Phylogeny of Comatulidae (Echinodermata: Crinoidea: Comatulida): a new classification and an assessment of morphological characters for crinoid taxonomy.

Comatulidae Fleming, 1828 (previously, and incorrectly, Comasteridae A.H. Clark, 1908a), is a group of feather star crinoids currently divided into four accepted subfamilies, 21 genera and approximately 95 nominal species. Comatulidae is the most commonly-encountered and species-rich crinoid group on shallow tropical coral reefs, particularly in the Indo-western Pacific region (IWP). We conducted a molecular phylogenetic analysis of the group with concatenated data from up to seven genes for 43 nominal species spanning 17 genera and all subfamilies. Basal nodes returned low support, but maximum likelihood, maximum parsimony, and Bayesian analyses were largely congruent, permitting an evaluation of current taxonomy and analysis of morphological character transformations. Two of the four current subfamilies were paraphyletic, whereas 15 of the 17 included genera returned as monophyletic. We provide a new classification with two subfamilies, Comatulinae and Comatellinae n. subfamily Summers, Messing, & Rouse, the former containing five tribes. We revised membership of analyzed genera to make them all clades and erected Anneissia n. gen. Summers, Messing, & Rouse. Transformation analyses for morphological features generally used in feather star classification (e.g., ray branching patterns, articulations) and those specifically for Comatulidae (e.g., comb pinnule form, mouth placement) were labile with considerable homoplasy. These traditional characters, in combination, allow for generic diagnoses, but in most cases we did not recover apomorphies for subfamilies, tribes, and genera. New morphological characters that will be informative for crinoid taxonomy and identification are still needed. DNA sequence data currently provides the most reliable method of identification to the species-level for many taxa of Comatulidae.

Fixed, free, and fixed: the fickle phylogeny of extant Crinoidea (Echinodermata) and their Permian-Triassic origin.

Although the status of Crinoidea (sea lilies and featherstars) as sister group to all other living echinoderms is well-established, relationships among crinoids, particularly extant forms, are debated. All living species are currently placed in Articulata, which is generally accepted as the only crinoid group to survive the Permian-Triassic extinction event. Recent classifications have recognized five major extant taxa: Isocrinida, Hyocrinida, Bourgueticrinina, Comatulidina and Cyrtocrinida, plus several smaller groups with uncertain taxonomic status, e.g., Guillecrinus, Proisocrinus and Caledonicrinus. Here we infer the phylogeny of extant Crinoidea using three mitochondrial genes and two nuclear genes from 59 crinoid terminals that span the majority of extant crinoid diversity. Although there is poor support for some of the more basal nodes, and some tree topologies varied with the data used and mode of analysis, we obtain several robust results. Cyrtocrinida, Hyocrinida, Isocrinida are all recovered as clades, but two stalked crinoid groups, Bourgueticrinina and Guillecrinina, nest among the featherstars, lending support to an argument that they are paedomorphic forms. Hence, they are reduced to families within Comatulida. Proisocrinus is clearly shown to be part of Isocrinida, and Caledonicrinus may not be a bourgueticrinid. Among comatulids, tree topologies show little congruence with current taxonomy, indicating that much systematic revision is required. Relaxed molecular clock analyses with eight fossil calibration points recover Articulata with a median date to the most recent common ancestor at 231-252mya in the Middle to Upper Triassic. These analyses tend to support the hypothesis that the group is a radiation from a small clade that passed through the Permian-Triassic extinction event rather than several lineages that survived. Our tree topologies show various scenarios for the evolution of stalks and cirri in Articulata, so it is clear that further data and taxon sampling are needed to recover a more robust phylogeny of the group.

Post-Paleozoic crinoid radiation in response to benthic predation preceded the Mesozoic marine revolution.

It has been argued that increases in predation over geological time should result in increases in defensive adaptations in prey taxa. Recent in situ and laboratory observations indicate that cidaroid sea urchins feed on live stalked crinoids, leaving distinct bite marks on their skeletal elements. Similar bite marks on fossil crinoids from Poland strongly suggest that these animals have been subject to echinoid predation since the Triassic. Following their near-demise during the end-Permian extinction, crinoids underwent a major evolutionary radiation during the Middle-Late Triassic that produced distinct morphological and behavioral novelties, particularly motile taxa that contrasted strongly with the predominantly sessile Paleozoic crinoid faunas. We suggest that the appearance and subsequent evolutionary success of motile crinoids were related to benthic predation by post-Paleozoic echinoids with their stronger and more active feeding apparatus and that, in the case of crinoids, the predation-driven Mesozoic marine revolution started earlier than in other groups, perhaps soon after the end-Permian extinction.

A revision of the genus Atelecrinus PH Carpenter (Echinodermata: Crinoidea).

The unusual bathyal comatulid crinoid genus Atelecrinus is widespread in the Atlantic and tropical Pacific Oceans and currently includes three recognized species. A re-assessment based on examination of new and existing specimens requires establishment of two new genera and five new species, and returns three junior synonyms to species-level status. Paratelecrinus is erected to accommodate Atelecrinus wyvilli PH Carpenter, A. conifer AH Clark, A. cubensis PH Carpenter, P. orthotriremis, new species, P. amenouzume new species, P. laticonulus new species and P. telo new species. Adelatelecrinus is erected to accommodate Atelecrinus sulcatus AH Clark and Adelatelecrinus vallatus new species. Atelecrinus retains A. balanoides PH Carpenter and A. helgae AH Clark, which restricts the genus to the Atlantic. In both Paratelecrinus and Adelatelecrinus, the basals articulate with the centrodorsal via ligament bundles anchored in deep ringlike interradial pits that project into the centrodorsal cavity, whereas in Atelecrinus the centrodorsal rim has shallow interradial concavities and attaches to the basals via a tight junction with no obvious ligament bundles. The spoon-shaped aboral fossa in the basals of Paratelecrinus appears to be unique among articulate crinoids and differs from the smooth fossa found in both Atelecrinus and Adelatelecrinus. New material extends the range of the family to the Indian Ocean. A few species are now known from enough specimens to identify some ontogenetic and distributional variations. Proximal ray morphology varies substantially with size in P. cubensis and P. orthotriremis. A. balanoides generally occurs in deeper water in the Lesser Antilles than in the Bahamas and Strait of Florida, while P. orthotriremis occurs in shallower water in the Lesser Antilles and deeper in the Bahamas.

Phylogeny and taxonomy of Himerometroidea (Echinodermata: Crinoidea).

Himerometroidea is a clade of chiefly shallow-water, tropical, feather-star crinoids that is currently divided, based on morphology, into four families comprising 119 extant species in 31 genera. Our molecular phylogenetic results, based on three mitochondrial (CO1, 16S, CytB) and two nuclear (ITS and 28S) markers for 55 accepted species in 23 of the extant genera, allow for six clades within Himerometroidea to be given family ranks. Maximum parsimony, maximum likelihood, and Bayesian inference analyses recovered largely congruent topologies with varying nodal support. A new classification revises generic placements among five families: Himerometridae, Colobometridae, and Mariametridae, all retained, and Pontiometridae and Stephanometridae both resurrected. Zygometridae is no longer accepted, since the type genus of the family, Zygometra, falls within Himerometridae. Catoptometra was recovered as a sister clade with respect to those treated herein as a new family, Catoptometridae. Two genera, Iconometra and Analcidometra, are retained within Himerometroidea but without family assignments pending further assessment of their positions. Currently published diagnostic and descriptive morphological features are noted where possible to support taxonomic names in the recovered phylogeny, although more examination of morphology is needed to identify synapomorphies and designate taxon names formally.

Light and vision in the deep-sea benthos: I. Bioluminescence at 500-1000 m depth in the Bahamian islands.

Bioluminescence is common and well studied in mesopelagic species. However, the extent of bioluminescence in benthic sites of similar depths is far less studied, although the relatively large eyes of benthic fish, crustaceans and cephalopods at bathyal depths suggest the presence of significant biogenic light. Using the Johnson-Sea-Link submersible, we collected numerous species of cnidarians, echinoderms, crustaceans, cephalopods and sponges, as well as one annelid from three sites in the northern Bahamas (500-1000 m depth). Using mechanical and chemical stimulation, we tested the collected species for light emission, and photographed and measured the spectra of the emitted light. In addition, in situ intensified video and still photos were taken of different benthic habitats. Surprisingly, bioluminescence in benthic animals at these sites was far less common than in mesopelagic animals from similar depths, with less than 20% of the collected species emitting light. Bioluminescent taxa comprised two species of anemone (Actinaria), a new genus and species of flabellate Parazoanthidae (formerly Gerardia sp.) (Zoanthidea), three sea pens (Pennatulacea), three bamboo corals (Alcyonacea), the chrysogorgiid coral Chrysogorgia desbonni (Alcyonacea), the caridean shrimp Parapandalus sp. and Heterocarpus ensifer (Decapoda), two holothuroids (Elasipodida and Aspidochirota) and the ophiuroid Ophiochiton ternispinus (Ophiurida). Except for the ophiuroid and the two shrimp, which emitted blue light (peak wavelengths 470 and 455 nm), all the species produced greener light than that measured in most mesopelagic taxa, with the emissions of the pennatulaceans being strongly shifted towards longer wavelengths. In situ observations suggested that bioluminescence associated with these sites was due primarily to light emitted by bioluminescent planktonic species as they struck filter feeders that extended into the water column.

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.

A revision of the unusual feather star genus Atopocrinus with a description of a new species (Echinodermata: Crinoidea).

Atopocrinus ojii new species, is described from deep water off Japan. The genus was previously monotypic and known only from the holotype of A. sibogae A. H. Clark, 1912, from Indonesia. Examination of all known specimens of Atopocrinus has added the following genus-level characters: cirrus sockets arising as tubes along the adoral surface of the centrodorsal, and faint, superficial, suture-like lines that cross the interradial ridges aboral to the centrodorsal/basal suture. The new species exhibits additional novel characters, some of which may be unique (e.g., deep pores around cirrus sockets) and others that may be shared with the incomplete holotype of A. sibogae (e.g., hollow pinnulars). Confusion about the structure of basal ossicles versus basal rays in Atopocrinus and other extant feather stars prompted inclusion of a review of the structure and development of these skeletal components. Video of a live specimen attributed to A. sibogae revealed that the distal half of the arms lack pinnules. The new species also shares several characters in common with Jurassic Spinimetra chesnieri Hess Thuy, 2017.

The Shallow-water Crinoid Fauna of Lakshadweep Atolls, North-western Indian Ocean.

A biodiversity survey carried out from 2016 to 2018 by the Department of Science and Technology in the Lakshadweep Atolls, India, recorded six species of shallow-water feather stars new to the archipelago (Comanthus wahlbergii, Comaster schlegelii, Himerometra robustipinna, Dichrometra palmata, Stephanometra indica, and Phanogenia typica). Himerometra sol A.H. Clark, 1912, previously known only from the Maldive Islands, is synonymized under Himerometra robustipinna (Carpenter, 1881). This study brings the total number of shallow-water crinoids recorded from Lakshadweep to ten species. Of the four species collected previously from the archipelago, only Comatella nigra was found in this survey. Of those not collected, Comatella stelligera and Oligometra serripinna are widespread in the Indo-western Pacific region, whereas Heterometra compta is known only from Lakshadweep, and its validity remains uncertain. The known shallow-water crinoid fauna of the archipelago is substantially less diverse than that of the adjacent and environmentally similar Maldive Islands, emphasizing the need for additional research in this island group, in particular, to determine whether the differences are actual or not, and whether they are based on natural conditions versus anthropogenic impacts.

The genera and species of Comatulidae (Comatulida: Crinoidea): taxonomic revisions and a molecular and morphological guide.

We provide a guide for identification of Comatulidae, a family of crinoid echinoderms, incorporating morphological and molecular evidence. A non-dichotomous key for all genera is included, as well as photographs of species most likely to be encountered in the tropical western Pacific Ocean. Based on sequencing of cytochrome oxidase subunit I (COI), and other genes when necessary, we identified four cases where taxonomic revision was needed. We synonymized Comaster nobilis under Comaster schlegelii, and Clarkcomanthus exilis under Clarkcomanthus comanthipinnus, and re-described Clarkcomanthus albinotus. We also showed the variation of COI sequences within and among species, which varies from lineage to lineage. In some cases, specimens with obvious morphological disparity possessed very little intraspecific molecular diversity (<1%). In others, specimens with nearly identical external appearances exhibited quite divergent COI sequences (up to 6%). These results, combined with the non-dichotomous key herein, offer guidelines for identification and discussion of existing and new species of Comatulidae.

Porphyrocrinus daniellalevyae n. sp. (Echinodermata: Crinoidea), a sea lily from the tropical western Atlantic with a unique crown pattern.

Porphyrocrinus daniellalevyae, new species, is described from irregular, hard-substrate, deep island slope habitats in the tropical western Atlantic Ocean (northwestern Bahamas and, probably, Honduras). It represents the first record of the genus from the western Atlantic, and is the first crinoid, living or fossil, known to gradually increase its number of rays with increasing size and, ostensibly, growth. The four known specimens have 8, 12, 14 and 15 radial ossicles, which give rise to undivided arms. The method of augmentation is unknown, although the possibility of autotomy of one and regrowth of two in its place is discussed. Augmentation of radial number may also account for the absence of an aboral nerve ring associated with the radial ring, unlike the arrangement in almost all living crinoids in which the nervous system has been examined. Ligamentary articulations (trifascial synarthries) exhibit fulcral ridge-and groove architecture between arm ossicle pairs that remains to be described for other members of the genus. The species also exhibits a filamentous arm tip lacking pinnules that has only been described for one other species of Porphyrocrinus.