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Numerous faunistic and ecological studies have been conducted throughout the Indo-Pacific Ocean to assess its biodiversity. Despite the abundance of research, studies on the species that inhabit the Indo-Pacific are still necessary due to its extent and high species richness. The major species richness of the genus Okenia Menke, 1830 (Nudibranchia, Goniodorididae) is found in the Indo-Pacific Ocean, including 38 of 60 valid species. Nevertheless, this number does not represent the real biodiversity, since at least 20 more species are already reported in field-guides as undescribed species belonging to this genus. The systematics of the genus Okenia are still unclear since it has been the subject of only a few and incomplete studies. In the present paper, we describe five new Okenia species from the coastlines of Japan, Mozambique and Australia: Okenia aurorapapillata sp. nov., Okenia elisae sp. nov., Okenia nakanoae sp. nov., Okenia siderata sp. nov. and Okenia tenuifibrata sp. nov. Moreover, anatomical details not previously described of Okenia atkinsonorum, Okenia barnardi, Okenia cf. echinata, Okenia hallucigenia, Okenia hiroi, Okenia japonica, Okenia pellucida, Okenia pilosa and Okenia rhinorma are provided. New partial sequences of standard markers (COI, 16S rRNA and H3) were obtained and a phylogenetic analysis that included all species with available data was performed. ZOOBANK urn:lsid:zoobank.org:pub:28AE2536-A264–4194–8AE3-C430620572E7

Sponges in the subfamily Phyllospongiinae are important components of coral reefs. However, significant taxonomic inconsistencies exist in this group due to the lack of useful morphological characters for species delineation. This study assesses the systematics of some common phyllospongiinids in the genera Carteriospongia, Phyllospongia and Strepsichordaia from tropical Australia and the Red Sea, by using a multigene approach that utilizes the Internal Transcribed Spacer 2, the complete ribosomal 18S rRNA and three 28s rRNA gene regions (D1‒D2, D3‒D5 and D6‒D8), which produced a phylogenetic framework in which complementary morphological taxonomic assessments were performed. Type specimens were included, where available, and six species clades were recovered, including the well-established Phyllospongia papyracea and Strepsichordaia lendenfeldi. Carteriospongia foliascens, the type species for the genus Carteriospongia, is transferred to the genus Phyllospongia, resulting in Carteriospongia becoming a synonym of Phyllospongia. Consequently, Carteriospongia flabellifera is removed from Carteriospongia and is reinstated to its original designation of Polyfibrospongia flabellifera. Two new species, Phyllospongia bergquistae sp. nov. and Polyfibrospongia kulit sp. nov., are described. With phyllospongiinid sponges increasingly used as models for assessing the effects of climate change and anthropogenic stressors, this study provides a reliable systematics framework for the accurate identification of common phyllospongiinids across the Indo-Pacific.

Background Marine invertebrates are abundant and diverse on the continental shelf in Antarctica, but little is known about their parasitic counterparts. Endoparasites are especially understudied because they often possess highly modified body plans that pose problems for their identification. Asterophila, a genus of endoparasitic gastropod in the family Eulimidae, forms cysts in the arms and central discs of asteroid sea stars. There are currently four known species in this genus, one of which has been described from the Antarctic Peninsula (A. perknasteri). This study employs molecular and morphological data to investigate the diversity of Asterophila in Antarctica and explore cophylogenetic patterns between host and parasite. Results A maximum-likelihood phylogeny of Asterophila and subsequent species-delimitation analysis uncovered nine well-supported putative species, eight of which are new to science. Most Asterophila species were found on a single host species, but four species were found on multiple hosts from one or two closely related genera, showing phylogenetic conservatism of host use. Both distance-based and event-based cophylogenetic analyses uncovered a strong signal of coevolution in this system, but most associations were explained by non-cospeciation events. Discussion The prevalence of duplication and host-switching events in Asterophila and its asteroid hosts suggests that synchronous evolution may be rare even in obligate endoparasitic systems. The apparent restricted distribution of Asterophila from around the Scotia Arc may be an artefact of concentrated sampling in the area and a low obvious prevalence of infection. Given the richness of parasites on a global scale, their role in promoting host diversification, and the threat of their loss through coextinction, future work should continue to investigate parasite diversity and coevolution in vulnerable ecosystems.

The cold waters of Antarctica are known to harbor a rich biodiversity. Our continuing interest in the chemical analysis of Antarctic invertebrates has resulted in the isolation of friomaramide (1), a new, highly modified hexapeptide, from the Antarctic sponge Inflatella coelosphaeroides. The structure of friomaramide was determined using spectroscopic methods and its configuration established by Marfey's method. Friomaramide, which bears the unusual permethylation of the amino acid backbone and is the longest polypeptide bearing a tryptenamine C-terminus, blocks >90% of Plasmodium falciparum liver-stage parasite development at 6.1 μM.

An anomalous El Nino-associated sea surface temperature stress event was predicted to affect tropical Australian reefs, including those in North Western Australia in the summer of 2015/2016. Thermal stress events are well known to result in widespread hard coral mortality events, but other symbiotic organisms such as soft corals, giant clams and sponges can also be affected. Here, we examine whether the 2016 thermal stress event deleteriously impacted coral reef communities in the remote Bonaparte Archipelago, central inshore Kimberley bioregion, North West Australia. Our results confirm the region experienced a thermal stress event of similar magnitude to other regional localities (i.e., southern Kimberley and Scott Reef), but contrary to those locations that experienced widespread bleaching events, we find no evidence to suggest widespread mortality events occurred among photosymbiotic organisms in the Bonaparte Archipelago. Photosymbiotic organisms in this region are assumed to be well adapted to fluctuating environmental conditions; however, in this instance, a greater magnitude of night-time cooling may have driven variability in regional susceptibility to thermal stress. The Bonaparte Archipelago is emerging as a globally significant ecological refuge for photosymbiotic benthic fauna that are threatened by cumulative anthropogenic and climate stressors in other parts of their distribution.

A recent taxonomic revision split the circumglobal sea hare Aplysia parvula into 10 constituent taxa, of which only three are likely to be found in the Southern Pacific. This prompted an investigation of animals previously identified as A. parvula from Australia. Specimens collected from Eastern Australia and Hunter Island, east of New Caledonia, could not be satisfactorily identified with any of the currently accepted taxa based on morphological diagnostic features listed in the revision, however, the presence of a highly concave shell is diagnostic. Quantification of genetic divergence using Cytochrome Oxidase I (COI) supports the delineation of this species as a distinct taxon, and a phylogenetic reconstruction based on concatenated COI, 16S and H3 markers reveals a sister relationship with the newly described Aplysia ghanimii from the Atlantic and Western Indian Oceans and an undescribed species from Japan. As a result, the name Aplysia concava G. B. Sowerby, I, 1833 is resurrected for this species. As the original description was based solely on a shell, a redescription is provided here with photographs of living animals and microscope images of internal anatomical structures.

Sponges represent one of the most species-rich hosts for commensal barnacles yet host utilisation and diversity have not been thoroughly examined. This study investigated the diversity and phylogenetic relationships of sponge-inhabiting barnacles within a single, targeted host group, primarily from Western Australian waters. Specimens of the sponge order Dictyoceratida were surveyed and a total of 64 host morphospecies, representing four families, were identified as barnacle hosts during the study. Utilising molecular (COI, 12S) and morphological methods 42 molecular operational taxonomic units (MOTUs) of barnacles, representing Acasta, Archiacasta, Euacasta and Neoacasta were identified. Comparing inter- and intra-MOTU genetic distances showed a barcode gap between 2.5% and 5% for COI, but between 1% and 1.5% in the 12S dataset, thus demonstrating COI as a more reliable barcoding region. These sponge-inhabiting barnacles were demonstrated to show high levels of host specificity with the majority being found in a single sponge species (74%), a single genus (83%) or a single host family (93%). Phylogenetic relationships among the barnacles were reconstructed using mitochondrial (12S, COI) and nuclear (H3, 28S) markers. None of the barnacle genera were recovered as monophyletic. Euacasta was paraphyletic in relation to the remaining Acastinae genera, which were polyphyletic. Six well-supported clades of molecular operational taxonomic units, herein considered to represent species complexes, were recovered, but relationships between them were not well supported. These complexes showed differing patterns of host usage, though most were phylogenetically conserved with sister lineages typically occupying related hosts within the same genus or family of sponge. The results show that host specialists are predominant, and the dynamics of host usage have played a significant role in the evolutionary history of the Acastinae.

International audience; Members of the trochoidean genus Margarella (Calliostomatidae) are broadly distributed across Antarctic and sub-Antarctic ecosystems. Here we used novel mitochondrial and nuclear gene sequences to clarify species boundaries and phylogenetic relationships among seven nominal species distributed on either side of the Antarctic Polar Front (APF). Molecular reconstructions and species-delimitation analyses recognized only four species: M. antarctica (the Antarctic Peninsula), M. achilles (endemic to South Georgia), M. steineni (South Georgia and Crozet Island) and the morphologically variable M. violacea (=M. expansa, M. porcellana and M. pruinosa), with populations in southern South America, the Falkland/Malvinas, Crozet and Kerguelen Islands. Margarella violacea and M. achilles are sister species, closely related to M. steineni, with M. antarctica sister to all these. This taxonomy reflects contrasting biogeographic patterns on either side of the APF in the Southern Ocean. Populations of Margarella north of the APF (M. violacea) showed significant genetic variation but with many shared haplotypes between geographically distant populations. By contrast, populations south of the APF (M. antarctica, M. steineni and M. achilles) exhibited fewer haplotypes and comprised three distinct species, each occurring across a separate geographical range. We hypothesize that the biogeographical differences may be the consequence of the presence north of the APF of buoyant kelps – potential long-distance dispersal vectors for these vetigastropods with benthic-protected development – and their near-absence to the south. Finally, we suggest that the low levels of genetic diversity within higher-latitude Margarella reflect the impact of Quaternary glacial cycles that exterminated local populations during their maxima.

Tudivasum Rosenberg & Petit, 1987 is a morphologically distinct gastropod genus of low diversity. All but one species are known from Australia and they occur from the intertidal zone down to hundreds of metres on the continental shelf. These carnivorous gastropods are thought to have intracapsular development. The six currently recognized extant species are reviewed here and their geographical ranges clarified. Two new species, Tudivasum chaneyi n. sp. and T. ashmorense n. sp., are described from Ashmore Reef, Western Australia, and are characterized by differences in protoconch colour and shell sculpture. The third new species, T. westrale n. sp., is described from the mid-west coast of Western Australia, where it has long been misidentified as T. spinosum (H. Adams & A. Adams, 1864). We generated a molecular phylogeny based on mitochondrial DNA sequence data to test morphological species concepts and reconstruct relationships among four of the described species. High levels of divergence within one of the new species could indicate an additional cryptic species.

During the Last Glacial Maximum (LGM), global sea levels were 120-130 m lower than today, resulting in the emergence of most continental shelves and extirpation of subtidal organisms from these areas. During the interglacial periods, rapid inundation of shelf regions created a dynamic environment for coastal organisms, such as the charismatic leafy seadragon (Phycodurus eques, Syngnathidae), a brooder with low dispersal ability inhabiting kelp beds in temperate Australia. Reconstructions of the palaeoshoreline revealed that the increase of shallow areas since the LGM was not uniform across the species' range and we investigated the effects of these asymmetries on genetic diversity and structuring. Using targeted capture of 857 variable ultraconserved elements (UCEs, 2,845 single nucleotide polymorphisms) in 68 individuals, we found that the regionally different shelf topographies were paralleled by contrasting population genetic patterns. In the west, populations may not have persisted through sea-level lows because shallow seabed was very limited. Shallow genetic structure, weak expansion signals and a westward cline in genetic diversity indicate a postglacial recolonization of the western part of the range from a more eastern location following sea-level rise. In the east, shallow seabed persisted during the LGM and increased considerably after the flooding of large bays, which resulted in strong demographic expansions, deeper genetic structure and higher genetic diversity. This study suggests that postglacial flooding with rising sea levels produced locally variable signatures in colonizing populations.

The taxonomy and diversity of chromodorid nudibranchs (Mollusca: Gastropoda: Nudibranchia) are relatively well-studied, although molecular analyses have only been applied to test few species-level groups. Within Chomodorididae, the genus Goniobranchus Pease, 1866 contains many different groups based on external colouration, including the red-reticulate group. Previously, most of the red-reticulate group has been considered as either a single variable species, identified as either Goniobranchus reticulatus Pease, 1866 or Goniobranchus tinctorius (Ruppell & Leuckart, 1828), or as a species complex. In this study, red-reticulate nudibranchs were collected from different locations spanning the Indo-Pacific. From these specimens, DNA data were derived, consisting of mitochondrial cytochrome c oxidase subunit I (COI) and 16S ribosomal DNA (16S rDNA) sequences. COI data were analysed using a distance-based species delimitation approach (ABGD), and the combined dataset (COI+16S rDNA) was analysed using maximum likelihood (ML) and Bayesian inference (BI) methods. The analyses identified five species-level clades, but none of the clades’ morphotypes matched exactly with the original descriptions of G. tinctorius or G. reticulatus. The non-reticulate, spotted species Goniobranchus splendidus (Angas, 1864) falls inside this complex. The existence of multiple species-level clades in the Goniobranchus red-reticulate group suggests this is a group of closely related species, comprising at least five undescribed taxa.

Maintaining the accretion potential and three dimensional structure of coral reefs is a priority but reef-building scleractinian corals are highly threatened and retreating. Hence future reefs are predicted to be dominated by non-constructional taxa. Since the Late Triassic however, other non-scleractinian anthozoans such as Heliopora have contributed to tropical and subtropical reef-building. Heliopora is an ancient and highly conserved reef building octocoral genus within the monospecific Family Helioporidae, represented by a single extant species – H. coerulea, Pallas, 1766. Here we show integrated morphological, genomic and reproductive evidence to substantiate the existence of a second species within the genus Heliopora. Importantly, some individuals of the new species herein described as Heliopora hiberniana sp. nov. feature a white skeleton indicating that the most diagnostic and conserved Heliopora character (the blue skeleton) can be displaced. The new species is currently known only from offshore areas in north Western Australia, which is a part of the world where coral bleaching events have severely impacted the scleractinian community over the last two decades. Field observations indicate individuals of both H. coerulea and H. hiberniana sp. nov. were intact after the 2016 Scott Reef thermal stress event, and we discuss the possibility that bleaching resistant non-scleractinian reef builders such as Heliopora could provide new ecological opportunities for the reconfiguration of future reefs by filling empty niches and functional roles left open by the regression of scleractinian corals.

Predicting biotic resistance to highly invasive strains of “killer algae” (Caulerpa spp.) requires understanding the diversity and feeding preferences of native consumers, including sea slugs in family Oxynoidae. Past studies reported low algal host specificity for Oxynoe (6 spp.) and Lobiger (4 spp.), but these taxonomically challenging slugs may represent species complexes of unrecognized specialists that prefer different Caulerpa spp. Here, we assess global diversity of these genera by integrating gene sequences with morphological data from microscopic teeth and internal shells, the only hard parts in these soft-bodied invertebrates. Four delimitation methods applied to datasets comprising mtDNA and/or nuclear alleles yielded up to 16 species hypotheses for samples comprising five nominal taxa, including five highly divergent species in Lobiger and five in Oxynoe. Depending on the analysis, a further four to six species were recovered in the O. antillarum-viridis complex, a clade in which mitochondrial divergence was low and nuclear alleles were shared among lineages. Bayesian species delimitation using only morphological data supported most candidate species, however, and integrative analyses combining morphological and genetic data fully supported all complex members. Collectively, our findings double the recognized biodiversity in Oxynoidae, and illustrate the value of including data from traits that mediate fast-evolving ecological interactions during species delimitation. Preference for Caulerpa spp. and radular tooth characteristics covaried among newly delimited species, highlighting an unappreciated degree of host specialization and coevolution in these taxa that may help predict their role in containing outbreaks of invasive algae.

During a 2013 cruise in the Southern Ocean we collected specimens of the octocoral Plumarella delicatissima between 800 and 950 m depth. Five new furanocembranoid diterpenes, keikipukalides A-E (1-5), the known diterpene pukalide aldehyde (6), and the known norditerpenoid ineleganolide (7) were isolated from the coral. These Plumarella terpenes lack mammalian cytotoxicity, while 2-7 display activity against Leishmania donovani between 1.9 and 12 μM. Structure elucidation was facilitated by one- and two-dimensional NMR spectroscopy and mass spectrometry, and keikipukalides A and E were confirmed by X-ray crystallography.

Rhodopemorphs are small, interstitial or psammobiotic heterobranch slugs, which have been troubling to place phylogenetically. Their small size and habit of living in or among sediment have led to a correlated reduction and simplification of morphology, and consequently to contradictory phylogenetic signal from anatomical features. When morphological data have previously been used to generate phylogenetic hypotheses, these were vulnerable to the effects of homoplasy. We collected multiple species of Rhodope, along with another rhodopemorph genus Helminthope, to produce DNA sequence data for the first time to test their monophyly. We sequenced mitochondrial and nuclear genes, and analysed the data under maximum likelihood and Bayesian inference. By analysing these data in a broader heterobranch dataset, we also examined the placement of Rhodopemorpha within Heterobranchia. Despite rhodopemorphs showing aspects of a euthyneurous (and pentaganglionate) condition, their placement in a molecular phylogeny occurs outside the taxon defined by this term (i.e. Euthyneura = Pentaganglionata). Instead, model-based inferences placed rhodopemorphs among basal heterobranch taxa, usually as a clade sister to a shelled group (Murchisonellidae) that was recently removed from the Pyramidelloidea. The Rhodopemorpha + Murchisonellidae clade is herein termed Allomorpha. Three-dimensional reconstruction methods have elucidated potential morphological homologies for Rhodopemorpha and comparisons with Murchisonellidae have also uncovered morphological support for this placement. Thus, we consider the phylogenetic placement of Rhodopemorpha solved, although relationships among lower Heterobranchia lineages remain challenging.

The cyanobacteriosponge Terpios hoshinota has been reported throughout the Indo-Pacific including the Great Barrier Reef, Australia. The species encrusts live coral, giant clams, and other benthos and can be a threat to benthic communities on coral reefs. The Kimberley region of Western Australia has some of the least impacted reefs globally. We report for the first time the presence of T. hoshinota in the eastern Indian Ocean on Kimberley inshore coral reefs. Given its invasive potential, reef health surveys should include this species, and monitoring approaches developed to audit the remote Kimberley for this and other invasive species.

The genus Alveopora is a scleractinian coral taxon whose phylogenetic classification has recently changed from the family Poritidae to Acroporidae. This change, which was made based on single-locus genetic data, has led to uncertainty about the placement of Alveopora and the ability for deep evolutionary relationships in these groups to be accurately recovered and represented by limited genetic datasets. We sought to characterize the higher-level position of Alveopora using newly available transcriptome data to confirm its placement within Acroporidae and resolve its closest ancestor. Here we present an analysis of a new 2031 gene dataset that confirms the placement of Alveopora within Acroporidae corroborating other single-locus (COI, 16S and ITS) analyses and a mitogenome dataset. We also resolve the position of Alveopora as sister to the genus Montipora. This has allowed the re-interpretation of morphology, and a rediagnosis of the family Acroporidae and the genus Alveopora.

The Splanchnotrophidae is a family of highly modified endoparasitic copepods known to infest nudibranch or sacoglossan sea slug hosts. Most splanchnotrophid species appear to be specific to a single host, but some were reported from up to nine different host species. However, splanchnotrophid taxonomy thus far is based on external morphology, and taxonomic descriptions are, mostly, old and lack detail. They are usually based on few specimens, with intraspecific variability rarely reported. The present study used molecular data for the first time to test (1) the current taxonomic hypotheses, (2) the apparently strict host specificity of the genusIsmailaand (3) the low host specificity of the genusSplanchnotrophuswith regard to the potential presence of cryptic species. Phylogenetic analyses herein used sequences of the barcoding region of the cytochrome oxidase I (COI) gene from 40 specimens representing 13 species of five genera. Species delimitation approaches include distance and barcoding gap analyses, haplotype networks and diagnostic nucleotides. Molecular results are largely compatible with the commonly accepted, morphology-based taxonomy of the Splanchnotrophidae. Strict host specificity could be confirmed for twoIsmailaspecies. COI analyses also supported the idea thatSplanchnotrophus angulatusis host-promiscuous. InIsmaila, morphology seems more suitable than barcoding to display speciation events via host switches in a recent Chilean radiation. InSplanchnotrophus, some genetic structure suggests ongoing diversification, which should be investigated further given the inadequate morphology-based taxonomy. The present study thus supports the presence of two different life history strategies in splanchnotrophids, which should be explored integratively.

The common seadragon is an iconic fish with presumed limited dispersal, because juveniles hatch directly from the tail of the male parent. Nothing is presently known of their phylogeographic structure, despite conservation concerns and a distribution spanning southern Australia. Here, we sequenced mitochondrial genes from 201 common seadragons in Western Australia, South Australia, Victoria, Tasmania and New South Wales. We show that common seadragon populations are highly structured geographically, and that genetic variation varies significantly. The historical Bassian Isthmus appears to have left a strong imprint on population structure. Populations east of the Bassian Isthmus are low in diversity and appear more connected than those in the west, although this is likely caused by a recent expansion from a common source after the Last Glacial Maximum. All individuals from Eden, Bicheno and Hobart are represented by only two haplotypes. Populations west of the Bassian Isthmus are more diverse, with the highest diversity indices shown by Western Australian and Spencer Gulf populations. A large sampling gap across the Great Australian Bight is yet to be resolved; the west versus east break here may be an artifact of this gap. Almost all sampled populations can be inferred to have limited gene flow among them, which has implications for recovery after local extinction. Populations thought to be in decline (Sydney, Hobart) showed low genetic diversity, which may make them vulnerable to further reductions.

The discovery of four new Xenoturbella species from deep waters of the eastern Pacific Ocean is reported here. The genus and two nominal species were described from the west coast of Sweden, but their taxonomic placement remains unstable. Limited evidence placed Xenoturbella with molluscs, but the tissues can be contaminated with prey. They were then considered deuterostomes. Further taxon sampling and analysis have grouped Xenoturbella with acoelomorphs (=Xenacoelomorpha) as sister to all other Bilateria (=Nephrozoa), or placed Xenacoelomorpha inside Deuterostomia with Ambulacraria (Hemichordata + Echinodermata). Here we describe four new species of Xenoturbella and reassess those hypotheses. A large species (>20 cm long) was found at cold-water hydrocarbon seeps at 2,890 m depth in Monterey Canyon and at 1,722 m in the Gulf of California (Mexico). A second large species (~10 cm long) also occurred at 1,722 m in the Gulf of California. The third large species (~15 cm long) was found at ~3,700 m depth near a newly discovered carbonate-hosted hydrothermal vent in the Gulf of California. Finally, a small species (~2.5 cm long), found near a whale carcass at 631 m depth in Monterey Submarine Canyon (California), resembles the two nominal species from Sweden. Analysis of whole mitochondrial genomes places the three larger species as a sister clade to the smaller Atlantic and Pacific species. Phylogenomic analyses of transcriptomic sequences support placement of Xenacoelomorpha as sister to Nephrozoa or Protostomia.

The small, green sea hare, Phyllaplysia viridis, originally described from Indonesia in 1905, has rarely been recorded since. Because the taxonomic status of Phyllaplysia is uncertain, there remains a need for clarification of the currently accepted species. To do so, we used contemporary specimens from eastern Australia and Indonesia to redescribe P. viridis using photographs and scanning electron imagery of its external and internal morphology. Although all examined specimens were small, consistently green animals, images revealed some variation in colour and pattern among adults. Molecular data confirmed the presence of P. viridis in the Philippines, Indonesia and eastern Australia. A phylogenetic tree based on available COI barcode sequences illustrate a sister relationship with other Phyllaplysia. The construction of a haplotype network failed to show any marked geographic structure among specimens.

Soft coral diversity in tropical northern Australia remains relatively understudied compared with other parts of the world. As a result of ongoing biodiversity surveys, we describe here a new genus of Octocorallia, Anastromvos, gen. nov., and two new species, A. aldersladei, sp. nov. and A. catherinae, sp. nov., collected from waters off the Pilbara, Kimberley and Darwin. To test the validity of the new genus, we used traditional morphological approaches combined with a molecular phylogeny using three mitochondrial genes (COI, mtMutS, ND2) and nuclear 28S. The markers did not amplify for the colony of A. catherinae, sp. nov., which was described on the basis of morphology only. The new genus, belonging to the family Alcyoniidae, is dimorphic, possessing autozooids and siphonozooids, and is characterised by its unique capitate growth form, stone-like colony consistency, heavy autozooid polyp armature and the possession of clubs, tuberculated spindles and/or oval-shaped sclerites and crosses. The molecular phylogeny shows the new genus as the sister group to Sarcophyton+Lobophytum, and forms a unique clade among other alcyoniid clades. The Sarcophyton–Lobophytum group of taxa can be ecologically dominant in shallow-water coral reef communities but there is still much taxonomic refinement needed for these and related genera. http://zoobank.org/urn:lsid:zoobank.org:pub:71E96A7A-A24D-4485-AF3B-834CEF959578

Increased exploration of northeastern Pacific deep-sea habitats has revealed a diverse and often poorly-known invertebrate community, including a number of undescribed species of nudibranchs studied herein. We used morphology to distinguish several new species from their congeners, and generated data where possible for mitochondrial (COI, 16S) and nuclear markers (H3) to place them in a phylogenetic context. We described here Tritonia nigritigris sp. nov., Dendronotus claguei sp. nov., Ziminella vrijenhoeki sp. nov., Cuthona methana sp. nov., Aeolidia libitinaria sp. nov. and redescribed Zeusia herculea (Bergh, 1894). Another species of Tritonia is described but not named due to the absence of reproductive system information. Although there are difficulties in collection from deep-sea habitats, only two of our new species are known from single specimens. As with many other deep-sea regions, we expect the number of new species from this region to increase with further exploration. Because the deep regions of the northeast Pacific are particularly vulnerable to the effects of decreasing oxygenation due to climate change, we consider that documenting this fauna has some level of urgency.

Note: In lieu of an abstract, this is an excerpt from the first page.Excerpt The authors wish to make the following corrections to this paper

Mimicry of warning signals is common, and can be mutualistic when mimetic species harbour equal levels of defence (Müllerian), or parasitic when mimics are undefended but still gain protection from their resemblance to the model (Batesian). However, whether chemically defended mimics should be similar in terms of toxicity (i.e. causing damage to the consumer) and/or unpalatability (i.e. distasteful to consumer) is unclear and in many studies remains undifferentiated. In this study, we investigated the evolution of visual signals and chemical defences in a putative mimicry ring of nudibranch molluscs. First, we demonstrated that the appearance of a group of red spotted nudibranchs molluscs was similar from the perspective of potential fish predators using visual modelling and pattern analysis. Second, using phylogenetic reconstruction, we demonstrated that this colour pattern has evolved multiple times in distantly related individuals. Third, we showed that these nudibranchs contained different chemical profiles used for defensive purposes. Finally, we demonstrated that although levels of distastefulness towards Palaemon shrimp remained relatively constant between species, toxicity levels towards brine shrimp varied significantly. We highlight the need to disentangle toxicity and taste when considering chemical defences in aposematic and mimetic species, and discuss the implications for aposematic and mimicry signal evolution.

Gutt, Julian ... et al.-- 17 pages, 4 figures, The biodiversity, ecosystem services and climate variability of the Antarctic continent and the Southern Ocean are major components of the whole Earth system. Antarctic ecosystems are driven more strongly by the physical environment than many other marine and terrestrial ecosystems. As a consequence, to understand ecological functioning, cross-disciplinary studies are especially important in Antarctic research. The conceptual study presented here is based on a workshop initiated by the Research Programme Antarctic Thresholds – Ecosystem Resilience and Adaptation of the Scientific Committee on Antarctic Research, which focussed on challenges in identifying and applying cross-disciplinary approaches in the Antarctic. Novel ideas and first steps in their implementation were clustered into eight themes. These ranged from scale problems, through risk maps, and organism/ecosystem responses to multiple environmental changes and evolutionary processes. Scaling models and data across different spatial and temporal scales were identified as an overarching challenge. Approaches to bridge gaps in Antarctic research programmes included multi-disciplinary monitoring, linking biomolecular findings and simulated physical environments, as well as integrative ecological modelling. The results of advanced cross-disciplinary approaches can contribute significantly to our knowledge of Antarctic and global ecosystem functioning, the consequences of climate change, and to global assessments that ultimately benefit humankind, In addition to the employers of the authors this study was funded by the SCAR SRPs AnT-ERA, AntClim21 and AntEco

For 40 years, paleontological studies of marine gastropods have suggested that species selection favors lineages with short-lived (lecithotrophic) larvae, which are less dispersive than long-lived (planktotrophic) larvae. Although lecithotrophs appeared to speciate more often and accumulate over time in some groups, lecithotrophy also increased extinction rates, and tests for state-dependent diversification were never performed. Molecular phylogenies of diverse groups instead suggested lecithotrophs accumulate without diversifying due to frequent, unidirectional character change. Although lecithotrophy has repeatedly originated in most phyla, no adult trait has been correlated with shifts in larval type. Thus, both the evolutionary origins of lecithotrophy and its consequences for patterns of species richness remain poorly understood. Here, we test hypothesized links between development mode and evolutionary rates using likelihood-based methods and a phylogeny of 202 species of gastropod molluscs in Sacoglossa, a clade of herbivorous sea slugs. Evolutionary quantitative genetics modeling and stochastic character mapping supported 27 origins of lecithotrophy. Tests for correlated evolution revealed lecithotrophy evolved more often in lineages investing in extra-embryonic yolk, the first adult trait associated with shifts in development mode across a group. However, contrary to predictions from paleontological studies, species selection actually favored planktotrophy; most extant lecithotrophs originated through recent character change, and did not subsequently diversify. Increased offspring provisioning in planktotrophs thus favored shifts to short-lived larvae, which led to short-lived lineages over macroevolutionary time scales. These findings challenge long-standing assumptions about the effects of alternative life histories in the sea. Species selection can explain the long-term persistence of planktotrophy, the ancestral state in most clades, despite frequent transitions to lecithotrophy.

The aeolid Pteraeolidia ianthina (Angas, 1864) is a strikingly-coloured aeolid nudibranch, informally known as the ‘Blue Dragon’. It is recognised as an unusually widespread Indo-Pacific species, with variation in colouration and morphology, and biogeographic differences in zooxanthellae (dinoflagellate symbionts of the genus Symbiodinium). This variation hints at possible cryptic species, which was tested here using phylogenetic analyses of mitochondrial DNA data (COI, 16S). Our results showed multiple well-supported clades with slight but consistent differences in radular morphology and colouration, and thus we clarify one of the three available names. A temperate NSW clade showed a more elongate and pointed central radular tooth and lacked white body colouration, in comparison to a more variable tropical clade, which had a shorter and more blunt central tooth. The type locality of Pteraeolidia ianthina is Sydney Harbour, New South Wales (NSW), Australia, and according to our study, does not occur outside NSW. Pteraeolidia semperi (Bergh, 1870) and P. scolopendrella (Risbec, 1928) are removed from synonymy with P. ianthina. Wider phylogeographic sampling is required before resolving the availability of the two remaining names, and subclades within the tropical clade, but there is evidence to suggest multiple cryptic species exist. The biogeographic differences in symbionts, and the importance of their role in life history, suggests that changes in symbiosis may have helped drive divergence via local adaptation in the host nudibranchs. © 2015 The Linnean Society of London

Many plants and animals store toxic or unpalatable compounds in tissues that are easily encountered by predators during attack. Defensive compounds can be produced de novo, or obtained from dietary sources and stored directly without selection or modification, or can be selectively sequestered or biotransformed. Storage strategies should be optimized to produce effective defence mechanisms but also prevent autotoxicity of the host. Nudibranch molluscs utilize a diverse range of chemical defences, and we investigated the accumulation and distribution of defensive secondary metabolites in body tissues of 19 species of Chromodorididae nudibranchs. We report different patterns of distribution across tissues, where: 1) the mantle had more or different (but structurally related) compounds than the viscera; 2) all compounds in the mantle were also in the viscera; and 3) the mantle had fewer compounds than the viscera. We found no further examples of species that selectively store a single compound, previously reported in Chromodoris species. Consistent with other studies, we found high concentrations of metabolites in mantle rim tissues compared to the viscera. Using bioassays, compounds in the mantle were more toxic than compounds found in the viscera for Glossodoris vespa Rudman, 1990 and Ceratosoma brevicaudatum Abraham, 1876. In G. vespa, compounds in the mantle were also more unpalatable to palaemonid shrimp than compounds found in the viscera. This indicates that these species may modify compounds to increase bioactivity for defensive purposes and/or selectively store more toxic compounds. We highlight clear differences in the storage of sequestered chemical defences, which may have important implications for species to employ effective defences against a range of predators.

Warning signal variation is ubiquitous but paradoxical: low variability should aid recognition and learning by predators. However, spatial variability in the direction and strength of selection for individual elements of the warning signal may allow phenotypic variation for some components, but not others. Variation in selection may occur if predators only learn particular colour pattern components rather than the entire signal. Here, we used a nudibranch mollusc,Goniobranchus splendidus, which exhibits a conspicuous red spot/white body/yellow rim colour pattern, to test this hypothesis. We first demonstrated that secondary metabolites stored within the nudibranch were unpalatable to a marine organism. Using pattern analysis, we demonstrated that the yellow rim remained invariable within and between populations; however, red spots varied significantly in both colour and pattern. In behavioural experiments, a potential fish predator,Rhinecanthus aculeatus, used the presence of the yellow rims to recognize and avoid warning signals. Yellow rims remained stable in the presence of high genetic divergence among populations. We therefore suggest that how predators learn warning signals may cause stabilizing selection on individual colour pattern elements, and will thus have important implications on the evolution of warning signals.

Chromodoris is a genus of colourful nudibranchs that feed on sponges and is found across the Indo-Pacific. While this was once the most diverse chromodorid genus, recent work has shown that the genus should be restricted to a monophyletic lineage that contains only 22 species, all of which exhibit black pigmentation and planar spawning behaviour. Earlier phylogenies of this group are poorly resolved and thus additional work is needed to clarify species boundaries within Chromodoris. This study presents a maximum-likelihood phylogeny based on mitochondrial loci (COI, 16S) for 345 Chromodoris specimens, including data from 323 new specimens and 22 from GenBank, from across the Indo-Pacific. Species hypotheses and phylogenetic analysis uncovered 39 taxa in total containing 18 undescribed species, with only five of 39 taxa showing stable colour patterns and distinct morphotypes. This study also presents the first evidence for regional mimicry in this genus, with C. colemani and C. joshi displaying geographically-based variation in colour patterns which appear to match locally abundant congenerics, highlighting the flexibility of these colour patterns in Chromodoris nudibranchs. The current phylogeny contains short branch lengths, polytomies and poor support at interior nodes, which is indicative of a recent radiation. As such, future work will employ a transcriptome-based exon capture approach for resolving the phylogeny of this group. In all, this study included 21 of the 22 described species in the Chromodoris sensu stricto group with broad sampling coverage from across the Indo-Pacific, constituting the most comprehensive sampling of this group to date. This work highlights several cases of undocumented diversity, ultimately expanding our knowledge of species boundaries in this group, while also demonstrating the limitations of colour patterns for species identification in this genus.

Multi-locus phylogenetic studies of echinoderms based on Sanger and RNA-seq technologies and the fossil record have provided evidence for the Asterozoa-Echinozoa hypothesis. This hypothesis posits a sister relationship between asterozoan classes (Asteroidea and Ophiuroidea) and a similar relationship between echinozoan classes (Echinoidea and Holothuroidea). Despite this consensus around Asterozoa-Echinozoa, phylogenetic relationships within the class Asteroidea (sea stars or starfish) have been controversial for over a century. Open questions include relationships within asteroids and the status of the enigmatic taxon Xyloplax. Xyloplax is thought by some to represent a newly discovered sixth class of echinoderms - and by others to be an asteroid. To address these questions, we applied a novel workflow to a large RNA-seq dataset that encompassed a broad taxonomic and genomic sample. This study included 15 species sampled from all extant orders and 13 families, plus four ophiuroid species as an outgroup. To expand the taxonomic coverage, the study also incorporated five previously published transcriptomes and one previously published expressed sequence tags (EST) dataset. We developed and applied methods that used a range of alignment parameters with increasing permissiveness in terms of gap characters present within an alignment. This procedure facilitated the selection of phylogenomic data subsets from large amounts of transcriptome data. The results included 19 nested data subsets that ranged from 37 to 4,281loci. Tree searches on all data subsets reconstructed Xyloplax as a velatid asteroid rather than a new class. This result implies that asteroid morphology remains labile well beyond the establishment of the body plan of the group. In the phylogenetic tree with the highest average asteroid nodal support several monophyletic groups were recovered. In this tree, Forcipulatida and Velatida are monophyletic and form a clade that includes Brisingida as sister to Forcipulatida. Xyloplax is consistently recovered as sister to Pteraster. Paxillosida and Spinulosida are each monophyletic, with Notomyotida as sister to the Paxillosida. Valvatida is recovered as paraphyletic. The results from other data subsets are largely consistent with these results. Our results support the hypothesis that the earliest divergence event among extant asteroids separated Velatida and Forcipulatacea from Valvatacea and Spinulosida.

The Murchisonellidae are a small taxon of minute snails with a high-spired shell that occur in shallow marine habitats. Molecular phylogenetics recently revealed that they are not members of the externally similar yet phylogenetically derived Pyramidellidae, but instead potentially one of the oldest clades among the heterobranch Gastropoda. Furthermore, current data surprisingly indicate a sister-group relationship with Rhodopemorpha, highly aberrant marine slugs with previously unclear affinities. Murchisonellidae are characterized by a specialized pincer-like radula, but very little further data exist on soft-body anatomy for most species, and there are only a few observations of living animals. Investigation of the anatomy of Murchisonellidae may thus yield new data providing insights into early heterobranch evolution and that of enigmatic Rhodopemorpha. We collected live specimens of the murchisonellid Koloonella cf. minutissima (Laseron, 1951), a member of a genus known mainly from eastern Australia. We provide detailed live photographs and interactive 3D data on all major organ systems, based on serial histological sections. The mantle cavity is shown to contain several distinct glands, a pair of which is conspicuously similar to glands found in Rhodopemorpha. The anterior digestive system contains a unique four-toothed radula, a feeble pharynx and a special, vacuolated oesophageal bulb. The reproductive system is complex and diaulic, and contains unusual structures. These results highlight structural diversity among minute lower Heterobranchia. Soft-body characters do not contradict, and may even support, the counterintuitive sister-group relationship with shell-less, wormshaped Rhodopemorpha. The classification of Murchisonellidae is discussed and a revised scheme is proposed.

An Antarctic coral belonging to the order Pennatulacea, collected during the 2013 austral autumn by trawl from 662 to 944 m depth, has yielded three new briarane diterpenes, bathyptilone A-C (1&ndash, 3) along with a trinorditerpene, enbepeanone A (4), which bears a new carbon skeleton. Structure elucidation was facilitated by one- and two-dimensional NMR spectroscopy, mass spectrometry and confirmed by X-ray crystallography. The three compounds were screened in four cancer cell lines. Bathyptilone A displayed selective nanomolar cytotoxicity against the neurogenic mammalian cell line Ntera-2.

Although variance in life history parameters is well known from comparisons among broad phylogenetic groups of marine invertebrates, there is still an outstanding need to increase empirical studies that compare closely related species. If the species under study share a recent common ancestor and developmental strategy, there is an opportunity to contrast maternal investment against interspecific variation while controlling for evolutionary distance. Furthermore, when these species co-occur, it allows for exploration of potential character displacement. We examined egg size and other factors related to reproduction in four closely related species, co-occurring nudibranchs belonging to the monophyletic Chromodoris planar spawning clade. The duration of oviposition appeared to be conserved and may be phylogenetically constrained in these four co-occurring species. In contrast, egg size differed significantly among species, but was not influenced by parental body length or position within the egg mass for an...

Until recently, only two species of seadragon were known, Phycodurus eques (the leafy seadragon) and Phyllopteryx taeniolatus (the common seadragon), both from Australia. In 2015, we described a new species of seadragon, Phyllopteryx dewysea (the ruby seadragon). Although the leafy and common seadragons are well known and commonly seen in aquarium exhibits world-wide, the ruby seadragon was known only from four preserved specimens, leaving many aspects of its biology unknown. Based on specimen records, it was speculated that the ruby seadragon normally lives at depths beyond recreational SCUBA diving limits, which may also explain why it went undiscovered for so long. The ruby seadragon also bears a superficial resemblance to the common seadragon, with a number of specimens misidentified in museum collections. The only recent live-collected specimen was trawled from the Recherche Archipelago, a cluster of over 100 islands in Western Australia. We took a small remotely operated vehicle (miniROV) to this locality to obtain the first images of live ruby seadragons. We made observations on the seadragon habitat and behavior, including feeding. We also provide new key observations on their morphology, notably that they lack dermal appendages and have a prehensile tail. We recommend that the ruby seadragon be protected as soon as practicable.

Background Although recent studies have greatly advanced understanding of deep molluscan phylogeny, placement of some taxa remains uncertain as different datasets support competing class-relationships. Traditionally, morphologists have placed Monoplacophora, a group of morphologically simple, limpet-like molluscs as sister group to all other conchiferans (shelled molluscs other than Polyplacophora), a grouping that is supported by the latest large-scale phylogenomic study that includes Laevipilina. However, molecular datasets dominated by nuclear ribosomal genes support Monoplacophora + Polyplacophora (Serialia). Here, we evaluate the potential of mitochondrial genome data for resolving placement of Monoplacophora. Results Two complete (Laevipilina antarctica and Vema ewingi) and one partial (Laevipilina hyalina) mitochondrial genomes were sequenced, assembled, and compared. All three genomes show a highly similar architecture including an unusually high number of non-coding regions. Comparison of monoplacophoran gene order shows a gene arrangement pattern not previously reported; there is an inversion of one large gene cluster. Our reanalyses of recently published polyplacophoran mitogenomes show, however, that this feature is also present in some chiton species. Maximum Likelihood and Bayesian Inference analyses of 13 mitochondrial protein-coding genes failed to robustly place Monoplacophora and hypothesis testing could not reject any of the evaluated placements of Monoplacophora. Conclusions Under both serialian or aculiferan-conchiferan scenarios, the observed gene cluster inversion appears to be a convergent evolution of gene arrangements in molluscs. Our phylogenetic results are inconclusive and sensitive to taxon sampling. Aculifera (Polyplacophora + Aplacophora) and Conchifera were never recovered. However, some analyses recovered Serialia (Monoplacophora + Polyplacophora), Diasoma (Bivalvia + Scaphopoda) or Pleistomollusca (Bivalvia + Gastropoda). Although we could not shed light on deep evolutionary traits of Mollusca we found unique patterns of gene arrangements that are common to monoplacophoran and chitonine polyplacophoran species but not to acanthochitonine Polyplacophora. Graphical abstract Complete mitochondrial genome of Laevipilina antarctica Electronic supplementary material The online version of this article (doi:10.1186/s12862-016-0829-3) contains supplementary material, which is available to authorized users.

Sea cucumbers (Holothuroidea) are a morphologically diverse, ecologically important, and economically valued clade of echinoderms; however, the understanding of the overall systematics of the group remains controversial. Here, we present a phylogeny of extant Holothuroidea assessed with maximum parsimony, maximum likelihood, and Bayesian approaches using approximately 4.3kb of mt- (COI, 16S, 12S) and nDNA (H3, 18S, 28S) sequences from 82 holothuroid terminals representing 23 of the 27 widely-accepted family-ranked taxa. Currently five holothuroid taxa of ordinal rank are accepted. We find that three of the five orders are non-monophyletic, and we revise the taxonomy of the groups accordingly. Apodida is sister to the rest of Holothuroidea, here considered Actinopoda. Within Actinopoda, Elasipodida in part is sister to the remaining Actinopoda. This latter clade, comprising holothuroids with respiratory trees, is now called Pneumonophora. The traditional Aspidochirotida is paraphyletic, with representatives from three orders (Molpadida, Dendrochirotida, and Elasipodida in part) nested within. Therefore, we discontinue the use of Aspidochirotida and instead erect Holothuriida as the sister group to the remaining Pneumonophora, here termed Neoholothuriida. We found four well-supported major clades in Neoholothuriida: Dendrochirotida, Molpadida and two new clades, Synallactida and Persiculida. The mapping of traditionally-used morphological characters in holothuroid systematics onto the phylogeny revealed marked homoplasy in most characters demonstrating that further taxonomic revision of Holothuroidea is required. Two time-tree analyses, one based on calibrations for uncontroversial crown group dates for Eleutherozoa, Echinozoa and Holothuroidea and another using these calibrations plus four more from within Holothuroidea, showed major discrepancies, suggesting that fossils of Holothuroidea may need reassessment in terms of placing these forms with existing crown clades.

Euthyneuran gastropods represent one of the most diverse lineages in Mollusca (with over 30,000 species), play significant ecological roles in aquatic and terrestrial environments and affect many aspects of human life. However, our understanding of their evolutionary relationships remains incomplete due to missing data for key phylogenetic lineages. The present study integrates such a neglected, ancient snail family Ringiculidae into a molecular systematics of Euthyneura for the first time and is supplemented by the first microanatomical data. Surprisingly, both molecular and morphological features present compelling evidence for the common ancestry of ringiculid snails with the highly dissimilar Nudipleura—the most species-rich and well-known taxon of sea slugs (nudibranchs and pleurobranchoids). A new taxon name Ringipleura is proposed here for these long-lost sisters, as one of three major euthyneuran clades with late Palaeozoic origins, along with Acteonacea (Acteonoidea + Rissoelloidea) and Tectipleura (Euopisthobranchia + Panpulmonata). The early Euthyneura are suggested to be at least temporary burrowers with a characteristic ‘bubble’ shell, hypertrophied foot and headshield as exemplified by many extant subtaxa with an infaunal mode of life, while the expansion of the mantle might have triggered the explosive Mesozoic radiation of the clade into diverse ecological niches.

In contrast to many tropical animals expanding southwards on the Australian coast concomitant with climate change, here we report a temperate endemic newly found in the tropics. Chromodorid nudibranchs are bright, colourful animals that rarely go unnoticed by divers and underwater photographers. The discovery of a new population, with divergent colouration is therefore significant. DNA sequencing confirms that despite departures from the known phenotypic variation, the specimen represents northern Goniobranchus splendidus and not an unknown close relative. Goniobranchus tinctorius represents the sister taxa to G. splendidus. With regard to secondary defences, the oxygenated terpenes found previously in this specimen are partially unique but also overlap with other G. splendidus from southern Queensland (QLD) and New South Wales (NSW). The tropical specimen from Mackay contains extracapsular yolk like other G. splendidus. This previously unknown tropical population may contribute selectively advantageous genes to cold-water species threatened by climate change. Competitive exclusion may explain why G. splendidus does not strongly overlap with its widespread sister taxon.

Sampling at appropriate spatial scales in the Southern Ocean is logistically challenging and may influence estimates of diversity by missing intermediate representatives. With the assistance of sampling efforts especially influenced by the International Polar Year 2007-2008, we gathered nearly 1500 specimens of the crinoid species Promachocrinus kerguelensis from around Antarctica. We used phylogeographic and phylogenetic tools to assess its genetic diversity, demographic history and evolutionary relationships. Six phylogroups (A-F) identified in an earlier study are corroborated here, with the addition of one new phylogroup (E2). All phylogroups are circumpolar, sympatric and eurybathic. The phylogeny of Promachocrinus phylogroups reveals two principal clades that may represent two different cryptic species with contrasting demographic histories. Genetic diversity indices vary dramatically within phylogroups, and within populations, suggesting multiple glacial refugia in the Southern Ocean: on the Kerguelen Plateau, in the East Weddell Sea and the South Shetland Islands (Atlantic sector), and on the East Antarctic continental shelf in the Dumont d'Urville Sea and Ross Sea. The inferences of gene flow vary among the phylogroups, showing discordant spatial patterns. Phylogroup A is the only one found in the Sub-Antarctic region, although without evident connectivity between Bouvet and Kerguelen populations. The Scotia Arc region shows high levels of connectivity between populations in most of the phylogroups, and barriers to gene flow are evident in East Antarctica.

Cephalaspid gastropod molluscs are increasingly being utilized in studies of tradeoffs associated with conflicts in simultaneous hermaphroditic reproduction. Chelidonura sandrana exhibits a wide range of colour forms, from black to yellow-spotted or marked with white. One of these colour morphs was described as Chelidonura tsurugensis Baba & Abe, 1959, while others have been accepted as natural variation within C. sandrana Rudman, 1973. Previous work has mostly assumed (1) that colour forms represent the same species and (2) that reproductive behaviour does not differ among colour forms. Mating behaviour between three co-occurring colour forms (simplified to ‘black’, ‘yellow’ and ‘white’) of the Chelidonura tsurugensis–sandrana species complex was investigated to test the above assumptions. Individuals of each colour form were crossed with individuals of the same form, and then with each of the two other colour forms, and with additional species as controls. Mating frequency, duration and an individual’s role of sperm donor or receiver were recorded. The results show that there was no evidence for positive assortative mating with respect to colour form. Instead, at least one colour form was shown to mate significantly more often, and for longer, with individuals of colour forms other than its own. There was no evidence for trading of intromissions or intromission duration among colour forms. Although all three colour forms readily mate and behave like a single biological species, the questionable identity of C. tsurugensis prevents synonymization of C. sandrana with this species. Conservatively, we suggest that until a systematic revision of Chelidonura is carried out, the application of the name C. tsurugensis should be restricted to Japan, and elsewhere the yellow-spotted colour form should be referred to as C. sandrana.

The turbellarian-like, radula-lacking Rhodope has been a mystery to taxonomists for over 160 years and was considered a specialized off-shoot of either opisthobranch or pulmonate Euthyneura. Occasionally reported from intertidal waters and sand habitats from all continents, most species of these minute slugs are poorly known and characterized mainly by differences in pigmentation. To understand the evolution of heterobranch microslugs, we established a morphological dataset for Rhodope by describing a new species found in the temperate waters of southern Australia. To set a standard for rhodopids, all major organ systems of R. rousei n. sp. are reconstructed three-dimensionally from series of semithin sections using the software Amira. Microanatomy confirms the loss of many general gastropod features such as foot, cephalic tentacles, shell, radula, mantle cavity, gill and heart. Excretory and digestive systems are heavily modified, with free rhogocytes in the presumed position of the heart, and a secondary buccal bulb replacing the function of the vestigial pharynx. Structural details of the monaulic but hermaphroditic genital system suggest cutaneous fertilization via spermatophores formed in specialized glands. The highly concentrated central nervous system is compared to those of other species of the genus and targets of all detectable nerves are summarized. These characters are compared with adaptations shown by other interstitial gastropods.

Some living organisms produce visible light (bioluminescence) for intra- or interspecific visual communication. Here, we describe a remarkable bioluminescent adaptation in the marine snailHinea brasiliana. This species produces a luminous display in response to mechanical stimulation caused by encounters with other motile organisms. The light is produced from discrete areas on the snail's body beneath the snail's shell, and must thus overcome this structural barrier to be viewed by an external receiver. The diffusion and transmission efficiency of the shell is greater than a commercial diffuser reference material. Most strikingly, the shell, although opaque and pigmented, selectively diffuses the blue-green wavelength of the species bioluminescence. This diffusion generates a luminous display that is enlarged relative to the original light source. This unusual shell thus allows spatially amplified outward transmission of light communication signals from the snail, while allowing the animal to remain safely inside its hard protective shell.

Wilson, N. G. & Rouse, G. W. (2010). Convergent camouflage and the non-monophyly of ‘seadragons’ (Syngnathidae: Teleostei): suggestions for a revised taxonomy of syngnathids. —Zoologica Scripta, 39, 551–558. The phylogeny and classification of the charismatic Syngnathidae (e.g. pipefish, seahorses) has not been comprehensively examined to date. In particular, we assessed morphological hypotheses that previously suggested the three ‘seadragon’ genera (Phycodurus, Phyllopteryx, Haliichthys) do not form a monophyletic group. We used three mitochondrial markers to investigate evolutionary relationships within Syngnathidae, and demonstrated that Phycodurus + Phyllopteryx formed a clade that excluded Haliichthys, indicating the elaborate appendages used for camouflage have evolved independently. A time-calibrated tree revealed the divergence of true seadragons as coincident with other kelp-associated fauna. We found evidence for the resurrection of neglected subfamily names, and recovered Doryrhampinae, Nerophinae, Soleganthinae, Phyllopteryginae, Sygnathoidinae and Haliichthyinae as clades. Even after removing these groups from what is currently recognized as Syngnathinae, we showed that the remaining members of Syngnathinae are not monophyletic. In the light of this information, some conclusions about the diversity of reproductive strategies found within ‘Syngnathinae’ need to be re-examined and further revision of syngnathid classification is needed.

Strong currents and deep passages of water can be barriers for larval dispersal of continental marine animals, but potential effects on direct developers are under-investigated. We examined the genetic structure of Doris kerguelenensis, a directly developing sea slug that occurs across the Drake Passage, the body of water separating Antarctica from South America. We found deep mitochondrial divergences within populations on both sides of the Drake Passage, and South American animals formed multiple sister-group relationships with Antarctic animals. A generalised molecular clock suggested these trans-Drake pairs diverged during the Pliocene–Pleistocene, after the formation of the Drake Passage. Statistical parsimony methods recovered 29 separate haplotype networks (many sympatric) that likely correlate with allopatric events caused by repeated glacial cycles. Data from 16S were congruent but more conserved than COI, and the estimated ancestral 16S haplotype was widespread. The marked difference in the substitution rates between these two mitochondrial genes results in different estimates of connectivity. Demographic analyses on networks revealed some evidence for selection and expanding populations. Contrasting with the Northern Hemisphere, glaciation in Antarctica appears to have increased rather than reduced genetic diversity. This suggests orbitally forced range dynamics based on Northern Hemisphere phylogeography do not hold for Antarctica. The diverse lineages found in D. kerguelenensis point towards a recent, explosive radiation, likely reflecting multiple refuges during glaciation events, combined with limited subsequent dispersal. Whether recognised as cryptic species or not, genetic diversity in Antarctic marine invertebrates appears higher than expected from morphological analyses, and supports the Antarctic biodiversity pump phenomenon.

We report observations on spawning and early development in bone-eating worms of the genus Osedax. Individual females of Osedax rubiplumus were observed at 1820 m depth freely spawning hundreds of oocytes, and females of an undescribed species, Osedax “orange collar”, were observed spawning in laboratory aquaria. Cytological and molecular analysis of the spawned oocytes of two Osedax species revealed no evidence for the bacterial endosymbionts that the female worms require for their nutrition, suggesting that the bacteria must be acquired later from the environment, as they are in other siboglinids. Individual O. “orange collar” females released an average of 335 (±130) eggs per day, but the number of oocytes spawned per day varied greatly, suggesting that not all the females spawned daily. Fertilization rates of the spawned oocytes varied from 0 to 100%, though most females showed nearly 100% fertilization rates. Oocytes spawned in the laboratory at 4–6°C were negatively buoyant. If fertilized, these oocytes extruded polar bodies and then after at least four hours cleaved unequally. Subsequent cleavages occurred in a spiral pattern at roughly 2-h intervals, resulting in free-swimming trochophore larvae after 24 h. These lecithotrophic trochophores swam for 9–16 days before settling with several hooked chaetae, similar to those of dwarf Osedax males. The larval life span of the Osedax species studied in the laboratory appears to be shorter than in closely related Vestimentifera. Osedax rubiplumus, on the other hand, has much larger oocytes and so may have greater dispersal potential than these other Osedax species. The high fecundity and apparently continuous reproduction of Osedax boneworms permits the opportunistic exploitation of sunken vertebrate bones.

Holland, N. D., Campbell, T. G., Garey, J. R., Holland, L. Z. and Wilson, N. G. 2009. The Florida amphioxus (Cephalochordata) hosts larvae of the tapeworm Acanthobothrium brevissime : natural history, anatomy and taxonomic identification of the parasite. — Acta Zoologica (Stockholm) 90 : 75-86. Plerocercoid larvae of a tapeworm are frequently found in the hindgut lumen of the Florida amphioxus ( Branchiostoma floridae ) in central west Florida. About three-quarters of the adult amphioxus are parasitized. On average, each adult amphioxus hosts about five tapeworm larvae. The residence time of the parasites in the amphioxus gut appears to be in the order of several months, which is considerably shorter than the potential lifetime of the host. The living larvae range in length (when fully extended) from 300 to 850 ∝ m and are approximately cone-shaped, tapering to a point posteriorly and bearing a single large sucker anteriorly. Toward the anterior end of the body are four hookless bothridia, each indented by three loculi plus an inconspicuous accessory sucker. The larvae initiate the early stages of hook formation when they are cultured for a few days in urea-saline (mimicking the gut fluid of the definitive host, which is an elasmobranch). The tapeworm larvae are identifiable to genus and species on the basis of correspondences between their nuclear ribosomal DNA genes and those of adult specimens of Acanthobothrium brevissime recovered from the spiral valve of a stingray from the same environment.