The Genus Teliocrinus (Crinoidea, Echinodermata): a Key Taxon Among Pentacrinid Stalked Crinoids
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Zoological Journal of the Linnean Society, 2009, 155, 22–39. With 10 figures The genus Teliocrinus (Crinoidea, Echinodermata): a key taxon among pentacrinid stalked crinoids MICHEL ROUX1, NADIA AMÉZIANE2* and MARC ELEAUME3 1UMR CNRS 5561 Biogéosciences, 22 rue de Bourgogne, 51350 Cormontreuil, France Downloaded from https://academic.oup.com/zoolinnean/article/155/1/22/2674285 by guest on 23 November 2020 2Département Milieux et Peuplements Aquatiques, USM 0401-UMR CNRS 5178, Muséum national d’Histoire naturelle, 55 rue Buffon, 75005 Paris, France 3Département Milieux et Peuplements Aquatiques, USM 0401-UMR CNRS 5178, Muséum national d’Histoire naturelle, 55 rue Buffon, 75005 Paris, France Received 24 March 2007; accepted for publication 25 July 2007 The main characters of the stalked crinoids of the family Pentacrinitidae attributed to the genus Teliocrinus are re-evaluated from a quantitative study of phenotype variation, new observations on arm and stalk articulations, and observation of ontogenetic trends. All of the specimens collected in the northern Indian Ocean belong to the same species, i.e. Teliocrinus springeri (Clark, 1909). However, two phenotypes living at different depths remain valid as subspecies: Teliocrinus springeri springeri (Clark, 1909) and Teliocrinus springeri liliaceus (Clark, 1909). Teliocrinus shares several ontogenetic trends with Endoxocrinus, especially in nonfunctional brachial articulations and stalk symplexies. Its assignment to the Diplocrininae is confirmed. A discussion of its affinities with pentacrinid fossil genera in which the crown is well preserved suggests that Diplocrininae could have first appeared during the Lower Cretaceous. A shortening of brachitaxes and a paedomorphic trend of stalk symplexies are the main other evolutionary traits. Nonfunctional articulations are frequently found at the paedomorphic pole of the heterochronic gradient, without clear derived characters. Classification of pentacrinids mainly based on such symplesiomorphy or paedomorphic characters must be definitively abandoned. However, in post-Palaeozoic stalked crinoids the scarcity of well-preserved fossils, the high frequency of paedomorphy, and convergent adaptive characters makes phylogenetic reconstruction only based on morphological characters very difficult and specula- tive. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 22–39. ADDITIONAL KEYWORDS: Echinodermata – evolution – extant – fossil – ontogeny – Pentacrinitidae – taxonomy. INTRODUCTION suggested close affinities between H. springeri, Hypalocrinus naresianus (Clark, 1908), and ‘Isocri- Among the pentacrinid stalked crinoids, the genus nus’ decorus (Wyville-Thomson, 1864). Three years Teliocrinus is remarkable in having primibrachials later, Clark (1912) proposed the genus Comastrocri- united by syzygies, and secundibrachials united by nus to accommodate for H. springeri, Hypalocrinus transverse ligamentary synarthries, synostoses (or ornatus Clark, 1909a, and Hypalocrinus liliaceus syzygies), or muscular synarthries (Roux, Messing Clark, 1909b. Clark (1912) emphasized the similari- & Améziane, 2002). Teliocrinus is the only extant ties in arm divisions between Comastrocrinus genus restricted to the northern Indian Ocean. The species and comatulid species in the subfamily type species, i.e. Hypalocrinus springeri (Clark, Capillasterinae as being ‘the result of the develop- 1909), was described by Clark (1909a). Using trans- ment under the same conditions’, i.e. adaptive char- verse ligamentary synarthries in proximal brachial acters. Döderlein (1912) attributed a number of articulations as a shared character, Clark (1909a) pentacrinid specimens to a new species and new genus Teliocrinus asper. Clark (1923), considering *Corresponding author. E-mail: [email protected] T. asper as a synonym of Comastrocrinus 22 © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 22–39 TELIOCRINUS (ECHINODERMATA): A KEY TAXON 23 springeri, transferred species in the genus Comas- (1909a, b) and deposited in the Indian Museum in trocrinus to Teliocrinus. Clark (1928) described Calcutta. We examined specimens described by Clark Teliocrinus monarthrus as a new species. Clark (1932): the specimens dredged by HMS Investigator (1946) synonymized T. monarthrus and Teliocrinus [deposited in the National Museum of Natural springeri. Clark (1932: 390–391) synonymized Telioc- History (USNM), Smithsonian Institution, Washing- rinus ornatus and Teliocrinus liliaceus with T. ton DC, USA], and the specimen collected by the springeri. This view was adopted by Oji (1990) and Cable [deposited in the Natural History Museum by Roux et al. (2002). (NHM), London, UK] (Table 1). Rasmussen (1961, 1978), Sieverts-Doreck (1971), Phenotypic variation was explored using data from Oji (1985), and Simms (1988) based their penta- direct observations of specimens, and from descrip- crinid classification on plesiomorphic characters – tions and figures from the literature. SEM observa- Downloaded from https://academic.oup.com/zoolinnean/article/155/1/22/2674285 by guest on 23 November 2020 mainly arm transverse synarthries – without any tions were conducted with the Service of Electronic information on ontogeny (Améziane-Cominardi & Microscopy of the IFR-BI at the Pierre & Marie Curie Roux, 1994). With such a view, Neocrinus blakei University (UPMC). For descriptions we refer to the (Carpenter, 1882) was transferred to the fossil genus terminology summarized by Roux et al. (2002). Bra- Isocrinus (Rasmussen, 1961), Neocrinus decorus was chitaxes are series of brachials between branch transferred to the Jurassic genus Chladocrinus points, either following a radial and including the (Sieverts-Doreck, 1971), and members of the family first ossicle at which the arm branches (axillary), or Pentacrinitidae s.l. were separated into several new following an axillary and including the next. Brachi- families (Simms, 1988). The genus Teliocrinus was taxes, beginning immediately following the radials, referred to the family Cainocrinidae, and Endoxocri- are often specified as primibrachial, secundibrachial, nus was referred to the family Isselicrinidae. Roux tertibrachial etc. composed of primi-, secundi- and (1981) distinguished five subfamilies, mostly based tertibrach ossicles, respectively. So, for example, the on the study of extant taxa and using characters third secundibrachial is IIBr3, or IIBr3ax if axillary. from both proximal arm pattern and stalk symplexy: Clark (1909a, b) described two different species: (1) Metacrininae (Metacrinus and Saracrinus), (2) T. springeri and T. liliaceus. The former has serrated Diplocrininae (Diplocrinus – now a synonym of arms and the latter has smooth arms (Figs 1, 2). As Endoxocrinus –andTeliocrinus), (3) Isocrininae demonstrated by David et al. (2006) on Endoxocrinus, (Cenocrinus as a recent representative of the Isocri- the character states ‘serrated’ versus ‘smooth’ distin- nus lineage), (4) Balanocrininae (Neocrinus and guish between infraspecific taxa. Therefore, in the Hypalocrinus as recent representatives of the Bal- first phase of the study we analyse without prejudg- anocrinus lineage), and (5) Pentacrinitinae (extinct ing the taxonomic status of these two phenotypes, and since the Late Jurassic). In the views of both Roux call them ‘phenotype springeri’ and ‘phenotype lili- (1981) (see also detailed historical introduction in aceus’. David et al., 2006) and Simms (1988), the general In this study synostosis and syzygy are used as term ‘isocrinids’ became inappropriate because it defined in Roux et al. (2002). SEM observations should be restricted to one family or subfamily that (Macurda, Meyer & Roux, 1978) have clarified the includes the genus Isocrinus. In their revision of the distinction between synostosis and syzygy using the genus Endoxocrinus, David et al. (2006) clarified the different types of stereom on facet articulation in hierarchy of characters in Diplocrininae. This led to pentacrinids (Roux, 1981; Améziane-Cominardi & the distinction of different ecophenotypes as subspe- Roux, 1994; Roux et al., 2002). The facets of a synos- cies related to depth and hydrodynamics, showing tosis are mainly covered by a stereom of regular singular adaptive characters mainly in their stalk. small meshes (synostosial stereom). In crinoid arms David et al. (2006) also suggested that Teliocrinus a synostosis transforms into a syzygy when an could be the oldest living representative of the sub- irregular and thick meshwork (syzygial stereom) family Diplocrininae. predominates, and can develop a vermiculated We offer a complementary description of Teliocrinus pattern or radial crenularium. During ontogeny, based on a re-examination of the characters in all synostosis is a primary stage and syzygy is a known specimens. We examine the place of this genus derived (secondary) stage. In intermediate stages, a in pentacrinid evolution and why there are difficulties syzygial stereom and crenularium may appear on to establish monophyletic groups. facets, whereas the suture remains synostosis-like in external morphology. In pentacrinid arms this stage is classically refered to as cryptosyzygy (Moore MATERIAL AND METHODS & Teichert, 1978). As a consequence, the term cryp- Holotypes of the crinoid species dredged during the tosyzygy is inappropriate and confusing, and must Investigator cruises were briefly described by Clark be dropped. © 2009 The Linnean Society of London, Zoological Journal of the