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Sea Urchins of the Genus Gracilechinus Fell & Pawson, 1966

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Sea Urchins of the Genus Gracilechinus Fell & Pawson, 1966 This article was downloaded by: [Kirill Minin] On: 02 October 2014, At: 07:19 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Marine Biology Research Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/smar20 Sea urchins of the genus Gracilechinus Fell & Pawson, 1966 from the Pacific Ocean: Morphology and evolutionary history Kirill V. Minina, Nikolay B. Petrovb & Irina P. Vladychenskayab a P. P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Moscow, Russia b A. N. Belozersky Research Institute of Physico-Chemical Biology, Moscow State University, Moscow, Russia Published online: 29 Sep 2014. Click for updates To cite this article: Kirill V. Minin, Nikolay B. Petrov & Irina P. Vladychenskaya (2014): Sea urchins of the genus Gracilechinus Fell & Pawson, 1966 from the Pacific Ocean: Morphology and evolutionary history, Marine Biology Research, DOI: 10.1080/17451000.2014.928413 To link to this article: http://dx.doi.org/10.1080/17451000.2014.928413 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http:// www.tandfonline.com/page/terms-and-conditions Marine Biology Research, 2014 http://dx.doi.org/10.1080/17451000.2014.928413 ORIGINAL ARTICLE Sea urchins of the genus Gracilechinus Fell & Pawson, 1966 from the Pacific Ocean: Morphology and evolutionary history KIRILL V. MININ1*, NIKOLAY B. PETROV2 & IRINA P. VLADYCHENSKAYA2 1P. P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Moscow, Russia, and 2A. N. Belozersky Research Institute of Physico-Chemical Biology, Moscow State University, Moscow, Russia Abstract Gracilechinus is a widely distributed genus of regular echinoids with a centre of species diversity in the North Atlantic (five of 10 species). Only three species of this genus are known from the Pacific Ocean: Gracilechinus multidentatus, G. euryporus and G. lucidus. The first two species were previously assigned to the genus Echinus, which differs from Gracilechinus in ambulacral tuberculation. According to our data, all three Pacific species possess ambulacral tuberculation characteristic of the genus Gracilechinus and differ from each other in the structure of the valves of tridentate pedicellariae and primary spines. Assignment of these three species to the genus Gracilechinus is also confirmed by phylogenetic analysis of the 657 bp fragment of the COI gene. The COI tree provides evidence for the validity of the genera Gracilechinus and Echinus and agrees in general with morphological data. Among species of Gracilechinus, two lineages are distinguished on the COI tree, differing in the shape of the tridentate pedicellariae valves. One lineage comprises only the North Atlantic species Gracilechinus alexandri, the other one includes the North Atlantic, South Atlantic and Pacific species. The genus Gracilechinus apparently originated in the North Atlantic and only representatives of the second lineage have penetrated the South Atlantic and the Pacific. Key words: Biogeographic history, COI, Echinus, Gracilechinus, Pacific, phylogeny Introduction other species of this genus, E. elegans Düben & Koren, 1844, E. alexandri Danielssen & Koren, Gracilechinus and Echinus are very closely related 1883, E. affinis Mortensen, 1903, E. atlanticus genera of sea urchins belonging to the family Echinidae. According to Mortensen (1943), the Mortensen, 1903, E. stenoporus Mortensen, 1942, genus Echinus included 17 species. Fell & Pawson and E. lucidus Döderlein, 1885, were transferred (1966) split Echinus and founded the new genus, to Gracilechinus (Smith & Kroh 2011; Kroh & Gracilechinus, to accommodate the species Echinus Hansson 2013). Downloaded by [Kirill Minin] at 07:19 02 October 2014 acutus Lamarck, 1816 and E. gracilis A. Agassiz, The assignment of E. multidentatus and E. eur- 1869. The new genus comprised species with a yporus H. L. Clark, 1912 to Echinus still remains primary tubercle on each ambulacral plate. Other questionable. Both for E. multidentatus (McKnight species were retained in the genus Echinus, charac- 1967) and E. euryporus (Clark 1925; Mortensen terized by having a primary tubercle on every second 1943) it was shown that they have a primary tubercle or third, or second and third ambulacral plate. on every ambulacral plate, which is the main McKnight (1967) transferred another Echinus spe- diagnostic feature of Gracilechinus. cies, Echinus multidentatus H. L. Clark, 1925, to the Gracilechinus has its centre of species diversity in genus Gracilechinus on the basis of the ambulacral the North Atlantic (five of eight species listed in tuberculation pattern of the type specimen and Smith & Kroh (2011) and Kroh (2012a)) with specimens from the Chatham Rise. Recently species occurring in a wide range of habitats and G. multidentatus was moved back to Echinus, but six depths from intertidal to the deep sea. According to *Corresponding author: Kirill V. Minin, P. P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Nakhimovsky Prospekt, 36, Moscow 117997, Russia. E-mail: [email protected] Published in collaboration with the Institute of Marine Research, Norway (Accepted 9 May 2014) © 2014 Taylor & Francis 2 K. V. Minin et al. Smith and Kroh (2011) and Kroh (2012a), Graci- to the Pacific Ocean from 1958 to 1987. Echinoids lechinus is represented in the Pacific Ocean by the were collected from the Southeast Pacific by the RV single species, G. lucidus (Döderlein, 1885). Two Akademik Kurchatov and RV Professor Shtokman, Echinus species, E. multidentatus and E. euryporus, from the South Pacific by RV Dmitry Mendeleev, possibly belonging to Gracilechinus, also occur in the and from the Northwest Pacific by RV Vityaz and Pacific Ocean (Mortensen 1943; Clark 1912; RV Odissey (Table I). In total 103 specimens of McKnight 1967, 1968; Kroh 2012b, 2012d). The Gracilechinus were obtained from six stations. All morphology and distribution of these three Pacific specimens were preserved in 96% ethanol on board species remain poorly known. the ships and then stored in the collections at the P. A study of the biogeographic history of Gracilechi- P. Shirshov Institute of Oceanology. Spine muscles nus was performed by Tyler et al. (1995). These and tube feet of one specimen of Echinus euryporus authors considered species of Gracilechinus in the H. L. Clark, 1912 (RV Professor Shtokman, St. 1964; genus Echinus. It was assumed that Echinus has a EspStock1964) and two specimens of Gracilechinus North Atlantic origin and that colonization of the lucidus (Döderlein, 1885) (RV Vityaz, St. 3768; different regions of the Pacific occurred separately. ElucV3768-25 and ElucV3768-27) were used for One possible pathway passed through the South the DNA extraction. Specimens of E. multidentatus Atlantic, turning eastward towards the Southwest H. L. Clark, 1925 (RV Dmitry Mendeleev, St. 1268) Pacific. This route coincides with the flow of the were unsuitable for DNA extraction because of poor North Atlantic Deep Water (NADW) capable of preservation condition. transporting larvae from the North Atlantic to the South Atlantic and then to the Southwest and Northwest Pacific. Because the NADW does not Morphological examination enter the Southeast Pacific, two possible pathways of Spines and pedicellariae for examination were colonization of this region were proposed: from the removed from the test and placed in sodium hypo- West Atlantic through the Straits of Panama or chlorite solution (household bleach), then washed through the Drake Passage (Tyler et al. 1995). with water and studied under a binocular dissecting In the present paper, redescriptions of the three microscope. The ultrastructure of cleaned spines and Pacific species of Gracilechinus are given. Phylogen- pedicellariae was studied using scanning electron etic relationships between the Pacific and the microscopy (SEM). Spines, pedicellariae and soft Atlantic Gracilechinus species are discussed and a tissue were removed from the test with a hard brush biogeographic history of this genus is proposed to make the plating and tuberculation patterns visible. based on the molecular phylogenetic tree generated Character terminology of the test follows Morten- using a fragment of the mitochondrial COI
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