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The Genus Amphiroa (Lithophylloideae, Corallinaceae, Rhodophyta) from the Temperate Coasts of the Australian Continent, Including the Newly Described A

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The Genus Amphiroa (Lithophylloideae, Corallinaceae, Rhodophyta) from the Temperate Coasts of the Australian Continent, Including the Newly Described A Phycologia (2009) Volume 48 (4), 258–290 Published 6 July 2009 The genus Amphiroa (Lithophylloideae, Corallinaceae, Rhodophyta) from the temperate coasts of the Australian continent, including the newly described A. klochkovana 1 1 2 ADELE S. HARVEY *, WILLIAM J. WOELKERLING AND ALAN J.K. MILLAR 1Department of Botany, La Trobe University, Bundoora, Vic. 3083, Australia 2Royal Botanic Gardens, Mrs Macquaries Road, Sydney, NSW 2000, Australia HARVEY A.S., WOELKERLING W.J. AND MILLAR A.J.K. 2009. The genus Amphiroa (Lithophylloideae, Corallinaceae, Rhodophyta) from the temperate coasts of the Australian continent, including the newly described A. klochkovana. Phycologia 48: 258–290. DOI: 10.2216/08-84.1 Studies of Amphiroa (Lithophylloideae, Corallinaceae, Rhodophyta) from the temperate coasts of Australia provide new evidence that differences in tetrasporangial conceptacle pore canal anatomy are diagnostically significant in delimiting species within the genus. Differences in overall morphology and genicular anatomy are also reliable for delimiting species. These data are supported by examination of relevant type specimens. Four species occur in temperate Australian waters. Three (Amphiroa anceps, Amphira beauvoisii, and the newly described Amphiroa klochkovana) occur in southeastern Australia, and three (A. anceps, A. beauvoisii,andAmphiroa gracilis) occur in southern and southwestern Australia. Comparisons of A. beauvoisii and A. anceps have shown that they cannot be separated at species level morphologically but clearly differ in tetrasporangial conceptacle pore canal anatomy. This has important flow-on implications concerning specimen identification, reported geogeographic distribution and putative heterotypic synonymy of the two species. Relevant historical data, a species key and a synoptic description of Amphiroa also are included. INTRODUCTION Gabo Island, Victoria roughly comprises the Peronian province (Womersley 1990, Millar 2007). Lord Howe Island Amphiroa, one of six currently recognized genera in the (see Millar & Kraft 1993) and Norfolk Island (see Millar Corallinaceae, subfamily Lithophylloideae, is widespread in 1999), although politically part of New South Wales, were not tropical and warm temperate waters, also occurs in cool included because they are 600 and 1400 km respectively from temperate waters, and is known from the fossil record. the Australian mainland and show only partial biogeographic Woelkerling et al. (2002, p. 370, table 2; p. 372, table 3) links to the Peronian province (Millar 2007). The southern summarized the diagnostic characters of and provided a coast of Australia, including Tasmania, constitutes the dichotomous key to the six known genera of Lithophylloi- Flindersianprovince,whichissaidtodiffusenorthward deae with living species [Amphiroa, Ezo, Lithophyllum along the west coast to somewhere between Cape Leeuwin (including Titanoderma), Lithothrix, Paulsilvella and Te- and Geraldton (Huisman 2007; Millar 2007). Following narea]. Womersley (1984, p. 13), warmer-water species that occur This account contains a new taxonomic treatment of along the southwestern coast of Australia but are not also species of Amphiroa (Corallinaceae, Corallinales, Rhodo- definitely known from the south coast are not dealt with. phyta) confirmed to occur in temperate Australia and In addition to the species accounts and new data on provides new evidence that differences in tetrasporangial relevant types, the taxonomic implications of the results are conceptacle pore canal anatomy are diagnostically signif- considered in relation to the diagnostic value at species level icant in separating species within the genus. Our study of of differences in tetrasporangial conceptacle pore canal new and historical collections, including types, has led to anatomy and differences in genicular anatomy, and in the conclusion that four species occur in temperate relation to species biogeography. Relevant historical data, a Australian waters. Three (Amphiroa anceps, Amphiroa species key, and a synoptic description of Amphiroa also are beauvoisii, and the newly described Amphiroa klochkovana) included. occur in southeastern Australia and three (A. anceps, A. beauvoisii, and Amphiroa gracilis) occur in southern and southwestern Australia. We also have concluded that the MATERIAL AND METHODS status of 21 taxa previously considered synonyms of A. anceps or A. beauvoisii requires reassessment. Data are based on type material, newly collected material In the context of this study, temperate Australia encom- and herbarium collections from Universite´ de Caen, Caen, passes two biogeographic provinces. The southeastern coastal France (CN), Laboratoire de Cryptogamie, Muse´um area from the Queensland–New South Wales border south to national d’Histoire naturelle, Paris, France (PC), Depart- ment of Botany, La Trobe University, Victoria, Australia * Corresponding author ([email protected]). (LTB), National Herbarium of Victoria, Royal Botanic 258 Harvey et al.: Amphiroa from temperate Australia 259 Gardens, South Yarra, Victoria, Australia (MEL) and endophytic base or lacking a recognizable base or with National Herbarium of New South Wales, Royal Botanic multiple attachment points apparently formed secondarily Gardens, Sydney, New South Wales, Australia (NSW). and erect to recumbent axes composed of calcified segments Specimens originally in LTB and transferred to MEL have (intergenicula) and secondarily decalcified segments (genic- both LTB and MEL numbers cited. Herbarium abbrevia- ula); branching of axes variously dichotomous, alternate tions follow the Index Herbariorum, formerly in print pinnate, irregular, trichotomous or clustered, with mixtures (Holmgren et al. 1990), now online electronically (Holm- and with adventitious branches occurring in some species; gren and Holmgren 1998). The International Code of branch segments flattened to compressed to cylindrical. Botanical Nomenclature (ICBN) cited is the Vienna edition Thallus construction in upright axes monomerous, (McNeill et al. 2006). consisting of a single system of branched, laterally coherent Field procedures, species identification and microtech- filaments organized into alternating intergenicula and nique are detailed in Harvey et al. (2006). Morphological genicula. and anatomical terminology follows Woelkerling (1988). Intergenicula (as seen in median longitudinal section) Synoptic descriptions contain information on all diagnostic composed of a central core (medullary) region in which characters and other significant features, including those thin-walled, calcified, elongate cells of adjacent filaments useful for specimen identification. become more or less aligned to form longer tiers of arching cells interspersed by shorter tiers, and a peripheral (cortical) region where distal portions of calcified core filaments or their derivatives bend outward and terminate at the thallus HISTORICAL BACKGROUND surface in epithallial cells with rounded or flattened outer walls; branch apices usually composed only of core Published records of species Amphiroa from temperate filaments that terminate in meristematic initials. Cells of Australia begin with Lamarck (1815: 238), who described adjacent intergenicular filaments linked by secondary pit A. anceps as a species of Corallina on the basis of specimens connections; cell fusions absent. from ‘…les mers Australes ou de la Nouvelle-Hollande’ Genicula arising secondarily behind branch apices; collected by F. Peron & C.A. Lesueur during the French composed (as seen in median longitudinal section) of one expedition of 1800–1804 under the command of T.N. or usually more arching tiers of transformed, thick-walled, Baudin. (Ducker 1979b). decalcified core-region cells, and in some species also Since then, at least six additional species [A. beauvoisii consisting of associated decalcified peripheral-region cells. Lamouroux, Amphiroa dilatata Lamouroux; Amphiroa Transition from decalcified genicula to calcified interge- ephedraea (Lamarck) Decaisne; Amphiroa galaxauroides nicula occurring within single tiers of cells or between Sonder; A. gracilis W.H. Harvey; Amphiroa nobilis Ku¨tzing] adjacent tiers of cells. have been reported from the region; published records are Life history, where known, triphasic with haploid summarized by Womersley & Johansen (1996) for southern gametangial plants, diploid carposporophytes and diploid and southwestern Australia and by Millar & Kraft (1993) tetrasporangial plants. Bisporangial plants of unknown for southeastern Australia. ploidy level also found in some species. Gametangial plants, In conjunction with a detailed study of species of tetrasporangial plants and bisporangial plants all isomor- Amphiroa collected during the Siboga Expedition (see phic; carposporophytes developing within old female Weber 1902 for details), Weber-van Bosse (1904, p. 93) conceptacles after presumed karyogamy. concluded that A. dilatata, A. galaxauroides, and A. nobilis Male and female gametangia, tetrasporangia and bispor- were conspecific with A. anceps, and that a W.H. Harvey angia formed in separate uniporate conceptacles; male and specimen of A. ephedraea from New South Wales (explicit female conceptacles occurring on the same, or in most locality not indicated) numbered 458 was a misidentified species, on different plants. All mature conceptacles specimen of A. anceps. situated on the surfaces of intergenicula, formed laterally More recently, Womersley & Johansen (1996) concluded in peripheral (cortical)
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