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Journal of Natural History This article was downloaded by:[Canadian Research Knowledge Network] On: 5 October 2007 Access Details: [subscription number 770938029] 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 Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713192031 Revision of the nudibranch gastropod genus Tyrinna Bergh, 1898 (Doridoidea: Chromodorididae) Michael Schrödl; Sandra V. Millen Online Publication Date: 01 August 2001 To cite this Article: Schrödl, Michael and Millen, Sandra V. (2001) 'Revision of the nudibranch gastropod genus Tyrinna Bergh, 1898 (Doridoidea: Chromodorididae)', Journal of Natural History, 35:8, 1143 - 1171 To link to this article: DOI: 10.1080/00222930152434472 URL: http://dx.doi.org/10.1080/00222930152434472 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf This article maybe used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Journal of Natural History, 2001, 35, 1143–1171 Revision of the nudibranch gastropod genus Tyrinna Bergh, 1898 (Doridoidea: Chromodorididae) MICHAEL SCHROÈ DL² and SANDRA V. MILLEN³ ² Zoologische Staatssammlung MuÈ nchen, MuÈ nchhausenstr 21, 81247 MuÈ nchen, Germany ³ Department of Zoology, University of British Columbia, 6270 University Boulevard, Vancouver, British Columbia V6T 1Z4, Canada (Accepted 1 March 2000) The rediscovered holotype of Doris delicata Abraham, 1877 from Chile is shown to belong to the genus Tyrinna Bergh, 1898. The type specimens of the Chilean congeners T. nobilis Bergh, 1898, and T. pusae Marcus, 1959, as well as of Chromodori s juvenca Bergh, 1898 are re-examined and are here considered to be synonyms of T. delicata. Following the new code of nomenclature, retention of the commonly used name T. nobilis instead of its replacement by the senior but disregarded synonym T . delicata is proposed. In this study, T. nobilis Bergh, 1898 is redescribed anatomically and histologically using Chilean and Argentinian specimens. They show a wide range of morphological variation, regarding radular characters and proportions of digestive and genital organs. A vestibular gland around the vagina is absent in T. nobilis. In Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 contrast to previous descriptions of Tyrinna, spicules have been found in histological sections of T. nobilis. The Panamic and amphiatlantic species T. evelinae (Marcus, 1958) is redescribed anatomically and histologically using specimens from the Gulf of California, northern Peru and Jamaica. Tyrinna nobilis and T. evelinae form a natural group, the genus Tyrinna, due to the synapomorphies: (1) oral tentacles digitiform, with a deep median longitudinal groove (`enrolled’); (2) long, narrow and convoluted vagina; and (3) two or three submarginal rows of mantle dermal formations. The placement of the incompletely described Cadlina burnayi Ortea, 1988 from Cape Verde Islands in Tyrinna by MuniaõÂ n et al. (1996 ) seems justi® ed due to its possession of longitudinally `enrolled’, digitiform tentacles. Three poorly known Japanese species placed in Tyrinna by Rudman (1984) and MuniaõÂ n et al. (1996) diŒer in their tentacle shape: Cadlina japonica Baba, 1937, for which a lectotype is herein designated, and Cadlina sagamiensis Baba, 1937, are transferred to Cadlinella due to their spiculose, tuberculate notum. Eliot (1913) includes `Cadlina (or Tyrinna) sp.’ which does not belong to Tyrinna, but was too incompletely described to allow certain generic placement. A phylogenetic discussion of basal Chromodorididae reveals a high degree of parallelism and uncertainty. Keywords: Mollusca, Nudibranchia, Chromodorididae, Tyrinna, taxonomy, anatomy, phylogeny. Introduction The chromodorid genus Tyrinna Bergh, 1898 is a small but controversial taxon. The main diagnostic character of Tyrinna nobilis Bergh, 1898, the type species of Journal of Natural History ISSN 0022-2933 print/ISSN 1464-5262 online Ñ 2001 Taylor & Francis Ltd http://www.tandf.co.uk/journals 1144 M. Schro¨dl and S. V. Millen the genus, was the `presence of transversal furrows on the inner side of the oral tentacles’ (Bergh, 1898). Rudman (1984) characterized Tyrinna as having `enrolled’ oral tentacles, a smooth mantle with rounded bumps and two rows of marginal mantle glands, bi® d jaw elements, a wide radular ribbon, a long and unarmed vagina and vas deferens, and vaginally inserted sperm receptacles with the insemination duct leaving the vagina distal to these ducts. In a redescription of T . nobilis, Muniaõ Â n et al. (1996) added two diagnostic characters for Tyrinna, the absence of notal spicules and the presence of a vestibular gland, and they discussed the phylogenetic consequences of the latter character. The anatomy, taxonomy and systematic placement of Tyrinna species is problem- atic. No vestibular gland covering the vagina of T. nobilis was mentioned in the detailed original descriptions of T. nobilis, nor for its Chilean congener Tyrinna pusae Marcus, 1959 (Bergh, 1898; Marcus, 1959). Both species were regarded synonyms by MuniaõÂ n et al. (1996) although the main distinguishing features used by Marcus (1959), i.e. structure and position of the stomach, and denticulations of lateral radular teeth, were not considered. The relationships of T. nobilis and T. pusae with Chromodoris juvenca Bergh, 1898 and Doris delicata Abraham, 1877, two poorly known Chilean species (see SchroÈ dl, 1996), were uncertain. A second species, Cadlina evelinae Marcus, 1958, was transferred to Tyrinna by Rudman (1984). Due to its wide geographic distribution in both Paci® c and Atlantic tropical waters and to diŒerences in the specimen descriptions, Ortea (1988) sus- pected a `little puzzle’ of cryptic species around T. evelinae. Muniaõ Â n et al. (1996) additionally assigned the incompletely described Cadlina burnayi Ortea, 1988 to the Downloaded By: [Canadian Research Knowledge Network] At: 23:15 5 October 2007 genus Tyrinna. This was due to two rows of mantle dermal formations (MDFs), a smooth mantle, `enrolled’ oral tentacles and a broad radula. Two poorly known Japanese species, Cadlina sagamiensis Baba, 1937 and Cadlina japonica Baba, 1937, had already been considered to belong to Tyrinna by Rudman (1984). MuniaõÂ n et al. (1996) con® rmed this placement, despite their spiculose, tuberculate notum and auriculate oral tentacles (Baba, 1937, 1949) which are in contrast to the diagnosis of the genus Tyrinna within the same study. The present study is the continuation of earlier work on Chilean Tyrinna species (SchroÈ dl, 1996; 1997a) with the endeavour (1) to describe in detail the anatomy of species related to T. nobilis and T. evelinae, i.e. regarding the structure of the reproductive and digestive system; (2) to investigate the variability of these characters; and (3) to revise critically the genus Tyrinna. To these purposes specimens of T . nobilis and T . evelinae have been collected and examined anatomically as well as histologically by serial sections. To clarify synonymies, the type specimens of T. nobilis, T . pusae, Chromodoris juvenca and Doris delicata have been re-examined. Additionally, museum specimens of T. nobilis det. Muniaõ Â n, T. evelinae det. Thompson, C. sagamiensis and C. japonica have been re-examined; the results are used to revise the genus Tyrinna, and to discuss its monophyly and systematic placement. Material and methods During the years 1991± 1998, scuba diving was used to look for nudibranchs at several localities on the Chilean and Argentinian coasts. Specimens of Tyrinna nobilis were found in the localities given in table 1. In total, 15 specimens were collected and described in living condition. Measurements of living specimens re¯ ect the maximal values of length, width and height of the crawling specimens (table 2). The Revision of the nudibranch genus T yrinna 1145 nudibranchs were relaxed using a 10% MgCl2 solution and ® xed in 70% ethanol. Ten of the larger specimens were dissected and described anatomically. One specimen from the Bay of Coliumo (no. 8), parts of the notum of specimen no. 1 and distal genitalia of specimen no. 9 were embedded in hydroxyethylmethacrylat e ( Kulzer), serially sectioned (2.5 mm) and stained with toluidine-blue for histological examina- tion. Voucher specimens of T . nobilis from Los Hornos and the Bay of Coliumo were deposited in the Zoologische Staatssammlung MuÈ nchen (ZSM, nos 19960723, 19960724 ). The undissected holotype of Doris delicata Abraham, 1877 (British Museum of Natural History, BMNH), no. 1995038, was re-examined externally. The Swedish Museum of Natural History kindly made available
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