Morphological cladistic analysis as a model for character evaluation in primitive living chitons (Polyplacophora, Lepidopleurina)

Chitons are often referred to as "living fossils" in part because they are proposed as one of the earliest-diverging groups of living molluscs, but also because the gross morphology of the polyplacophoran shell has been conserved for hundreds of millions of years. As such, the analysis of evolution and radiation within polyplacophorans is of considerable interest not only for resolving the shape of pan-molluscan phylogeny but also as model organisms for the study of character evolution. This study presents a new, rigorous cladistic analysis of the morphological characters used in taxonomic descriptions for chitons in the living suborder Lepidopleurina Thiele, 1910 (the earliest-derived living group of chitons). Shell-based characters alone entirely fail to recover any recognized subdivisions within the group, which may raise serious questions about the application of fossil data (from isolated shell valves). New analysis including characters from girdle armature and gill arrangements recovers some genera within the group but also points to the lack of monophyly within the main genus Leptochiton Gray, 1847. Additional characters from molecular data and soft anatomy, used in combination, are clearly needed to resolve questions of chiton relationships. However, the data sets currently available already provide interesting insights into the analytical power of traditional morphology as well as some knowledge about the early evolution and radiation of this group.

New records for the shallow-water chiton fauna (Mollusca, Polyplacophora) of the Azores (NE Atlantic)

Published records, original data from recent field work on all of the islands of the Azores (NE Atlantic), and a revision of the entire mollusc collection deposited in the Department of Biology of the University of the Azores (DBUA) were used to compile a checklist of the shallow-water Polyplacophora of the Azores. Lepidochitona cf. canariensis and Tonicella rubra are reported for the first time for this archipelago, increasing the recorded Azorean fauna to seven species.

Do chitons have a compass? Evidence for magnetic sensitivity in Polyplacophora

Several animals and microbes have been shown to be sensitive to magnetic fields, though the exact mechanisms of this ability remain unclear in many animals. Chitons are marine molluscs which have high levels of biomineralised magnetite coating their radulae. This discovery led to persistent anecdotal suggestions that they too may be able to navigationally respond to magnetic fields. Several researchers have attempted to test this, but to date there have been no large-scale controlled empirical trials. In the current study, four chiton species (Katharina tunicata, Mopalia kennerleyi, Mopalia muscosa and Leptochiton rugatus, n=24 in each) were subjected to natural and artificially rotated magnetic fields while their movement through an arena was recorded over four hours. Field orientation did not influence the position of the chitons at the end of trials, possibly as a result of the primacy of other sensory cues (i.e. thigmotaxis). Under non-rotated magnetic field conditions, the orientation of subjects when they first reached the edge of an arena was clustered around 309-345 degrees (north-north-west) in all four species. However, orientations were random under the rotated magnetic field, which may indicate a disruptive effect of field rotation. This pattern suggests that chitons can detect and respond to magnetism.

Consensus and Confusion in Molluscan Trees: Evaluating Morphological and Molecular Phylogenies

Mollusks are the most morphologically disparate living animal phylum, they have diversified into all habitats, and have a deep fossil record. Monophyly and identity of their eight living classes is undisputed, but relationships between these groups and patterns of their early radiation have remained elusive. Arguments about traditional morphological phylogeny focus on a small number of topological concepts but often without regard to proximity of the individual classes. In contrast, molecular studies have proposed a number of radically different, inherently contradictory, and controversial sister relationships. Here, we assembled a dataset of 42 unique published trees describing molluscan interrelationships. We used these data to ask several questions about the state of resolution of molluscan phylogeny compared to a null model of the variation possible in random trees constructed from a monophyletic assemblage of eight terminals. Although 27 different unique trees have been proposed from morphological inference, the majority of these are not statistically different from each other. Within the available molecular topologies, only four studies to date have included the deep-sea class Monoplacophora; but 36.4% of all trees are not significantly different. We also present supertrees derived from 2 data partitions and 3 methods, including all available molecular molluscan phylogenies, which will form the basis for future hypothesis testing. The supertrees presented here were not constructed to provide yet another hypothesis of molluscan relationships, but rather to algorithmically evaluate the relationships present in the disparate published topologies. Based on the totality of available evidence, certain patterns of relatedness among constituent taxa become clear. The internodal distance is consistently short between a few taxon pairs, particularly supporting the relatedness of Monoplacophora and the chitons, Polyplacophora. Other taxon pairs are rarely or never found in close proximity, such as the vermiform Caudofoveata and Bivalvia. Our results have specific utility for guiding constructive research planning in order to better test relationships in Mollusca as well as other problematic groups. Taxa with consistently proximate relationships should be the focus of a combined approach in a concerted assessment of potential genetic and anatomical homology, while unequivocally distant taxa will make the most constructive choices for exemplar selection in higher-level phylogenomic analyses.

How the mollusc got its scales: Convergent evolution of the molluscan scleritome

Radiation of dramatically disparate forms among the phylum Mollusca remains a key question in metazoan evolution, and requires careful evaluation of homology of hard parts throughout the deep fossil record. Enigmatic early Cambrian taxa such as Halkieria and Wiwaxia (in the clade Halwaxiida) have been proposed to represent stem-group aculiferan molluscs (Caudofoveata+Solenogastres+Polyplacophora), as complex scleritomes were considered to be unique to aculiferans among extant molluscs. The 'scaly-foot gastropod' (Neomphalina: Peltospiridae) from hydrothermal vents of the Indian Ocean, however, also carries dermal sclerites and thus challenges this inferred homology. Despite superficial similarities to various mollusc sclerites, the scaly-foot gastropod sclerites are secreted in layers covering outpockets of epithelium and are largely proteinaceous, while chiton (Polyplacophora: Chitonida) sclerites are secreted to fill an invaginated cuticular chamber and are largely calcareous. Marked differences in the underlying epithelium of the scaly-foot gastropod sclerites and operculum suggest that the sclerites do not originate from multiplication of the operculum. This convergence in different classes highlights the ability of molluscs to adapt mineralized dermal structures, as supported by the extensive early fossil record of molluscs with scleritomes. Sclerites of halwaxiids are morphologically variable, undermining the assumed affinity of specific taxa with chitons, or the larger putative clade Aculifera. Comparisons with independently derived similar structures in living molluscs are essential for determining homology among fossils and their position with respect to the enigmatic evolution of molluscan shell forms in deep time.

The heart of a dragon: 3D anatomical reconstruction of the 'scaly-foot gastropod' (Mollusca: Gastropoda: Neomphalina) reveals its extraordinary circulatory system

Introduction: The 'scaly-foot gastropod' (Chrysomallon squamiferum Chen et al., 2015) from deep-sea hydrothermal vent ecosystems of the Indian Ocean is an active mobile gastropod occurring in locally high densities, and it is distinctive for the dermal scales covering the exterior surface of its foot. These iron-sulfide coated sclerites, and its nutritional dependence on endosymbiotic bacteria, are both noted as adaptations to the extreme environment in the flow of hydrogen sulfide. We present evidence for other adaptations of the 'scaly-foot gastropod' to life in an extreme environment, investigated through dissection and 3D tomographic reconstruction of the internal anatomy.

Results: Our anatomical investigations of juvenile and adult specimens reveal a large unganglionated nervous system, a simple and reduced digestive system, and that the animal is a simultaneous hermaphrodite. We show that Chrysomallon squamiferum relies on endosymbiotic bacteria throughout post-larval life. Of particular interest is the circulatory system: Chrysomallon has a very large ctenidium supported by extensive blood sinuses filled with haemocoel. The ctenidium provides oxygen for the host but the circulatory system is enlarged beyond the scope of other similar vent gastropods. At the posterior of the ctenidium is a remarkably large and well-developed heart. Based on the volume of the auricle and ventricle, the heart complex represents approximately 4 % of the body volume. This proportionally giant heart primarily sucks blood through the ctenidium and supplies the highly vascularised oesophageal gland. Thus we infer the elaborate cardiovascular system most likely evolved to oxygenate the endosymbionts in an oxygen poor environment and/or to supply hydrogen sulfide to the endosymbionts.

Conclusions: This study exemplifies how understanding the autecology of an organism can be enhanced by detailed investigation of internal anatomy. This gastropod is a large and active species that is abundant in its hydrothermal vent field ecosystem. Yet all of its remarkable features-protective dermal sclerites, circulatory system, high fecundity-can be viewed as adaptations beneficial to its endosymbiont microbes. We interpret these results to show that, as a result of specialisation to resolve energetic needs in an extreme chemosynthetic environment, this dramatic dragon-like species has become a carrying vessel for its bacteria.

A juvenile recruitment prediction model for Ruditapes decussatus (L.) (Bivalvia : Mollusca)

Despite the importance of larval abundance in determining the recruitment of benthic marine invertebrates and as a major factor in marine benthic community structure, relating planktonic larval abundance with post-settlement post-larvae and juveniles in the benthos is difficult. It is hampered by several methodological difficulties, including sampling frequency, ability to follow larval and post-larval or juvenile cohorts, and ability to calculate growth and mortality rates. In our work, an intensive sampling strategy was used. Larvae in the plankton were collected at weekly intervals, while post-larvae that settled into collectors were analysed fortnightly. Planktonic larval and benthic post-larval/juvenile cohorts were determined, and growth and mortality rates calculated. Integration of all equations allowed the development of a theoretical formulation that, based on the abundance and planktonic larval duration, permits an estimation of the future abundance of post-larvae/juveniles during the first year of benthic life. The model can be applied to a sample in which it was necessary only to measure larval length.

Lolliguncula tydeus BRAKONIECKI, 1980 (Mollusca:Cephalopoda) registrado en Perú

Se da a conocer el primer registro del calmar Lollinguncula tydeus Brakoniecki, 1980, para el Perú. Su presencia esta asociada con Fenómenos El Niño. Se proporciona una breve descripción, datos morfométricos y merísticos e índices de esta especie, que se compara con Lolliguncula panamensis.

INTRACELLULAR OSMOREGULATION IN THE ESTUARINE MOLLUSC VILLORITA CYPRINOIDES VAR. COCHINENSIS (Mollusca: Bivalvia) Hanley

The present investigation is dedicated to understanding various mechanisms of salinity tolerance in the estuarine clam V. cyprinoides var. cochinensis. Even though V. cyprinoids var. cochinensis and V. cyprinoides are found to coexist in the same area, V. cyprinoids is reported to tolerate higher salinities than variety cochinenesis. Variations in the salinity of sea water may affect the aquatic organisms through specific gravity control and variations in osmotic pressure. The specific gravity of most soft tissues is close to that of normal seawater. Many bottom living forms, both attached and motile, have very high specific gravities eg.villorita cyprinoids. Villorita spp. Occurs abundantly in the reaches of the estuary and backwaters of Kerala. In both marine and estuarine forms, it is observed that mantle employs a lesser quantity of amino acids compared to adductor and foot. The regulation of cell volume is not carried out equally in all types of tissues. The capability of salinity tolerance is an aggregate of both the capabilities of extra cellular anisosmotic and intracellular isosmotic regulations in osmoconforming animals. The ultimate aim of water regulation is to regulate the cell volume.T here are slight changes occur in cell volume even in osmoregulators. These studies can also help in revealing the changes brought about in the cellular organelles like lysosomes, which were found to have a role in the osmoregulatory process. The osmoregulatory machinery of estuarine animals is more streamlined for a successful life in the estuarine regime.

The Epipalaeolithic (Iberomaurusian) at Grotte des Pigeons (Taforalt), Morocco: a preliminary study of the land Mollusca

Land snail middens occur widely throughout the circum-Mediterranean, particularly in the Maghreb and Libya, but are relatively understudied when compared with the better known coastal marine middens. The site of Taforalt (Grotte des Pigeons) in north east Morocco contains thick ashy deposits with considerable numbers of land snails, dating between 13,000 and 11,000 BP. The site has been excavated periodically since its discovery in 1908, but little attention has been paid to the land snail component. Recent excavations at the site as part of the Cemeteries and Sedentism Project has facilitated reinvestigation of the Iberomaurusian layers, including the upper ashy midden unit. This investigation addresses questions surrounding chronological variation in the lithic assemblage, subsistence strategies, population health and mortuary practices. Preliminary work on the land snails has identified a low species diversity in the grey series midden layers, characterised by the large edible species Alabastrina alabastrites, Helix cf aspersa, Dupotetia dupotetiana, Otala punctata and Cernuella sp, which appear to represent an abrupt intensification in the use of land snails as a food source. Significant quantities of associated plant material and animal bone suggest that land snails are part of a broader based diet which is adopted from around 13,000 BP.

Lysossomal membrane stability of the brown mussel Perna perna (Linnaeus) (Mollusca, Bivalvia) exposed to the anionic surfactant Linear Alkylbenzene Sulphonate (LAS).

The effects of the Linear Alkylbenzene Sulphonate (LAS) were evaluated on the mussel Perna perna (Linnaeus, 1758), using a cellular level biomarker. The Neutral Red Retention Time (NRRT) assay was used to estimate effects at cellular levels. Significant effects were observed for the NRRT assay, even in low concentrations. The effects at cellular level were progressive, suggesting that the organisms are not capable to recover of such increasing effects. Additionally, the results show that the levels of LAS observed for Brazilian coastal waters may chronically affect the biota.

Faecal bacteria in Perna perna (Linnaeus, 1758) (Mollusca: Bivalvia) for biomonitoring coastal waters and seafood quality

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)