Marine of Amchitka Island (Aleutian Islands). II. '

MICHAEL J. WYNNE2

ABSTRACT: Pleuroblepbaris stichidophora gen. et sp. nov., from Amchitka Island in the Aleutian Island s, is described as new to science. This taxon is the only repre­ sentative of the Bonnemaisoniaceae (Nemaliales, Rhodophyta) collected at Am­ chitka. It is distinguished from other members of the family by the presence of macroscopic tetrasporophytes with compound tetrasporangial stichidia arising along the margins of laminate axes. These tetrasporic branchlets are homologous to in­ determinate branches. Gland cells with brownish contents are present over the sur­ face of the laminate axes and also on the stichidia. Although numerous specimens have been collected, tetrasporic plants are the only fertile stages observed so far.

T HE FAMILY BONNEMAISONIACEAE (Nemali­ axibus processuum determinatorum nascuntur; ales, Rhodophyta) is represented at Amchitka filamentum primarium ramulorum tetraspori­ Island in the Aleutian Archipelago by a single corum uniseriatum, e cellulis trapezoideis quae of an undescribed genus. The first paper duos ordines stichidiorum alternantium efficiunt in this series (Wynne, 1970 ) has furn ished compositum. Tetrasporangia cruciate divisa 50­ the histor ical background of phycological ac­ 55 fA. diam ., 2 (interdum 3) ad omnen altitudi­ tivity in the Aleutians and environs, and has nem stichidii nisi segmentis infimis superioribus­ also reported on the location of Amchitka Is­ que reperta. Glandicellulae nencon in stichidio land and of the various collection sites men­ repertae. tioned in the present account. Thalli consisting of branched, flattened, rib­ Pleuroblepharis stichidophora gen. et sp. nov. bon-like axes, 5 to 10 (to 15 ) em long, to 4 mm broad ; growth by means of a single apical Figs. 1-12 cell with two cutting surfaces which cut off seg­ Thalli ex axibus ramosis applanatis vittafor­ ments obliquely and alternately, each segment mibus, 5-10 (-15) em long ., usque ad 4 mm cutting off two opposite pericentral cells, one lat. constantes; crescentia per singulam cellulam of which gives rise to a determinate subulate apicalem duas superficiales ad exsectionendum branchlet (1 mm long) and the other represents quae segmenta oblique alterneque exsectiunt the primordium of a potential indeterminate habentem effecta; omne segmentum duas cellulas branch; most of the possible indeterminate pericentrales oppositas exsectiens, quarum una branches remain undeveloped in the axils of ramulum (1 mm long.) determinatum subula­ the determinate processes. Surface (Fig. 4) is tum efficit, altera primordium rami indetermi­ covered by cortical cells intermingled with con­ nati potentis fingit; plurimi rami indeterminati spicuous brownish gland cells. Cortical cells possibiles in axibus processuum determinatorum (Figs. 5, 6) with 12 to 20 platelike parietal aborti manent. Superficies cellulis corticalibus chromatophores. Tetrasporic plants are the cum glandicellulis conspicuis subbruneis inter­ only fertile stages known so far ; tetrasporangia mixtis tecta. Plantae tetrasporicae solus status develop in branchlets, or "compound stichidia," fert ilis adhuc notae ; tetrasporangia in ramulis which are to 3 mm in length; tetrasporic branch­ usque ad 3 mm long. effecta; ramuli tetra­ lets are homologous to indeterminate branches sporici ramis indeterminatis homologi, itaque in and thus arise in the axils (Figs. 2, 9, 10) of the determ inate processes; tetrasporic branch­ 1 Manuscript received April 24, 1970. lets with a uniseriate main filament composed of 2 Department of Botany, The University of Texas at Austin, Austin, Texas 78712. trapezoidal cells, giving rise to two rows of al- 433 434 PACIFIC SCIENCE, Vol. 24, October 1970

FIG. 1. Pleuroblepbaris stichidophora. Holotype specimen ( WTU 242770) . Scale =3 em. ternating stichidia (Figs. 11, 12) . Tetrasporan­ 3-10 m). Square Bay: ix.1967, W einmann 33, gia cruciately divided, 50 to 55 !.t in diameter, Weinmann 263 (on Ptilota); viii.1968, W ein­ with 2 (at times 3) tetrasporangia occurring at mann 703A. Sea Otter Pen Cove: vi.1969, Le­ each level of the stichidium (Figs. 3, 7, 8), bednik 372 (on Odonthalia; 7-8 m) . South except for the lowermost and the upp er seg­ Bight : xii.1968, Lebednik 125A . St. Makarius ments. Gland cells also present on the stichid­ Bay: iii.1968, W ynne 1287 (tetrasporic); ium. Wynne 1529 (tetrasporic) .Rifle Range Point : iii.1968, Wynne 1490 (tetrasporic) . HOLOTYPUS : A tetrasporic specimen (Wynne 135 7) (Fig. 1), collected at St. Makarius Bay, Specimens of Pleuroblepbaris stichidophora Amchitka Island, March 16, 1968; deposited in are invariably epiphytic on the red algal genera the University of Washington Herbarium Ptilota and Odonthalia and are collected either (WTU 242770). Isotypes deposited in the her­ on the shore where the host has been cast up baria of the University of British Columbia, the or in the sublittoral by diving . Although this University of California, and the University of alga has been collected during most of the year, Texas. fertile plants were encountered only in March ADDITIONAL AMCHITKA COLLECTIONS : Con­ and April, and all of the numerous specimens stantine Harbor: iii.1968, Wynne 1348 ( tetra­ collected during these two months were tetra­ sporic) , Wynne 1510 (tetrasporic); iv.1969, sporic. Lebednik 312 (tetrasporic) . Constantine H ar­ The distinctive gland cells, which are charac­ bor dock: iv.1969, Lebednik 256 (tetrasporic; teristic of most members of the Bonnemaison- Marine Algae of Amchitka-WYNN E 435

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5

7

40jL 5,6 FIGS. 2- 8. P/eurob/epharis sticbidopbora. 2, Young tetrasporic branchlet. 3, Ind ividual stichid ium with immature tetrasporangia (shaded) . 4, Surface of laminate axis (gland cells are shaded). 5, 6, Cortical cells of laminate axis with chromatophores (shaded) . 7, Individual stichidium of mature tetrasporic branchlet, arising from uniseriate axis. 8, Portion of stichidium. 436 PACIFIC SCIENCE, Vol. 24, October 1970

FIGS. 9-12. Pleuroblepbaris sticbidopbora. 9, Young tetrasporic branchlet arising in axil of determinate branch; a sterile determinate branch has been developed at the lower left portion of the branchlet; X 128. 10, Young tetrasporic branchlet with some tetrasporangia being differentiated; X 120. 11, Mid-region of ma­ ture tetrasporic branchlet; X 114 . 12, Same as 11;X 110 . iaceae, indicated the affinity of the new genus Preliminary Culturing Studies for that family. The brownish color of their Both foliar axes and tetrasporic branchlets contents is like that of the iodine cells is As­ were excised and placed in a culture medium of paragopsis (Feldmann and Feldmann, 1942) supplemented natural seawater. The foliar axes rather than that of the colorless glandular cells continued to grow as broad leafy thalli and in . maintained their normal structure. The uniseri- Marine Algae of Amchitka-WYNNE 437 ate axes, on the other hand, grew indefinitely as and G) and Delisen fimbriata3 (Levring 1953, filamentous tufts, often reaching 2 em in length, fig. 55 B), which are the type species of these as contrasted with a length of about 3 mm on genera, tetrasporangia occur in sori located di­ the naturally occurring plants. These tufts even­ rectly on the flattened axes (see also Kylin, tually ceased producing tetrasporangia but grew 1956) . Thus, Plearoblepbaris, with tetraspo­ as vegetative uniseriate axes. Iodine cells contin­ rangia developing in lateral filaments which ued to be cut off alternately between adjacent arise from the margins of laminate axes, is strik­ cells, resembling the pattern in the "Trailliella­ ingly different from the other members of this stage" of (see Chihara, family. 1961; Taylor, 1957). In culture the iodine cells Feldmann and Feldmann (1942) established become colorless and refractile. The mature the as a distinct order to em­ vegetative cells of the axes are highly vacuolate phasize the heteromorphic nature of the game­ and have numerous parietal chromatophores. tophytic and sporophytic phases of Bonnemai­ Eventually the cultures died. At no time were sonia and . However, as has been sexual stages observed in nature or in culture. pointed out above, this trait does not occur in Discussion all members of the family. Papenfuss (1966) argued against the family's separation from the This remarkable new genus of the Bonnernai­ Nemaliales despite Fan's (1961) supplemental soniaceae is distinguished from other genera of information concerning the origin of the goni­ the family by the compound tetrasporangial moblast. The reports of heteromorphic life stichidia, which arise along the margins of the histories in other families of the Nemaliales (see flat axes. Until now the family has consisted of Martin , 1969; Ramus, 1969; West, 1968) have five genera (Kylin, 1956): Bonnemaisonia C. provided additional support for the retention Agardh , A sparagopsis Montagne, J. of Bonnemaisoniaceae within the Nemaliales . Agardh, Lamouroux, and Leptophyllis The presence of macroscopic tetrasporophytes J. Agardh. in Pleuroblepbaris stichidophora would at least Attention has been focused on this family suggest that it is another example of a species since an alternation of heteromorphic phases in the Bonnemaisoniaceae which does not have was demonstrated in certain members. The a heteromorphic life history. It will be of great tetrasporophyte of Asparagopsis is a polysiph­ interest to learn the nature of the sexual stages onous filamentous phase referred to as the in this new genus. "Falkenbergia-stage" (Feldmann and Feld­ mann, 1939, 1942, 1952; Feldmann, 1965; Chi­ hara, 1961) . Bonnemaisonia hamifera and B. ACKNOWLEDGMENTS nootkana have dissimilar filamentous tetrasporo­ I am grateful to my colleagues Dr. R. E. Nor­ phytes known as the "Trailliella-stage" (Harder ris, Mr. P. A. Lebednik, and Mr. F. C. We in­ and Koch, 1949; Chihara, 1961, 1965) ; how­ mann for making available their Amchitka col­ ever, the type species of the genus, B. aspara­ lections, made under US-AEC Contract No . AT goMes, apparently lacks a tetrasporic stage (26-1) -171. I also wish to thank the US-AEC­ (Feldmann and Feldmann, 1942, 1946; Feld­ Nevada Operations Office, which provided air mann, 1966). This latter type of life history was transportation and field support enabling me to also demonstrated in Ptilonia okadai by Chihara visit Amchitka to collect specimens. This study (1962) . By culturing carpospores he obtained was begun during the tenure of a National Sci­ a "Hymenoclonium-stage" as the basal disc ence Foundation postdoctoral fellowship at the which returned the apparent gametophyte. The University of Washington. Dr. R. E. Norris single report (Lucas and Perrin, 1947) of a kindly gave me access to the culturing facilities tetrasporic specimen of Ptilonia australasica must be regarded as a dubious record. An alternation of isomorphic phases has been 3 Contrary to Levring's observations of tetrasporo­ phytes among Australian specimens of this species, reported in Leptophyllis and Delisea. In both Chihara (196 2) reported that this species in Japan Leptopbyllis conferta (Levring, 1953, figs. 55 C has a haplobiontic life history. 438 PACIFIC SCIENCE, Vol. 24, October 1970 in his laboratory. I think Dr. H. Croasdale for la Societe d'histoire naturelle de 1'Afrique du providing the Latin diagnosis. Nord, vol. 37, pp. 35-38. --- 1952. Nouvelles recherches sur Ie cycle des Bonnemaisoniacees: Ie developpement des LITERATURE CITED tetraspores du Falkenbergia rttflanosa CHIHARA, M. 1961. Life cycle of the bonne­ (Harv.) Schmitz. Revue generale de botan­ maisoniaceous algae in Japan (1) . Scientific ique, vol. 59, pp. 313-323. Reports, Tokyo Kyoiku Daigaku, sect. B, vol. HARDER, R., and W . KOCH. 1949. Life history 10, pp. 121-153. of Bonnemaisonia hamifera (Trailliella in­ --- 1962. Life cycle of the bonnernaisoni­ tricata). Nature, vol. 163, p. 106. aceous algae in Japan (2) . Scientific Reports, KYLIN, H. 1956. Die Gattungen der Rhodophy­ Tokyo Kyoiku Daigaku, sect. B, vol. 11, pp. ceen. Lund, Gleerup. xv + 673 pp. 27-53. LEVRING, T . 1953. The marine algae of Aus­ --- 1965. Germination of the carpospores tralia. I. Rhodophyta: Goniotrichales, Bangi­ of Bonnemaisonia nootkana, with special ref­ ales and Nemalionales. Arkiv for Botanik, erence to the life cycle. Phycologia, vol. 5, A. S., vol. 2, pp. 457-566. pp.71-79. LUCAS, A. H. S., and F. PERRIN. 1947. The sea­ FAN, K.-c. 1961. Morphological studies of the weeds of South Australia. Part II . The red Gelidiales . University of California Publica­ seaweeds, pp. 113-458. Adelaide, Govern­ ment Printer. tions in Botany, vol. 32, pp. 315-368. MARTIN, M. T. 1969. A review of life-histo­ FELDMANN, G. 1965 . Le developpement des ries in the Nemalionales and some allied tetraspores de Falkenbergia ruflanosa et Ie genera. British Phycological Journal, vol. 4, cycle des Bonnemaisoniales. Revue generale pp. 145-158. de botanique, vol. 62, pp. 621-626. PAPENFUSS, G. F. 1966. A review of the pres­ --- 1966. Sur le cycle haplobiontique du ent system of classification of the Floride­ Bonnemaisonia asparagoides (Woodw.) Ag. ophycidae. Phycologia, vol. 5, pp. 247-255. Compte rendu de I'Academic des sciences, RAMUS, J. 1969. The developmental sequence Paris, vol. 262D, pp. 1695-1698. of the marine red alga Pseudogloiphloea in FELDMANN, J., and G. FELDMANN. 1939. Sur culture. University of California Publications le developpement des carpospores et l'alter­ in Botany, vol. 52, 42 pp. nance de generations de I'Asparagopsis ar­ TAYLOR, W. R. 1957. Marine algae of the mata Harvey. Compte rendu de l'Academie northeastern coast of North America. Rev. des sciences, Paris, vol. 208, pp. 1240-1242. ed. University of Michigan Studies, Scientific --- 1942. Recherches sur les Bonnemaison­ Series, 13, viii + 509 pp. iacees et leur alternance de generations. An­ WEST, J. A. 1968. Morphology and reproduc­ nales des sciences naturelles, Botanique, ser. tion of the red alga Acrochaetium pectinatum 11, vol. 3, pp. 75-175. in culture. J. Phycology 4 :89-99. --- 1946. A propos d'un recent travail du WYNNE, M. J. 1970. Marine algae of Am­ Prof. H. Kylin sur 1'alternance de generations chitka Island (Aleutian Islands) . I. Deles­ du Bonnemaisonia asparagoides. Bulletin de seriaceae. Syesis, vol. 3, pp. 95-144.