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Eudoxia Macra Is the Eudoxid Stage of Lensia Cossack (Siphonophora, Cnidaria) Mary M

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Eudoxia Macra Is the Eudoxid Stage of Lensia Cossack (Siphonophora, Cnidaria) Mary M This article was downloaded by: [Smithsonian Institution Libraries] On: 17 October 2013, At: 08:20 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 Systematics and Biodiversity Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tsab20 The end of an enigmatic taxon: Eudoxia macra is the eudoxid stage of Lensia cossack (Siphonophora, Cnidaria) Mary M. Grossmann a b , Dhugal J. Lindsay a b & Allen G. Collins c a Yokohama City University , Yokohama , 236-0027 , Japan b Japan Agency for Marine–Earth Science and Technology (JAMSTEC) , Yokosuka , 237-0061 , Japan c National Museum of Natural History , Washington , DC , 20013-7012 , USA Published online: 03 Oct 2013. To cite this article: Mary M. Grossmann , Dhugal J. Lindsay & Allen G. Collins (2013) The end of an enigmatic taxon: Eudoxia macra is the eudoxid stage of Lensia cossack (Siphonophora, Cnidaria), Systematics and Biodiversity, 11:3, 381-387, DOI: 10.1080/14772000.2013.825658 To link to this article: http://dx.doi.org/10.1080/14772000.2013.825658 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. 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. 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Authors also have the option of publishing an Open Select article under the Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0/. It is essential that you check the license status of any given Open and Open Select article to confirm conditions of access and use. Downloaded by [Smithsonian Institution Libraries] at 08:20 17 October 2013 Systematics and Biodiversity (2013), 11(3): 381–387 Research Article The end of an enigmatic taxon: Eudoxia macra is the eudoxid stage of Lensia cossack (Siphonophora, Cnidaria) MARY M. GROSSMANN1,2, DHUGAL J. LINDSAY1,2 & ALLEN G. COLLINS3 1Yokohama City University 236-0027, Yokohama, Japan 2Japan Agency for Marine–Earth Science and Technology (JAMSTEC) 237-0061, Yokosuka, Japan 3National Museum of Natural History, 20013-7012, Washington, DC, USA (Received 15 March 2013; revised 25 June 2013; accepted 12 July 2013) The greatest challenge in the study of the cnidarian order Siphonophora lies in the identification of all parts of the fragile colonies usually separated during net sampling. Species are usually established based on the morphology of at least one swimming bell of the polygastric stage, and a single genus, Eudoxia, has housed the description of free sexual eudoxid stages until these can be reliably linked to a polygastric stage. This genus, extensively used until the early 20th century, has progressively been emptied and, at the present day, contains only one form for which the identification of the polygastric stage has remained completely enigmatic: Eudoxia macra Totton, 1954. DNA barcoding techniques using the mitochondrial 16S gene allowed this eudoxid stage to finally be linked with its polygastric stage, Lensia cossack Totton, 1941. http://zoobank.org/urn:lsid:zoobank.org:pub:F253E52B-C14D-4440-9BA5-EB9EAC6B42E0 Key words: DNA barcoding, Eudoxia macra, Lensia cossack, geographic divergence, mitochondrial 16S, Siphonophora Introduction which may well be conspecific with Pyrostephos vanhoef- The greatest challenge in the study of the cnidarian or- feni Moser, 1925 (Grossmann et al., 2013). der Siphonophora lies in the identification of all parts of The genus Eudoxia (and its precursor Ersaea), exten- the fragile colonies usually separated during net sampling. sively used until the early twentieth century, has progres- The identification of different life stages of species under sively been emptied and, at the present day, contains only different names is not uncommon in the history of bino- one completely enigmatic form: Eudoxia macra Totton, mial taxonomy. Within the Cnidaria, the Anthomedusae and 1954. The association of eudoxid and polygastric stages Athecata orders, and the Leptomedusae and Thecata orders is primarily done through morphological and distributional separated the medusoid and polypoid forms, respectively, similarities. Indeed, the phyllocyst of the bract of many of species presently regrouped into the Anthoathecata and eudoxid stages resembles, in shape, the somatocyst of the Downloaded by [Smithsonian Institution Libraries] at 08:20 17 October 2013 Leptothecata. anterior nectophore of the polygastric stage (Totton, 1965; In the order Siphonophora, species are usually estab- Mapstone, 2009); and polygastric stages having nec- lished based on the morphology of at least one swimming tophores with marked ridges tend to have eudoxid stages bell of the polygastric stage, and a single genus, Eudoxia, with ridged bracts (personal observation: e.g. 5-ridged has housed the description of free sexual eudoxid stages Lensia, genera Diphyes, Eudoxoides, Muggiaea, etc.). Al- until these can be reliably linked to a polygastric stage. No though this identification of eudoxid stages has been per- other life stages have been described unless they could be formed successfully many times in the Mediterranean Sea ´ linked to a known polygastric stage (e.g. Russell, 1938; (e.g. Totton, 1932; Gamulin, 1966; Gamulin & Krsiniˇ c, 2000), where the diversity of Siphonophora is relatively D. Carre,´ 1972; C. Carre,´ 1979), except for the small post-larval physonect stage Mica micula Margulis, 1982, low, the extremely high biodiversity of Japanese waters (Fujikura et al., 2012) would not normally lend itself to this type of enterprise. Applying DNA barcoding techniques to the cnidarian or- Correspondence to: Mary M. Grossmann. E-mail: [email protected] der Siphonophora, the position of Eudoxia macra within the ISSN 1477-2000 print / 1478-0933 online C 2013 The Author(s). Published by Taylor & Francis. This is an Open Access article. Non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly attributed, cited, and is not altered, transformed, or built upon in any way, is permitted. The moral rights of the named author(s) have been asserted. http://dx.doi.org/10.1080/14772000.2013.825658 382 M. M. Grossmann et al. family Diphyidae Quoy & Gaimard, 1827 and, more specif- Twenty-two specimens of seven species of the genus ically, amongst the non-ridged Lensia species is discussed, Lensia, and four Eudoxia macra gonophores were col- using both morphological and genetic data from specimens lected using plankton nets and preserved in 99.5% ethanol caught in south-east Asian and Antarctic waters. or frozen at –20◦C onboard after identification to species level (Table 1). Frozen specimens were transferred to 99.5% Material and methods ethanol before DNA extraction. For 15 specimens (access The formalin-seawater preserved material included four numbers KC78253434, 40–53), total DNA was extracted complete Eudoxia macra eudoxids, nine bracts and 14 using the Qiagen (Valencia, California, USA) DNEasy gonophores collected in 330 μm mesh IONESS (Kitamura Blood & Tissue kit, and a 623 bp segment of the mitochon- et al., 2001) tows in March 2006 off south-eastern Japan; drial 16S gene amplified and sequenced using ‘primer 1’ seven gonophores collected with a 1.13 m diameter fry net and ‘primer 2’ from Cunningham & Buss (1993) with the (330 μm mesh) between October and December 2010 off TaKaRa (Otsu, Shiga, Japan) ExTaq and BigDye kits. Se- the north-eastern coast of Japan;
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