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Corallinaceae, Rhodophyta) Based on Studies of Type and Other · Critical Specimens*

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Corallinaceae, Rhodophyta) Based on Studies of Type and Other · Critical Specimens* Phycologia (1985) Volume 24 (3), 317-337 A taxonomic and nomenclatural reassessment of Tenarea, Titanoderma and Dermatolithon (Corallinaceae, Rhodophyta) based on studies of type and other· critical specimens* WM. J. WOELKERLING', YVONNE M. CHAMBERLAIN2,4 AND PAUL C. SILVA3 'Department 0/ Botany, La Trobe University, Bundoora, Vic. 3083, Australia 2The Marine Laboratory, Portsmouth Polytechnic, Ferry Road, Hayling Island POll ODG, England 3Herbarium, Department 0/ Botany, University o/ California, Berkeley, CA 94720, USA WM. J. WOELKERLING, Y.M. CHAMBERLAIN AND P.C. SILVA. 1985. A taxonomic and nomenclatural reassessment of Tenarea, Titanodermaand Dermatolithon (Corallinaceae, Rhodophyta) ba��d on studies of type and other critical specimens. Phyc% gia 24: 317-337. The taxonomy and nomenclature of genera of the TenarealDermatolithon complex has been reas­ sessed on the basis of a study of generitype and other critical specimens. The type specimen of Millepora tortuosa Esper, which is the type of Tenarea Bory 1832, proves to be con specific with the specimen on which Bory based his description of the genus. Tenarea undu/osa Bory is an illegitimate name for T.tortuosa (Esper) Lemoine rather than an independent species. Since 1898, M. tortuosa has been incorrectly associated with the alga that is the principal component of the 'trottoirs' (coralline pavements) of the western Mediterranean. This alga is referable to Lithophyllum lichenoides Philippi. TitanodermaNageli 1858 is an earlier homotypic synonym of Dermatolithon Foslie 1898, both being based on Me/obesia pustu/ata Lamouroux. Titanoderma,in view of its clear record, is adopted in preference to proposing Dermatolithon for conservation. Tenarea comprises a single species of Lithophylloideae in which the thallus is composed of erect or ascending lamellae whose vegetative tissues are organized in an isobilateral manner and include a medulla of two layers of palisade cells. Titanoderma (syn. Dermatolithon) encompasses those Lithophylloideae in which the thallus consists largely or entirely of a prostrate crust whose tissues are organized in a dorsiventral manner and include a unistratose hypothallium composed of palisade-like cells. Titanoderma includes at least 16 species. INTRODUCTION ically. The controversy was settled in favour of the second alternative (in the absence of an over­ The circumscription and naming of genera in the whelming case for conservation) in 1981 by the TenarealDermatolithon complex (Corallina­ Nomenclature Section of the International Bo­ ceae, Rhodophyta) has been attended by uncer­ tanical Congress at Sydney and is incorporated tainties as is indicated by Johansen (1981, pp. in Articles 10.2 and 10.3 of the current Inter­ 42, 217, 223, 225, 226) in his recent survey of national Code of Botanical Nomenclature coralline algae. In addition to·diff erences in taxo­ (ICBN-see Voss 1983). Second, no attempt has nomic opinion, two factors are responsible for been made to examine the type specimens of these uncertainties. First, it has been a long­ these generic names and thus the nomenclature standing controversy whether a generic name of this complex has lacked the foundation that must be typified with material in the hands of is essential to stability. the author (that is, material upon which the ge­ During the course of studies on the nongenic­ neric description was primarily based) or, alter­ ulate Corallinaceae of the British Isles (by Y.M.C.) natively, with the type of a cited species, whether and of southern Australia (by W.J.W.), indepen­ or not the citation proves to be correct taxonom- dent visits were made by these authors to CN (Universite de Caen, France) in 1979 and 1980 * Dedicated to the memory of Mme Marie Lemoine to examine Lamouroux's specimens of Melobes­ (1887-1984) in recognition of her generous help over ia pustulata, Der­ 76 years (1909-1984) to all people interested in cor­ on which the generic name allines. matolithon is based. Subsequently (1981), Y.M.C. • Mrs Y.M. Butler. and W.J.W. concurrently studied material on 317 318 Phycologia, Vol. 24 (3), 1985 which Bory based his description of Tenarea, abbreviations are taken from Holmgren et al housed at PC (Laboratoire de Cryptogamie, Mu­ (1981). seum National d'Histoire Naturelle, Paris). The whereabouts of Esper's specimen of Millepora tortuosa, on which the concept of Tenarea must HISTORICAL BACKGROUND be based in accordance with the Sydney decision, remained unknown until 1983 when one of us Tenarea (Y.M.C.) discovered that it was at FR (For­ Bory (1832, p. 207, pI. 54, fig. 3) established the schungsinstitut Senckenberg, Frankfurt, Ger­ genus Tenarea to accommodate a 'polypier' many). [Dr M. Grasshoff from FR has indicated (plant-like animal) collected at Cap Tenare (Cape in correspondence (with W.J.W.) that it was Tainaron), Greece. The organisms were de­ transferred from ER (Botanisches Institut der scribed as calcareous, canary yellow when grow­ Universitiit Erlangen, Germany) to FR in 1970.] ing but bleached when dry and composed of Comparative examination of the Lamouroux, lamellae that were undulate-tortuouse, anasto­ Bory, and Esper specimens together with studies mosing, and folded together. Bory was explicit of more recently collected materials enabled us in indicating that it was not new to science at the to resolve various uncertainties in the Tenareal species level and cited Millepora tortuosa Esper Dermatolithon complex at the generic level. Type (1796, p. 118, Millepora pI. xxii) in synonymy. specimens have been designated and relevant This name had been applied by Esper to a more historical background data are provided. or less globular calcareous organism with thin, ascending, variously twisted lamellae which were MATERIALS AND METHODS closely united with one another and which had scattered hemispherical 'pores' (conceptacles). Data were obtained from type and other critical Esper's specimen presumably came from the specimens housed at CN, FR, L (Rijksherba­ Mediterranean, and he compared his organism rium, Leiden, Netherlands) and PC and from with several others, including Corallina offici­ additional collections housed at L TB (Depart­ nalis Linnaeus and Millepora lichenoides (Ellis) ment of Botany, La Trobe University, Bundoora, Ellis & Solander. Unfortunately, in an action that Victoria, Australia). Microtechnique follows complicated subsequent nomenclature, Bory Jones & Woelkerling (1984) and permanent slides (1832) changed the specific epithet to undulosa. have been retained at LTB. Fractures for scan­ The ICBN always has proscribed unnecessary ning electron microscopy were prepared by changing of epithets and if Bory had not estab­ breaking small pieces of thallus, mounting these lished a new genus at the same time that he on aluminium stubs with 'Fotobond' acrylic ad­ changed the epithet, no problem would have aris­ hesive (Agfa-Gevaert Limited), and double-coat­ en. Establishing a new genus, however, allowed ing the specimens with carbon and then with gold for the interpretation of Tenarea undulosa as a prior to viewing in a Siemens ETEC Autoscan new species and the type of its genus, an inter­ microscope. Wherever cell measurements are pretation which has been held by many botanists given, 'L' denotes length, 'D' denotes diameter, but which was ruled out by the Sydney decision and 'LID' denotes the ratio of length to diameter. in 1981 (ICBN, Arts 10.2, 10.3; see Voss 1983). Measurements include the cell walls. Herbarium T'enareawas overlooked until Hariot (1895) Figs 1-6. Lectotype of Tenarea tortuosa (Esper) Lemoine. Fig. 1. Lectotype specimen (housed at FR). Fig. 2. Surface of lectotype showing numerous, tortuouse, intertwined, branched lamellae. Fig. 3. Apex of lamella showing characteristic groove. Fig. 4. Close-up of portion of groove (arrows). Note distinct cell row on opposite sides of groove. Fig. 5. Surface of lamella showing early stage of branch development, three co nceptacles, and numerous co n­ cavities denoting the positions of epithallial cells. Fig. 6. Fracture showing one lamella (a) abutting against the surface of another lamella (b). Both lamellae are fractured in a transverse plane. Woelkerling et al: Tenarea, Titanoderma and Dermatolithon 319 320 Phycologia, Vol. 24 (3), 1985 Figs 7-11. Tenarea tortuosa. Lectotype, Figs 7- 10; LTB 14645, Fig. II. Fig. 7. Penetration of one lamel1a (a) by another lamel1a (b); the latter is transversely fractured above the region of penetratio n. Woelkerling et al: Tenarea, Titanodermaand Dermatolithon 321 re-examined Bory's specimen and confirmed that hypothallium that is unistratose and palisade­ it was a nongeniculate coralline alga. He noted like (i.e. with vertically elongate, more or less Bory's citation of Esper's diagnosis of Millepora obliquely arranged cells) and thus generically dis­ tortuosa, but chose to retain the epithet undulosa. tinct from Lithophyllum; and Considering mainly external morphology, Ha­ (2) that plants identified by Lemoine and oth­ riot concluded that Tenarea Bory (1832) was ers as Tenarea tortuosa are truly referable to congeneric with and had priority over Litho­ Lithophyllum, belonging to the species L. tor­ phyllum Philippi (1837). He did not examine tuosum (Esper) Foslie. Huve's concept of Lith­ Philippi's collections, however, and the first ophyllum tortuosum, however, was not based on modem account of this critical material was not an examination of Esper's type. published until 88 years later (Woelkerling 1983). Dermatolithon Without examining the type
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