An underwater imagery identification guide for shallow, mesophotic and deep-sea benthos in Maldives.

Background: During the 2022 Nekton Maldives Mission, we deployed a variety of platforms (snorkelling, remotely-operated vehicles and manned submersibles) to conduct video surveys of the biodiversity and composition of shallow (< 30 m), mesophotic (30-150 m) and deep-sea (> 150 m) benthos found in the Maldives' central and southern atolls. In total, ~ 80 hrs of stereo-video footage were collected during the benthic transect surveys, which were subsequently processed using annotation software in order to evaluate benthic biodiversity and community composition. Here, we present a photographic guide for the visual, in situ identification of reef benthos encountered, including corals, sponges and other invertebrates that inhabit Maldives' nearshore habitats. We hope that this identification guide will aid future imagery-based surveys or observations of organisms during fieldwork. New information: A total of 283 morphotypes were identified, including those belonging to Octocorallia (61), Scleractinia (57), Porifera (38), Asteroidea (22), Antipatharia (15), Decapoda (13), Hydrozoa (12), Holothuroidea (10), Actiniaria (9), Echinoidea (8), Annelida (6), Chlorophyta (5), Gastropoda (4), Bivalvia (4), Ascidiacea (3), Crinoidea (3), Bryozoa (2), Cyanobacteria (2), Zoantharia (2), Cephalopoda (1), Ceriantharia (1), Corallimorpharia (1), Ctenophora (1), Ophiuroidea (1), Rhodophyta (1) and to an unknown category (1). Out of these, we identified 40 to species level, 120 to genus, 47 to family, 14 to order and suborder, 58 to class and subclass, two to phylum and one was of unknown phylum. This represents the first attempt to catalogue the mesophotic and deep-sea benthic megafaunal diversity in the Maldives using underwater imagery.

New Taxa and Occurrences of Mesophotic and Deep-sea Goniasteridae (Valvatida, Asteroidea) from Okinawa and adjacent regions.

Thirteen new records of Goniasteridae showing mesophotic occurrence are presented, including a new species of Pseudoceramaster doto n. sp. from southern Japan and a new genus and species, Churaumiastra hoshi nov. gen. nov. sp. which occurs in southern Japan, the Philippines and Western Australia.

Systematics of deep-sea starfish order Brisingida (Echinodermata: Asteroidea), with a revised classification and assessments of morphological characters.

Brisingida Fisher 1928 is one of the seven currently recognised starfish orders, and one of the least known taxa as being exclusive deep-sea inhabitants. Modern deep-sea expeditions revealed their common occurrences in various deep-sea settings including seamounts, basins and hydrothermal vent peripheral, underlining the necessity of clarifying their global diversity and phylogeny. In this study, we present a comprehensive molecular phylogeny of Brisingida which encompasses the highest taxonomic diversity to date. DNA sequences (COI, 16S, 12S and 28S) were obtained from 225 specimens collected in the global ocean, identified as 58 species spanning 15 of the 17 extant genera. Phylogenetic relationship was inferred using both maximum likelihood and Bayesian inference methods, revealing polyphyletic families and genera and indicating nonnegligible bias in prior morphology-based systematics. Based on the new phylogeny, a novel classification of the order, consisting of 5 families and 17 genera, is proposed. Families Odinellidae, Brisingasteridae and Novodiniidae (sensu Clark and Mah, 2001) were resurrected to encompass the genera Odinella, Brisingaster and Novodinia. Brisingidae and Freyellidae were revised to include 11 and 3 genera, respectively. A new genus and species, two new subgenera and seven new combinations are described and a key to each genus and family is provided. Transformations of morphological traits were evaluated under the present phylogenetic hypothesis. A series of paedomorphic characters were found in many genera and species, which led to a high degree of homoplasy across phylogenetically distant groups. Our results provide new insights in the phylogeny and ontogeny of the order, and highlight the necessity to evaluate character convergence under sound phylogenetic hypothesis.

New Genera, Species, and observations on the biology of Antarctic Valvatida (Asteroidea).

Assessment of Antarctic taxonomic diversity for asteroids and other taxa for poorly studied or unknown regions, such as the deep-sea, will be important for our understanding of these understudied habitats. Eleven new species and a new genus are described from three families (Asterinidae, Goniasteridae, Solasteridae) within the Valvatida, nearly all of which were collected from deep-sea settings below 1000 m by the US Antarctic Research Program in the 1960s. A new subfamily, the Kampylasterinae subfam. nov. is designated for Kampylaster and Astrotholus nov. gen. which were supported as sister taxa on a monophyletic clade within the Asterinidae. Astrotholus nov. gen. is described to accommodate "Anseropoda" antarctica and four new bathyal and abyssal species, which are a significant morphological divergence from the typological definition of Anseropoda. New species of the goniasterid Notioceramus and the solasterid Paralophaster are also described from bathyal depths (2000-3000 m). Paralophaster ferax n. sp. is among the deepest asteroids known to brood, is the first brooding species within Paralophaster and the second species in the Solasteridae known to brood. Following examination of the type and molecular data, Lophaster densus is found to be included within Paralophaster. A review of Antarctic Lophaster species shows additional specimens of Lophaster abbreviatus which support it as a distinct species from Lophaster stellans. New occurrence data for bathyal Antarctic Asteroidea as well as unusual-gut content observations of shallower-water species are also included.

Eyes, Vision, and Bioluminescence in Deep-Sea Brisingid Sea Stars.

AbstractSea stars are a major component of the megabenthos in most marine habitats, including those within the deep sea. Being radially symmetric, sea stars have sensory structures that are evenly distributed along the arms, with a compound eye located on each arm tip of most examined species. Surprisingly, eyes with a spatial resolution that rivals the highest acuity known among sea stars so far were recently found in Novodinia americana, a member of the deep-sea sea star order Brisingida. Here, we examined 21 species across 11 brisingid genera for the presence of eyes; where eyes were present, we used morphological characteristics to evaluate spatial resolution and sensitivity. This study found that eyes were present within 43% of the examined species. These brisingid eyes were relatively large compared to those of other deep-sea sea stars, with a high number of densely packed ommatidia. One of the examined species, Brisingaster robillardi, had more than 600 ommatidia per eye, which is the highest number of ommatidia found in any sea star eye so far. Combined, the results indicate that brisingid eyes are adapted for spatial resolution over sensitivity. Together with results showing that many brisingids are bioluminescent, this relatively high spatial resolution suggests that the group may use their eyes to support visually guided intraspecific communication based on bioluminescent signals. Phylogenetic analysis indicated that the common ancestor of brisingids had eyes (P = 0.72) and that eyes were lost once within the clade.

New Genera, Species and Occurrences of Deep-Sea Asteroidea (Valvatacea, Forcipulatacea, Echinodermata) collected from the North Pacific Ocean by the CAPSTONE Expedition.

Between 2015 and 2018, the NOAA ship Okeanos Explorer embarked on the CAPSTONE expedition, that collected video and specimen data from the Hawaiian Islands region and other American territories in the Central and South Pacific Ocean. Ultimately 20 species, 12 of which are newly described, from 14 genera in 3 families are described herein with one further undescribed species of Pythonaster currently in preparation. Okeanos Explorer collected a total of 17 sea star specimens, of which 9 represent new species and 4 new genera. Additionally, one species imaged by Okeanos Explorer during the campaign was discovered in a museum collection is described herein. Supplementary collections from the tropical Pacific by the R/V Nautilus resulted in a further 11th newly described species which had been originally observed by Okeanos Explorer. A total of 21 specimens were involved as part of this effort. Heligmaster n. gen. is the first new genus of the rarely encountered deep-sea Myxasteridae since Fisher (1906). The Goniasteridae, particularly the corallivorous subfamily Hippasterinae were most heavily represented in the collected material. Video imagery was extensive and provided new occurrence data for rarely seen species as well as several new records that likely represent undescribed species. Many of the new species collected were observed preying on of deep-sea octocorals and sponges.

A new genus and species of Goniasteridae, Peedeeaster sandersoni, and the first occurrence of Sclerasterias (Asteriidae) from the Cretaceous Peedee Formation of North Carolina.

Two new fossil asteroids are described from the Maastrichtian (Cretaceous) Peedee Formation in North Carolina. The first, a member of the Asteriidae, is identified as Sclerasterias sp. based primarily on the superomarginals, spination, and presence of transverse ribs on the distalmost armtip. This is the first occurrence of Sclerasterias in the Cretaceous of North America. The second asteroid taxon is based on a relatively well-preserved series of specimens described as a new genus and species, Peedeeaster sandersoni (Goniasteridae) which displays a pentagonal body shape similar to those of modern Peltaster and Sphaeriodiscus and is also comparable to such Cretaceous goniasterids such as Metopaster.

Megafauna of the German exploration licence area for seafloor massive sulphides along the Central and South East Indian Ridge (Indian Ocean).

BACKGROUND: The growing interest in mineral resources of the deep sea, such as seafloor massive sulphide deposits, has led to an increasing number of exploration licences issued by the International Seabed Authority. In the Indian Ocean, four licence areas exist, resulting in an increasing number of new hydrothermal vent fields and the discovery of new species. Most studies focus on active venting areas including their ecology, but the non-vent megafauna of the Central Indian Ridge and South East Indian Ridge remains poorly known.In the framework of the Indian Ocean Exploration project in the German license area for seafloor massive sulphides, baseline imagery and sampling surveys were conducted yearly during research expeditions from 2013 to 2018, using video sledges and Remotely Operated Vehicles. NEW INFORMATION: This is the first report of an imagery collection of megafauna from the southern Central Indian- and South East Indian Ridge, reporting the taxonomic richness and their distribution. A total of 218 taxa were recorded and identified, based on imagery, with additional morphological and molecular confirmed identifications of 20 taxa from 89 sampled specimens. The compiled fauna catalogue is a synthesis of megafauna occurrences aiming at a consistent morphological identification of taxa and showing their regional distribution. The imagery data were collected during multiple research cruises in different exploration clusters of the German licence area, located 500 km north of the Rodriguez Triple Junction along the Central Indian Ridge and 500 km southeast of it along the Southeast Indian Ridge.

Reef benthos of Seychelles - A field guide.

BACKGROUND: During the 2019 First Descent: Seychelles Expedition, shallow and deep reef ecosystems of the Seychelles Outer Islands were studied by deploying a variety of underwater technologies to survey their benthic flora and fauna. Submersibles, remotely operated vehicles (ROVs) and SCUBA diving teams used stereo-video camera systems to record benthic communities during transect surveys conducted at 10 m, 30 m, 60 m, 120 m, 250 m and 350 m depths. In total, ~ 45 h of video footage was collected during benthic transect surveys, which was subsequently processed using annotation software in order to assess reef biodiversity and community composition. Here, we present a photographic guide for the visual identification of the marine macrophytes, corals, sponges and other common invertebrates that inhabit Seychelles' reefs. It is hoped that the resulting guide will aid marine biologists, conservationists, managers, divers and naturalists with the coarse identification of organisms as seen in underwater footage or live in the field. NEW INFORMATION: A total of 184 morphotypes (= morphologically similar individuals) were identified belonging to Octocorallia (47), Porifera (35), Scleractinia (32), Asteroidea (19), Echinoidea (10), Actiniaria (9), Chlorophyta (8), Antipatharia (6), Hydrozoa (6), Holothuroidea (5), Mollusca (2), Rhodophyta (2), Tracheophyta (2), Annelida (1), Crinoidea (1), Ctenophora (1), Ochrophyta (1) and Zoantharia (1). Out of these, we identified one to phylum level, eight to class, 14 to order, 27 to family, 110 to genus and 24 to species. This represents the first attempt to catalogue the benthic diversity from shallow reefs and up to 350 m depth in Seychelles.

The East Pacific/South Pacific Boundary: New taxa and occurrences from Rapa Nui (Easter Island), New Caledonia and adjacent regions.

Recent expeditions to Rapa Nui (also known as Easter Island) and New Caledonia have revealed undescribed species from mesophotic and deeper depths. This includes three new species from Rapa Nui, Hacelia raaraa, Linckia profunda (Ophidiasteridae), Uokeaster ahi (Asterodiscididae) and two new species from New Caledonia, Astroglypha pyramidata n. gen. and Ophidiaster colossus (Ophidiasteridae). The new genus Astroglypha is described for A. pyramidata but the genus also includes the Atlantic Tamaria passiflora, which is reassigned herein. Pauliastra n. gen. is designated as a replacement for the homonym issue with Pauliella. New occurrences and synonymies are addressed for taxa related to New Caledonia, Rapa Nui and adjacent regions. A morphology based phylogenetic analysis agrees with prior work which placed Goniaster among the Asterodiscididae and posits biogeographic relationships among asterodiscidid genera. Implications for the Goniasteridae and placement of Goniaster among asterodiscidid genera are discussed. Biogeography and relationships among taxa from Rapa Nui and New Caledonia are reviewed. In situ observations from species observed from Rapa Nui are included.

The first complete mitochondrial genome of the Northern Pacific deep-sea goniasterid sea star Ceramaster japonicus (Sladen, 1889) determined using NGS-based shotgun sequencing.

The full mitogenome of an ethanol-preserved museum specimen of Ceramaster japonicus was determined using the NGS Illumina MiSeq platform. The specimen was collected from Tosa Bay, Japan, facing the Pacific Ocean (33.0781 N 134.0601 E), at 700 m depth in 2011. The mitogenome shows a typical metazoan genomic structure, with all of the 37 genes included in its 16,370 base-long mitogenome. We conducted phylogenetic analyses using a data set including 18 publicly available asteroids rooted against five ophiuroids as outgroups. The result confirms the position of C. japonicus in the order Valvatida. The complete mitogenome of C. japonicus reported here is the first reported for the family Goniasteridae Forbes, 1841.

New species, occurrence records and observations of predation by deep-sea Asteroidea (Echinodermata) from the North Atlantic by NOAA ship Okeanos Explorer.

Exploratory cruises by the NOAA Ship Okeanos Explorer have resulted in a substantial contribution in our understanding of deep-sea echinoderm biodiversity, biology, and ecology in the North Atlantic. This includes the description and in situ feeding observations of two, new corallivorous goniasterid species, Evoplosoma nizinskiae n. sp. and Sibogaster bathyheuretor n. sp. Significant in situ observations include a synchronous feeding event including multiple goniasterid asteroids and a cidaroid urchin on a large demosponge, providing new data for understanding echinoderm feeding behavior, including agonistic behavior, in deep-sea settings and new, in situ feeding observations for 28 deep-sea species including the myxasterid Pythonaster atlantidis, the korethrasterid Remaster palmatus and the poorly understood hippasterine goniasterids, Gilbertaster caribaea and Sthenaster emmae.

New species and occurrence records of Japanese Solasteridae and Ganeriidae including a new species of Paralophaster from the North Pacific with an overview of Hyalinothrix.

Japanese waters contain a highly diverse echinoderm fauna including many undiscovered species, especially in deeper-water settings. Molecular phylogenetic data has shown a close relationship between three established families of asteroids, the Asterinidae, the Solasteridae and the Ganeriidae, which are three groups that are well-represented in the region. We present two undescribed species of solasterids, Lophaster cactorum n. sp. and Paralophaster gomo n. sp., the latter represents a genus which had previously been known primarily from Antarctic settings. New occurrence records of the tropical deep-sea Pacific solasterid Rhipidaster vannipes Sladen, 1889 has led to synonymy of Xenorias polyctenius Fisher, 1913. Three undescribed species of Hyalinothrix, Hyalinothrix diversus n. sp., Hyaliothrix enoki n. sp. and Hyalinothrix virtrispinum n. sp. are described. Nepanthia grangei McKnight, 2001 is transferred to Hyalinothrix. A new subfamily, the Hyalinothricinae is established within the Asterinidae, including Tarachaster and Seriaster, which was formerly assigned to the Solasteridae. Knightaster is reassigned to the Asterinidae from the Ganeriidae.

Additive Manufacturing of Optical Quality Germania-Silica Glasses.

Direct ink writing (DIW) three-dimensional (3D) printing provides a revolutionary approach to fabricating components with gradients in material properties. Herein, we report a method for generating colloidal germania feedstock and germania-silica inks for the production of optical quality germania-silica (GeO2-SiO2) glasses by DIW, making available a new material composition for the development of multimaterial and functionally graded optical quality glasses and ceramics by additive manufacturing. Colloidal germania and silica particles are prepared by a base-catalyzed sol-gel method and converted to printable shear-thinning suspensions with desired viscoelastic properties for DIW. The volatile solvents are then evaporated, and the green bodies are calcined and sintered to produce transparent, crack-free glasses. Chemical and structural evolution of GeO2-SiO2 glasses is confirmed by nuclear magnetic resonance, X-ray diffraction, and Raman spectroscopy. UV-vis transmission and optical homogeneity measurements reveal comparable performance of the 3D printed GeO2-SiO2 glasses to glasses produced using conventional approaches and improved performance over 3D printed TiO2-SiO2 inks. Moreover, because GeO2-SiO2 inks are compatible with DIW technology, they offer exciting options for forming new materials with patterned compositions such as gradients in the refractive index that cannot be achieved with conventional manufacturing approaches.

A framework for the development of a global standardised marine taxon reference image database (SMarTaR-ID) to support image-based analyses.

Video and image data are regularly used in the field of benthic ecology to document biodiversity. However, their use is subject to a number of challenges, principally the identification of taxa within the images without associated physical specimens. The challenge of applying traditional taxonomic keys to the identification of fauna from images has led to the development of personal, group, or institution level reference image catalogues of operational taxonomic units (OTUs) or morphospecies. Lack of standardisation among these reference catalogues has led to problems with observer bias and the inability to combine datasets across studies. In addition, lack of a common reference standard is stifling efforts in the application of artificial intelligence to taxon identification. Using the North Atlantic deep sea as a case study, we propose a database structure to facilitate standardisation of morphospecies image catalogues between research groups and support future use in multiple front-end applications. We also propose a framework for coordination of international efforts to develop reference guides for the identification of marine species from images. The proposed structure maps to the Darwin Core standard to allow integration with existing databases. We suggest a management framework where high-level taxonomic groups are curated by a regional team, consisting of both end users and taxonomic experts. We identify a mechanism by which overall quality of data within a common reference guide could be raised over the next decade. Finally, we discuss the role of a common reference standard in advancing marine ecology and supporting sustainable use of this ecosystem.

Antarctic and Sub-Antarctic Asteroidea database.

The present dataset is a compilation of georeferenced occurrences of asteroids (Echinodermata: Asteroidea) in the Southern Ocean. Occurrence data south of 45°S latitude were mined from various sources together with information regarding the taxonomy, the sampling source and sampling sites when available. Records from 1872 to 2016 were thoroughly checked to ensure the quality of a dataset that reaches a total of 13,840 occurrences from 4,580 unique sampling events. Information regarding the reproductive strategy (brooders vs. broadcasters) of 63 species is also made available. This dataset represents the most exhaustive occurrence database on Antarctic and Sub-Antarctic asteroids.

New genera, species and occurrence records of Goniasteridae (Asteroidea; Echinodermata) from the Indian Ocean.

Modern goniasterids are the most numerous of living asteroids in terms of described genera and species and they have important ecological roles from shallow to deep-water marine habitats. Recent MNHN expeditions and historical collections in the USNM have resulted in the discovery of 18 new species, three new genera and multiple new occurrence records from the western Indian Ocean region including Madagascar, Glorioso and Mayotte islands, Walters Shoal, South Africa, and Somalia. This report provides the first significant contribution to knowledge of deep-sea Asteroidea from the Indian Ocean since the late 20th Century. Several deep-sea species, previously known from the North Pacific are now reported from the western Indian Ocean. Gut contents from Stellaster and Ogmaster indicate deposit feeding. Feeding modes of this and other deep-sea species are discussed. Comments are made on fossil members of included taxa. A checklist of Indian Ocean Goniasteridae is also included.

The phylogeny of extant starfish (Asteroidea: Echinodermata) including Xyloplax, based on comparative transcriptomics.

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.

Megafauna of the UKSRL exploration contract area and eastern Clarion-Clipperton Zone in the Pacific Ocean: Echinodermata.

BACKGROUND: There is growing interest in mining polymetallic nodules from the abyssal Clarion-Clipperton Zone (CCZ) in the tropical Pacific Ocean. Despite being the focus of environmental studies for decades, the benthic megafauna of the CCZ remain poorly known. In order to predict and manage the environmental impacts of mining in the CCZ, baseline knowledge of the megafauna is essential. The ABYSSLINE Project has conducted benthic biological baseline surveys in the UK Seabed Resources Ltd polymetallic-nodule exploration contract area (UK-1). Prior to these research cruises in 2013 and 2015, no biological studies had been done in this area of the eastern CCZ. NEW INFORMATION: Using a Remotely Operated Vehicle and Autonomous Underwater Vehicle, the megafauna within the UKSRL exploration contract area (UK-1) and at a site ~250 km east of the UK-1 area were surveyed, allowing us to make the first estimates of megafaunal morphospecies richness from the imagery collected. Here, we present an atlas of the abyssal echinoderm megafauna observed and collected during the ABYSSLINE cruises to the UK-1 polymetallic-nodule exploration contract area in the CCZ. There appear to be at least 62 distinct morphospecies (13 Asteroidea, 5 Crinoidea, 9 Echinoidea, 29 Holothuroidea and 6 Ophiuroidea) identified mostly by imagery but also using molecular barcoding for a limited number of animals that were collected. This atlas will aid the synthesis of megafaunal presence/absence data collected by contractors, scientists and other stakeholders undertaking work in the CCZ, ultimately helping to decipher the biogeography of the megafauna in this threatened habitat.