Deep-sea origin and in-situ diversification of chrysogorgiid octocorals

The diversity, ubiquity and prevalence in deep waters of the octocoral family Chrysogorgiidae Verrill, 1883 make it noteworthy as a model system to study radiation and diversification in the deep sea. Here we provide the first comprehensive phylogenetic analysis of the Chrysogorgiidae, and compare phylogeny and depth distribution. Phylogenetic relationships among 10 of 14 currently-described Chrysogorgiidae genera were inferred based on mitochondrial (mtMutS, cox1) and nuclear (18S) markers. Bathymetric distribution was estimated from multiple sources, including museum records, a literature review, and our own sampling records (985 stations, 2345 specimens). Genetic analyses suggest that the Chrysogorgiidae as currently described is a polyphyletic family. Shallow-water genera, and two of eight deep-water genera, appear more closely related to other octocoral families than to the remainder of the monophyletic, deep-water chrysogorgiid genera. Monophyletic chrysogorgiids are composed of strictly (Iridogorgia Verrill, 1883, Metallogorgia Versluys, 1902, Radicipes Stearns, 1883, Pseudochrysogorgia Pante & France, 2010) and predominantly (Chrysogorgia Duchassaing & Michelotti, 1864) deep-sea genera that diversified in situ. This group is sister to gold corals (Primnoidae Milne Edwards, 1857) and deep-sea bamboo corals (Keratoisidinae Gray, 1870), whose diversity also peaks in the deep sea. Nine species of Chrysogorgia that were described from depths shallower than 200 m, and mtMutS haplotypes sequenced from specimens sampled as shallow as 101 m, suggest a shallow-water emergence of some Chrysogorgia species.     

A Comprehensive Phylogenetic Analysis of the Scleractinia (Cnidaria,Anthozoa) Based on Mitochondrial CO1 Sequence Data

Background

Classical morphological taxonomy places the approximately 1400 recognized species of Scleractinia (hard corals) into 27 families, but many aspects of coral evolution remain unclear despite the application of molecular phylogenetic methods. In part, this may be a consequence of such studies focusing on the reef-building (shallow water and zooxanthellate) Scleractinia, and largely ignoring the large number of deep-sea species. To better understand broad patterns of coral evolution, we generated molecular data for a broad and representative range of deep sea scleractinians collected off New Caledonia and Australia during the last decade, and conducted the most comprehensive molecular phylogenetic analysis to date of the order Scleractinia.

Methodology

Partial (595 bp) sequences of the mitochondrial cytochrome oxidase subunit 1 (CO1) gene were determined for 65 deep-sea (azooxanthellate) scleractinians and 11 shallow-water species. These new data were aligned with 158 published sequences, generating a 234 taxon dataset representing 25 of the 27 currently recognized scleractinian families.

Principal Findings/Conclusions

There was a striking discrepancy between the taxonomic validity of coral families consisting predominantly of deep-sea or shallow-water species. Most families composed predominantly of deep-sea azooxanthellate species were monophyletic in both maximum likelihood and Bayesian analyses but, by contrast (and consistent with previous studies), most families composed predominantly of shallow-water zooxanthellate taxa were polyphyletic, although Acroporidae, Poritidae, Pocilloporidae, and Fungiidae were exceptions to this general pattern. One factor contributing to this inconsistency may be the greater environmental stability of deep-sea environments, effectively removing taxonomic “noise” contributed by phenotypic plasticity. Our phylogenetic analyses imply that the most basal extant scleractinians are azooxanthellate solitary corals from deep-water, their divergence predating that of the robust and complex corals. Deep-sea corals are likely to be critical to understanding anthozoan evolution and the origins of the Scleractinia.     

Microboring organisms in living stylasterid corals (Cnidaria,Hydrozoa)

Microboring or euendolithic microorganisms, which colonize and penetrate various carbonate substrates, are abundant in coral reef ecosystems and play a major role in reef carbonate dissolution. A few studies reported the presence of euendoliths in stylasterid coral skeletons but the biological identity, distribution and abundance of these microorganisms remain largely unknown. Observations of over 100 stylasterid colonies, collected in the Indo-Pacific area, revealed for the first time that the association between these corals and euendolith organisms appears to be quite common in shallow tropical waters. The most abundant euendolith was identified as a cryptic stage in the development of the rhodophyte Porphyra (Conchocelis stage). The euendoliths were observed in the skeletons of seven species of three genera (four Stylaster, two Distichopora and one Lepidotheca). The presence of euendoliths inside skeletons conferred a particular colour to the studied stylasterid corals. Distribution and abundance of microborings varied significantly among stylasterid species and among branches of a single colony and so did the colour of their skeletons. Colonization of skeletons and the associated colour distribution were almost uniform in some stylasterids, forming an upward gradually diminishing or sharply limited gradient. This study shows that patterns of euendolith colonization and growth in stylasterid skeletons may depend on the stage of the euendolith development as well as on their environmental requirements such as light exposure.     

Facies analysis and sequence stratigraphy of an Upper Jurassic carbonate ramp in the Eastern Alborz range and Binalud Mountains, NE Iran

Upper Jurassic (Oxfordian-Kimmeridgian-Tithonian?) strata of NE Iran (Lar Formation) are composed of medium- to thick-bedded, mostly grainy limestones with various skeletal (bivalves, foraminifera, algae, corals, echinoderms, brachiopods, and radiolaria) and nonskeletal (peloids, ooids, intraclasts, and oncoids) components. Facies analysis documents low- to high-energy environments, including tidal-flat, lagoonal, barrier, and open-marine facies. Because of the wide lateral distribution of facies and the apparent absence of distinct paleobathymetric changes, the depositional system likely represents a westward-deepening homoclinal ramp. Four third-order depositional sequences can be distinguished in each of five stratigraphic measured sections. Transgressive system tracts (TST) show deepening-upward trends, in which shallow-water (tidal flat and lagoonal) facies are overlain by deeper-water (barrier and open-marine) facies. Highstand systems tracts (HST) show shallowing-upward trends in which deep-water facies are overlain by shallow-water facies. All sequence boundaries in the study area (except at the top of the stratigraphic column) are of the nonerosional (SB2) type. Correlation of depositional sequences in the studied sections show that relatively shallow marine (tidal-flat, lagoonal, barrier, and shallow open-marine) conditions dominated in the area. These alternated with deep-water open-marine wackestone and mudstones representing zones of maximum flooding (MFZ).     

The Biology of Lophelia pertusa (Linnaeus 1758) and Other Deep-Water Reef-Forming Corals and Impacts from Human Activities.

Over the last twenty years, human exploitation has begun to have an impact in the deep sea, especially in the upper bathyal zone. This has mainly taken the form of deep-sea fishing but more recently oil exploration has extended beyond the continental shelf. Deep-water coral reefs occur in the upper bathyal zone throughout the world. These structures, however, are poorly studied with respect to their occurrence, biology and the diversity of the communities associated with them. In the North-East Atlantic the coral Lophelia pertusa has frequently been recorded. The present review examines the current knowledge on L. pertusa and discusses similarities between its biology and that of other deep-water, reef-forming, corals. It is concluded that L. pertusa is a reef-forming coral that has a highly diverse associated fauna. Associated diversity is compared with that of tropical shallow-water reefs. Such a highly diverse fauna may be shared with other deep-water, reef-forming, corals though as yet many of these are poorly studied. The main potential threats to L. pertusa in the North-East Atlantic are considered to be natural phenomena, such as slope failures and changes in ocean circulation and anthropogenic impacts such as deep-sea fishing and oil exploration. The existing and potential impacts of these activities on L. pertusa are discussed. Deep-sea fishing is also known to have had a significant impact on deep-water reefs in other parts of the world.     

Out of Their Depth? Isolated Deep Populations of the Cosmopolitan Coral Desmophyllum dianthus May Be Highly Vulnerable to Environmental Change

Deep sea scleractinian corals will be particularly vulnerable to the effects of climate change, facing loss of up to 70% of their habitat as the Aragonite Saturation Horizon (below which corals are unable to form calcium carbonate skeletons) rises. Persistence of deep sea scleractinian corals will therefore rely on the ability of larvae to disperse to, and colonise, suitable shallow-water habitat. We used DNA sequence data of the internal transcribed spacer (ITS), the mitochondrial ribosomal subunit (16S) and mitochondrial control region (MtC) to determine levels of gene flow both within and among populations of the deep sea coral Desmophyllum dianthus in SE Australia, New Zealand and Chile to assess the ability of corals to disperse into different regions and habitats. We found significant genetic subdivision among the three widely separated geographic regions consistent with isolation and limited contemporary gene flow. Furthermore, corals from different depth strata (shallow <600 m, mid 1000-1500 m, deep >1500 m) even on the same or nearby seamounts were strongly differentiated, indicating limited vertical larval dispersal. Genetic differentiation with depth is consistent with the stratification of the Subantarctic Mode Water, Antarctic Intermediate Water, the Circumpolar Deep and North Pacific Deep Waters in the Southern Ocean, and we propose that coral larvae will be retained within, and rarely migrate among, these water masses. The apparent absence of vertical larval dispersal suggests deep populations of D. dianthus are unlikely to colonise shallow water as the aragonite saturation horizon rises and deep waters become uninhabitable. Similarly, assumptions that deep populations will act as refuges for shallow populations that are impacted by activities such as fishing or mining are also unlikely to hold true. Clearly future environmental management strategies must consider both regional and depth-related isolation of deep-sea coral populations.     

The importance of habitat and life history to extinction risk in sharks, skates, rays and chimaeras

We compared life-history traits and extinction risk of chondrichthyans (sharks, rays and chimaeras), a group of high conservation concern, from the three major marine habitats (continental shelves, open ocean and deep sea), controlling for phylogenetic correlation. Deep-water chondrichthyans had a higher age at maturity and longevity, and a lower growth completion rate than shallow-water species. The average fishing mortality needed to drive a deep-water chondrichthyan species to extinction (Fextinct) was 38-58% of that estimated for oceanic and continental shelf species, respectively. Mean values of Fextinct were 0.149, 0.250 and 0.368 for deep-water, oceanic and continental shelf species, respectively. Reproductive mode was an important determinant of extinction risk, while body size had a weak effect on extinction risk. As extinction risk was highly correlated with phylogeny, the loss of species will be accompanied by a loss of phylogenetic diversity. Conservation priority should not be restricted to large species, as is usually suggested, since many small species, like those inhabiting the deep ocean, are also highly vulnerable to extinction. Fishing mortality of deep-water chondrichthyans already exploited should be minimized, and new deep-water fisheries affecting chondrichthyans should be prevented.     

Repeated invasions into the twilight zone: evolutionary origins of a novel assemblage of fishes from deep Caribbean reefs

Mesophotic and deeper reefs of the tropics are poorly known and underexplored ecosystems worldwide. Collectively referred to as the ‘twilight zone’, depths below ~30–50 m are home to many species of reef fishes that are absent from shallower depths, including many undescribed and endemic species. We currently lack even a basic understanding of the diversity and evolutionary origins of fishes on tropical mesophotic reefs. Recent submersible collections in the Caribbean have provided new specimens that are enabling phylogenetic reconstructions that incorporate deep‐reef representatives of tropical fish genera. Here, we investigate evolutionary depth transitions in the family Gobiidae (gobies), the most diverse group of tropical marine fishes. Using divergence‐time estimation coupled with stochastic character mapping to infer the timing of shallow‐to‐deep habitat transitions in gobies, we demonstrate at least four transitions from shallow to mesophotic depths. Habitat transitions occurred in two broad time periods (Miocene, Pliocene–Pleistocene), and may have been linked to the availability of underutilized niches, as well as the evolution of morphological/behavioural adaptations for life on deep reefs. Further, our analysis shows that at least three evolutionary lineages that invaded deep habitats subsequently underwent speciation, reflecting another unique mode of radiation within the Gobiidae. Lastly, we synthesize depth distributions for 95 species of Caribbean gobies, which reveal major bathymetric faunal breaks at the boundary between euphotic and mesophotic reefs. Ultimately, our study is the first rigorous investigation into the origin of Caribbean deep‐reef fishes and provides a framework for future studies that utilize rare, deep‐reef specimens.     

Cold-water coral reef frameworks,megafaunal communities and evidence for coral carbonate mounds on the Hatton Bank,north east Atlantic

Offshore banks and seamounts sustain diverse megafaunal communities, including framework reefs formed by cold-water corals. Few studies have quantified environmental effects on the alpha or beta diversity of these communities. We adopted an interdisciplinary approach that used historical geophysical data to identify topographic highs on Hatton Bank, which were surveyed visually. The resulting photographic data were used to examine relationships between megafaunal communities and macrohabitat, the latter defined into six categories (mud, sand, cobbles, coral rubble, coral framework, rock). The survey stations revealed considerable small-scale variability in macrohabitat from exposed Late Palaeocene lava flows to quiescent muddy habitats and coral-built carbonate mounds. The first reported evidence for coral carbonate mound development in UK waters is presented, which was most pronounced near present-day or former sites of topographic change, suggesting that local current acceleration favoured coral framework growth and mound initiation. Alpha diversity varied significantly across macrohabitats, but not between rock and coral rubble, or between smaller grain sized categories of cobbles, sand and mud. Community composition differed between most macrohabitats, and variation in beta diversity across Hatton Bank was largely explained by fine-scale substratum. Certain megafauna were clearly associated with particular macrohabitats, with stylasterid corals notably associated with cobble and rock habitats and coral habitats characterized by a diverse community of suspension-feeders. The visual surveys also produced novel images of deep-water megafauna including a new photographic record of the gorgonian coral Paragorgia arborea, a species not previously reported from Rockall Plateau. Further interdisciplinary studies are needed to interpret beta diversity across these and other environmental gradients on Hatton Bank. It is clear that efforts are also needed to improve our understanding of the genetic connectivity and biogeography of vulnerable deep-water ecosystems and to develop predictive models of their occurrence that can help inform future conservation measures. An erratum to this article can be found at     

Illuminating the evolution of bioluminescence in sharks

The evolutionary context in which shark bioluminescence originated is poorly understood, despite it being critical to uncovering influential factors in the evolutionary history and diversity of living chondrichthyans as well as the mechanisms of deep-water colonization by vertebrates. This study provides the first joint reconstruction of the habitats, lifestyles, and occurrence of bioluminescence in the evolution of squalomorph sharks using ancestral state estimation analysis to resolve the timing of deep-sea colonization, the evolutionary origin of bioluminescence and the ancestral ecologies of this group. The results suggest that most squalomorphs originated in neritic environments from where they colonized deep waters on several independent occasions during the Late Jurassic and Early Cretaceous, predating most of the previous estimates of the timing of this event. The colonization of the deep sea took place via the benthic zone, in contrast to the view that an intermediate mesopelagic stage occurred during this ecological transition. Finally, the analyses accounting for uncertainty of the presence of bioluminescence strongly support that this trait evolved only once among sharks in a bathydemersal ancestor. This study reveals that shark bioluminescence evolved in a complex scenario that combines elements of several previous proposals, and enriches our perspective on the sequence of events that characterized the vertebrate conquest of the deep sea.     

Phylogenetic Relationships among Deep-Sea and Chemosynthetic Sea Anemones: Actinoscyphiidae and Actinostolidae (Actiniaria: Mesomyaria)

Sea anemones (Cnidaria, Actiniaria) are present in all marine ecosystems, including chemosynthetic environments. The high level of endemicity of sea anemones in chemosynthetic environments and the taxonomic confusion in many of the groups to which these animals belong makes their systematic relationships obscure. We use five molecular markers to explore the phylogenetic relationships of the superfamily Mesomyaria, which includes most of the species that live in chemosynthetic, deep-sea, and polar sea habitats and to test the monophyly of the recently defined clades Actinostolina and Chemosynthina. We found that sea anemones of chemosynthetic environments derive from at least two different lineages: one lineage including acontiate deep-sea taxa and the other primarily encompassing shallow-water taxa.     

Recently discovered landlocked basins in Indonesia reveal high habitat diversity in anchialine systems

In this article, the variability of physical settings of anchialine systems in Indonesia is discussed together with the consequences these settings have for the environment and biota within the systems. Exploration in two karstic areas (Berau, East Kalimantan and Raja Ampat, West Papua) has resulted in the discovery of 20 previously unknown anchialine systems in Indonesia. Based on parameters such as bathymetry, size, coastline, salinity, water temperature, pH, degree of connection to the sea, and the presence-absence of selected key taxa we distinguish three types of (non-cave) anchialine systems in the Indo-Pacific: (1) Marine lakes with large and deep basins containing brackish to almost fully marine waters. Marine lakes show a range in the degree of connection to the sea with the result that the higher the connection the more the lake resembles a lagoon in both water chemistry and biota, while the more isolated lakes have brackish water and contain unique species that are rarely found in the adjacent sea. (2) Anchialine pools with small and shallow basins containing brackish water and low diversity of macrofauna. (3) Blue pools in chasms that contain water with a clear halocline and are possibly connected to anchialine caves. Study of the many unique features of anchialine systems will enhance our understanding of the physical and ecological processes responsible for diversification in tropical shallow marine environments.     

Facies and synsedimentary tectonics on a Badenian carbonate platform in the southern Vienna Basin (Austria, Central Paratethys)

The Mannersdorf quarries at the northeastern edge of the Leitha Mountains (Lower Austria) preserve a record of pre-, syn- and post-tectonical phases of a Badenian carbonate platform in the Vienna Basin. The pre-tectonic phase is reported by a marine transgression with the development of a coastal slope scree and subsequent prograding of a Gilbert-type fan delta, overlain by very heterogeneous corallinacean limestones. A fault divides the study area into two independent tectonic blocks, which have been logged and subjected to detailed investigation and sampling. The corallinacean limestones of the first block indicate shallow-water environments (i.e., seagrass meadows) and gradual transitions from shallower to deeper environments, while the second block shows an unconformity, which is linked to a rapid facies change from relatively deeper environments (i.e., indicated by the abundance of in situ Pholadomya) to shallow waters (indicated by corals). Contrary to coral-bearing limestones of the same age at the southwestern part of the Leitha Mountains, corals are generally rare in the limestones of the Mannersdorf quarries, which represent mostly deeper environments with conspicuous differences in faunal associations. The onlap of limestones on a tectonic-caused flexure indicates syn-tectonical movements. Paleostress analyses verify a normal-fault reactivated as a dextral strike-slip fault. The temporal character of this fault is indicated by a post-tectonical phase with a marine transgression, a burial of the fault and neptunian dyke development.     

Diversity of Saint Helena Island and zoogeography of zoantharians in the Atlantic Ocean: Jigsaw falling into place

Diversity surveys in isolated sites, such as oceanic islands, provide biogeographic data that can improve our analyses and knowledge of evolutionary processes in the oceans. Zoantharians (Cnidaria: Anthozoa) are common and widespread components of shallow-water reefs, but distributional analyses are scarce for this group. In this study, we collected Zoantharia specimens from around Saint Helena Island (STH) in the mid-Atlantic and identified species using external morphology and molecular data. Moreover, we compiled and analysed the most comprehensive distributional data for zoantharian species in the subtropical and tropical Atlantic Ocean to date. Our results show eight zoantharian species in STH, which includes seven new records for STH waters. Furthermore, all families and genera of the suborder Brachycnemina recorded are widespread in the Atlantic Ocean, including at least four amphi-Atlantic species. The Caribbean is the richness centre in the Atlantic Ocean for zoantharian species, a pattern similar to that observed for many other subtropical/tropical marine taxa. However, Zoantharia may have a lower endemism rate in some areas than other common reef animals, for example zooxanthellate scleractinian corals and reef fishes. Moreover, zoantharian species have a more extensive distribution than close-related taxa such as zooxanthellate scleractinian corals and hydrocorals in the Atlantic Ocean.     

Threatened Corals Provide Underexplored Microbial Habitats

Contemporary in-depth sequencing of environmental samples has provided novel insights into microbial community structures, revealing that their diversity had been previously underestimated. Communities in marine environments are commonly composed of a few dominant taxa and a high number of taxonomically diverse, low-abundance organisms. However, studying the roles and genomic information of these “rare” organisms remains challenging, because little is known about their ecological niches and the environmental conditions to which they respond. Given the current threat to coral reef ecosystems, we investigated the potential of corals to provide highly specialized habitats for bacterial taxa including those that are rarely detected or absent in surrounding reef waters. The analysis of more than 350,000 small subunit ribosomal RNA (16S rRNA) sequence tags and almost 2,000 nearly full-length 16S rRNA gene sequences revealed that rare seawater biosphere members are highly abundant or even dominant in diverse Caribbean corals. Closely related corals (in the same genus/family) harbored similar bacterial communities. At higher taxonomic levels, however, the similarities of these communities did not correlate with the phylogenetic relationships among corals, opening novel questions about the evolutionary stability of coral-microbial associations. Large proportions of OTUs (28.7–49.1%) were unique to the coral species of origin. Analysis of the most dominant ribotypes suggests that many uncovered bacterial taxa exist in coral habitats and await future exploration. Our results indicate that coral species, and by extension other animal hosts, act as specialized habitats of otherwise rare microbes in marine ecosystems. Here, deep sequencing provided insights into coral microbiota at an unparalleled resolution and revealed that corals harbor many bacterial taxa previously not known. Given that two of the coral species investigated are listed as threatened under the U.S. Endangered Species Act, our results add an important microbial diversity-based perspective to the significance of conserving coral reefs.     

An ecologically mixed brachiopod fauna from Changhsingian deep-water basin of South China: consequence of end-Permian global warming

The Late Permian Shaiwa Group of the Ziyun area of Guizhou, South China is a deep-water facies succession characterized by deep-water assemblages of pelagic radiolarians, foraminifers, bivalves, ammonoids and brachiopods. Here we report 20 brachiopod species in 18 genera from the uppermost Shaiwa Group. This brachiopod fauna is latest Changhsingian in age and dominated by productides. The palaeoecologic and taphonomic analysis reveals that the brachiopod fauna is preserved in situ. The attachment modes and substratum preference demonstrate that the Shaiwa brachiopod fauna comprises admixed elements of deep-water and shallow-water assemblages. The presence of the shallow-water brachiopods in the Shaiwa faunas indicates the involuntary settlement of shallow-water brachiopods. The stressed ecologic pressure, triggered by warming surface waters, restricted ecospace and short food sources, may have forced some shallow-water elements to move to hospitable deep-water settings and others to modify their habiting behaviours and exploit new ecospace in deep-water environments. We infer that the end-Permian global warming and subsequent transgression event may have accounted for the stressed environmental pressure in the shallow-water communities prior to the end-Permian mass extinction.     

Major adaptive radiation in neritopsine gastropods estimated from 28S rRNA sequences and fossil records

A well-supported phylogeny of the Neritopsina, a gastropod superorder archaic in origin, radiated ecologically and diverse in morphology, is reconstructed based on partial 28S rRNA sequences. The result (Neritopsidae (Hydrocenidae (Helicinidae + Neritiliidae) (Neritidae + Phenacolepadidae))) is highly congruent with the fossil records and the character distribution of reproductive tracts in extant taxa. We suggest that the Neritopsina originated in subtidal shallow waters, invaded the land and became fully terrestrial at least three times in different clades, by the extinct Dawsonellidae in the Late Palaeozoic and by the Helicinidae and Hydrocenidae in the Mesozoic. Invasion of fresh- and brackish waters is prevalent among the Neritopsina as the Jurassic and freshwater ancestory is most probable for helicinids. The Phenacolepadidae, a group exclusively inhabiting dysoxic environments, colonized deep-sea hydrothermal vents and seeps in the Late Cretaceous or Early Cenozoic. Submarine caves have served as refuges for the archaic Neritopsidae since the Early to Middle Cenozoic, and the marine neritopsine slug Titiscania represents a highly specialized but relatively recent offshoot of this family. The Neritiliidae is another clade to be found utilizing submarine caves as shelter by the Oligocene; once adapted to the completely dark environment, but some neritiliids have immigrated to surface freshwater habitats.     

Phylogenetic measures reveal eco-evolutionary drivers of biodiversity along a depth gradient

Energy and environmental stability are positively correlated with species richness along broad-scale spatial gradients in terrestrial ecosystems, so their relative importance in generating and preserving diversity cannot be readily disentangled. This study seeks to exploit the negative correlation between energy and stability along the oceanic depth gradient to better understand their relative contribution in shaping broadscale biodiversity patterns. We develop a conceptual framework by simulating speciation and extinction along energy and stability gradients to generate expected patterns of biodiversity for a suite of complementary phylogenetic diversity metrics. Using a time-calibrated molecular phylogeny for New Zealand marine ray-finned fishes and a replicated community ecological sampling design, we then modelled these metrics along large-scale depth and latitude gradients. Our results indicate that energy-rich shallow waters may be an engine of diversity for percomorphs, but also suggest that recent speciation occurs in ancient fish lineages in the deep sea, hence questioning the role of energy as a key driver of speciation. Despite potentially facing high extinction early in their evolution, ancient phylogenetic lineages specialized for the deep-sea were likely preserved by environmental stability during the Cenozoic. Furthermore, intermediate depths might be a ‘museum’ (or zone of overlap) for distinct lineages that occur predominantly in either shallow or deep-sea waters. These intermediate depths (500–900 m) may form a ‘phylogenetic diversity bank’, perhaps providing a refuge during ancient (Mesozoic) extreme anoxic events affecting the deep sea and more recent (Pliocene–Pleistocene) climatic events occurring in shallow ecosystems. Finally, the phylogenetic structures observed in fish communities at intermediate depths suggest other processes might restrict the co-occurrence of closely related species. Overall, by combining a conceptual framework with models of empirical phylogenetic diversity patterns, our study paves the way for understanding the determinants of biodiversity across the largest habitat on earth.     

Diversity of Tanaidacea (Crustacea: Peracarida) in the World's Oceans – How Far Have We Come?

Tanaidaceans are small peracarid crustaceans which occur in all marine habitats, over the full range of depths, and rarely into fresh waters. Yet they have no obligate dispersive phase in their life-cycle. Populations are thus inevitably isolated, and allopatric speciation and high regional diversity are inevitable; cosmopolitan distributions are considered to be unlikely or non-existent. Options for passive dispersion are discussed. Tanaidaceans appear to have first evolved in shallow waters, the region of greatest diversification of the Apseudomorpha and some tanaidomorph families, while in deeper waters the apseudomorphs have subsequently evolved two or three distinct phyletic lines. The Neotanaidomorpha has evolved separately and diversified globally in deep waters, and the Tanaidomorpha has undergone the greatest evolution, diversification and adaptation, to the point where some of the deep-water taxa are recolonizing shallow waters. Analysis of their geographic distribution shows some level of regional isolation, but suffers from inclusion of polyphyletic taxa and a general lack of data, particularly for deep waters. It is concluded that the diversity of the tanaidomorphs in deeper waters and in certain ocean regions remains to be discovered; that the smaller taxa are largely understudied; and that numerous cryptic species remain to be distinguished. Thus the number of species currently recognized is likely to be an order of magnitude too low, and globally the Tanaidacea potentially rival the Amphipoda and Isopoda in diversity.     

The Halomonhystera disjuncta population is homogeneous across the Håkon Mosby mud volcano (Barents Sea) but is genetically differentiated from its shallow‐water relatives

The deep sea has a high biodiversity and a characteristic bathyal fauna. Earlier evidence suggested that at least some shallow‐water species invaded the ecosystem followed by radiation leading to endemic deep‐sea lineages with a genetic and/or morphological similarity to their shallow‐water counterparts. The nematode Halomonhystera disjuncta has been reported from shallow‐water habitats and the deep sea [Håkon Mosby mud volcano (HMMV)], but the morphological features and the phylogenetic relationships between deep‐sea and shallow‐water representatives remain largely unknown. Furthermore, nothing is known about the genetic structure of the H. disjuncta population within the HMMV. This study is the first integrative approach in which the morphological and phylogenetic relationships between a deep‐sea and shallow‐water free‐living nematode species are investigated. To elucidate the phylogenetic relationships, we analysed the mitochondrial gene Cytochrome oxidase c subunit I (COI) and three nuclear ribosomal genes (Internal Transcribed Spacer region, 18S and the D2D3 region of 28S). Our results show that deep‐sea nematodes comprise an endemic lineage compared to the shallow‐water representatives with different morphometric features. COI genetic divergence between the deep‐sea and shallow‐water specimens ranges between 19.1% and 25.2%. Taking these findings into account, we conclude that the deep‐sea form is a new species. amova revealed no genetic structure across the HMMV, suggesting that nematodes are able to disperse efficiently in the mud volcano.