The temperate Australasian genus Papawera Oskars and Malaquias, 2019 (Gastropoda: Cephalaspidea: Haminoeidae), with a redescription of P. zelandiae and P. maugeansis

The genus Papawera includes two species of haminoeid snails found only in temperate waters of New Zealand and southeastern Australia. In this work, we redescribe the Papawera species based on characters of their external morphology, shells, and anatomical features such as radulae, jaws, gizzard plates, and male reproductive systems, using for the first time, scanning electron microscopy. A multi-locus phylogenetic hypothesis and the species delimitation method Automatic Barcode Gap Discovery based on DNA sequences of the cytochrome c oxidase subunit I gene were used to corroborate species status. The type species of the genus, P. zelandiae, is restricted to New Zealand and P. maugeansis is only known from South Australia, Victoria, and Tasmania. These species are easily distinguished externally by the shape of the shell, colouration of the living animal, and morphology of the cephalic shield. Anatomically they have differences in the number of marginal teeth, distribution of rods in the gizzard plates, and anatomy of the fundus in the male reproductive system.publishedVersio

The systematics of Cephalaspidea (Mollusca: gastropods) revisited, with a study on the diversity of deep sea Philinidae sensu lato from the West Pacific

The Cephalaspidea are the second most diverse marine clade of the Euthyneura gastropods, after the Nudipleura with many groups still known largely from shells or little anatomical data. These marine snails occur worldwide across all latitudes and depths. The definition of the group and the relationships between members has been hampered by the difficulty of establishing sound synapomorphies, but the advent of molecular phylogenetics in recent times has helped changed significantly this situation. Yet, because of reduced taxon sampling and few genetic markers employed in previous studies many questions about the sister relationships and monophyletic status of several families remained open. In this study over 100 taxa were included covering 100% of the traditionally recognized valid families and about 50% of the genera. The goals were to (1) produce a new hypothesis of relationships for the Cephalaspidea gastropods including for the first time representatives from all families by means of multilocus phylogenetic analyses (2) to infer relationships between families and test their monophyly, and (3) to study the diversity and systematics of the elusive deep sea West Pacific Philinidae sensu lato cephalaspids. Bayesian molecular phylogenetics based on two mitochondrial (COI, 16S rRNA) and two nuclear gene markers (28S rRNA and Histone-3) was used to infer the relationships of Cephalaspidea and to aid on species delimitation within Philinidae s. l.. Additionally, species of the Philinidae s. l. were studied by fine anatomical dissection and scanning electron microscopy. The monophyly of the Cephalaspidea without Runcinacea was confirmed. The families Cylichnidae, Diaphanidae, Haminoeidae, Philinidae, Retusidae, and Scaphandridae were found to be not monophyletic. This result suggests that the family level taxonomy of the Cephalaspidea warrants a profound revision and several new family and genera names are required to reflect the new phylogenetic hypothesis presented in this work. Philinidae s. l. was polyphyletic with four independent main lineages (family level) and seven genera. The generic names Hermania, Laona, Philinorbis, and Praephiline are reinstated as valid. Sixteen species of deep sea West Pacific Philinidae s. l were recognized of which 13 are putatively new to Science, emphasising how little is still known about the biodiversity of the deep sea in the tropical Indo-Pacific

The systematics and phylogeny of Haminoeidae (Mollusca, Cephalaspidea), with special focus on the taxonomy and phylogeny of the genus Haloa sensu lato

The family Haminoeidae consist of herbivorous snails found worldwide in tropical to temperate shallow waters on reefs, seagrass beds, rocky shores and mangroves. The family is the most diverse within the Order Cephalaspidea, but it has been plagued by systematic and taxonomic confusion due to many species and genera being vaguely defined only on shell or inconsistently on a few anatomical features. In this thesis I aim to resolve the taxonomy and systematics of the family and genera based on an integrative approach using a combination of molecular phylogenetic analyses using the mitochondrial genes cytochrome c oxidase subunit I, 12S rRNA, 16S rRNA, and the nuclear genes 18S rRNA, 28S rRNA, and Histone 3 coupled with morpho-anatomical data, the latter resulting from a revision of anatomical characters based in literature and dedicated anatomical dissections. Several of the traditional Haminoeidae genera were synonymized (Austrocylichna, Limulatys, Micraenigma, Nipponatys, Tepidatys) resurrected (Haloa, Haminella, Lamprohaminoea, Roxaniella, Weinkauffia) or showed to not belong in the family (Cylichnium, Hamineobulla, Micratys, Mimiatys, Mnestia, Osorattis, Roxania, Spissitydeus). The results showed that the family is composed of 17 genera including the new genus here described Vellicolla and the informal clade “Minihaminoeids”. Further, the type genus Haminoea was shown to be non-monophyletic, with three main radiations, namely Haminoea (Atlantic + eastern Pacific), Haloa sensu lato (Indo-West Pacific), and Smaragdinella. A comprehensive study of Haloa s. l. based on an expanded taxon sampling including representatives from putatively all species demonstrated that this clade consist of four clades warranting generic status with unique ecological, biogeographic, and morphological features, namely Haloa sensu stricto (13 species with dull colours distributed across the Indo-West Pacific), Lamprohaminoea (5 species with bright colours distributed across the Indo-West Pacific), and the two new genera here described Bakawan (4 species restricted to mangrove habitats) and Papawera (2 species restricted to temperate waters of Australasia). Systematic revisions were conducted for the four genera and 7 new species were found and described (Haloa [2], Lamprohaminoea [3], Bakawan [2])

A new phylogeny of the Cephalaspidea (Gastropoda: Heterobranchia) based on expanded taxon sampling and gene markers

The Cephalaspidea is a diverse marine clade of euthyneuran gastropods with many groups still known largely from shells or scant anatomical data. The definition of the group and the relationships between members has been hampered by the difficulty of establishing sound synapomorphies, but the advent of molecular phylogenetics is helping to change significantly this situation. Yet, because of limited taxon sampling and few genetic markers employed in previous studies, many questions about the sister relationships and monophyletic status of several families remained open. In this study 109 species of Cephalaspidea were included covering 100% of traditional family-level diversity (12 families) and 50% of all genera (33 genera). Bayesian and maximum likelihood phylogenetics analyses based on two mitochondrial (COI, 16S rRNA) and two nuclear gene markers (28S rRNA and Histone-3) were used to infer the relationships of Cephalaspidea. The monophyly of the Cephalaspidea was confirmed. The families Cylichnidae, Diaphanidae, Haminoeidae, Philinidae, and Retusidae were found non-monophyletic. This result suggests that the family level taxonomy of the Cephalaspidea warrants a profound revision and several new family and genus names are required to reflect the new phylogenetic hypothesis presented here. We propose a new classification of the Cephalaspidea including five new families (Alacuppidae, Colinatydidae, Colpodaspididae, Mnestiidae, Philinorbidae) and one new genus (Alacuppa). Two family names (Acteocinidae, Laonidae) and two genera (Laona, Philinorbis) are reinstated as valid. An additional lineage with family rank (Philinidae “Clade 4”) was unravelled, but no genus and species names are available to reflect the phylogeny and formal description will take place elsewhere

A molecular phylogeny of the Indo-West Pacific species of Haloa sensu lato gastropods (Cephalaspidea: Haminoeidae): Tethyan vicariance, generic diversity, and ecological specialization

The cephalaspidean gastropod genus Haminoea has been considered a worldwide radiation with species living in intertidal and shallow areas with algae, seagrass sandy-mud, mangroves, and coral reefs. Recently this monophyletic status was questioned and it was suggested that Haminoea proper only included Atlantic plus eastern Pacific species, whereas the Indo-West Pacific species were a separate radiation belonging to the genus Haloa. In this work we used an extended dataset of Haminoea sensu lato including to our best knowledge representatives of all Indo-West Pacific species, plus a large representation of Atlantic and eastern Pacific species. Bayesian and maximum likelihood molecular phylogenetic analyses using three mitochondrial (COI, 12S rRNA, 16S rRNA) and two nuclear genes (28S rRNA and Histone-3) were employed to study relationships and diversification including also the closely related genus Smaragdinella. Our results support a monophyletic Atlantic + eastern Pacific Haminoea clade and a monophyletic Indo-West Pacific radiation with five genera all defined by unique morphological characters and ecological requirements, namely Haloa proper with tropical and sub-tropical species spread across the Indo-West Pacific, Lamprohaminoea including only colourful species of tropical and sub-tropical affinity, Bakawan gen. nov. with species associated with mangrove habitats across the tropical eastern Indian Ocean and western Pacific, Papawera gen. nov. restricted to temperate waters of Australasia, and Smaragdinella the only cephalaspidean genus inhabiting intertidal hard substrates across sub-tropical and tropical regions of the Indo-West Pacific. This result suggests the role of the closure of the Tethys seaway structuring the phylogeny of worldwide “Haminoea” snails and of ecology driving the phylogenetic structure of the Indo-West Pacific radiation

A molecular phylogeny of the Indo-West Pacific species of Haloa sensu lato gastropods (Cephalaspidea: Haminoeidae): Tethyan vicariance, generic diversity, and ecological specialization

The cephalaspidean gastropod genus Haminoea has been considered a worldwide radiation with species living in intertidal and shallow areas with algae, seagrass sandy-mud, mangroves, and coral reefs. Recently this monophyletic status was questioned and it was suggested that Haminoea proper only included Atlantic plus eastern Pacific species, whereas the Indo-West Pacific species were a separate radiation belonging to the genus Haloa. In this work we used an extended dataset of Haminoea sensu lato including to our best knowledge representatives of all Indo-West Pacific species, plus a large representation of Atlantic and eastern Pacific species. Bayesian and maximum likelihood molecular phylogenetic analyses using three mitochondrial (COI, 12S rRNA, 16S rRNA) and two nuclear genes (28S rRNA and Histone-3) were employed to study relationships and diversification including also the closely related genus Smaragdinella. Our results support a monophyletic Atlantic + eastern Pacific Haminoea clade and a monophyletic Indo-West Pacific radiation with five genera all defined by unique morphological characters and ecological requirements, namely Haloa proper with tropical and sub-tropical species spread across the Indo-West Pacific, Lamprohaminoea including only colourful species of tropical and sub-tropical affinity, Bakawan gen. nov. with species associated with mangrove habitats across the tropical eastern Indian Ocean and western Pacific, Papawera gen. nov. restricted to temperate waters of Australasia, and Smaragdinella the only cephalaspidean genus inhabiting intertidal hard substrates across sub-tropical and tropical regions of the Indo-West Pacific. This result suggests the role of the closure of the Tethys seaway structuring the phylogeny of worldwide “Haminoea” snails and of ecology driving the phylogenetic structure of the Indo-West Pacific radiation

The temperate Australasian genus Papawera Oskars and Malaquias, 2019 (Gastropoda: Cephalaspidea: Haminoeidae), with a redescription of P. zelandiae and P. maugeansis

The genus Papawera includes two species of haminoeid snails found only in temperate waters of New Zealand and southeastern Australia. In this work, we redescribe the Papawera species based on characters of their external morphology, shells, and anatomical features such as radulae, jaws, gizzard plates, and male reproductive systems, using for the first time, scanning electron microscopy. A multi-locus phylogenetic hypothesis and the species delimitation method Automatic Barcode Gap Discovery based on DNA sequences of the cytochrome c oxidase subunit I gene were used to corroborate species status. The type species of the genus, P. zelandiae, is restricted to New Zealand and P. maugeansis is only known from South Australia, Victoria, and Tasmania. These species are easily distinguished externally by the shape of the shell, colouration of the living animal, and morphology of the cephalic shield. Anatomically they have differences in the number of marginal teeth, distribution of rods in the gizzard plates, and anatomy of the fundus in the male reproductive system

New observations of the enigmatic West African Cellana limpet (Mollusca: Gastropoda: Nacellidae)

Background: Identification of limpets is often hampered by highly variable within-species shell morphologies and colour patterns. Since pre-Linnean times this has produced complex taxonomies with confusing nomenclatorial histories and uncertain distribution patterns. This is the case for a complex of taxa associated with Cymbula safiana (Lamarck, 1819) and with the rejected name Patella nigra. We DNA sequenced limpets from Nigeria that were originally identified as C. safiana. Comparisons with available cox1 data of patellogastropods show that the specimens actually belong to the genus Cellana Adams, 1869 which has been recorded only once before in the Atlantic Ocean with the finding of specimens from Ghana. Results: We are reporting findings of Cellana sp. from the Gulf of Guinea for the second time. Specimens from Nigeria are 100 % similar to previously published cox1 sequences from Ghana. Due to variable shell characteristics we suspect that this species may have been confused with Cymbula safiana (Lamarck, 1819) in previous records. Inspection of the radula sack and radula demonstrates clear similarities with other Cellana species and contrasting differences in the organization of the teeth in Cymbula. Conclusions: Because Cellana is a possible candidate of invasive species in West Africa and Cymbula is considered as endangered, is seems particularly important to be able to distinguish between the two without being dependent on DNA analysis. When shell morphology seems to be of questionable diagnostic value, examination of the radula will help in future mapping and monitoring of these two species. A cox1 gene tree with Nigerian sequences included is in line with findings of previous authors and restates the need for taxonomic revision of the species clustering with Cellana toreuma (Reeve, 1854) and parts of a polyphyletic Cellana radiata von Born, 1778

Diversity and systematics of philinid snails (Gastropoda: Cephalaspidea) in West Africa with remarks on the biogeography of the region

West Africa is often considered one of the least studied regions of the world concerning marine biodiversity. Knowledge about the philinid snails of the region has largely been based on shells, but shells can be insufficient to discriminate between species. In this paper, we review the diversity and revise the systematics of the West African species of Philinidae sensu lato based on a comprehensive literature review and the study of shells, anatomy and DNA sequence data of novel specimens obtained by the R/V Dr Fridtjof Nansen (2005–2012) between Morocco and Angola. We use scanning electron microscopy together with a cytochrome c oxidase subunit I molecular phylogeny and molecular species delimitation methods (ABGD and SDP Geneious Plug In) to define species. Twenty species (19 formally named) were found to occur in the region, including four new species we describe here: Laona nanseni sp. nov., Philine cerebralis sp. nov., Philine schrammi sp. nov. and Spiniphiline caboverdensis sp. nov. We identified a complex of four cryptic species under the name P. quadripartita (P. guineensis, P. quadripartita, Philine schrammi sp. nov., Philine sp.). A biogeographical break is observed around Cape Verde and the Sahelian upwelling system coinciding with the West African Transition Province; eight species have their northern geographical limit or are restricted to this Province, while ten species have their southern limit here or just further north. Only one species seems to span across this biogeographical breaking point (P. scabra), but we speculate that this might result from taxonomic misidentifications