Phycoiogicai Researcli 1999; 47: 97-100
Research note First report of Gelidlella ligulata (Gelidiales, Rhodophyta) in Japan
Satoshi Shimada and Michio Masuda* Division of Biological Sciences, Graduate School of Science, Hokkaido University, Sapporo 060-0810, Japan.
Herbarium of the Graduate School of Science, SUMMARY Hokkaido University, Sapporo (SAP 063883-063886). Some plants were transported live to Hokkaido Univer- A gelidialean red alga that was newly found in Japan- sity for culture studies. Uniatgal cultures were estab- ese waters is referred to as Geiidieiia iiguiata Dawson, lished from excised apical tips of creeping axes of It is characterized by erect lanceolate blades develop- plants collected at Izu-misaki and Benkene-misaki and ing from a creeping axis that are relatively large in the were grown in Tris-buffered medium (Van Der Meer and genus Geiidieiia. Among the 22 species currently rec- Patwary 1991) at both 15 and 20X, with a 16:8 h LD ognized in the genus, Geiidieiia indica Sreenivasa Rao photoperiod and photon flux of 15-25 pE m ^s \ is most similar to G. iiguiata and a further comparative The following observations are based on field collected study is needed to elucidate the status of G. indica. and cultured materials. The plants form tufts on bedrock Geiidieiia iiguiata and the type species of the genus, in the middle intertidal zone of sheltered shores or m Geiidieiia acerosa (Forsskal) Feldmann et Hamel. have tidal pools. They are up to 4,5 cm tall (Fig, 2) and are the unicellular independent attachments that are com- dark red to purplish red in colour. Individual plants mon to the members of Geiidieiia investigated to date. consist of a creeping axis and erect blades. The creeping This type of attachment is unique in the Gehdiales axis attaches to the substratum by unicellular indepen- and this feature may be a useful taxonomic criterion in dent attachments (Figs 3,4) that are 50-240 nm in distinguishing Geiidieiia from other genera. length and 10 |jm in diameter. The creeping axis is sub- terete, 300-500 |jm in diameter and is branched irregu- Key words: Gelidiales, Geiidieiia indica, Geiidieiia larly. Erect blades arise trom the creeping axis. They are iiguiata, Japan, morphology, Pacific, Rhodophyta, terete (250-350 pm in diameter) at the proximal portion, secondary rhizoidal attachment. gradually expanding and become flattened. The blades are fan-shaped when young (Fig, 5), but become lan- ceolate with age (1-3 mm wide, 100-270 pm thick). Geiidieiia iiguiata Dawson 1953: p, 81, pi, 3, figs 3-5, They are usually simple, but are sometimes irregularly to Holotype: Dawson 6808 (11,in.1949) on sheet dichotomously branched (Fig. 6), Blade margins are 54721 in the Herbarium of the Allan Hancock Foun- undulate and sometimes ruffled, Subterete to lanceolate dation (HAHF). proliferations issue from both sides of blades pmnately Type locality: Cabeza Ballena, Baja California, (Fig, 6), injured (perhaps grazed) ends of blades (Fig, 7) Distribution: Baja California {Dawson 1953), Solomon and blade surfaces, A dome-shaped apical cell is evident Islands (Womersley and Bailey 1969) and Japan (present at the apices of creeping axes (Fig, 8) and erect blades, paper). as is typical of the Gelidiales. Both creeping axes and Japanese name: Sasaba-shimatengusa, erect blades (Fig, 9) consist of a medulla composed of Geiidieiia iiguiata Dawson is a gelidialean red alga that 10-18 layers of cells 6-40 pm in diameter and a cortex was established in 1953 by Dawson, Until now, there composed of 2-3 layers of smaller cells 3-5 pm in diam- have been only two reports of this species, one from Baja eter. Rhizines (slender, thick-walled, internal, hypha-like California (Dawson 1953) and another from the Soiomon filaments) are absent throughout creeping axes and erect Islands (Womersley and Bailey 1969}, We report here the blades. Reproductive structures were not found in the occurrence of this species in Japanese waters. present specimens. Plants of Geiidieiia iiguiata were collected from Miyake Island (Fig. 1) at Izu-misaki (13 July 1998) and Benkene-misaki (14 July 1998), The majority *To whom correspondence shouid be addressed. of these materials were fixed and preserved in 10% Email:
Izu-misaki
2 km
Fig. 1, Map showing the locations on Miyake Island where Geli- Fig. 2 Formalin seawater-preserved specimen of Gelidiella ligu- diella ligulata vjas collected. iata collected at Izu-misaki, Miyake Island,
The genus currently includes 22 species that are blades becoming closely pinnately branched near distinguished from members of other genera in the the extremities and readily deciduous and apparently Gelidiales by the absence of rhizines, Kraft and Abbott serving as a vegetative means of reproduction, whereas (1998} enumerated 23 species of Gelidlella but Sreenivasa Rao (1970) reported that lateral branches included G. calcicola Maggs et Guiry (Maggs and Guiry of G. indica are not generally deciduous. Pinnate 1987), which had been transferred earlier to Gelidium branches that are very similar to those of Dawson's alga as G. calcicola (Maggs et Guiry) R, E. Norris (1992) on (Dawson 1953. pi, 6, fig, 5) have been found in the the basis of having rhizines only at the attachment Japanese material (Fig, 6), but they are not deciduous. points. The species of Gelidiella have been character- At present, the deciduous nature of Dawson's (1953) ized by thallus habit, thallus size (height and diameter/ material, whether those deciduous branches function width), axis symmetry, tetrasporangial location and as propagules or are artefacts during preservation, has tetrasporangial arrangement (Kraft and Abbott 1998), not been confirmed, Gelidiella Indica seems to rep- A few species are known to have flattened thalii: resent matured stages of G, ligulata and the latter G, bornetil Weber-van Bosse) Feldmann et Hamel (Dawson 1953) has nomenclatural priority over the (Feldmann and Hamel 1934), G. feldmannii Baardseih former (Sreenivasa Rao 1970). However, it is prudent (1941), G. irdica Sreenivasa Rao (1970) and G. iigu- to maintain G. ligulata and G. indica as separate lata. Of these species. G. bornetii and G, feldmannii species until fully matured plants of the former alga differ from the Japanese material by having very narrow from its type locality are collected and these two algae blades up to 300 pm wide (Weber-/an Bosse 1926; as are thoroughly compared. We refer our material to Gelidium bornetii V>leber-\jar\ Bosse) and 500 pm wide G. ligulata {\\di\. has nomenclatural priority over G. indica. (Baardseth 1941). respectively, Gelidiella indica and In the molecular analyses of small subunit rDNA G. ligulata are more similar to the Japanese material. and rbcL sequences, G, ligulata from Japan clustered These two species have lanceolate erect blades, with Gelidieila acerosa (Forsskal) Fetdmann et Hamel Dawson (1953) did not describe tetrasporangia, but with 100% bootstrap values (Shimada et al. unpubl. Womersley and Bailey (1959) reported tetrasporangial data, 1999), lateral branches of their material from the Solomon The genus Gelidietla has been primarily distin- Islands, Similarly, G. indica was described to have guished from other genera in the Gelidiales by the tetrasporangial lateral branches in the upper to middle absence of rhizines throughout the thalii (Feldmann part of the erect axes (Sreenivasa Rao 1970; fig, 4c), and Hamel 1934). It has been traditionally character- One marked difference between these two species ized by the lack of a sexual generation (Fan 1951; is the presence/absence of deciduous branches. For Santelices 1997). However, the discovery of a single G. ligulata, Dawson (1953; p, 81) described older male gametophyte in Gelidiella acerosa from Malaysia Gelidiella liguiata 99
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Figs 3-9. Gelidiella ligulata collected at Izu-misaki. Formalin-seawater-preserved material unless otherwise indicated. 3. Secondary rhizoidal attachments. 4. Secondary rhizoidal attachments (cultured plant grown at 20"C, 15:8 h LD tor 1 month}. 5. Young fan-shaped blade5 issued from a creeping axis. 6. Uppermost portion of an erect blade forming branches irregularly to pinnately. 7. Proliferations from an injured (perhaps grazed) end of a blade. 8. Dome-shaped apical cell at the apex ot a creeping axis. 9. Transverse section of a blade showing the absence of rhizines m the cortex and medulla.
(Santelices 1997) requires reexamination of previous 1994). The peg type of secondary rhizoidal attach- collections and further research on the presence of ments has been found in Pterociadia lucida (Brown et gametophytic stages in the genus. Furthermore, the Turner) J. Agardh (Shimada and Masuda, unpubl, obs, genus can be characterized by the exclusive production based on material from Scarborough, Perth, 7 Dec- of the unicellular independent type of secondary rhi- ember 1997) and Pterocladielia (Perrone 1994; for zoidal attachment (Perrone 1994). Three types of sec- P. capiilacea (Gmelin) Santelices et Hommersand and ondary attachments are known in the Gelidiales: P. melanoidea (Schousboe ex Bornet) Santelices et (i) unicellular independent type; (ii) peg type; and Hommersand). The brush type occurs in other genera, (iii) brush type. Of these types, only the unicellular such as Gelidium (Perrone 1994; for G. iatifolium independent type has been reported for Geiidielia in (Greville) Bornet et Thuret and G. pusilium (Stack- G. lubrica (Kutzing) Feldmann et Hamel, G. nigrescens house) Le Jolis; Shimada and Masuda. unpubl. obs. for (Feldmann) Feldmann et Hamel, G. rameliosa{KiJizmg) G. vagum Okamura from Jodogahama, Iwate Prefecture, Feldmann et Hamel. G. pannosa (Feldmann) Feldmann 11 June 1997 and G. eiegans Kutzing from Awaji Island, et Hamel and G. anf/pa/Celan (De Gregorio and Perrone Hyogo Prefecture, 16 May 1996), Acanthopeitis japon- 100 S. Shimada and M, Masuda
/ca Okamura (Shimada and Masuda unpubl. obs. from REFERENCES material from Shimoda, Shizuoka Prefecture, 25 Sep- tember 1996) and Ptilophora subcostata (Okamura) Baardseth, E, 1941, The marine algae of Tristan da Cunha. Norris (Shimada and Masuda, unpubl. obs., material Res. Norw. Sci. Exped. Tristan Da Cunha 9: 1-173. from Naminoura, Wakayama Prefecture, 29 November Dawson, E. Y. 1953. Marine red algae of Pacific Mexico. Part 1996). Perrone (1994) suggested that the unicellular 1. Bangiales to Corallinaceae subf. Corallinoideae. Allan independent attachments can be used as a diagnostic Hancock Pacif. Exped. 17: 1 171. feature at the generic levei of Gelidiella, Pteroctadia De Gregorio, S. and Perrone, C. 1994. Rhizoid ontogenesis and Gelidium. This is confirmed for Gelidiella ligulata in Ge//c'/e//a Feldmann et Hamel (Gelidiales, Rhodophyta), in this study and the type species in this genus, Giorn. Bot. Ital. 128: 1085-7, Gelidiella acerosa (Shlmada and Masuda, unpubl. obs. Fan, K-C. 1951. Morphological studies of the Gelidiales, Univ. based on materials from Pulau Gulisaan, Sandakan, Calif. Publ. Bot. 32: 315-68. Sabah, Malaysia, 16 May 1998 and Ginowan, Okinawa Feldmann, J. and Hamel, G. 1934, Observations sur queiques Prefecture, 14 June 1998). Thus, the members of Geli- Gelidiacees. Rev. Gen. Bot. 46: 528-49. diella investigated so far have the unicellular indepen- Kraft, G, T and Abbott, I. A. 1998. Gelidiella womersleyana dent attachments that are unique in the Gelidiales. (Gelidiales, Rhodophyta), a diminutive new species from Kraft and Abbott (1998, figs 4,6,7), however, describe the Hawaiian Islands, Bot. Mar. 41: 51-61, and illustrate the attachment of Gelidiella womers- Maggs, C. A. and Guiry, M, D. 1987. Gelidiella calcicola sp. leyana Kraft et Abbott as rhizoids issuing as consoii- nov. (Rhodophyta) from the British Isles and northern dated cables on prostrate axes that look like the peg France. Br. Rhycol. J. 22: 417-34. type. Gelidiella womersleyana has erect axes of unique Norris, R. E. 1992. A proposed phylogenetic scheme for morphology, which are composed of a slender poly- the Gelidiales. In Abbott, I, A, (Ed.), Taxonomy of Eco- stromatic central region surrounded on both sides by nomic Seaweeds, Vol. 3. California Sea Grant Gollege, Uni- monostromatic wings and shows no clear affinities to versity of California, La Jolla, pp. 151-71, any other Gelidiella (Kraft and Abbott 1998). There Perrone, G. 1994, Diagnostic and taxonomic value ot the is a strong possibility that this species is actually not rhizoids in the Gelidiales: Some considerations, Giorn. a member of Gelidiella. As the genus Gelidiella is Bot. Ital. 128; 1088-91, believed to not be a natural taxon and requires taxo- Santelices, B. 1997. The spermatangial sorus of Gelidiella nomic revision (Kraft and Abbott 1998), molecular acerosa (Gelidiellaceae, Gelidiales). /n Abbott, I. A, (Ed.). analyses of the members of the genus is apparently Taxonomy of Economic Seaweeds, Vol, 6, California Sea needed. Grant Gollege Program, La Jolla, pp. 77-87. Sreenivasa Rao, P. 1970. Systematics of Indian Geiidiales. Phycos9: 63-78. ACKNOWLEDGEMENTS Van Der Meer, J. P. and Patwary, M, U, 1991. Genetic allevia- tion of the self-fertilization complication when hybridizing We thank Dr J. Huisman of Murdoch University and monoecious Gelidium vagum. Hydrobiologia 221: 167-79, Dr S. Kawaguchi of Kyushu University for providing Weber-van Bosse, A. 1926. Papers from Dr. Th- Mortensen's some of the materials. We also thank Dr K. Kogame of Pacific Expedition 1914-16. XXXIII. Algues de I'Expedi- Hokkaido University for his technical assistance and tion danoise aux Ties Kei. Vidensk. Medd. Dan. Naturhist. helpful discussion. This study was supported, in part, Foren. Kobenhavn SI: 57-155, by a Grant-in-Aid for International Scientific Research Womersley, H. B. S. and Bailey, A- 1969, Marine algae of (Field Research; No. 09041134) from the Ministry of the Solomon Islands. Pfiil. Trans. Roy. Soc. Lond. B 259: Education, Science, Sports and Culture, Japan. 257-352.