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The genus Haloa includes dull-coloured species of haminoeid snails inhabiting tidal and shallow waters of the tropical and sub-tropical Indo-West Pacific. This paper reports on the diversity and systematics of Haloa based on the phylogenetic hypothesis generated by Oskars and Malaquias (2019) and on the morphological study of specimens. Shells, external features and anatomical characters from the jaws, radula, gizzard plates and male reproductive system were studied by optical and scanning electron microscopy. The molecular species delimitation method Automatic Barcode Gap Discovery, using DNA sequences of the cytochrome c oxidase subunit I gene, was implemented to aid delimiting candidate species. Thirteen species were recognised as valid, including two new to science, namely Haloa cobbi, sp. nov. and Haloa eora, sp. nov., and one unnamed species herein referred to as Haloa sp. 1. Species depict intraspecific variability in their colouration and little character displacement in their anatomy. Shells are of limited taxonomic use; only details of the male reproductive system (e.g. fundus) and the DNA were truly diagnostic. Three species have an Indo-West Pacific distribution and three are restricted to the Indian Ocean. The remaining seven species occur in the Western and Central Pacific Ocean, three of them having geographical ranges restricted to offshore islands.

The genus Bakawan includes species of haminoeid snails associated with mangrove habitats and mud flats in the Indo-West Pacific. Here, we revise the diversity and systematics of the genus Bakawan based on our recent molecular phylogeny (Oskars & Malaquias, 2019) and detailed analysis of morphological characters. We examined a range of morphological characters (the shells, external morphology of the animal, jaw, radula, gizzard plate and male reproductive system) using light and scanning electron microscopy. We also carried out a species delimitation analysis (we used the automatic barcode gap discovery method) based on cytochrome c oxidase subunit I DNA sequence data. Four distinct species were recognized: Bakawan rotundata (A. Adams, 1850), which is the type species of the genus and ranges from the eastern Indian Ocean to the western Pacific; B. fusca (A. Adams, 1850), currently known only from the Philippines; and two species new to science, B. puti n. sp., known only from the Philippines, and B. hedleyi n. sp., restricted to tropical eastern Australia.

Scaphander Montfort, 1810, is a genus of deep-sea, soft-bottom gastropods composed of approximately 23 species distributed worldwide. The systematics of the genus in the Atlantic was recently revised and eight species were recognized. The present study describes a new species (Scaphander meridionalis sp. n.) from the Argentine continental slope using morphological and molecular data. Shells, gizzard plates, radulae, and male reproductive systems were studied by optical and scanning electron microscopy and compared with data from all other known Scaphander species. Bayesian molecular phylogenetics based on two mitochondrial (cytochrome c oxidase subunit I and 16S rRNA) and one nuclear (28S rRNA) genes together with the molecular species delimitation method Automatic Barcode Gap Discovery were used to compare the novel samples with all known Atlantic species. Our results revealed that the Argentinian specimens have a distinct shell and penial papilla and were molecularly 6.35–6.53%, 6.9–7.44%, and 7.99–8.89% distinct (COI uncorrected p-distance) from their closest relatives, the Pacific species Scaphander grandis and Scaphander mundus, and the northern Atlantic species Scaphander nobilis, respectively. Scaphander meridionalis sp. n. is the first species only known from the South Atlantic and its occurrence off Argentina represents a new southern geographical limit of the distribution of the genus in the Atlantic Ocean.

In this paper current knowledge on the diversity of the gastropod cephalaspidean genus Haminoea from India is revised. A comprehensive literature review was undertaken and novel specimens were coll...

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. publishedVersion

The genus Lamprohaminoea includes species of colourful haminoeid snails associated with coral reefs and rocky shores in the tropical Indo-West Pacific. In this work, we revise the diversity and systematics of Lamprohaminoea species based on a phylogenetic hypothesis and on a detailed morphological analysis of specimens. Shells, external features of the animals and anatomical characters from the jaws, radula, gizzard plates, and male reproductive system were studied by optical and scanning electron microscopy. Additionally, a molecular species delimitation analysis based on the Automatic Barcode Gap Discovery method using DNA sequences of the cytochrome c oxidase subunit I was implemented. Five species were recognised, three of them new to science, namely L. cymbalum, L. ovalis, L. vamiziensis sp. nov., L. evelinae sp. nov., and L. mikkelsenae sp. nov. Morphologically, these species can be separated by subtle differences of their external colouration and by features of the male reproductive system. Lamprohaminoea vamiziensis sp. nov. is known only from the western Indian Ocean and L. evelinae sp. nov., from the west Pacific, whereas the other three occur across the Indo-West Pacific realm.

Haminoeidae is the most diverse family of Cephalaspidea with 13 to 17 genera commonly recognised as valid and with 46 genera that historically have been moved back and forth between Haminoeidae and other families. Due to poor definition of most genera the family is plagued by extensive taxonomic confusion and its generic composition and internal relationships remain uncertain. In this work we have integrated the study of type material, original descriptions, shells, morpho-anatomical data, and molecular phylogenetics (Bayesian, maximum likelihood, and maximum parsimony) based on five genetic markers (the mitochondrial genes cytochrome c oxidase subunit I and 16SrRNA and the nuclear genes 18SrRNA, 28SrRNA, and histoneH3) to delimit the valid genera, define synapomorphic traits, and establish synonym lists. Three hundred and ninety novel sequences were generated. In total 14 genera were recognised; one genus (Vellicolla gen. nov.) is here described as new and an additional fifteenth group was identified, but no species could be formally ascribed to it and therefore remains unnamed (here designated informally as mini-haminoeids). The relationships of genera are discussed and seven deep clades have been identified but are not formally named because of lack of recognisable synapomorphies for several of them. A new classification for Haminoeidae is proposed including 14 valid genera and one informal group.

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. publishedVersion

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. publishedVersion

Oskars, Trond Roger, Mifsud, Constantine, Malaquias, Manuel António E. (2017): Redescription of the Cephalaspidea gastropod Atys jeffreysi (Weinkauff, 1866) (Haminoeidae), with a discussion on the phylogenetic affinities of the Mediterranean species of the genus. Journal of Natural History 51 (27-28): 1593-1608, DOI: 10.1080/00222933.2017.1340526, URL: http://dx.doi.org/10.1080/00222933.2017.1340526

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. publishedVersion

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. publishedVersion