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The Morphology and Taxonomy of the Red Alga Sarconema (Gigartinales: Solieriaceae)

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The Morphology and Taxonomy of the Red Alga Sarconema (Gigartinales: Solieriaceae) Phycologia (1974) Volume 13 (1), 31-44 The morphology and taxonomy of the red alga Sarconema (Gigartinales: Solieriaceae) GEORGE F. PAPENFUSS* AND TrKVAH EOELSTEINt * Department of Botany, University of California, Berkeley, California 94720, U.S.A. t Atlantic Regional Laboratory, National Research Council of Canada, Halifax, Nova Scotia, Canada A study of the structure and reproduction of two species of Sarcollemafrom East Africa has shown that the genus is correctly placed in the Solieriaceae. The genus agrees with Solieria in that the auxiliary cell is indistinguishable before diploidization. A sheath of pericar pic filaments is not produced about the gonimoblast as it is in CallophyclIs, Solieria, Agardhiella and Meristotheca. Only two species of Sarcollemaare recognized hy us, S. filiforme(Sander) Kylin and S. scillaioidesB0rgesen. S. Jurcellatum Zanardini, S. montagnei (Grunow) Kylin, S. indicum (J. Agardh) Kylin, S. Jurcatum B0rgesen and S. filiformef. gracillima Rayss are regarded as representative of S. filiforme. Dicranema setaceum Sonder from Queensland, Australia, and D. setaceum var. "polensis Grunow from Samoa were also found to be S. filiforme. Introduction (now known as Callophycus Trevisan, 1848; Silva, 1957). The same seven genera were placed in the The history of the family Solieriaceae, to which Solieriaceae by Schmitz & Hauptfleisch (1896). Sarconema belongs, begins with J. Agardh, who in However, in both these accounts the family was 1852 (pp. 718 and 721) erected the tribe Solierieae for treated as a subfamily of the RhodophyJIidaceae. the genus Solieria J. Agardh (1842). Harvey (1853, De Toni (1897, 1924) adopted the classification of & p. 115) elevated the Solierieae to the rank of sub­ Schmitz HauptfJeisch.. family (except that he called it suborder) and included Kylin in 1925 (p. 25) erected the family Rhab­ in it the genus Eucheuma J. Agardh (1847) in addition doniaceae for the reception of Rhabdonia, Agardhiella to Solieria. In 1872 J. Agardh included in the Schmitz (Schmitz & Hauptfleisch, 1896), Anatheca Solierieae his new genus Meristotheca and Rhabdonia Schmitz (Schmitz & Hauptfleisch, 1896), Flahaliltia Harvey (Hooker & Harvey, 1847), in addition to Bornet (1892) and Solieria. (The inclusion of Solieria, Solieria and Eucheuma, and in 1876 he added: the type genus of the Solieriaceae, in the Rhab­ Gelinaria Sonder (with a query; the genus is now doniaceae by Kylin illegitimatizes the family.) In placed in the Cryptonemiaceae), Carpococcus 1932 Kylin removed Solieria, Anatheca, Flahaultia J. Agardh (currently treated as asynonym of Sarcodia and Agardhiella to the Solieriaceae (the existence of J. Agardh of the family Sarcodiaceae), and Catenella which he apparently had overlooked in 1925) and he Greville (1830) (with a query; the genus is now referred Catenella, Erythroclonium and Areschougia, referred to the Rhabdoniaceae). in addition to Rhabdonia, to the Rhabdoniaceae. In Hauck in 1883 (pp. 17 and 186) elevated the sub­ his Gattungen Kylin (1956) added Catenellocolax family Solierieae to the rank of famiIy and in 1886 he Weber-van Bosse (1928) to the Rhabdoniaceae and placed Sarconema Zanardini (1858) in the family. dates the family as of 1932. Searles (1968, pp. 44 and Schmitz in 1889 (p. 442) placed the following seven 63) has provisionally also assigned Caulacanthus genera in the Solieriaceae (J. Agardh) Hauck: Kiitzing (1843), Taylorophyclls Dawson (1961) and Rhabdonia Harvey, Erythroclonium Sonder (1852), Heringia J. Agardh (1844) to the Rhabdoniaceae. Areschougia Harvey (1855, nomen conservandum), Joly & Alveal (1969) and Augier & Boudouresque Solieria J. Agardh, EllcheumaJ. Agardh, Sarconema (1971) have provisionally also placed the new genera Zanardini, and Thysanocladia (Endlicher) Lindley Montemaria and Feldmannophycus, respectively, in the Rhabdoniaceae. As an appendix to the present Issued as NRCC No. 13699. paper, the family is proposed for conservation. 31 32 Phyc% gia, Vol. 13 (1), 1974 In 1932 Kylin placed the following ten genera in In view of our very meagre knowledge of the the Solieriaceae: Thysanocladia (= Callophycus), development of the female reproductive apparatus Flahaultia, Sarcodiotheca Kylin, gen. nov., Agard­ and the carposporophyte in the Solieriaceae and the hiella, Solieria, Sarconema, Eucheuma, Anatheca, availability of fertile material of Sarconema from Meristotheca, and Euryomma Schmitz (Schmitz & East Africa, the present study was undertaken with Hauptfleisch, 1896). In 1934 Kylin included Opun­ the hope of contributing to a better understanding tiella Kylin (1925) and Turnerella Schmitz (Schmitz of this genus and the family. Two species (the only & Hauptfleisch, 1896) in the family and in 1956 he two in the genus in our opinion, as will be pointed credited the family with two additional genera, out in later pages), S. filiforme(Sonder) Kylin and namely, Predaea G. DeToni (1936) and Gardneriella S. scinaioides B0rgesen were studied. Both liquid Kylin (194 1). preserved and herbarium specimens were examined. Dawson (1949) added the monotypic new genus Reticulobotrys to the family and B0rgesen (1953) added the new genus Tenaciphyllum. Feldmann Structure of the thallus (1942) removed Predaea to the Nemastomaceae, in which family the genus had been placed by Setchell Thalli of Sarconema (Figs. 1-6, 19, 20 and 23-25) are & Gardner (1930) when they described it (as terete, solid, 0,8-2 mm in diameter near the base, up Clarionea gen. nov.). Dawson (1961) also placed to 20 cm long, dichotomously to subdichotomously Predaea in the Nemastomaceae and Kraft & Abbott branched, and attached by a branched, hapteroid (1971) have also convincingly demonstrated that holdfast. The branches taper gradually toward the this genus belongs in the Nemastomaceae rather than tip. Regeneration occurs from branches whose tips in the Solieriaceae. Kraft & Abbott also reduced have been lost. One or several new branches may be Yadranella Ercegovic (1949) to asynonym of Predaea. produced from such tips, which frequently results in The features whereby the Solieriaceae may be a disturbance of the dichotomous habit. characterized are: (1) the thallus is multiaxial in The thallus is muItiaxial in construction and construction and has a filamentous medulla; (2) consists of three clearly defined tissues (Figs. 19 and procarps are lacking; (3) the auxiliary cell initiates 20); an outer cortex of one or two layers of small, only one gonimoblast cell; (4) the gonimoblast at tightly packed pigmented cells (Figs. 8 and 20); an first develops inwardly; (5) the cystocarps are inner cortex of several layers of irregularly shaped, embedded in the thallus or project from the thallus pseudoparenchymatous cells, which in size decrease surface and are usually provided with a distinct toward the outer surface (Figs. 19 and 20); and a ostiole; (6) a pericarp is usually, but not always, prod­ central medulla of interwoven thick-walled filaments, uced; and (7) the tetrasporangia are zonately divided. composed of elongated cells with lateral pit connec­ The family Rhabdoniaceae which has been tions between adjacent filaments (Figs. 19 and 20). confused with the Solieriaceae differs from the latter especially in that (1) the thallus is uniaxial in construction, and (2) the cystocarp of all genera Development of the female reproductive system lacks a special pericarp. Information of a greater or lesser amount on the As in other members of the Solieriaceae, procarps Sarconema. ontogeny of the cystocarp in the Solieriaceae is are not formed in Carpogonial branches available for the following genera: Sarcodiotheca are produced in large numbers in the tips of the * (Kylin, 1925, as Anatheca), Turnerella (Kylin, 1934; thallus branches. The carpogonial branch is com- Tokida & Masaki, 1957),* Sarconema (Kylin, 1932; * Young stages were best observed in material of S. scinaioides Rayss, 1963), Opuntiella (Kylin, 1934), Agardhiella that was collected by Papenfuss & Scagel (PR-YIl-\02) near (Osterhout, 1896; Kylin, 1928), Solieria (Bornet, in Dar-es-Salaam, Tanganyika, in October 1962. Rayss's (1963) cystocarpic material of Sarconema from the eastern Mediter­ Bornet & Thuret, 1880; Kylin, 1932), Meristotheca ranean was collected in July and October. Hauck's (1886) (Kylin, 1932), Gardneriella (Kylin, 1941) and material of this genus, collected by Hildebrandt in Somalia, Callophyclls(Hewitt, 1960). contained cystocarps in March, Newton's (1953) plant from the Hanish Islands, Red Sea, contained tetrasporangia in * Tetrasporophytes have been unknown in Tlirnerella. September, and Aleem (1948), who was the first to report the Recently South et al. (1972) showed that T. pennyi) of genus from the Mediterranean, obtained plants with tetra­ the north Atlantic Ocean possesses a crustose, Crlloria-like sporangia in October. B0rgesen (1932) had tetrasporic tetrasporic phase. specimens gathered (in Pakistan or India) in January. Papenfuss and Edelstein: Morphology and taxonomy of Sarconema 33 2em � ,' 2 em .2 I....-..J FIGS. 1-3. Sarcol1ema /ili/orme. FIG. 1. Habit of a liquid preserved cystocarpic specimen collected by Papenfuss & Scagel (PR·YUJ-48) near Dar·es·Salaam, Tanganyika, on 11-13 October 1962. FIG. 2. Habit of herbarium specimen (showing the dichotomously branched nature of the species) collected by Papenfuss & Friedmann (E62/20036) at Entedebir lslet, Dahlak Archipelago, Ethiopia, on 13 March 1962. FIG. 3. Voucher specimen of Borgensen's 1934 record of S. jill'carul1I Borgesen from Pakistan (NY). c 34 Phyc% gia, Vol.
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