New Records and Notes on Hawaiian Marine Benthic , including Pseudochlorodesmis abbreviata (Gilbert), n. comb. (Udoteaceae) and Cladophora luxurians (Gilbert), n. comb. (Cladophoraceae)l

Isabella A. Abbott and John M. Huisman 2

Abstract: Morphology, , and nomenclature of three species of Ha­ waiian (Chlorophyta) are examined. Udotea? abbreviata Gilbert is shown to be incorrectly placed in that genus and more appropriately allied to Pseudochlorodesmis. The complex nomenclatural relationships ofCladophora tilden­ iae Brand in Tilden, Cladophora tildeniae Brand, and Cladophora hawaiiana Til­ den are described, with the latter deemed the appropriate name and Microdietyon japonicum var. laxum Gilbert regarded as a synonym. An examination of Cla­ dophoropsis luxurians Gilbert has shown it to have delayed formation oftransverse walls at the bases of lateral branches, a feature not consistent with inclusion in Cladophoropsis but rather with Cladophora. The new combinations Pseudochloro­ desmis abbreviata (Gilbert) Abbott & Huisman and Cladophora luxurians (Gilbert) Abbott & Huisman are made, and nine species of marine benthic Chlorophyta are newly recorded for the Hawaiian Islands.

THE MARINE BENTHIC algal flora of the Ha­ MATERIALS AND METHODS waiian Islands has, historically and recently, received a great deal of attention, both from were preserved either in formalin! indigenous Hawaiians as a source offood and seawater or as dried herbarium specimens from phycologists as constituting a particu­ and on microscope slides. Materials for slide larly rich biodiversity and as a source ofmany preparations were stained with aniline blue interesting and novel taxa. A recent outcome and mounted in water for photography, and of this attention was the publication of Ma­ then infused with corn syrup (Karo, CPC International) solution to make permanent rine of the Hawaiian Islands (Abbott 1999), the first major algal flora for the cen­ mounts. Pyrenoids were observed by staining tral Pacific and the first of a series that even­ with Lugol's solution. Specimens were exam­ tually also will include the brown and green ined with a microscope (Olympus BX3) and seaweeds. photographed using a digital camera (Olym­ During preparatory research for the pus D11). Herbarium abbreviations follow Chlorophyta volume we encountered three Holmgren et al. (1990) with the exception of species with a confused nomenclatural and lA, which refers to the 1. Abbott collection housed at the Department of Botany, Uni­ taxonomic past, as well as documented nine new records for the Hawaiian Islands, all of versity of Hawai'i at Manoa. which we report here. RESULTS Pseudochlorodesmis abbreviata (Gilbert) Abbott

I Research supported by the David and Lucile Pack­ & Huisman, n. comb. ard Foundation. Manuscript accepted 17 October 2002. Basionym: Udotea? abbreviata Gilbert, Pac. Sci. 2 Department of Botany, University of Hawai'i at 16: 142, fig. 8a-c (1962). Manoa, 3190 Maile Way, Honolulu, Hawai'i 96822. TYPE SPECIMEN: Gilbert 9511, on cal­ careous rocks, 1-1.5 m below surface on reef Pacific Science (2003), vol. 57, no. 3:275-285 opposite Marine Biological Laboratory, Wai­ © 2003 by University of Hawai'i Press kiki, Honolulu, O'ahu, 21 April 1959 (n.v., All rights reserved fide Gilbert 1962). This specimen cannot be

275 276 PACIFIC SCIENCE· July 2003 located and in its absence we lectotypify the multistratose blades, with lateral appendages species with Gilbert's fig. 8a-c (reproduced as on the siphons of many blades and all stalks our Figure 1). (Littler and Littler 1990), and, for the reasons SPECIMENS EXAMINED: Moloka'i: Kalau­ just given, also does not belong in Rhipidosi­ papa, BISH 653286 (Figure 2). phon. DISTRIBUTION: Hawaiian Islands. We have examined specimens (Figure 2) HABIT AND VEGETATIVE STRUCTURE: The collected in recent years from Moloka'i Island following description is based on the recently that seem be in almost every way referable to collected specimens. Dimensions given in Udotea? abbreviata, although the dimensions parentheses are from Gilbert (1962). of the recent collections vary slightly from Plants filamentous, siphonous; with a those given by Gilbert. The upright portions prostrate basal portion of limited extent and of the plants are comparable with those de­ upright, dichotomously branched fronds in scribed by Gilbert, but the new specimens which the branches lie in one plane; prostrate have a slightly more extensive basal portion portion irregularly branched and coralloid; that is irregularly swollen and has occasional upright portion to 5 mm (5 mm) tall, with a constrictions. monosiphonous stalk to 2.6 mm in length and Two species, in separate genera, are rele­ 75-135 (72) /lm diam., with upper siphons vant to a consideration of the proper place­ 45-65 /lm diam., 45-50 (25-38) /lm diam. ment of U? abbreviata. Its basal portion is near apices; branching above strictly dichoto­ strongly reminiscent of that found in Pseu­ mous, with 100-650 /lm between successive dochlorodesmis parva Gilbert (1962), the type dichotomies; chloroplasts numerous, ovoid to specimen of which was collected from the lenticular, approximately 5 by 2 /lm. same locality as U? abbreviata. The genus REMARKS: Gilbert (1962) described Udo­ Pseudochlorodesmis was originally described tea? abbreviata from specimens collected from by B0rgesen (192 5:77) for P. furcellata (Za­ WaikIkI, O'ahu Island. The plants were si­ nardini) B0rgesen, based on Bryopsis furcellata phonous, with an irregularly branched basal Zanardini (1843) from the Mediterranean, portion and upright fronds consisting of a although B0rgesen based the genus concept monosiphonous stipe and dichotomously di­ on material from the Canary Islands. Defin­ vided upper portion, with the forks all lying ing characters of the genus include a basal in one plane (Gilbert 1962: fig. 8a-c, repro­ portion that is irregularly swollen and with duced here as Figure la-c). Gilbert was constrictions, and an upright frond that lacks clearly uncertain as to the correct generic constrictions at the infrequent branch points. placement of this species, comparing it first These features are also found in U? ab­ with representatives of Flabellaria (= Udotea) , although it differs in that the up­ and Udotea before settling on the latter with right siphons are distinctly dichotomously "real uncertainty" (Gilbert 1962:143) and branched and the constrictions are not noting in particular that the basal stalk and present at the bases of the upright fronds. branching in one plane were similar to fea­ However, Segawa (1941) reported regularly tures of Udotea javensis (Montagne) Gepp & dichotomously branched P. furcellata from Gepp (now Rhipidosiphon javensis Montagne). Japan, and Kraft (2000) recorded the same The calcified thallus and constricted siphons species from Lord Howe Island, southwest­ of U javensis, however, were sufficiently dif­ ern Pacific, that also seemingly lacked basal ferent in his opinion to warrant the descrip­ constrictions. tion of a new species for the WaikIkI plants. The dichotomously branched siphons of Udotea? abbreviata has not been reported since U? abbreviata are also strongly reminiscent of its original collection and thus has not been those of Derbesia? fastigiata Taylor (1928:94, critically appraised; the apparent loss of type plate 11: figs. 1-3), a species originally de­ material is an additional difficulty. The spe­ scribed from Florida but also reported from cies is clearly incorrectly placed in Udotea, a several other tropical locations, including genus whose members are calcified and have Hawai'i (Gilbert 1962) and the Southwest Hawaiian Chlorophyta . Abbott and Huisman 277

1 2

a c

FIGURES 1-2. Pseudochlorodesmis abbreviata (Gilbert) Abbott & Huisman, n. comb. 1. Gilbert's illustrations from the protologue: a,b, Upper and lower portions of the same . c, Upper portion of a plant to illustrate that dichotomies may be close or distant (caption from Gilbert 1962); scale = 1 mm. 2. Comparable specimen from Moloka'i, BISH 653286; scale = 500 Ilm.

Pacific Ocean (Silva et al. 1996, Phillips ity that U. abbreviata represents a growth 1997). This species was included in Derbesia form ofP. fureellata, but until more specimens with some doubt by Taylor (1928). Derbesia can be examined we prefer to keep them fastigiata differs from u.? abbreviata in form­ separate. ing an extensive prostrate system composed ofinterwoven siphons that are not irregularly Cladophora hawaiiana Tilden swollen (Gilbert 1962). The genus Derbesia American algae 7, nos. 615, 616 (1909). differs from Pseudochlorodesmis in the lack of Synonyms: Cladophora tildenii Brand, Beih. amyloplasts (colorless plastids) in the former Bot. Centralbl. 18: 186-187, plate VI, figs. and their presence in the latter, but we were 24-27 (1904); nom. inval., non C. tildenii unable to ascertain with certainty whether the Brand in Tilden, American algae 6, no. 541 recent specimens of u.? abbreviata contained (1902) [= C. canalicularis (Roth) Kiitzing, amyloplasts. Some colorless plastids were ob­ fide Tilden 1909]. served, but these were mixed among more Microdictyon japonicum var. laxum Gilbert, numerous green plastids and could possibly Pac. Sci. 16: 139, figs. 5C,D; 6 (1962). be artifacts of preservation. LECTOTYPE: Brand, 1904, plate VI, figs. From our examination of u.? abbreviata, 24-27 (reproduced as our Figure 3). we have concluded that its uncalcified thallus SPECIMENS EXAMINED: O'ahu: Ka'ena and unadorned, dichotomous siphons pre­ Point, BISH 651438; Keawa'ula (Yokohama clude its placement in Udotea, and we feel that Bay), BISH 653055; Lana'i lookout, Halona, the species is more appropriately placed in BISH 648526. Moloka'i: Kalaekapu Point, Pseudochlorodesmis. There is a strong possibil- BISH 628824; Moku Islet site LVII, BISH 3

5 '

6 Hawaiian Chlorophyta . Abbott and Huisman 279

625020; Kalaupapa, north of lighthouse, BISH features that he considered diagnostic of the 653276. Hawai'i: Lili'uokalani Gardens Pond, species: (1) a fan-shaped thallus; (2) conspic­ BISH 636077. uous rhizoids that are not produced laterally DISTRIBUTION: Hawaiian Islands. from the axis but as basipetal projections from HABIT AND VEGETATIVE STRUCTURE: Plants lower cells of the main axis; and (3) scattered 1.5-3 em tall, dark green, with several fanlike adhesions of apical cells to lower-order fronds arising from a common rhizoidal branch cells, as in Microdietyon. Unfortu­ holdfastj each frond with lower cells bearing nately, none of these features is displayed by strong, nonseptate, uniseriate rhizoids (Fig­ the specimens that were circulated in Tilden's ure 5) from the basal poles, these intertwining American algae exsiccatae (as no. 541) under and directed downward for 3-4 mm; lower the name of Cladophora tildenii Brand. In fact, axial cells cylindrical, 80-200 Ilm diam., this number represents a freshwater species of length: width = 2-6: 1, with apical ends fre­ Cladophora collected from Pearl City, O'ahu, quently widening and becoming clavate (Fig­ that Brand (1904) designated as Cladophora ure 3); upper cells of axes 50-80 Ilm diam.j (Spongomorpha) longiarticulata Nordstedt var. apical cells with broadly rounded ends. valida Brand but that was later identified by Branching opposite or flabellate, with most Tilden as C. canalicularis (Roth) Klitzing, a nodes initially bearing 2 or 4 opposite late­ species that is currently considered to be a ral branches (Figures 3, 4), these eventually synonym of C. glomerata (Linnaeus) Klitzing becoming indistinguishable from primary (Guiry and Nic Dhonncha 2002). A specimen axes belonging to a flabellate node of five of this entity from Tilden's collection (no. branches; lateral branches bearing 1- to 4­ 541) is preserved in the Bishop Museum (no. celled branchlets either adaxially or abaxially, 531869), Honolulu, and has been examined. with each of the penultimate orders bearing It is a sparsely branched plant, with sub­ 1- to 2-celled branchlets, either oppositely or dichotomies every 1-9 cells; the cells are occasionally secundly. Intercalary divisions in 270-540 Ilm long by 135-160 Ilm diam. and primary axes present, resulting in internodes tapering to 45 Ilm diam. near the apex. The 2 cells long, but these obscured by rapid ini­ morphology of no. 541 is thus clearly not the tiation of adventitious laterals that often re­ same as that described by Brand for Clado­ main short, 1-2 cells long, with those in the phora tildeniae (1904), and the specimen is inner, lower portions of some plants occa­ probably correctly identified as C. glomerata, a sionally forming terminal adhesions to a cell species that is common in Hawaiian streams of an adjacent branch (Figure 6). (A. R. Sherwood, pers. comm.). We conclude REPRODUCTION: Brand (1904) illustrated therefore that Cladophora tildeniae Brand in fertile zooidangia, each with a raised, distal Tilden (1902) is not the same as Cladophora pore (see our Figure 3). Identical reproduc­ tildeniae Brand (1904). Moreover, because tive structures were observed in several Cladophora tildeniae Brand (1904) is a later specimens (e.g., BISH 653276, BISH 648426) homonym of Cladophora tildeniae Brand in from recent collections. Tilden (1902), the name has to be rejected. REMARKS: The name Cladophora tildeniae Tilden realized that an error had been Brand (as "tildenii") first appeared in Tilden's made, and in Fascicle 7 of American algae American algae, no. 541 (1902), but it was 2 yr (1909) she placed C. tildeniae Brand (1904:86, later before a complete description was given. referred to as "in manuscript" by Tildenj non Brand (1904:186, figs. 24-27, reproduced as C. tildeniae Brand in Tilden 1902) in synon­ our Figure 3) highlighted the following three ymy with her new Cladophora hawaiiana Til- .. FIGURES 3-6. Cladophora hawaiiana Tilden. 3. The lectotype illustrations, reproduced from Brand (1904). 4. Apex of plant showing initiation of lateral branches, BISH 648426; scale = 500 11m. 5. Descending rhizoids forming a dense, entwined mass, BISH 648426; scale = 250 !!ill. 6. Lateral branches forming attachments with nearby filaments, BISH 625020; scale = 250 !!ill.

Hawaiian Chlorophyta . Abbott and Huisman 281 den (nos. 615-616). Although not explicitly dophora hawaiiana. In addition, the lateral stated, the implication of Tilden's treatment branches of Rhipidophyllon reticulatum appar­ of this species is that C. hawaiiana was a sub­ ently always arise in one plane (B0rgesen stitute name for C. tildeniae sensu Brand. The 1924:252), whereas occasional branches of C. type of this species is therefore the Brand hawaiiana do not. specimens, but unfortunately these seem to Cladophora hawaiiana also displays many of be lost, so here we lectotypify the species with the features of the widespread Willeella ordi­ Brand's figures (1904, figs. 24-27, repro­ nata B0rgesen, 1930 (see Itono 1970, van den duced as our Figure 3), which are sufficiently Hoek and de Rios 1972). Willeella was re­ detailed to characterize the species. duced to a (possibly monospecific) section of To further complicate matters, the speci­ Cladophora by van den Hoek (1982), but re­ mens distributed by Tilden as C. hawaiiana cently Silva et al. (1996) maintained it at the (no. 615) do not agree with Brand's descrip­ genus level, in anticipation of future molecu­ tion (P. Silva, pers. comm.), and their identity lar data that they suggest will support the is yet to be ascertained. These specimens recognition of segregate genera. Similari­ were collected at La'ie, O'ahu Island, a ma­ ties between the two species include their rine locality. branching pattern and the production of Two additional taxa are relevant to a con­ closely adpressed, descending rhizoids from sideration of C. hawaiiana. The first, Miero­ the bases of lower lateral branches. However, dictyon japonicum var. laxum, was described by whereas Cladophora hawaiiana does produce Gilbert (1962:139, figs. 5C,D; 6C) for speci­ several laterals of equal length from each mens from Maui, Hawaiian Islands. Gilbert's node, these do not arise synchronously (as do variety is considered by us to represent C. those of Willeella). Kraft and Millar (in Kraft hawaiiana. Although the planar branching 2000:548) recently described a species of and secondary adhesions of this variety un­ Cladophora (c. willeelloides Kraft & Millar) that doubtedly led Gilbert to place it in Miero­ has characteristics similar to those of C. ha­ dictyon, we feel that the rarity of secondary waiiana, and we follow their interpretation adhesions more strongly allies it to Clado­ that such plants should be maintained in Cla­ phora. A second species with some similarities dophora. In addition to the nonsynchronous to C. hawaiiana is Rhipidophyllon reticulatum production of laterals, those authors high­ (Askenasy) Heydrich (type locality: Dirk lighted several additional characters of C. Hartog Island, Western Australia). B0rge­ willeelloides that differed from Willeella ordi­ sen's (1924) depiction of this species from nata, including paired, nonrostrate ostioles Easter Island, however, indicates that the lat­ in zooidangia. Zooidangia in Cladophora ha­ eral filaments are issued from the apices of waiiana, as in C. willeelloides, produce raised the subtending cells and are always upwardly pores, further reinforcing the distinction be­ directed, unlike the spreading laterals of Cla- tween the two species and Willeella. Clado- .. FIGURES 7-19. 7. Herbarium specimen of Cladophora luxuriam (Gilbert) Abbott & Huisman, n. comb., BISH 515693; scale = 1 cm. 8. Cladophora luxuriam, detail showing initiation oflateral branches and delayed formation of cross-walls; scale = 2 mm. 9. Ulvella lem P. Crouan & H. Crouan, showing rougWy circular outline (specimens depicted in Figures 9-13 are all growing on the surface of Rhizocloniztnz grande, BISH 771278); scale = 40 /lm. 10. Ulvella setchellii Dan­ geard, with elongate cells; scale = 40 /lm. 11. Uronema marina Womersley, showing unbranched arcuate filament; scale = 40 /lm. 12. Stromatella monostromatica (Dangeard) Kornmann & Sahling, with filaments coalescing in the center but free at margins; scale = 40 J.lffi. 13. Entocladia viridis Reinke, with free filaments and cells with pyrenoids; scale = 40 /lm. 14. Ulvo toenioto (Setchell) Setchell & Gardner, showing milled margins of thallus, IA 17613; scale = 2 cm. 15. Codium pictumtwn F. F. Pedroche & P. C. Silva, growing closely adpressed to the substratum, IA 28794; scale = 2 mm. 16. Codium picturotzlm utricles, IA 28794; scale = 100/lm. 17. Derbesio tenuissimo (Moris & De Notaris) Crouan, thallus habit, IA 28048; scale = 300 /lm. 18. Derbesio tenuissimo, stained with Lugol's solution, showing pyrenoids in the plas­ tids, IA 28048; scale = 40 J.lffi. 19. Derbesio tC1Zuissimo, detail of gametangium with double septa at base, IA 28048; scale = 100 /lm. 282 PACIFIC SCIENCE· July 2003

phora hawaiiana differs from C. willeelloides in cell. An examination of C. luxurians reveals the presence of occasional anastomoses be­ that, although initially in open connection to tween branches and absence of long inter­ the bearing cell, cross-walls are subsequently nodes. Intercalary divisions in C. hawaiiana formed at the bases of lateral branches are followed rapidly by the initiation oflateral (Figure 8), a feature also illustrated by Gil­ branches, such that internodes are generally bert (1962: fig. 3) and by Brand (1904) in only one-celled (rarely up to four cells long); his description of C. montagnei var. waia­ in contrast, internodes in C. willeelloides can naeana, which we consider to be synonymous be up to 14 cells long (Kraft 2000). with C. luxurians. Van den Hoek and Womersley (1984) reported that in many Cladophora luxurians (Gilbert) Abbott & species of Cladophora section Repentes cross­ Huisman, n. comb. wall formation is delayed, as is also the case in Basionym: Cladophoropsis luxurians Gilbert, C. luxurians. Pac. Sci. 16: 136 (1962). Cladophoropsis luxurians seems to be closely Synonym: Cladophora (Aegagropila) montagnei related to Cladophora catenata (Linnaeus) Kiitzing var. waianaeana Brand, Beih. Bot. Kiitzing as described and illustrated by van Centralbl. 18: 185-186, plate V, figs. 21­ den Hoek and Chihara (2000: fig. 18). That 23 (1904). species also forms dense cushions, with stiff, SPECIMENS EXAMINED: Kaua'i: KIpl1 Kai, branched, often curved stolonlike filaments. BISH 531172. O'ahu: Kane'ohe Bay, BISH Secondary rhizoids and attachments are also 525117; Ka'ena Point, BISH 516991. Molo­ formed at the apices. Cladophora catenata ap­ ka'i: (as C. luxurians) BISH 515671. Maui: pears to differ from C. luxurians in the smaller (Gilbert as C. luxurians) BISH 515693; Hana diameter of cells, with those of the former Bay, IA 14564. reaching a maximum diameter of 510 J..lm (van DISTRIBUTION: Hawaiian Islands; possi­ den Hoek and Chihara 2000:46), whereas bly more widespread. cells of C. luxurians can be up to 700 J..lm HABIT AND VEGETATIVE STRUCTURE: Plants diam. Payri et al. (2000) recorded Cladophor­ in mostly decumbent clumps 1-5 cm across opsis luxurians from French Polynesia, but (Figure 7), composed of coarse, branched fil­ their description indicates that cross-walls aments; branching sparse or profuse, usually are not formed and thus does not agree secund and irregular; cross-wall formation at with C. luxurians as known from Hawaiian base of laterals delayed (Figure 8); apices collections. Hawaiian specimens clearly re­ often becoming decumbent and reattached, ferable to C. luxurians were also identified as some with small specialized hapteroid cells at Cladophora montagneana var. waianaeana by apices; filaments (250) 400-700 J..lm diam.; Brand (1904), but the species is not at all rhizoids, where developed laterally, about similar to Cladophora montagneana Kiitzing one-half the diameter of adjacent branch, [as depicted by van den Hoek and W omers­ long, mostly nonseptate but with basal cross­ ley (1984)] and that placement cannot be walls. supported. Our results indicate that C. lux­ REMARKS: Cladophoropsis luxurians is a urians cannot be maintained in Cladophoropsis common and conspicuous species on Ha­ and is more appropriately allied with Clado­ waiian reef flats, where it forms mats of phora. imbricating, bright green filaments. Char­ acteristic features of the species include its New Records ofChlorophyta for the Hawaiian arcuate filaments with often-secund lateral Islands branches (Figures 7, 8) and the formation of secondary attachments at the apices of The following species are newly recorded for branches. The species does not belong in the Hawaiian Islands. In some cases they Cladophoropsis, however, in which the defining represent widespread but cryptic species, feature is the persistent open connection be­ generally only noticed during microscopical tween lateral branches and the subtending examinations of larger algae. Hawaiian Chlorophyta . Abbott and Huisman 283

Ulvella lens P. Crouan & H. Crouan several pyrenoids (as opposed to one) per Ann. Sci. Nat. Bot. 12: 288 (1859). plastid. Figure 9 DISTRIBUTION: Widespread in tropical Ulva taeniata (Setchell) Setchell & Gardner and warmer seas. Univ. Calif. Publ. Bot. 8: 286 (1920). Figure 14 Ulvella setchellii Dangeard SPECIMENS EXAMINED: Kaua'i: Po'ipl1 Bull. Soc. Bot. Fr. 78: 318 (1931). Beach, IA 13514. Maui: Hina Bay, IA 14568. Figure 10 DISTRIBUTION: Widespread in the Pacific DISTRIBUTION: Widespread in tropical to and Indian Oceans. colder seas. REMARKS: Ulva taeniata is a distinctive species with narrow, straplike branches and Uronema marina W omersley conspicuously ruffled margins. Although the Marine Benthic Flora Southern Australia 1: 131 common (in the Hawaiian Islands) Ulva fas­ (1984). ciata has similarly narrow branched axes and Figure 11 may appear to have ruffled margins, the latter DISTRIBUTION: Hawai'i; Australia. feature is not characteristic of the majority of populations; moreover, the bases of U. fasciata Stromatella monostromatica (Dangeard) Korn­ are formed by a broadened portion (the rem­ mann & Sahling nants of the original flat blade when the plant Helgol. Meeresunters. 36: 12 (1983). was young) from which the divided portions Figure 12 project like fingers, whereas in U. taeniata DISTRIBUTION: Widespread in tropical to there are no broadened portions of the single colder seas. blades, nor are there any unruffled margins.

Entocladia viridis Reinke Cladophoropsis sundanensis Reinbold Bot. Z. 37: 476 (1879). Nuova Notar. ser. 16, p. 147 (1905). Figure 13 SPECIMENS EXAMINED: Kaua'i: KIpl1 Kai, DISTRIBUTION: Widespread; recorded BISH 53113. O'ahu: Waikiki, IA 19812. Mo­ from the Tropics to polar regions. loka'i: BISH 91964. Kaho'olawe: Pu'u Koa'e, REMARKS: These five species form crusts IA 21116a. Maui: Ka'uiki Head, IA 18027. or filaments that grow on or course through Hawai'i: South Kona, BISH 515727. the cuticles of larger, generally green, algae. DISTRIBUTION: Widespread in the Pacific They are probably widespread but are com­ and Indian Oceans. monly overlooked due to their habitat and REMARKS: The specimens reported here microscopic size. These records are of speci­ are included in Cladophoropsis sundanensis mens observed on a single host plant, Rhizo­ based on their dimensions (with filaments clonium grande B0rgesen (BISH 771278), but generally 80-100 /lm in diam.) and thallus the species are known to occur on a variety structure. In Cladophoropsis the lateral of hosts (Kraft 2000). Ulvella lens (Figure 9) branches remain in open connection with and U. setchellii (Figure 10) form roughly cir­ the bearing cell (i.e., no cellular cross-wall cular thalli on the surface of the host cuticle, is formed). Cladophoropsis sundanensis is com­ differing in the radially elongate cells of the monly reported in the Indo-Pacific, but Kraft latter. Uronema marina (Figure 11) occurs (2000) questioned whether many of these as short, unbranched filaments. Stromatella records refer to the same species because not monostromatica (Figure 12) has filaments co­ all agree with the type description. alesced near the center but free at the mar­ gins, and Entocladia viridis (Figure 13) has Codium picturatum F. F. Pedroche & P. C. mostly free filaments that grow within the Silva cuticle of the host. In addition, Entocladia Acta Bot. Mex. 35 :2, figs. 1-5 (1996). differs from Stromatella in the presence of Figures 15, 16 284 PACIFIC SCIENCE· July 2003

SPECIMENS EXAMINED: Hawai'i: Puna­ marina (without). The specimens reported lu'u, IA 28794. here were preserved in formalin and the DISTRIBUTION: Hawai'i; southern part of pyrenoids were not obvious, but staining with Gulf of California; tropical Pacific Mexico; Lugol's solution clearly showed pyrenoids Panama; Colombia; Galapagos. within each plastid (Figure 18). The plastids REMARKS: Codium picturatum was de­ were smaller than generally described, but scribed by Pedroche and Silva (1996) for this is most likely due to preservation, which plants from the Pacific coast of tropical can cause the plastids to become rounded America. It is a distinctive species due to its when they are usually lenticular (Womersley extremely thin crustose habit (Figure 15), 1984). Other vegetative and reproductive unusual utricle morphology (Figure 16), and features are compatible with D. tenuissima. tortuous medullary filaments. The Hawaiian plants were closely adpressed to basalt rock at ACKNOWLEDGMENTS less than 1 m depth and could easily have been overlooked or mistaken for the more We thank Paul Silva (University of California common Codium arabicum Ktitzing. The at Berkeley), who identified the specimen of identity of the Hawaiian specimens was as­ Codium picturatum, and Lynn Hodgsen (Uni­ certained by P. Silva (pers. comm., 2002). versity of Hawai'i), who collected it. Gerry Kraft (University of Melbourne) critically Derbesia tenuissima (Moris & De Notaris) read the manuscript and suggested numerous Crouan improvements. Fl. Finist., p. 133 (1867). Figures 17-19 Literature Cited SPECIMENS EXAMINED: Maui: Makena, IA 28048. Abbott, I. A. 1999. Marine red algae of the DISTRIBUTION: Northeast Atlantic; Med­ Hawaiian Islands. Bishop Museum Press, iterranean; western Pacific Ocean; Indian Honolulu. Ocean. B0rgesen, F. 1924. Marine algae from Easter REMARKS: The specimens reported here Island. Pages 247-309 in C. Skottsberg, are included under Derbesia tenuissima due to ed. The natural history ofJuan Fernandez their similarities with the descriptions given and Easter Island. Vol. 2. 50 figs. Uppsala. by Kobara and Chihara (1981), Womersley ---. 1925. Marine algae from the Canary (1984), and Burrows (1991). The double septa Islands, especially from Teneriffe and below the sporangia (Figure 19) are consid­ Gran Canaria. I. CWorophyceae. K. Dan. ered to be distinctive of D. tenuissima accord­ Vidensk. Selsk. BioI. Medd. 5 (3): 1-123, ing to Womersley (1984), but various authors 49 figs. have also included specimens displaying such ---. 1930. Some Indian green and brown a condition in the closely related Derbesia algae especially from the shores of the marina (Lyngbye) Solier (e.g., Kraft 2000) Presidency of Bombay. J. Indian Bot. Soc. and thus the character is not regarded as 9:151-174. critical in species determination. W omersley Brand, F. 1904. Uber die Anheftung der (1984) regarded the presence of double septa Cladophoraceen und tiber verschiedene at the bases ofvegetative laterals as distinctive polynesische Formen dieser Familie. Beih. of D. marina, and such septa occasionally are Bot. Centralbl. 18 (Abt. 1): 165-193, plates present in the Hawaiian plants, which there­ V,VI. fore display apparently characteristic features Burrows, E. M. 1991. Seaweeds of the British of two species. Both Kobara and Chihara Isles. Vol. 2. Chlorophyta. Natural History (1981) and Womersley (1984), however, re­ Museum Publications, London. gard the presence or absence of pyrenoids in Gilbert, W. J. 1962. Contribution to the ma­ the chloroplast as the critical feature for sep­ rine CWorophyta of Hawaii, I. Pac. Sci. arating D. tenuissima (with pyrenoids) from D. 16:135-144. Hawaiian CWorophyta . Abbott and Huisman 285

Guiry, M. D., and E. Nic Dhonncha. 2002. Howe Island, south-western Pacific. Aust. AlgaeBase. Worldwide Web electronic Syst. Bot. 13:509-648. publication. www.algaebase.org [3 October Littler, D. S., and M. M. Littler. 1990. Sys­ 2002]. tematics of Udotea species (Bryopsidales, Hoek, C. van den. 1982. A taxonomic revi­ Chlorophyta) in the tropical western At­ sion of the American species of Cladophora lantic. Phycologia 29:206-252. (Chlorophyceae) in the North Atlantic Payri, c., A. de R. N'Yeurt, and J. Orempul­ Ocean and their geographic distribution. ler. 2000. Algae of French Polynesia. Au Verh. K. Ned. Akad. Wet. Md. Natuurk. Vent des TIes. Editions, Tahiti. Tweede Reeks Sect. 78. 236 pp. Pedroche, F. F., and P. C. Silva. 1996. Co­ Hoek, C. van den, and M. Chihara. 2000. A dium picturatum sp. nov. (Chlorophyta), taxonomic revision of the marine species una especie extraordinaria del Pacifico of Cladophora (Chlorophyta) along the tropical Mexicano. Acta Bot. Mex. 35: 1-8. coasts of Japan and the Russian Far-East. Phillips, J. A. 1997. Algae. Pages 223-240 in Nat. Sci. Mus. Tokyo, Monogr. 19:1-242. R. J. F. Henderson, ed. Queensland plants: Hoek, C. van den, and N. de Rios. 1972. Names and distribution. Queensland Her­ Willeella ordinata Boergesen, 1930 (Chloro­ barium, Department of Environment, phyceae, Cladophorales): First record for Indooroopilly, Queensland. America. J. Phycol. 8:207-208. Segawa, S. 1941. New or noteworthy algae Hoek, C. van den, and H. B. S. Womersley. from Izu I. Sci. Pap. Inst. Algol. Res. Fac. 1984. Genus Cladophora Kuetzing 1843: Sci. Hokkaido Imp. Univ. 2:251-271, 262, nom. cons. Pages 185-213 in H. B. S. plates LV-LVIII. W omersley. The marine benthic flora of Silva, P. c., P. Basson, and R. Moe. 1996. southern Australia. Part I. Government Catalogue of benthic marine algae of the Printer, South Australia. Indian Ocean. Univ. Calif. Publ. Bot. 79. Holmgren, P. K., N. H. Holmgren, and L. C. 1279 pp. Barnett. 1990. Index herbariorum. Part 1: Taylor, W. R. 1928. The marine algae of The herbaria of the world. 8th ed. New Florida with special reference to the York Botanical Garden, New York [Reg­ Dry Tortugas. Carnegie Inst. Washington num Vegetabile, Vol. 120]. Publ. 379: (v +) 219 pp. Itono, H. 1970. On the two species of genus Tilden, J. E. 1902. American algae. Century Willeella of southern Japan. Bull. Jpn. Soc. 6, nos. 501-600. Minneapolis. Phycol. 18:127-130. ---. 1909. American algae. Century 7, Kobara, T., and M. Chihara. 1981. Labora­ nos. 601-650. Minneapolis. tory culture of two species of Derbesia Womersley, H. B. S. 1984. The marine ben­ (Class Chlorophyceae) in Japan. Bot. Mag. thic flora of southern Australia. Part I. Tokyo 94:1-10. Government Printer, South Australia. Kraft, G. T. 2000. Marine and estuarine Zanardini, G. 1843. Saggio di classificazione benthic green algae (Chlorophyta) of Lord naturale delle Ficee. Venice.