Spawning behavior and larval development in Mopalia lignosa and Mopalia muscosa (Mollusca: Polyplacophora) in central California

Spawning behavior and external features of the larval development were studied in the chitons Mopalia muscosa and M. lignosa during the months of April-June, 1974, at Pacific Grove, California. ... The sequence of events in the development of the two species in the same, though some differences in timing exist.

First record of a chiton from the Palaeocene of Denmark (Polyplacophora : Leptochitonidae) and its phylogenetic affinities

Anew species of fossil polyplacophoran from the Danian (Lower Palaeocene) of Denmark is described from over 450 individual disarticulated plates. The polyplacophorans originate from the 'nose-chalk' in the classical Danish locality of Fakse Quarry, an unconsolidated coral limestone in which aragonitic mollusc shells are preserved through transformation into calcite. In plate architecture and sculpture, the new Danish material is similar to Recent Leptochiton spp., but differs in its underdeveloped apophyses and high dorsal elevation (height/width ca. 0.54). Cladistic analysis of 55 original shell characters coded for more than loo Recent and fossil species in the order Lepiclopleurida shows very high resolution of interspecific relationships, but does not consistently recover traditional genera or subgenera. Inter-relationships within the suborder Lepidopleurina are of particular interest as it is often considered the most 'basal' neoloricate lineage. In a local context, the presence of chitons in the faunal assemblage of Fakse contributes evidence of shallow depositional depth for at least some elements of this Palaeocene seabed, a well-studied formation of azooxanthellic coral limestones. This new record for Denmark represents a well-dated and ecologically well-understood fossil chiton with potential value for understanding the radiation of the Neotoricata.

Parasitic foraminifers on a deep-sea chiton (Mollusca, Polyplacophora, Leptochitonidae) from Iceland

Epibiotic foraminifers selectively settle on the most food-rich area of the host substrate, even when the species acts as a facultative ectoparasite in later life stages. In 398 specimens examined of the deep-sea chiton Leptochiton arcticus from Iceland, 46% show evidence of infestation by foraminifers, with many showing extensive shell damage from present and past bioeroding epibionts. Disturbances to the inner layer of the host shell are indicative of parasitism, as evidenced both by wound healing calcification and protrusions of the foraminiferan tubules. The epibionts employ different feeding strategies at different stages of their life cycle, taking advantage of nutrient availability from the posterior respiration currents and excrement of the chitons as juveniles, and feeding parasitically as adults. Epibiont persistence on individual hosts-through successive generations, or long-term continuous bioerosion by epibionts-allow larger adult parasitic foraminifers of Hyrrokkin sarcophaga to penetrate the thick tail valve of a chiton and feed parasitically on the host tissue. The proportion of chitons infested increases with host size, indicating that epibionts are accumulated through a chiton's life, seemingly without major detriment to host survivorship.

The deep-sea chiton Nierstraszella (Mollusca: Polyplacophora: Lepidopleurida) in the Indo-West Pacific: taxonomy, morphology and a bizarre ectosymbiont

This study investigated the taxonomy and distribution of the deep-sea polyplacophoran mollusc Nierstraszella Sirenko, 1992 in the Indo-West Pacific, based on a collection of 516 specimens collected in the Philippines and Solomon Islands. Although seven species names have historically been proposed in this group of chitons, all have been considered as synonyms of the monotypic N. lineata (Nierstrasz, 1905). Morphological examination of this new material reveals the presence of two species. N. lineata is distinct from N. andamanica (Smith, 1906), based on morphological characters given in the original species description and very distinctly different morphology of aesthete pores in the shell surface. Furthermore, populations of N. andamanica in the Philippines and Solomon Islands are locally colonized with the epibiotic (ectoparasitic) bryozoan Pseudobathyalozoon profundum d'Hondt, 2006. These bryozoans attach ventrally to the girdle of the host chiton and the erect zooids feed within the pallial cavity, among the chiton's gills.

Gross anatomy and positional homology of gills, gonopores, and nephridiopores in “basal” living chitons (Polyplacophora: Lepidopleurina)

Traditional shell characters are insufficient to differentiate taxa within the polyplacophoran order Lepidopleurida. Additional morphological character sets from soft anatomy (e.g., gamete morphology, gill arrangement, and locations of gonopores and nephidiopores) have previously been described from only a small number of taxa. This study reports for the first time, positions of the gonopores and nephridiopores for 17 species in the Lepidopleurina. The position of both types of pores on the longitudinal body axis varies within a generalized range of the posterior third of the body; however, the separation between the pores as a proportion of the specimen’s foot length varies from 3.7% to 17% in different species. Positions of pores relative to the serial gills are also variable within species, and future studies may require a new descriptive basis in order to resolve positional homology. The order Lepidopleurida occupies a critical position with respect to understanding larger-scale patterns in polyplacophoran (and molluscan) evolution.

The Irish fossil Polyplacophora

Three species of fossil polyplacophoran molluscs are known from Ireland. Two species were originally described in the nineteenth century: Helminthochiton griffithi Salter in M‘Coy, 1846 and Pterochiton thomondiensis (Baily, 1859), and an articulated specimen representing a third indeterminate species, described here for the first time. Previous work on the evolutionary context of these species has relied on published illustrations and descriptions without examination of the type material. as chitons are considered rare in the fossil record, these specimens represent an interesting and important aspect of Irish palaeobiology.

Evolution in the deep sea: a combined analysis of the earliest diverging living chitons (Mollusca : Polyplacophora : Lepidopleurida)

Lepidopleurida is the earliest diverged group of living polyplacophoran molluscs. They are found predominantly in the deep sea, including sunken wood, cold seeps, other abyssal habitats, and a few species are found in shallow water. The group is morphologically identified by anatomical features of their gills, sensory aesthetes, and gametes. Their shell features closely resemble the oldest fossils that can be identified as modern polyplacophorans. We present the first molecular phylogenetic study of this group, and also the first combined phylogenetic analysis for any chiton, including three gene regions and 69 morphological characters. The results show that Lepidopleurida is unambiguously monophyletic, and the nine genera fall into five distinct clades, which partly support the current view of polyplacophoran taxonomy. The genus Hanleyella Sirenko, 1973 is included in the family Protochitonidae, and Ferreiraellidae constitutes another distinct clade. The large cosmopolitan genus Leptochiton Gray, 1847 is not monophyletic; Leptochiton and Leptochitonidae sensu stricto are restricted to North Atlantic and Mediterranean taxa. Leptochitonidae s. str. is sister to Protochitonidae. The results also suggest two separate clades independently inhabiting sunken wood substrates in the south-west Pacific. Antarctic and other chemosynthetic-dwelling species may be derived from wood-living species. Substantial taxonomic revision remains to be done to resolve lepidopleuran classification, but the phylogeny presented here is a dramatic step forward in clarifying the relationships within this interesting group.

Deep-sea chitons from sunken wood in the West Pacific (Mollusca: Polyplacophora: Lepidopleurida): taxonomy, distribution, and seven new species

Natural deposits of sunken wood provide an important habitat for deep-sea invertebrates. Deep-sea chitons in the primitive order Lepidopleurida are typically collected rarely and as single specimens. However, these animals have been recovered in large densities associated with sunken wood in the tropical West Pacific, in groups of up to 50 individuals. Four deep- sea expeditions in the West Pacific, to the Philippines, Solomon Islands, and Vanuatu, recovered a large number of poly- placophorans. We have examined the morphology as well as the range and distribution of these species, based on the larg- est collection ever examined (more than 1300 individuals). These species show potentially adapted characters associated with exploitation of sunken wood as habitat, such as protruding caps on sensory shell pores (aesthetes) and large interseg- mental bristles with potential sensory function. In this study we investigated the twenty-two species recovered, including seven newly described here (Leptochiton consimilis n. sp., L. angustidens n. sp., L. dykei n. sp., L. samadiae n. sp., L. longisetosus n. sp., L. clarki n. sp., L. schwabei n. sp.), and provide the first identification key to the 34 lepidopleuran chitons known from sunken wood worldwide.

Variation in oxygen consumption among 'living fossils' (Mollusca: Polyplacophora)

Polyplacophoran molluscs (chitons) are phylogenetically ancient and morphologically constrained, yet multiple living species are often found co-occurring within widely overlapping ecological niches. This study used two sets of experiments to compare interspecific variation among co-occurring species in the North Atlantic (Ireland) and separately in the North Pacific (British Columbia, Canada) chiton faunas. A complementary review of historical literature on polyplacophoran physiology provides an overview of the high level of metabolic variability in this group of 'living fossils'. Species examined in de novo experiments showed significant variation in oxygen consumption both under air-saturated water conditions (normoxia), and in response to decreasing oxygen availability (hypoxia). Some species demonstrate an ability to maintain constant oxygen uptake rates despite hypoxia (oxyregulators), while others oxyconform, with uptake rate dependent on ambient oxygen tension. These organisms are often amalgamated in studies of benthic communities, yet show obvious physiological difference that may impact their response or tolerance to environmental change.

Extensive local-scale gene flow and long-term population stability in the intertidal mollusc Katharina tunicata (Mollusca: Polyplacophora)

The dispersal capabilities of intertidal organisms may represent a key factor to their survival in the face of global warming, as species that cannot adapt to the various effects of climate change will have to migrate to track suitable habitat. Although species with pelagic larval phases might be expected to have a greater capacity for dispersal than those with benthic larvae, interspecies comparisons have shown that this is not always the case. Consequently, population genetic approaches are being increasingly used to gain insights into dispersal through studying patterns of gene flow. In the present study, we used nuclear single-nucleotide polymorphisms (SNPs) and mitochondrial DNA (mtDNA) sequencing to elucidate fine-scale patterns of genetic variation between populations of the Black Katy Chiton, Katharina tunicata, separated by 15-150 km in south-west Vancouver Island. Both the nuclear and mitochondrial data sets revealed no genetic differentiation between the populations studied, and an isolation-with-migration analysis indicated extensive local-scale gene flow, suggesting an absence of barriers to dispersal. Population demographic analysis also revealed long-term population stability through previous periods of climate change associated with the Pleistocene glaciations. Together, the findings of the present study suggest that this high potential for dispersal may allow K. tunicata to respond to current global warming by tracking suitable habitat, consistent with its long-term demographic stability through previous changes in the Earth's climate. (C) 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106, 589597.

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.

Chiton phylogeny (Mollusca Polyplacophora) and the placement of the enigmatic species Choriplax grayi (H. Adams & Angas)

Shallow marine chitons (Mollusca:Polyplacophora:Chitonida) are widespread and well described from established morphoanatomical characters, yet key aspects of polyplacophoran phylogeny have remained unresolved. Several species, including Hemiarthrum setulosum Carpenter in Dall, 1876, and especially the rare and enigmatic Choriplax grayi (Adams & Angas, 1864), defy systematic placement. Choriplax is known from only a handful of specimens and its morphology is a mosaic of key taxonomic features from two different clades. Here, new molecular evidence provides robust support for its correct association with a third different clade: Choriplax is placed in the superfamily Mopalioidea. Hemiarthrum is included in Cryptoplacoidea, as predicted from morphological evidence. Our multigene analysis of standard nuclear and mitochondrial markers demonstrates that the topology of the order Chitonida is divided into four clades, which have also been recovered in previous studies: Mopalioidea is sister to Cryptoplacoidea, forming a clade Acanthochitonina. The family Callochitonidae is sister to Acanthochitonina. Chitonoidea is resolved as the earliest diverging group within Chitonida. Consideration of this unexpected result for Choriplax and our well-supported phylogeny has revealed differing patterns of shell reduction separating the two superfamilies within Acanthochitonina. As in many molluscs, shell reduction as well as the de novo development of key shell features has occurred using different mechanisms, in multiple lineages of chitons.

The Continuing Debate on Deep Molluscan Phylogeny: Evidence for Serialia (Mollusca, Monoplacophora + Polyplacophora)

Molluscs are a diverse animal phylum with a formidable fossil record. Although there is little doubt about the monophyly of the eight extant classes, relationships between these groups are controversial.We analysed a comprehensive multilocus molecular data set for molluscs, the first to include multiple species from all classes, including five monoplacophorans in both extant families. Our analyses of fivemarkers resolve two major clades: the first includes gastropods and bivalves sister to Serialia (monoplacophorans and chitons), and the second comprises scaphopods sister to aplacophorans and cephalopods. Traditional groupings such as Testaria, Aculifera, and Conchifera are rejected by our data with significant Approximately Unbiased (AU) test values. A new molecular clock indicates that molluscs had a terminal Precambrian origin with rapid divergence of all eight extant classes in the Cambrian. Therecovery of Serialia as a derived, Late Cambrian clade is potentially in line with the stratigraphic chronology of morphologically heterogeneous early mollusc fossils. Serialia is in conflict with traditional molluscan classifications and recent phylogenomic data. Yet our hypothesis, as others from molecular data, implies frequent molluscan shell and body transformations by heterochronic shifts in development and multiple convergent adaptations, leading to the variable shells and body plans in extant lineages.