In the Tropical Western Atlantic: I. Colombia and Venezuela

BULLETIN OF MARINE SCIENCE, 33(4): 881-898, 1983

GENUS POLYSIPHONIA (RHODOPHYTA, CERAMIALES) IN THE TROPICAL WESTERN ATLANTIC: I. COLOMBIA AND VENEZUELA

Donald F. Kapraun, Andres J. Lemus and German Bula-Meyer

ABSTRACT Keys, illustrations, and descriptions are provided for 10 species of Polysiphonia from the Caribbean coast of Colombia and Venezuela: p, atlantica. P. binneyi, P. denudata. p, eastwoodae, P. Jerulacea, P. jlaccidissima, P. havanensis. p, howei, P. sphaerocarpa and P. subtilissima. Five of these species are newly reported for the study area. Comparisons are made with descriptions for these species elsewhere, and taxonomic and phytogeographic conclusions given.

Twenty-two species of Polysiphonia. exclusive of cool temperate species which occur in the Carolinas and Bermuda, are known from the warm temperate and tropical waters of the western Atlantic (Taylor, 1960; 1976; Wynne and Edwards, 1970; Cordeiro-Marino and Oliveira F., 1970; Brauner, 1975; Oliveira F. and Ugadim, 1976; Kapraun, 1977b; 1979a; 1980a; Kapraun and Norris, 1982). De- spite the increased efforts of the past decade, there are still no published accounts of the taxonomically important morphological features for many of these species, including some of the most commonly reported. The present study of Polysiphonia from Colombia and Venezuela (part 1),along with a complementary investigation of this genus from other western Atlantic sites which will be published separately (part II), is intended to provide these details for both previously known and newly reported species in the western Atlantic.

MATERIALS AND METHODS

Polysiphonia material for this study was collected from a wide range of habitats in Venezuela (Fig. I) and Colombia (Fig. 2) by Andres J. Lemus and German Bula-Meyer, respectively, Specimens studied are deposited in the UNC-Wilmington (WNC) Herbarium as karo slides and/or dried specimens. Descriptions and camera lucida drawings herein presented are based solely on specimens from this study.

KEY TO THE SPECIES OF POLYSIPHONIA IN CARIBBEAN VENEZUELA AND COLOMBIA

Ia. Peri central cells 5 or more .. , 2 lb. Pericentral cells 4 ,...... 3 2a. Pericentral cells 5-6 .. P. denudata 2b. Pericentral cells 10-12 P. howei 3a. Branches arising in axils of trichoblasts 4 3b. Branches replacing trichoblasts 6 4a. Plants with a creeping base or extensive decumbent branch system; main axes less than 150 Itm diam 5 4b. Plants primarily erect; main axes 200-300 Itm diam .. P. binneyi 5a. Main axes 100-150 Itm diam; rhizoids in open connection with pericentral cells .. P. havanensis 5b. Main axes 80-100 Itm diam; rhizoids cut off from pericentral cells P. j/accidissima 6a. Rhizoids in open connection with pericentral cells 7 6b. Rhizoids cut off from pericentral cells _.._.._...... 8

881 882 BULLETIN OF MARINE SCIENCE. VOL. 33. NO.4. 1983

15 to 76 75

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00' 63' Figure I. (Left) Map of the northeast coast of Venezuela showing collection sites. I. Bahia de Mo- chima, 2. Cumana, 3. Patao, 4. Macuro, 5. Cariaquito, 6. Punta Piedras, 7. Punta Sabaneta. Figure 2. (Right) Map of the Colombian coast in the region of Cartagena-Santa Marta showing collection sites. I. Isla del Tesoro, 2. Cienaga Grande de Santa Marta, 3. La Lorna, 4. Punta Gloria, 5. Rodadero, 6. Santa Marta, 7. Ensenada de Concha, 8. Ensenada de Chengue, 9. Ensenada dc Gayraca, 10. Ensenada de Nenguange.

7a. Plants minute, less than I em tall, with main axes less than 60 !Lm diam; branches developing unilaterally from prostrate axes, giving a dorsiventral habit P. atlantica 7b. Plants to 4 em tall, main axes to 150 !Lm diam; branches developing radially from both prostrate and erect axes...... P. subtilissima Sa. Plants small, to 1.5 cm tall, main filaments less than 120 !Lm diam; often epiphytic, erect filaments arising from a discoid base P. sphaerocarpa 8b. Plants larger, to 5 cm tall, main axes to 250!Lm diam; erect filaments from entangled decumbent branches .. _...... 9 9a. Segments in main axes 0.5 diam long; branchlets short, distinctly spindle-shaped with constricted bases .. P. ferulacea 9b. Segments in main axes 1-·2 diam long; branchlets elongate, hardly narrowed at the base ...... P. eastwoodae

Polysiphonia atlantica Kapraun and Norris Figures 41 and 42

P. atlantica Kapraun and Norris 1982: 226. P. macrocarpa Harvey in MacKay 1836: 206 [non Polysiphonia macrocarpa (c. Agardh) Sprengel IS27: 350]. Description. - Plants minute to 1 cm tall, creeping, entangled with other algae, reddish-brown to black; erect filaments sparingly subdichotomously branched; branches in erect filaments replacing trichoblasts in development; segments to 60 JJmdiam, to 2 diam long; pericentral cells 4; trichoblasts scarce, with one division; apical cell conspicuous; prostrate axes to 60 JJm diam; erect axes arising endog- enously, or exogenously from apical primordia with delayed development; rhizoids abundant, one per segment, forming from the middle of pericentral cells and remaining in open connection with them. Type- Locality. - PortS1.ewart, Miltown Malbay, Ireland. Distribution. - Western Atlantic (Taylor, 1960; Dawes, 1974; Oliveira Filho and KAPRAUN ET AL.: POLYSIPIIONIA IN THE WESTERN ATLANTIC 883

Ugadim, 1976; Kapraun, 1977b; 1979b; Kapraun and Norris, 1982), temperate eastern Atlantic (Batten, 1923; Ardre, 1970). This species is newly reported from the study area.

Specimens Studied.- Venezuela: Bahia de Mochima (Lemus, 3 July 1979, WNC 14384). Irapa, Golfo de Paria (Lemus, 19 June 1976, WNC 12166). Cariaquito, Golfo de Pari a (Lemus, 17 June 1976, WNC 12167). Remarks.-Kapraun and Norris (1982) proposed a new name, P. atlantica, because the taxon P. macrocarpa Harvey in Mackay (1836) is invalid (Womersley, 1979), being a later homonym of P. macrocarpa (c. Agardh) Sprengel (1827: 350). Specimens obtained in this study are comparable with material from the Harvey Herbarium (Babbacom be, nearTorbay [Torquay], Devonshire, England, Sept. 1845, US 66490, and Batcombe, Dorsetshire, England, 1836, US 66491). However, a cystocarpic specimen issued with the Danicis Indiae Occidentalis exsiccata as P. macrocarpa (F. B0rgesen, St. Croix, Christiansted, 21 Jan. 1896, MICH s.n.) was determined to be Polysiphonia subtilissima. Although B0rgesen's illustrations for 'Po macrocarpa' (1918, figs. 272-276) are in close agreement with those of Harvey (1846, as P. pulvinata Spreng.) and Batten (1923, figs. 12-14) and ours for P. atlantica, his species description is in variance in some important respects (e.g., plant height and filament diameter, suggesting the possibility that P. subtilissima was included in his species concept of'P. macrocarpd' This supposition is further strengthened when one considers that B0rgesen (1918; 1924) never reported P. subtilissima from the West Indies where it is abundant and wide-spread. Womersley (1979) suggested that Polysiphonia atlantica (as P. macrocarpa) and P. subtilissima are closely related. Studies of these taxa in the western Atlantic, however, have shown them to have distinct developmental patterns. Whereas Polysiphonia subtilissima has radial development of branches in prostrate axes, P. atlantica gives rise to unilateral filaments from prostrate axes, producing a dorsiventral habit (Kapraun, 1977b; 1979b).

Polysiphonia binneyi Harvey Figures 3-6

P. binneyi Harvey 1853: 37. Description.-Plants small, to 3 em tall from a discoid base; branching pseudo- dichotomous below, alternate above; lower portions of main axes attached to the substratum by rhizoids which remain in open connection with the pericentral cells; pericentral cells 4; main axes 200-300 ~m diam; segments about 1 diam long below, 1.5-2.0 diam tall above; branches developing in axils oftrichoblasts; branch apices blunt, nearly obscured by numerous persistent highly branched trichoblasts; adventitious trichoblasts developing freely from scar cells in older branches; spermatangial branches cylindrical, about 50 x 200 Mm,often with 2- 3 sterile tip cells, and with subtending trichoblasts; pericarps spherical, about 250 ~m diam, opening by a wide ostiole; tetrasporangia swollen, to 70 ~m diam, in short spiral series. Type- Locality. - Key West, Florida. Distribution. - Tropical western Atlantic (Howe, 1920; Chapman, 1963; Taylor, 1960; 1969; 1976; Taylor and Bernatowicz, 1969; Mathieson and Dawes, 1975).

Specimens Studied.-Colombia: Ensenada de Concha (Bula-Meyer, 26 July 1979, WNC 12158). En- senada de Chengue (Bula-Meyer, IS June 1979, WNC 14380). 884 BULLETIN OF MARINE SCIENCE, VOL 33, NO, 4, 1983

Figures 3-6. Po{ysiphonia binneyi. Fig. 3. Discoidal base with rhizoids in open connection with pericentral cells. Fig. 4. Tetrasporangia. Fig. 5. Spermatangial branches. Fig. 6. Pericarp and adventitious trichoblasts. Figs. 3, 5, 6: Scale = 100 !Lm. Fig. 4: Scale = 50 !Lm.

Remarks. -Polysiphonia binneyi is clearly distinguished from P. havanensis, the other species encountered in this study which has branches developing in the axils of trichoblasts and continuous connection of rhizoids with pericentral cells. In contrast to Polysiphonia binneyi, P. havanensis develops erect axes from an extensive prostrate system, has spermatangial branches mostly without sterile tip cells, broadly oval pericarps, and tetrasporangia in long spiral series. KAPRAUN ET AL.: POt YSIPfIONIA IN THE WESTERN ATLANTIC 885

Polysiphonia denudata (Dillwyn) Greville ex Harvey in W. J. Hooker Figures 39 and 40

P. denudata (Dillwyn) Greville ex Harvey in W. J. Hooker 1833: 332. Conferva denudata Dillwyn 1809: 85. P. denudata (Dillwyn) Kuetzing 1849: 824. P. variegata (J. G. Agardh) Zanardini 1841: 60. Description. - Plants dark red, erect, to 5 cm tall from a discoid base; branching dichotomous; above branches widely divergent, below becoming decumbent and attached to the substratum by rhizoids cut offfrom the proximal end of peri central cells; main axes 200-300 !-tm diam; branches arising in axils of trichoblasts; pericentral cells 5 (-6); tetrasporangia to 50 !-tm diam, in long spiral series in branch tips; sexual structures not seen. Type- Locality. - Southampton, England. Distribution. - Temperate and tropical waters throughout the world: western At- lantic (Taylor, 1957; 1960; 1969; Edwards, 1970; de Rios, 1972; Kapraun, 1977b; 1978; 1979b; Schnetter and Bula-Meyer, 1977; Oliveira Filho, 1977; Kapraun and Norris, 1982), Atlantic Europe, Morocco, and the Mediterranean (Batten, 1923; Veldkamp, 1950; Ardre, 1970; Lauret, 1970) and West Africa (Lawson and John, 1977).

Specimens Studied.-Colombia: Ensenada de Nenguange (Bula-Meyer, 2 Sept. 1976, WNC 14381). Venezuela: Cumana (Lemus, IS Aug. 1976, WNC 14382). Cariaquito (Lemus s.n., 17 June 1976). Macuro (Lemus s.n., 16 June 1976).

Polysiphonia eastwoodae Setchell and Gardner Figures 25-29

P. eastwoodae Setchell in Gardner 1930: 161. P. snyderae Kylin 1941: 35. P. mol/is sensu Hollenberg 1961: 359 [non P. mol/is J. D. Hooker & Harvey 1847: 397]. Description. - Plants dull reddish brown, 2-3 (-5) cm tall from a discoid base; erect filaments dichotomously branched, replacing trichob1asts in development; older branches becoming decumbent and attached to the substratum by rhizoids cut off from the proximal end of peri central cells; peri central cells 4; segments 1- 2 diam long, to 250 !-tm diam, trichoblasts 1 per segment, soon decidous, leaving persistent scar cells; tetrasporangia swollen in spiral series in clustered branchlets; spermatangial branches 50 X 150-200 !-tm, with or without a solitary sterile tip cell, with or without a subtending trichoblast. Type- Locality. - Guadalupe Island, South Anchorage, off Baja California. Distribution. - Temperate and tropical Pacific (Kylin, 1941, as P. snyderae; Hol- lenberg, 1961; Meiiez, 1964; Abbott and Hollenberg, 1976; Hollenberg and Norris, 1977; all as P. moWs).

Specimens Studied.- Venezuela: Cariaquito (Lemus, 17 June 1976, WNC 14390). Remarks. - Womersley (1979) critically examined the type material of Polysi- phonia moWs J. D. Hooker in Harvey and determined that "almost certainly" branches develop in the axils of trichoblasts, in contrast to the concept of this taxon developed by Hollenberg (1961; 1968a). In the present study, specimens clearly showed branches replacing trichob1asts in development, and so are referred 886 BULLETIN OF MARINE SCIENCE. VOL. 33. NO.4. 1983 to Polysiphonia eastwoodae Setchell in Gardner, which seems to be the available epithet for this distinct entity. Specimens from this study are comparable with Polysiphonia eastwoodae from the type locality (Isotype: H. L. Mason #55, South Anchorage, Guadalupe Island, off Baja California, Mexico AHFH 8860) as well as specimens identified as P. moWs sensu Hollenberg from Pacific Mexico (EYD 2936 in USC 2788) and Fiji (WNC 10055, WNC 10894). Although specimens from Venezuela are consider- ably smaller than those from Pacific Ocean localities, they agree in all pertinent details. Apparently, this is the first report for this species from the tropical Atlantic. It is tempting to speculate that this taxon is a recent addition to the Caribbean flora, having gained access via the Panama Canal. Lipkin and Safriel (1971) documented similar dispersal and migration in historic times since the connection of the Mediterranean and Red Seas by the canal at Suez.

Polysiphoniaferulacea Suhr in J. Agardh Figures 7-10

p./erulacea Suhr in J. Agardh 1863: 980. Description. - Plants coarse, dark brown to black; filaments to 4 cm tall, initially from a discoid base; older plants forming an entangled fibrous base of decumbent branches, secondarily attached by rhizoids cut offfrom pericentral cells; peri central cells 4; segments in main axes about 0.5 diam long, to 200 Jlm diam; branch lets short, distinctly spindle-shaped with constricted bases; erect apices acute with conspicuous apical cells and persistent trichoblasts; decumbent apices devoid of trichoblasts; branches replacing trichoblasts in development; spermatangial branches cylindrical, about 50 X 150 Jlm, most often with 1-2 sterile tip cells; pericarps spherical, to 250 Jlm diam; tetrasporangia little swollen in short spiral series in branch tips. Type-Locality. -Atlantic coast of Mexico. Distribution. - Temperate and tropical western Atlantic (B0rgesen, 1918; Taylor, 1928; 1960; 1969; 1980; Taylor and Rhyne, 1970; de Rios, 1972; Richardson, 1975; Oliveira Filho, 1977; Kapraun, 1977a; b; 1980a; Kapraun and Norris, 1982), Portugal (Ardre, 1970), and East Africa (Jaasund, 1976).

Specimens Studied. -Colombia: Ensenada de Nenguange (Bula-Meyer, 5 Nov. 1976, WNC 12163). Rodadero (Bula-Meyer, 20 July 1976, WNC 12162). Ensenada de Nenguange (Bula-Meyer, 13 July 1978, WNC 12165). La Lorna (Bula-Meyer, 24 Aug. 1976, WNC 12159). Punta Gloria (Bula-Meyer, 10 July 1976, WNC 12164). Ensenada de Chengue (Bula-Meyer #350, 15 June 1979).

Polysiphonia jlaccidissima Hollenberg Figures 11-14

P. flaccidissima Hollenberg 1942: 783. Description.-Plants minute, erect filaments to 1.5 cm tall from an extensive creeping system; pericentral cells 4; prostrate axes to 100 Jlm diam, erect filaments to 80-100 Jlm; branches developing in axils of trichoblasts; scar cells giving rise to adventitious branches; rhizoids cut off from the proximal end of pericentral cells; tetrasporangia to 75 Jlm diam, in long spiral series; mature pericarps ovate, 150 X 175 Jlm. Type-Locality. -Laguna Beach, Orange County, California. KAPRAUN ET AL.: POL l'S/P1/(h\'/..I IN THE WESTERN ATLANTIC 887

Figures 7-10. Polysiphoniaferulacea. Fig. 7. Spermatangial branches on spindle-shaped branchlet. Fig. 8. Immature pericarps. Fig. 9. Rhizoids cut off from pericentral cells in decumbent branch. Fig. 10. Tetrasporangia. Fig. 7: Scale = 50 /lm. Figs. 8, 10: Scale = 100 /lm. Fig. 9: Scale = 200 /lm.

Distribution. - Temperate and tropical Pacific (Hollenberg, 1942; Dawson et al., 1964; Hollenberg and Norris, 1977) and western Atlantic (Brauner, 1975; Ka- praun, 1979b; 1980a; Kapraun and Norris, 1982). This species is previously unreported from the coasts of Venezuela and Colombia.

Specimens Studied. - Venezuela: Macuro (Lemus, 29 Sept. 1975, WNC 143983). Colombia: Isla del Tesoro (Bula-Meyer, I Dec. 1979, WNC 12161). 888 BULLETL'I OF MARINE SCIENCE, VOL. 33, NO.4. 1983

Figures 11-14, Polysiphoniaj/accidissima, Fig, II. Pericarp, Fig, 12, Apex showing branch arising in axil of trichoblast. Fig, 13, Tetrasporangia, Fig, 14, Rhizoids cut off from pericentral cells in decumbent axis, Fig, II: Scale = 50 /lm. Figs. 12-14: Scale = 100 /lm,

Remarks. - Womersley (J 979) compared south Australian and Mediterranean material identified as Polysiphonia sertularioides (Grateloup) J. Agardh with descriptions of P. jlaccidissima from the Pacific Ocean (Hollenberg, 1942, 1968a; Abbott and Hollenberg, 1976). He concluded that no satisfactory differences are apparent between these two taxa, and raised the query that P. jlaccidissima Hol- lenberg might be a synonym of P. sertularioides. Material from the present study is referred to Polysiphoniajlaccidissima until our investigation of the relationship of this taxon to P. sertularioides is complete. KAPRAUN ET AL.: POLYSIPHONIA IN THE WESTERN ATLANTIC 889

Figures 15-20. Polysiphonia havanensis. Fig. 15. Tetrasporangia. Fig. 16. Spermatangial branches. Fig. 17. Peri carp. Fig. 18. Apex with branch arising in axil oftrichob]ast. Fig. 19. Decumbent branch showing rhizoids remaining in open connection with pericentral cells. Fig. 20. Adventitious branch. Figs. ] 5-19: Scale = 50 11m. Fig. 20: Scale = 100 11m.

Polysiphonia havanensis Montagne Figures 15-20

?P. havanensis Montagne ]837: 352. Description. - Plants reddish-brown, with erect filaments to 3 em tall from an extensive creeping system; main axes to 100-150 I-Lm diam, with an abundance 890 BULLETIN OF MARINE SCIENCE, VOL. 33, NO.4, 1983 of adventitious branch lets; pericentral cells 4; apices with numerous highly branched trichoblasts; basally attenuated branches arising in axils of trichoblasts; rhizoids developing from the middle of pericentral cells to which they remain in open connection; spermatangial branches about 75 X 150 Ilm, mostly without sterile tip cells; pericarps ovate, 250 X 300 Ilm; tetrasporangia to 50 J.Lm diam, in long spiral series in upper branches, Type-Locality,-Havana, Cuba, Distribution. - Temperate and tropical western Atlantic (Borgesen, 1918; Taylor, 1960; 1976; Chapman, 1963; Oliveira Filho, 1977; Kapraun, 1979b).

Specimens Studied.-Colombia: ]sla de Baru (Bula-Meyer, 7 Dec. ]979, WNC 12]60). Ensenada de Gayraca (Bu]a-Meyer #383, 3 Aug. 1979). Venezuela: Bahia de Mochima (Lemus, 3 Ju]y 1979, WNC ]4385). Patao (Lemus, 29 Sept. ]975, WNC ]4387). Remarks, - The relationship of Polysiphonia havanensis sensu Borgesen to P. sertularioides was previously questioned because of the several similarities which exist between these two taxa (Kapraun, 1977b). No taxonomic conclusions were made, though, as the nature of rhizoids in the latter species was unknown. Re- cently, Womersley (1979) examined material identified as Polysiphonia sertularioides from the Mediterranean and found it to have rhizoids cut off from the proximal end of peri central cells, thus clearly distinguishing this taxon from P. havanensis. Although Polysiphonia havanensis sensu Borgesen is now seen to be a distinct entity, its taxonomic validity remains in question. Ardre (1970) examined the type specimens of Montagne for P. havanensis and found them to correspond to "P. macrocarpa Harvey." Since the Montagne specimens were sterile, she was unable to make exact determination but noted that Montagne material did not compare favorably with P. havanensis as described by Borgesen (1918). Specimens from the present study are referred to Polysiphonia havanensis until our investigation of the validity of Montagne's taxon is complete.

Polysiphonia howei Hollenberg in Taylor Figures 21-24

P. hOlVei Hollenberg in Taylor 1945: 302. Description. - Plants small, up to 2 cm tall from a spreading prostrate system; decumbent filaments 100-150 J.Lm diam; prostrate axes with 10-12 pericentral cells around a relatively large central cell, the pericentral cells shifting to offset position and forming secondary pit connections with the two pericentral cells of adjacent segments; erect branches arising exogenously at intervals of3-6 segments, first strongly curved toward the apex of prostrate branches; branches in erect filaments replacing trichoblasts in development; trichoblasts highly branched with characteristically short basal cells; rhizoids cut off from the proximal or distal end of pericentral cells, often several per segment; tetrasporangia in short spiral se- quence in the tips of little swollen erect branches; spermatangial branches 50 X 120-150 J.Lm, on 2-celled stalks, with or without sterile tip cells, lacking subtending trichoblasts. Type- Locality. - Berry Islands, Bahamas. Distribution. - Temperate and tropical western Atlantic (Hollenberg, 1958; Tay- lor, 1969; Taylor and Bernatowicz, 1969; Richardson, 1975; Bird and McIntosh, 1979; Kapraun, 1980a) and Pacific Oceans (Hollenberg, 1968b). This species is previously unreported from the study area. KAPRAUN ET AL.: POLYS/PHON/A IN THE WESTERN ATLANTIC 891

Figures 21-24. Polysiphonia howei. Fig. 21. Tetrasporangia. Fig. 22. Horizontal apex. Fig. 23. Sper- matangial branches. Fig. 24. Erect branch apex. Figs. 21, 22, 24: Scale = 100 ILm. Fig. 23: Scale = 50 ILm.

Specimens Studied.-Venezuela: Piedras de Soro, near Punta Piedras (Lemus, 28 Sept. 1975, WNC 14388). Punta Sabaneta (Lemus, 28 Sept. 1975; WNC 14389). Remarks. -Hollenberg (1958, 1968b) discussed the confusion over the relationship between Polysiphonia howei and Lophosiphonia obscura. The genus Lopho- siphonia, as currently circumscribed, includes plants with chiefly prostrate habit, endogenous origin of erect branches, and dorsiventral or bilateral apex development (Fig. 43) (Hollenberg, 1968a; Rueness, 1971). Hollenberg (1958) examined New World collections attributed to Lophosiphonia obscura and found them without exception to have exogenous branch development and spiral arrangement of branches and trichoblasts in erect filaments, and concluded that these specimens were Polysiphonia howei. He further remarked that it was doubtful if any western hemisphere plants previously identified with Lophosiphonia obscura (c. Ag.) Falk. would prove to be correctly determined. Consequently, it is unfortunate that Chapman (1963) included this taxon in his algal flora of Jamaica, and with a description apparently based on two separate entities, one with 6 and the other with 12-18 pericentral cells. Lophosiphonia obscura, which has 6 pericentral cells, is known from the Med- iterranean (Hollenberg, 1958) and adjacent Atlantic Europe (Ardre, 1970). How- ever, material from the Canary Islands attributed to this taxon by Bl2lrgesen(1930) is described as having trichoblasts in spiral arrangement and so clearly is not a Lophosiphonia. Since the number of pericentral cells was not indicated, no con- jecture can be made as to whether or not this material too belongs to Polysiphonia howei without a critical examination of the specimens. Two species of Polysiphonia that occur in the American tropics could be con- fused with P. howei: P. hemisphaerica var. boldii and P. exilis. Polysiphonia 892 BULLETIN OF MARINE SCIENCE. VOL. 33. NO.4. 1983

Figures 25-29. Polysiphonia eastwoodae. Fig. 25. Spermatangial branch. Fig. 26. Tetrasporangia. Fig. 27. Apex showing branch replacing trichoblasts in development. Fig. 28. Decumbent branch showing rhizoids cut off from pericentral cells. Fig. 29. Lateral branch. Figs. 25-29: Scale = 50 lim.

hemisphaerica var. boldii has 6 (-7) pericentral cells and so can be readily distinguished upon microscopic examination. Polysiphonia exilis has 8-10 pericentral cells, but differs from P. howei in several respects; P. exilis has rhizoids developing from the middle of pericentral cells, the pericentral cells in prostrate axes are in longitudinal rows rather than offset, and erect branches are straight rather than curved forward toward the apex (Hollenberg, 1968b). KAPRAUN ET AL.: POU'SIPI/ONI,·I IN THE WESTERN ATLANTIC 893

Figures 30-33. Polysiphonia sphaerocarpa. Fig. 30. Tetrasporangia. Fig. 31. Pericarp. Fig. 32. De- cumbent branch showing rhizoids cut off from peri central cells. Fig. 33. Branch apex. Figs. 30, 31, 32: Scale = 100 jlm. Fig. 33: Scale = 50 jlID.

Polysiphonia sphaerocarpa Bf2Jrgesen Figures 30-33

P. sphaerocarpa Bergesen 1918: 271. Description. -Plants small, epiphytic, to 1.5 cm tall from a discoid base; branches replacing trichoblasts in development; branches dichotomous, to 120 ,urn diam, segments 1-2 diam long; older branches becoming decumbent, attached to the substratum by rhizoids cut off from pericentral cells; pericentral cells 4; pericarps spherical, about 250 ,urn diam, with a flared ostiole of large cells; tetrasporangia in swollen spiral series in branch tips. Type-Locality.-Store Nordsidebugt, St. Thomas, U.S. Virgin Islands. Distribution. - Widespread in the tropical western Atlantic (Taylor, 1960, 1969; 894 BULLETIN OF MARINE SCIENCE. VOL. 33. NO.4. 1983

Figures 34-38. Polysiphonia subtilissima. Fig. 34. Tetrasporangia. Fig. 35. Branch apex. Figs. 36- 38. Variation in rhizoid development. Figs. 34, 36-38: Scale = 100 !Lm. Fig. 35: Scale = 50 !Lm.

Figures 39-40. Polysiphonia denudata. Fig. 39. Tetrasporangia. Fig. 40. Branch apex. Fig. 39: Scale = 100 !Lm. Fig. 40: Scale = 50 ,urn. Figures 41-42. Polysiphonia atlantica. Fig. 41. Branch apex with radial development oftrichoblasts. Fig. 42. Horizontal apex with unilateral branch development. Note rhizoids in open connection with pericentral cells. Fig. 41: Scale = 100 /lm. Fig. 42. Scale = 50 !Lm.

Taylor and Rhyne, 1970; de Rios, 1972; Kapraun, 1977b; Kapraun and Norris, 1982) and Pacific Oceans (Hollenberg, 1968b).

Specimens SlUdied.- Venezuela: Bahia de Mochima (Lemus, 3 July 1979, WNC 14391). Punta Piedras (Lemus, 28 Sept. 1975, WNC 14392). KAPRAUN ET AL.: I'OU"SII'IIO.\'I.·IIN THE WESTERN ATLANTIC 895

Figure 43. Lophosiphonia cristata. Horizontal apex with unilateral branch development and erect filaments with unilateral trichoblast development. Scale = 100 11m.

Polysiphonia subtilissima Montagne Figures 34-38

P. Sublilissima Montagne 1840: 199. Description.-Plants olive to purple, forming dense tufts to 4 cm tall; erect and prostrate axes with distinctive radial development of branch primordia; erect filaments subdichotomous; trichoblasts scarce with few dichotomies; branches replacing trichoblasts in development; pericentral cells 4, segments 1-2 diam long, to 150 .urn diam; prostrate axes 80-100 .urn diam; rhizoids developing from the middle of pericentral cells and remaining in open connection with them; tetrasporangia swollen, in long straight series. Type- Locality. -Cayenne, French Guiana. Distribution. - Tropical and subtropical western Atlantic (Taylor, 1928, 1960; 1969; Richardson, 1975; Oliveira Filho and Ugadim, 1976; Kapraun, 1979b; 1980a; Kapraun and Norris, 1982) and Pacific Oceans (Dawson et aI., 1964; Hollenberg, 1968a; Womersley, 1979). This species is previously unreported from the coasts of Venezuela and Colombia.

Specimens Studied.- Venezuela: Macuro (Lemus, 17 June 1976, WNC 14393). Punta Piedras (Lemus #14 & 15, ]7 June 1976). Patao (Lemus, 19 Sept. 1975, WNC 14394). Colombia: Cienaga Grande de Santa Marta (Bula-Meyer, 3 Oct. 1977, WNC 12157). Santa Marta (Bu]a-Meyer, ]5 Aug. 1975, WNC 12155). Ensenada de Nenguange (Bula-Meyer, 15 Sept. 1977, WNC ]2156). Remarks. - Womersley (1979) considers that Polysiphonia subtilissima and P. urceolata (Lightfoot) Greville, along with P. abscissa Hooker and Harvey and P. pacifica Hollenberg, comprise a complex of closely related taxa. Although no assessment of this claim can be made in the case of the two Pacific taxa, P. abscissa and P. pac{fica. the European-North Atlantic P. urceolata, itself an ill-defined complex (Kapraun, 1979a), is morphologically and physiologically distinct from 896 BULLETIN OF MARINE SCIENCE. VOL. 33. NO.4. 1983

New World P. subtilissima (Kapraun, 1980b). It should be further noted that the P. subtilissima in the western Atlantic has spermatangial branches replacing the entire trichoblast and lacks sterile tip cells (Kapraun, 1980b), in contrast to the description given for south Australian material which reportedly has spermatangial branches with prominent tips of 4-6 cells (Womersley, 1979).

LITERATURE CITED

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DATEACCEPTED: August 16, 1982.

ADDRESSES:(D.F.K.) Department of Biology. University of North Carolina, Wilmington, North Car- olina 28406; (A.J.L.) Departamento de Biologfa Marina, lnstituto Oceanografico, Universidad de Ori- ente, Cumana, Venezuela; (G.B.M.) Departamento de Biologfa, Universidad Tecnologica del Magda- lena. Santa Marta, Magdalena, Colombia.