotic species, guidance regarding habitat and invasive species management, and illustrates how native-exotic relationships can be misleading in the absence of experimental tests of such interactions., Competing Interests: The authors have declared that no competing interests exist.

The marine snail Littorina saxatilis exhibits extreme morphological variation between and within geographical regions and represents an excellent model for assessing local adaptation. Previous studies support the hypothesis of parallel evolution in sympatry of two morphologically different ecotypes (named as RB and SU) that co-inhabit different habitats from Galician rocky shores (NW Spain), and which are interrupted by sheltered areas inhabited by a different morph never studied before (named as SRB). Here, we use morphological and mitochondrial DNA (mtDNA) sequence data to test hypotheses on the origin and diversification of SRB snails and to assess their evolutionary relationships with RB and SU ecotypes. Our results show that the SRB morph displays the largest size and shell elongation and the smallest relative shell aperture, representing an extreme type of the RB vs. SU polymorphism, which has been linked to adaptation to sheltered ecological factors. Phylogenetic analysis shows that the SRB morph shares ancestry with RB and SU ecotypes, rejecting the hypothesis that the SRB morph marks relict populations from which these ecotypes evolved in Galician coasts. Our data support that genetic differentiation among SRB, RB and SU morphs results from a general pattern of restricted gene flow and isolation by distance linked to the colonization of Galician coasts by two independent mtDNA lineages, rather than from a random fragmentation of the initial distributional range. Therefore, the confinement of distinct lineages to specific geographical areas denote evident limits to the distances these snails can disperse. Morphological analysis indicates no association between mtDNA lineage and a specific morphotype, and suggests the independent gain of convergent morphological patterns within each mtDNA lineage in populations occupying contrasting habitats following the colonization of Galician coasts.

Quantitative analysis of organismal form is an important component for almost every branch of biology. Although generally considered an easily-measurable structure, the quantification of gastropod shell form is still a challenge because many shells lack homologous structures and have a spiral form that is difficult to capture with linear measurements. In view of this, we adopt the idea of theoretical modelling of shell form, in which the shell form is the product of aperture ontogeny profiles in terms of aperture growth trajectory that is quantified as curvature and torsion, and of aperture form that is represented by size and shape. We develop a workflow for the analysis of shell forms based on the aperture ontogeny profile, starting from the procedure of data preparation (retopologising the shell model), via data acquisition (calculation of aperture growth trajectory, aperture form and ontogeny axis), and data presentation (qualitative comparison between shell forms) and ending with data analysis (quantitative comparison between shell forms). We evaluate our methods on representative shells of the genera Opisthostoma and Plectostoma, which exhibit great variability in shell form. The outcome suggests that our method is a robust, reproducible, and versatile approach for the analysis of shell form. Finally, we propose several potential applications of our methods in functional morphology, theoretical modelling, taxonomy, and evolutionary biology.

The zebra mussel Dreissena polymorpha is a well-established invasive model organism. Although extensively used in environmental sciences, virtually nothing is known of the molecular process of its shell calcification. By describing the microstructure, geochemistry and biochemistry/proteomics of the shell, the present study aims at promoting this species as a model organism in biomineralization studies, in order to establish a bridge with ecotoxicology, while sketching evolutionary conclusions. The shell of D. polymorpha exhibits the classical crossed-lamellar/complex crossed lamellar combination found in several heterodont bivalves, in addition to an external thin layer, the characteristics of which differ from what was described in earlier publication. We show that the shell selectively concentrates some heavy metals, in particular uranium, which predisposes D. polymorpha to local bioremediation of this pollutant. We establish the biochemical signature of the shell matrix, demonstrating that it interacts with the in vitro precipitation of calcium carbonate and inhibits calcium carbonate crystal formation, but these two properties are not strongly expressed. This matrix, although overall weakly glycosylated, contains a set of putatively calcium-binding proteins and a set of acidic sulphated proteins. 2D-gels reveal more than fifty proteins, twenty of which we identify by MS-MS analysis. We tentatively link the shell protein profile of D. polymorpha and the peculiar recent evolution of this invasive species of Ponto-Caspian origin, which has spread all across Europe in the last three centuries.

The Ammonoidea is a group of extinct cephalopods ideal to study evolution through deep time. The evolution of the planispiral shell and complexly folded septa in ammonoids has been thought to have increased the functional surface area of the chambers permitting enhanced metabolic functions such as: chamber emptying, rate of mineralization and increased growth rates throughout ontogeny. Using nano-computed tomography and synchrotron radiation based micro-computed tomography, we present the first study of ontogenetic changes in surface area to volume ratios in the phragmocone chambers of several phylogenetically distant ammonoids and extant cephalopods. Contrary to the initial hypothesis, ammonoids do not possess a persistently high relative chamber surface area. Instead, the functional surface area of the chambers is higher in earliest ontogeny when compared to Spirula spirula. The higher the functional surface area the quicker the potential emptying rate of the chamber; quicker chamber emptying rates would theoretically permit faster growth. This is supported by the persistently higher siphuncular surface area to chamber volume ratio we collected for the ammonite Amauroceras sp. compared to either S. spirula or nautilids. We demonstrate that the curvature of the surface of the chamber increases with greater septal complexity increasing the potential refilling rates. We further show a unique relationship between ammonoid chamber shape and size that does not exist in S. spirula or nautilids. This view of chamber function also has implications for the evolution of the internal shell of coleoids, relating this event to the decoupling of soft-body growth and shell growth.

Probiotics are widely used in nutrition, and their mode of action is intensively studied in mammals and birds; however, it is almost unknown in reptiles. In the present study, Trachemys scripta scripta and Sternotherus odoratus were used to assess the effects of dietary probiotics on chelonian gastrointestinal tract microecology. In the first, 20-week experiment, 40 young T. s. scripta were randomly distributed to four experimental groups: 1st, (CON)--with no additives; 2nd, (SSPA) with Bacillus subtilis PB6; 3rd, (MSP)--with multiple strain probiotic; and 4th, (SSPB) with Bacillus subtilis C-3102. The first study has shown that SSPA and MSP decreased the numbers of total bacteria, Enterobacteriace, Staphylococcus sp. and Streptococcus sp. excreted to water and increased the villous height and mucosa thickness in duodenum. SSPB improved the duodenal microstructure; however, it also increased numbers of kanamycin and vancomycin resistant bacteria, Staphylococcus sp. and Streptococcus sp., in water. In the second, 52-week experiment, 30 S. odoratus were randomly assigned to three dietary treatments. CON, SSPA and MSP groups. The MSP preparation increased the body weight gain, crude ash, Ca and P share in the turtles' shells. Both probiotics affected duodenal histomorphology. SSPA decreased the villous height, while MSP increased the villous height and mucosa thickness, and decreased the crypt depth. SSPA decreased the concentrations of bacteria excreted to water. In the case of intestinal microbiota, bacteria suppressing effects were observed in the case of both probiotics. MSP increased the number of Bifidobacterium sp. and Lactobacillus sp./Enteroccoccus sp., and decreased the number of Clostridium perfringens and Campylobacter sp. in the small intestine. In the large intestine it lowered, amongst others, Bacteroides-Pervotella cluster, Clostridium leptum subgroup and Clostridium perfringens numbers. The above-mentioned results suggest that probiotics are useful in turtle nutrition due to their positive effects on growth performance, shell mineralization, duodenal histomorphology and microbiota.

The exploitation of mollusks by the first anatomically modern humans is a central question for archaeologists. This paper focuses on level 8 (dated around ∼ 100 ka BP) of El Harhoura 2 Cave, located along the coastline in the Rabat-Témara region (Morocco). The large quantity of Patella sp. shells found in this level highlights questions regarding their origin and preservation. This study presents an estimation of the preservation status of these shells. We focus here on the diagenetic evolution of both the microstructural patterns and organic components of crossed-foliated shell layers, in order to assess the viability of further investigations based on shell layer minor elements, isotopic or biochemical compositions. The results show that the shells seem to be well conserved, with microstructural patterns preserved down to sub-micrometric scales, and that some organic components are still present in situ. But faint taphonomic degradations affecting both mineral and organic components are nonetheless evidenced, such as the disappearance of organic envelopes surrounding crossed-foliated lamellae, combined with a partial recrystallization of the lamellae. Our results provide a solid case-study of the early stages of the diagenetic evolution of crossed-foliated shell layers. Moreover, they highlight the fact that extreme caution must be taken before using fossil shells for palaeoenvironmental or geochronological reconstructions. Without thorough investigation, the alteration patterns illustrated here would easily have gone unnoticed. However, these degradations are liable to bias any proxy based on the elemental, isotopic or biochemical composition of the shells. This study also provides significant data concerning human subsistence behavior: the presence of notches and the good preservation state of limpet shells (no dissolution/recrystallization, no bioerosion and no abrasion/fragmentation aspects) would attest that limpets were gathered alive with tools by Middle Palaeolithic (Aterian) populations in North Africa for consumption.

As typical biofoulers, barnacles possess hard shells and cause serious biofouling problems. In this study, we analyzed the protein component of the barnacle Amphibalanus (= Balanus) amphitrite shell using gel-based proteomics. The results revealed 52 proteins in the A. Amphitrite shell. Among them, 40 proteins were categorized into 11 functional groups based on KOG database, and the remaining 12 proteins were unknown. Besides the known proteins in barnacle shell (SIPC, carbonic anhydrase and acidic acid matrix protein), we also identified chorion peroxidase, C-type lectin-like domains, serine proteases and proteinase inhibitor proteins in the A. Amphitrite shell. The sequences of these proteins were characterized and their potential functions were discussed. Histology and DAPI staining revealed living cells in the shell, which might secrete the shell proteins identified in this study.

The freshwater pearl mussel family Margaritiferidae includes 13 extant species, which are all listed by IUCN as endangered or vulnerable taxa. In this study, an extensive spatial sampling of Margaritifera spp. across the Russian Far East (Amur Basin, Kamchatka Peninsula, Kurile Archipelago and Sakhalin Island) was conducted for a revision of their taxonomy and distribution ranges. Based on their DNA sequences, shell and soft tissue morphology, three valid species were identified: Margaritifera dahurica (Middendorff, 1850), M. laevis (Haas, 1910) and M. middendorffi (Rosén, 1926). M. dahurica ranges across the Amur basin and some of the nearest river systems. M. laevis is distributed in Japan, Sakhalin Island and the Kurile Archipelago. M. middendorffi was previously considered an endemic species of the Kamchatka. However, it is widespread in the rivers of Kamchatka, Sakhalin Island, the Kurile Islands (across the Bussol Strait, which is the most significant biogeographical boundary within the archipelago), and, likely, in Japan. The Japanese species M. togakushiensis Kondo & Kobayashi, 2005 seems to be conspecific with M. middendorffi because of similar morphological patterns, small shell size (<100 mm long) and overlapped ranges, but it is in need of a separate revision. Phylogenetic analysis reveals that two NW Pacific margaritiferid species, M. laevis and M. middendorffi, formed a monophyletic 18S rDNA clade together with the North American species M. marrianae and M. falcata. The patterns that were found in these Margaritifera spp. are similar to those of freshwater fishes, indicating multiple colonizations of Eastern Asia by different mitochondrial lineages, including an ancient Beringian exchange between freshwater faunas across the Pacific.

We explored variation in patterns of percussive stone-tool use on coastal foods by Burmese long-tailed macaques (Macaca fascicularis aurea) from two islands in Laem Son National Park, Ranong, Thailand. We catalogued variation into three hammering classes and 17 action patterns, after examining 638 tool-use bouts across 90 individuals. Hammering class was based on the stone surface used for striking food, being face, point, and edge hammering. Action patterns were discriminated by tool material, hand use, posture, and striking motion. Hammering class was analyzed for associations with material and behavioural elements of tool use. Action patterns were not, owing to insufficient instances of most patterns. We collected 3077 scan samples from 109 macaques on Piak Nam Yai Island's coasts, to determine the proportion of individuals using each hammering class and action pattern. Point hammering was significantly more associated with sessile foods, smaller tools, faster striking rates, smoother recoil, unimanual use, and more varied striking direction, than were face and edge hammering, while both point and edge hammering were significantly more associated with precision gripping than face hammering. Edge hammering also showed distinct differences depending on whether such hammering was applied to sessile or unattached foods, resembling point hammering for sessile foods and face hammering for unattached foods. Point hammering and sessile edge hammering compared to prior descriptions of axe hammering, while face and unattached edge hammering compared to pound hammering. Analysis of scans showed that 80% of individuals used tools, each employing one to four different action patterns. The most common patterns were unimanual point hammering (58%), symmetrical-bimanual face hammering (47%) and unimanual face hammering (37%). Unimanual edge hammering was relatively frequent (13%), compared to the other thirteen rare action patterns (<5%). We compare our study to other stone-using primates, and discuss implications for further research.

Philobryids (Bivalvia: Arcoida) are one of the most speciose marine bivalve families in the Southern Ocean and are common throughout the Southern Hemisphere. Considering this diversity and their brooding reproductive mode (limiting long-distance dispersal), this family may have been present in the Southern Ocean since its inception. However Philobrya and Adacnarca appear only in the Quaternary fossil record of the Antarctic, suggesting a much more recent incursion. Molecular dating provides an independent means of measuring the time of origin and radiation of this poorly known group. Here we present the first combined molecular and morphological investigation of the Philobryidae in the Southern Ocean. Two nuclear loci (18S and 28S) were amplified from 35 Southern Ocean Adacnarca and Philobrya specimens, with a combined sequence length of 2,282 base pairs (bp). Adacnarca specimens (A. nitens and A. limopsoides) were resolved as a strongly supported monophyletic group. Genus Philobrya fell into two strongly supported groups ('sublaevis' and 'magellanica/wandelensis'), paraphyletic with Adacnarca. The A. nitens species complex is identified as at least seven morpho-species through morphological and genetic analysis of taxon clustering. Phylogenetic analyses resolve Philobryidae as a strongly supported monophyletic clade and sister taxon to the Limopsidae, as anticipated by their classification into the superfamily Limopsoidea. Bayesian relaxed clock analyses of divergence times suggest that genus Adacnarca radiated in the Southern Ocean from the Early Paleogene, while P. sublaevis and P. wandelensis clades radiated in the late Miocene, following the formation of the Antarctic Circumpolar Current.

The biology of modern Conidae (cone snails)--which includes the hyperdiverse genus Conus--has been intensively studied, but the fossil record of the clade remains poorly understood, particularly within an evolutionary framework. Here, ultraviolet light is used to reveal and characterize the original shell coloration patterns of 28 species of cone snails from three Neogene coral reef-associated deposits from the Cibao Valley, northern Dominican Republic. These fossils come from the upper Miocene Cercado Fm. and lower Pliocene Gurabo Fm., and range in age from about 6.6-4.8 Ma. Comparison of the revealed coloration patterns with those of extant species allow the taxa to be assigned to three genera of cone snails (Profundiconus, Conasprella, and Conus) and at least nine subgenera. Thirteen members of these phylogenetically diverse reef faunas are described as new species. These include: Profundiconus? hennigi, Conasprella (Ximeniconus) ageri, Conus anningae, Conus lyelli, Conus (Atlanticonus?) franklinae, Conus (Stephanoconus) gouldi, Conus (Stephanoconus) bellacoensis, Conus (Ductoconus) cashi, Conus (Dauciconus) garrisoni, Conus (Dauciconus?) zambaensis, Conus (Spuriconus?) kaesleri, Conus (Spuriconus?) lombardii, and Conus (Lautoconus?) carlottae. Each of the three reef deposits contain a minimum of 14-16 cone snail species, levels of diversity that are similar to modern Indo-Pacific reef systems. Finally, most of the 28 species can be assigned to modern clades and thus have important implications for understanding the biogeographic and temporal histories of these clades in tropical America.

The loggerhead sea turtle (Caretta caretta) is found throughout the waters of the Atlantic, Pacific, and Indian Oceans. It is a protected species throughout much of its range due to threats such as habitat loss, fisheries interactions, hatchling predation, and marine debris. Loggerheads that occur in the southeastern U.S. are listed as "threatened" on the U.S. Endangered Species List, and receive state and federal protection. As part of an on-going population assessment conducted by the National Marine Fisheries Service, samples were collected from juvenile loggerhead sea turtles in Core Sound, North Carolina, between 2004 and 2007 to gain insight on the baseline health of the threatened Northwest Atlantic Ocean population. The aims of the current study were to establish hematologic and biochemical reference intervals for this population, and to assess variation of the hematologic and plasma biochemical analytes by season, water temperature, and sex and size of the turtles. Reference intervals for the clinical pathology parameters were estimated following Clinical Laboratory Standards Institute guidelines. Season, water temperature, sex, and size of the turtles were found to be significant factors of variation for parameter values. Seasonal variation could be attributed to physiological effects of decreasing photoperiod, cooler water temperature, and migration during the fall months. Packed cell volume, total protein, and albumin increased with increasing size of the turtles. The size-related differences in analytes documented in the present study are consistent with other reports of variation in clinical pathology parameters by size and age in sea turtles. As a component of a health assessment of juvenile loggerhead sea turtles in North Carolina, this study will serve as a baseline aiding in evaluation of trends for this population and as a diagnostic tool for assessing the health and prognosis for loggerhead sea turtles undergoing rehabilitation.

An approach is developed to estimate size of Iceland scallop shells from AUV photos. A small-scale camera based AUV survey of Iceland scallops was conducted at a defined site off West Iceland. Prior to height estimation of the identified shells, the distortions introduced by the vehicle orientation and the camera lens were corrected. The average AUV pitch and roll was 1.3 and 2.3 deg that resulted in <2% error in ground distance rendering these effects negligible. A quadratic polynomial model was identified for lens distortion correction. This model successfully predicted a theoretical grid from a frame photographed underwater, representing the inherent lens distortion. The predicted shell heights were scaled for the distance from the bottom at which the photos were taken. This approach was validated by height estimation of scallops of known sizes. An underestimation of approximately 0.5 cm was seen, which could be attributed to pixel error, where each pixel represented 0.24 x 0.27 cm. After correcting for this difference the estimated heights ranged from 3.8-9.3 cm. A comparison of the height-distribution from a small-scale dredge survey carried out in the vicinity showed non-overlapping peaks in size distribution, with scallops of a broader size range visible in the AUV survey. Further investigations are necessary to evaluate any underlying bias and to validate how representative these surveys are of the true population. The low resolution images made identification of smaller scallops difficult. Overall, the observations of very few small scallops in both surveys could be attributed to low recruitment levels in the recent years due to the known scallop parasite outbreak in the region.

A high degree of intraspecific variation, both genetic and in shell morphology, of the operculate land snail Cyclophorus fulguratus (Pfeiffer, 1854) suggests that its classification as a single species warrants reconsideration. We sequenced two nuclear (18S and 28S) and two mitochondrial (16S and COI) genes of 46 C. fulguratus specimens and used them to estimate the phylogeny and to determine the validity of species boundaries. Molecular phylogenetic analyses revealed the presence of three lineages corresponding to three geographically disjunctive populations of C. fulguratus in Thailand. Likelihood tests of topologies significantly supported the non-monophyly of the C. fulguratus-complex and Bayesian species delimitation analysis significantly supported the potential representation as distinct species of these three lineages. Discriminant function analysis based on geometric-morphometrics of shell shape allowed for significant distinction of these three candidate species, although they revealed a considerable degree of overlap of shell shape reflecting their crypsis morphologically. The diagnostic characters are provided by color pattern, pattern of protoconch and pattern of jaw. In conclusion, the results support that the C. fulguratus s.l., as currently recognized, consists of three distinct species in Thailand: C. fulguratus s.s., C. rangunensis and C. abditus sp.nov., which are described herein.

Shell beads appear to have been one of the earliest examples of personal adornments. Marine shells identified far from the shore evidence long-distance transport and imply networks of exchange and negotiation. However, worked beads lose taxonomic clues to identification, and this may be compounded by taphonomic alteration. Consequently, the significance of this key early artefact may be underestimated. We report the use of bulk amino acid composition of the stable intra-crystalline proteins preserved in shell biominerals and the application of pattern recognition methods to a large dataset (777 samples) to demonstrate that taxonomic identification can be achieved at genus level. Amino acid analyses are fast (<2 hours per sample) and micro-destructive (sample size <2 mg). Their integration with non-destructive techniques provides a valuable and affordable tool, which can be used by archaeologists and museum curators to gain insight into early exploitation of natural resources by humans. Here we combine amino acid analyses, macro- and microstructural observations (by light microscopy and scanning electron microscopy) and Raman spectroscopy to try to identify the raw material used for beads discovered at the Early Bronze Age site of Great Cornard (UK). Our results show that at least two shell taxa were used and we hypothesise that these were sourced locally.

Early life history stages of marine organisms are generally thought to be more sensitive to environmental stress than adults. Although most marine invertebrates are broadcast spawners, some species are brooders and/or protect their embryos in egg or capsules. Brooding and encapsulation strategies are typically assumed to confer greater safety and protection to embryos, although little is known about the physico-chemical conditions within egg capsules. In the context of ocean acidification, the protective role of encapsulation remains to be investigated. To address this issue, we conducted experiments on the gastropod Crepidula fornicata. This species broods its embryos within capsules located under the female and veliger larvae are released directly into the water column. C. fornicata adults were reared at the current level of CO2 partial pressure (pCO2) (390 μatm) and at elevated levels (750 and 1400 μatm) before and after fertilization and until larval release, such that larval development occurred entirely at a given pCO2. The pCO2 effects on shell morphology, the frequency of abnormalities and mineralization level were investigated on released larvae. Shell length decreased by 6% and shell surface area by 11% at elevated pCO2 (1400 μatm). The percentage of abnormalities was 1.5- to 4-fold higher at 750 μatm and 1400 μatm pCO2, respectively, than at 390 μatm. The intensity of birefringence, used as a proxy for the mineralization level of the larval shell, also decreased with increasing pCO2. These negative results are likely explained by increased intracapsular acidosis due to elevated pCO2 in extracapsular seawater. The encapsulation of C. fornicata embryos did not protect them against the deleterious effects of a predicted pCO2 increase. Nevertheless, C. fornicata larvae seemed less affected than other mollusk species. Further studies are needed to identify the critical points of the life cycle in this species in light of future ocean acidification.

A detailed investigation of the shell formation of the palaeoheterodont 'living fossil' Neotrigonia concentrated on the timing and manufacture of the calcified 'bosses' which stud the outside of all trigonioid bivalves (extant and fossil) has been conducted. Electron microscopy and optical microscopy revealed that Neotrigonia spp. have a spiral-shaped periostracal groove. The periostracum itself is secreted by the basal cell, as a thin dark pellicle, becoming progressively transformed into a thin dark layer by additions of secretions from the internal outer mantle fold. Later, intense secretion of the internal surface of the outer mantle fold forms a translucent layer, which becomes transformed by tanning into a dark layer. The initiation of calcified bosses occurred at a very early stage of periostracum formation, deep within the periostracal groove immediately below the initialmost dark layer. At this stage, they consist of a series of polycyclically twinned crystals. The bosses grow as the periostracum traverse through the periostracal groove, in coordination with the thickening of the dark periostracal layer and until, upon reaching the mantle edge, they impinge upon each other and become transformed into large prisms separated by dark periostracal walls. In conclusion, the initial bosses and the external part of the prismatic layer are fully intraperiostracal. With later growth, the prisms transform into fibrous aggregates, although the details of the process are unknown. This reinforces the relationships with other groups that have the ability to form intraperiostracal calcifications, for example the unionoids with which the trigonioids form the clade Paleoheterodonta. The presence of similar structures in anomalodesmatans and other euheterodonts raises the question of whether this indicates a relationship or represents a convergence. The identification of very early calcification within an organic sheet has interesting implications for our understanding of how shells may have evolved.

Ocean surface pH levels are predicted to fall by 0.3-0.4 pH units by the end of the century and are likely to coincide with an increase in sea surface temperature of 2-4 °C. The combined effect of ocean acidification and warming on the functional properties of bivalve shells is largely unknown and of growing concern as the shell provides protection from mechanical and environmental challenges. We examined the effects of near-future pH (ambient pH -0.4 pH units) and warming (ambient temperature +4 °C) on the shells of the commercially important bivalve, Mytilus edulis when fed for a limited period (4-6 h day(-1)). After six months exposure, warming, but not acidification, significantly reduced shell strength determined as reductions in the maximum load endured by the shells. However, acidification resulted in a reduction in shell flex before failure. Reductions in shell strength with warming could not be explained by alterations in morphology, or shell composition but were accompanied by reductions in shell surface area, and by a fall in whole-body condition index. It appears that warming has an indirect effect on shell strength by re-allocating energy from shell formation to support temperature-related increases in maintenance costs, especially as food supply was limited and the mussels were probably relying on internal energy reserves. The maintenance of shell strength despite seawater acidification suggests that biomineralisation processes are unaffected by the associated changes in CaCO3 saturation levels. We conclude that under near-future climate change conditions, ocean warming will pose a greater risk to shell integrity in M. edulis than ocean acidification when food availability is limited.

Ecophenotypes reflect local matches between organisms and their environment, and show plasticity across generations in response to current living conditions. Plastic responses in shell morphology and shell growth have been widely studied in gastropods and are often related to environmental calcium availability, which influences shell biomineralisation. To date, all of these studies have overlooked micro-scale structure of the shell, in addition to how it is related to species responses in the context of environmental pressure. This study is the first to demonstrate that environmental factors induce a bi-modal variation in the shell micro-scale structure of a land gastropod. Notodiscus hookeri is the only native land snail present in the Crozet Archipelago (sub-Antarctic region). The adults have evolved into two ecophenotypes, which are referred to here as MS (mineral shell) and OS (organic shell). The MS-ecophenotype is characterised by a thick mineralised shell. It is primarily distributed along the coastline, and could be associated to the presence of exchangeable calcium in the clay minerals of the soils. The Os-ecophenotype is characterised by a thin organic shell. It is primarily distributed at high altitudes in the mesic and xeric fell-fields in soils with large particles that lack clay and exchangeable calcium. Snails of the Os-ecophenotype are characterised by thinner and larger shell sizes compared to snails of the MS-ecophenotype, indicating a trade-off between mineral thickness and shell size. This pattern increased along a temporal scale; whereby, older adult snails were more clearly separated into two clusters compared to the younger adult snails. The prevalence of glycine-rich proteins in the organic shell layer of N. hookeri, along with the absence of chitin, differs to the organic scaffolds of molluscan biominerals. The present study provides new insights for testing the adaptive value of phenotypic plasticity in response to spatial and temporal environmental variations.

The majority of marine benthic invertebrates protect themselves from predators by producing calcareous tubes or shells that have remarkable mechanical strength. An elevation of CO2 or a decrease in pH in the environment can reduce intracellular pH at the site of calcification and thus interfere with animal's ability to accrete CaCO3. In nature, decreased pH in combination with stressors associated with climate change may result in the animal producing severely damaged and mechanically weak tubes. This study investigated how the interaction of environmental drivers affects production of calcareous tubes by the serpulid tubeworm, Hydroides elegans. In a factorial manipulative experiment, we analyzed the effects of pH (8.1 and 7.8), salinity (34 and 27‰), and temperature (23°C and 29°C) on the biomineral composition, ultrastructure and mechanical properties of the tubes. At an elevated temperature of 29°C, the tube calcite/aragonite ratio and Mg/Ca ratio were both increased, the Sr/Ca ratio was decreased, and the amorphous CaCO3 content was reduced. Notably, at elevated temperature with decreased pH and reduced salinity, the constructed tubes had a more compact ultrastructure with enhanced hardness and elasticity compared to decreased pH at ambient temperature. Thus, elevated temperature rescued the decreased pH-induced tube impairments. This indicates that tubeworms are likely to thrive in early subtropical summer climate. In the context of climate change, tubeworms could be resilient to the projected near-future decreased pH or salinity as long as surface seawater temperature rise at least by 4°C.

Freshwater snails in the family Bithyniidae are the first intermediate host for Southeast Asian liver fluke (Opisthorchis viverrini), the causative agent of opisthorchiasis. Unfortunately, the subtle morphological characters that differentiate species in this group are not easily discerned by non-specialists. This is a serious matter because the identification of bithyniid species is a fundamental prerequisite for better understanding of the epidemiology of this disease. Because DNA barcoding, the analysis of sequence diversity in the 5' region of the mitochondrial COI gene, has shown strong performance in other taxonomic groups, we decided to test its capacity to resolve 10 species/ subspecies of bithyniids from Thailand. Our analysis of 217 specimens indicated that COI sequences delivered species-level identification for 9 of 10 currently recognized species. The mean intraspecific divergence of COI was 2.3% (range 0-9.2 %), whereas sequence divergences between congeneric species averaged 8.7% (range 0-22.2 %). Although our results indicate that DNA barcoding can differentiate species of these medically-important snails, we also detected evidence for the presence of one overlooked species and one possible case of synonymy.

A scanty but varied ensemble of finds challenges the idea that Neandertal material culture was essentially static and did not include symbolic items. In this study we report on a fragmentary Miocene-Pliocene fossil marine shell, Aspamarginata, discovered in a Discoid Mousterian layer of the Fumane Cave, northern Italy, dated to at least 47.6-45.0 Cal ky BP. The shell was collected by Neandertals at a fossil exposure probably located more than 100 kms from the site. Microscopic analysis of the shell surface identifies clusters of striations on the inner lip. A dark red substance, trapped inside micropits produced by bioeroders, is interpreted as pigment that was homogeneously smeared on the outer shell surface. Dispersive X-ray and Raman analysis identify the pigment as pure hematite. Of the four hypotheses we considered to explain the presence of this object at the site, two (tool, pigment container) are discarded because in contradiction with observations. Although the other two ("manuport", personal ornament) are both possible, we favor the hypothesis that the object was modified and suspended by a 'thread' for visual display as a pendant. Together with contextual and chronometric data, our results support the hypothesis that deliberate transport and coloring of an exotic object, and perhaps its use as pendant, was a component of Neandertal symbolic culture, well before the earliest appearance of the anatomically modern humans in Europe.

We propose an integrative approach that explains patterns of recruitment to adult populations in sessile organisms by considering the numbers of individuals and their body size. A recruitment model, based on a small number of parameters, was developed for sessile organisms and tested using the barnacle Semibalanus balanoides, a marine invertebrate inhabiting North Atlantic intertidal shores. Incorporating barnacle body size improved model fit beyond that based on density alone, showing that growth played an important role in how resource limitation affected survival. Our approach uncovered the following: First, changes in the shape of the recruitment curve resulted from the balance between individual growth and mortality. Second, recruitment was limited by the least plastic trait used to characterise body size, operculum area. Basal area, a trait that responded to increases in barnacle density, did not contribute significantly to explain patterns of recruitment. Third, some temporal variation is explained by changes in the amount of space occupied by shells of dead barnacles: at high cover barnacles are densely packed and these shells remain long after death. Fourth, seasonal variation and spatial variation in survival can be separated from that resulting from resource limitation; survival was predicted for two different shores and four sampling times using a single recruitment model. We conclude that applying this integrative approach to recruitment will lead to a considerable advance in understanding patterns of mortality of early stages of sessile organisms.

Mollusk remains are abundant in archaeological sites in the Guanzhong Basin of Northwestern China, providing good opportunities for investigations into the use of mollusks by prehistoric humans. Here we report on freshwater gastropod and bivalve mollusks covering the time interval from about 5600 to 4500 cal. yrs BP from sites of Mid-Late Neolithic age. They are identified as Cipangopaludina chinensis and Unio douglasiae, both of which are currently food for humans. The shells are well preserved and have no signs of abrasion. They are all freshwater gastropods and bivalves found in pits without water-reworked deposits and have modern representatives which can be observed in rivers, reservoirs, and paddy fields in the studied region. Mollusk shells were frequently recovered in association with mammal bones, lithic artifacts, and pottery. These lines of evidence indicate that the mollusks are the remains of prehistoric meals. The mollusk shells were likely discarded into the pits by prehistoric humans after the flesh was eaten. However, these mollusk remains may not have been staple food since they are not found in large quantities. Mollusk shell tools and ornaments are also observed. Shell tools include shell knives, shell reaphooks and arrowheads, whereas shell ornaments are composed of pendants and loops. All the shell tools and ornaments are made of bivalve mollusks and do not occur in large numbers. The finding of these freshwater mollusk remains supports the view that the middle Holocene climate in the Guanzhong Basin may have been warm and moist, which was probably favorable to freshwater mollusks growing and developing in the region.

Drill holes made by predators in prey shells are widely considered to be the most unambiguous bodies of evidence of predator-prey interactions in the fossil record. However, recognition of traces of predatory origin from those formed by abiotic factors still waits for a rigorous evaluation as a prerequisite to ascertain predation intensity through geologic time and to test macroevolutionary patterns. New experimental data from tumbling various extant shells demonstrate that abrasion may leave holes strongly resembling the traces produced by drilling predators. They typically represent singular, circular to oval penetrations perpendicular to the shell surface. These data provide an alternative explanation to the drilling predation hypothesis for the origin of holes recorded in fossil shells. Although various non-morphological criteria (evaluation of holes for non-random distribution) and morphometric studies (quantification of the drill hole shape) have been employed to separate biological from abiotic traces, these are probably insufficient to exclude abrasion artifacts, consequently leading to overestimate predation intensity. As a result, from now on, we must adopt more rigorous criteria to appropriately distinguish abrasion artifacts from drill holes, such as microstructural identification of micro-rasping traces.

On the basis of data in the literature, the percentages of dextral versus sinistral species of snails have been calculated for western Europe, Turkey, North America (north of Mexico), and Japan. When the family of Clausiliidae is represented, about a quarter of all snail species may be sinistral, whereas less than one per cent of the species may be sinistral where that family does not occur. The number of single-gene speciation events on the basis of chirality, resulting in the origin of mirror image species, is not closely linked to the percentage of sinistral versus dextral species in a particular region. Turkey is nevertheless exceptional by both a high percentage of sinistral species and a high number of speciation events resulting in mirror image species. Shell morphology and genetic background may influence the ease of chirality-linked speciation, whereas sinistrality may additionally be selected against by internal selection. For the Clausiliidae, the fossil record and the recent fauna suggest that successful reversals in coiling direction occurred with a frequency of once every three to four million years.

We document differences in shell damage and shell thickness in a bivalve mollusc (Laternula elliptica) from seven sites around Antarctica with differing exposures to ice movement. These range from 60% of the sea bed impacted by ice per year (Hangar Cove, Antarctic Peninsula) to those protected by virtually permanent sea ice cover (McMurdo Sound). Patterns of shell damage consistent with blunt force trauma were observed in populations where ice scour frequently occurs; damage repair frequencies and the thickness of shells correlated positively with the frequency of iceberg scour at the different sites with the highest repair rates and thicker shells at Hangar Cove (74.2% of animals damaged) compared to the other less impacted sites (less than 10% at McMurdo Sound). Genetic analysis of population structure using Amplified Fragment Length Polymorphisms (AFLPs) revealed no genetic differences between the two sites showing the greatest difference in shell morphology and repair rates. Taken together, our results suggest that L. elliptica exhibits considerable phenotypic plasticity in response to geographic variation in physical disturbance.

Assessment of plastic and heritable components of phenotypic variation is crucial for understanding the evolution of adaptive character traits in heterogeneous environments. We assessed the above in relation to adaptive shell morphology of the rocky intertidal snail Nucella lapillus by reciprocal transplantation of snails between two shores differing in wave action and rearing snails of the same provenance in a common garden. Results were compared with those reported for similar experiments conducted elsewhere. Microsatellite variation indicated limited gene flow between the populations. Intrinsic growth rate was greater in exposed-site than sheltered-site snails, but the reverse was true of absolute growth rate, suggesting heritable compensation for reduced foraging opportunity at the exposed site. Shell morphology of reciprocal transplants partially converged through plasticity toward that of native snails. Shell morphology of F(2)s in the common garden partially retained characteristics of the P-generation, suggesting genetic control. A maternal effect was revealed by greater resemblance of F(1)s than F(2)s to the P-generation. The observed synergistic effects of plastic, maternal and genetic control of shell-shape may be expected to maximise fitness when environmental characteristics become unpredictable through dispersal.

Understanding the effects of environmental change on the distribution and abundance of strongly interacting organisms, such as intertidal macroalgae and their grazers, needs a thorough knowledge of their underpinning ecological relationships. Control of grazer-plant interactions is bi-directional on northwestern European coasts: grazing by limpets structures populations of macroalgae, while macroalgae provide habitat and food for limpets. Scottish shores dominated by the macroalga Fucus vesiculosus support lower densities and larger sizes of limpets Patella vulgata than shores with less Fucus. These patterns may be due to differences in inter-size-class competitive interactions of limpets among shores with different covers of Fucus. To examine this model, densities of small and large limpets were manipulated in plots with and without Fucus. Amounts of biofilm were measured in each plot. The presence of Fucus increased survival but hindered growth of small (15 mm TL) limpets, which were negatively affected by the presence of large limpets (31 mm TL). In contrast, large limpets were not affected by the presence of Fucus or of small limpets. This suggests the occurrence of asymmetric inter-size-class competition, which was influenced by the presence of macroalgae. Macroalgae and increased densities of limpets did not influence amounts of biofilm. Our findings highlight the role of interactions among organisms in generating ecological responses to environmental change.

Many plant seeds and invertebrates can survive passage through the digestive system of birds, which may lead to long distance dispersal (endozoochory) in case of prolonged retention by moving vectors. Endozoochorous dispersal by waterbirds has nowadays been documented for many aquatic plant seeds, algae and dormant life stages of aquatic invertebrates. Anecdotal information indicates that endozoochory is also possible for fully functional, active aquatic organisms, a phenomenon that we here address experimentally using aquatic snails. We fed four species of aquatic snails to mallards (Anas platyrhynchos), and monitored snail retrieval and survival over time. One of the snail species tested was found to survive passage through the digestive tract of mallards as fully functional adults. Hydrobia (Peringia) ulvae survived up to five hours in the digestive tract. This suggests a maximum potential transport distance of up to 300 km may be possible if these snails are taken by flying birds, although the actual dispersal distance greatly depends on additional factors such as the behavior of the vectors. We put forward that more organisms that acquired traits for survival in stochastic environments such as wetlands, but not specifically adapted for endozoochory, may be sufficiently equipped to successfully pass a bird's digestive system. This may be explained by a digestive trade-off in birds, which maximize their net energy intake rate rather than digestive efficiency, since higher efficiency comes with the cost of prolonged retention times and hence reduces food intake. The resulting lower digestive efficiency allows species like aquatic snails, and potentially other fully functional organisms without obvious dispersal adaptations, to be transported internally. Adopting this view, endozoochorous dispersal may be more common than up to now thought.