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Culture studies on Pedobesia ryukyuensis (Derbesiales, ), a new record in Brazil

EDISON J. DE PAULA' AND JOHN A. WEST2

, Departamento de Botdnica et Centro de Biologia Marinha, Universidade de Siio Paulo, Caixa Postal 11461, Siio Paulo. SP, Brazil 2 Department of Botany, University of California, Berkeley, California 94720, USA

E.J. DE PAULA ANDJ.A. WEST. 1986. Culture studies on Pedobesia ryukyuensis (Derbesiales, Chlorophyta), a new record in Brazil. Phycologia 25: 482-493.

Pedobesia ryukyuensiswas collected in 1982 and 1983 from the Centro de Biologia Marinha (CEBIMAR), Sao Sebastiao, SP, Brazil and placed in unialgal culture. These isolates exhibit a direct sporophytic recycling life history typical of Pedobesia with three developmental stages: an encrusting calcified basal disc; branched rugose filaments arising from the base; and smooth filaments bearing sporangia. Comparisons of the Brazilian material with the known species of Pedobesia revealed the greatest morphological affinity with P. ryukyuensis(yamada et Tanaka) Kobara & Chihara and P. feldmannii Abelard, but difficulties in distinguishing the two entities are discussed. Although the life history is considered as a direct sporophytic type, specialized intracellular meiotic and gametic events may exist.

INTRODUCTION Codomier (1974) and Feldmann et al (1975) de­ scribed the morphology, fine structure and cell The Pedobesia was established by MacRaild wall chemistry of Pedobesia lamourouxii (J. & Womersley (1974) based on culture studies of Agardh) Feldmann from the Mediterranean. In Derbesia clavaeformis (J. Agardh) De Toni iso­ 1982 Abelard described a new species, P. feld­ lated in Australia. Pedobesia differs from Der­ mannii Abelard, from the Galapagos Islands. besia and Bryopsidella in two major ways: Chihara & Kobara (1982) and Kobara & Chihara (1) Life history: Pedobesia has a direct spo­ (1981, 1984) have also investigated the culture rophytic recycling, whereas Derbesia and and of P. lamourouxii and P. ryu­ Bryopsidellaspecies usually exhibit a hetero­ kyuensis (Yamada & Tanaka) Kobara & Chihara morphic life history with the diploid sporo­ in Japan. Womersley (1981) and Hawkes (1983) phyte as a branched filamentous stage, bearing deal with the occurrence of P. clavaeformis (J. sporangia that produce stephanokont meio­ Agardh) MacRaild & Womersley in New Zea­ zoospores, and the gametophyte as vesicular land. coenocyte (Halicystis) or a plumose branched Pedobesia is not known with assurance in the phase (Bryopsidella).However, some Derbesia western Atlantic and the first author's initial dis­ marina isolates (Kornmann 1966, 1970; Sears covery of plants in the aquarium at the Centro & Wilce 1970) also exhibit direct sporophytic de Biologia Marinha (CEBIMAR) da Universi­ recycling. For a complete discussion see Tan­ dade de Sao Paulo, and later in the field at Praia ner (1981). do Segredo, Sao Sebastiao, in the State of Sao (2) Morphology: Pedobesia has a unique dis­ Paulo in Brazil, resulted in the present paper. coid basal system with cell walls composed partially of calcium carbonate (aragonite) from which an intermediate non-calcified filamen­ MATERIALS AND METHODS tous stage arises before the upright, sporangia­ bearing filaments are produced. The first specimens were obtained as fragments Since 1974 there has been a sharp assurgency of the basal disc from an aquarium at CEBIMAR of interest in this distinctive genus. Feldmann & in early April 1982. These were cultured in 125

482 de Paula and West: Culture studies on Pedobesia 483

ml Erlenmeyer flasks with unenriched sterile sea­ cause of overgrowth by adjacent discs. These water, 20DC 16: 8 h light-dark (LD) cycle and ca. plants died in an aquarium accident in December 10-1 5/Lmol m-2 S-1 daylight fluorescent lighting. 1982, but other discs were observed in another With fortnightly changes of the medium, upright aquarium in April 1983. rugose filaments5-10 mm long were formed in In late April and early May 1983, living plants 2 months. were obtained in the lower intertidal zone of the Apices were excised to obtain unialgal cultures exposed shore at Praia do Segredo. This locality but the coccolithophorid Ochrosphaera sp. was is characterized by intense grazing by the sea a serious contaminant tending to overgrow Pe­ urchin Echinometra lucunter and molluscs, Ac­ dobesia. In June 1982, isolation efforts were maea subrugosa Orbigny and Astrea olfersii Phi­ continued using Provasoli's ES medium-PES lippi, resulting in poor algal growth except for (McLachlan 1973) diluted to Yto strength (2 mliL Paragoniolithon solubile (Foslie & Howe in Howe) sterile seawater) in 2.0 x 4.5 cm Petri dishes, 20- Adey, Townsend & Boykins. Pedobesia is re­ 40 /Lmol m-2 S-1 daylight fluorescent illumina­ stricted to crevices, Acmaea shells and shells of tion, 12: 12 LD at 23DC. Unialgal cultures were dead Balanus. In most instances it grows as small obtained only in experiments where diatoms and (5-8 mm), irregular discs over Paragoniolithon a filamentous tuft of Pedobesia were inoculated and is commonly associated with Derbesia sp., in Erlenmeyer flaskswith bubbling aeration in Halicystis sp. and Bryopsis pennata Lamouroux. 125 ml of Yto PES medium. The diatoms ap­ The discs were seen only after careful examina­ parently outcompeted the other contaminant tion of the shells under the stereoscopic micro­ Ochrosphaera. After 1 week, apices were excised scope. No adult sporangium-bearing thalli were and transferred to a fresh culture medium with found in the field despite a bimonthly survey of Ge0 (1 mg/L) to eliminate the diatoms, thus the area. 2 producing a unialgal culture of Pedobesia.

Unialgal cultures used for the life history stud­ Culture studies ies were maintained in 4.5 x 2.0 cm Petri dishes, 10-20 and 30-45/Lmol m-2 S-I, 16: 8 LD at 20- Although various conditions were tested, the 25DC, with weekly to biweekly changes of the complete life history was observed only under culture medium (PES, 1";0 to 'h strength). 10-20 and 30-45 /Lmol m-2 S-1 daylight flu­ Adult sporangium-bearing filaments were pro­ orescent illumination, 20-25DC, 16: 8 LD. Both duced 15-30 days after PES enrichment was in­ basal discs and rugose filaments (Fig. 1) grew in creased from Yto to 'h strength. When apical seg­ all conditions tested, but the discs were initiated ments were excised and returned to 1";0 strength at higher PFD (30-45 /Lmol m-2 S-I) and the enrichment PES medium and low photon flux filaments were initiated at low PFD (l0-20 /Lmol density (PFD), 10-20 /Lmol m -2 s-', they con­ m-2 S-I). Conversion from the discoid to the tinued to grow without sporangium formation. filamentous habit was induced by reversing the However, segments returned to higher PFD (30- culture conditions. 45 /Lmol m-2 S-I), '/4 to 'h strength enrichment Apices of these filaments were used as inoc­ medium, continued to grow vegetatively, and also ulum and in low PFD they continued to grow formed sporangia again in 2-7 days. and branch irregularly in all directions. Those growing upward were more uniform in shape than those growing downward. The growth of these filaments was slow, only a few millimetres per RESULTS month. When transferred to higher PFD they showed a sharp negative phototropism growing General and field observations downward and attaching to the substrate. The In April 1982, numerous green crustose discs calcified basal discs developed from these at­ (20-30 mm diam.) of Pedobesia were seen at­ tached apices (Fig. 1). tached to the glass wall and the white PVC tubing Upon initial contact, filament apices became in a display aquarium at CEBIMAR. The con­ more tortuous and produced a digitate process. figuration of these discs was semicircular to ir­ Whenever one process contacted another, cir­ regular, possibly because of grazing by the sea cular to ovoid pores developed between them. urchin Echinometra lucunter Linnaeus or be- No septation occurred and the disc protoplast 484 Phyc% gia,Vol. 25 (4), 1986

lOOIJM ; --

4

Figs 1-6.Pedobesia ryukyuensis. Fig. 1. Rugose branched filaments and discs. Fig. 2. Young fan-shaped disc. Fig. 3. Discs showing lines of growth, lines of regrowth and radial lines. Fig. 4. Individual disc showing new young discs that overgrow the older parts. Fig. 5. Portion of disc showing the pores. Fig. 6. Portion of disc showing the pores and elevated translucent patches (pillars) near the pore.

was continuous with that of the filaments. Con­ and later became semicircular in outline (Figs 3, tinued growth produced a fiat, prostrate, perfo­ 14). rate stage that was initially fan-shaped (Fig. 2) Periodic changes of the culture medium may de Paula and West: Culture studies on Pedobesia 485

have resulted in the concentric growth lines in (Fig. 8) they were rare and appeared singly on the discs (Fig. 3). Marginal growth of the disc the unbranched erect siphon. However, sporan­ was either uniform or variable, resulting in con­ gial development varies a great deal. A reduction vex segments that contacted adjacent segments series of the sporangial development is shown in forming radial lines between them (Fig. 3). From Fig. 21, in comparison with other species of Pe­ the disc margins or regrowth regions new discs dobesia and Derbesia. arose that overgrew the older parts and new ru­ The normal, mature sporangia measured were gose filaments also developed (Figs 4, 14). Discs obovoid, 139-192 JLmlong and 60-100 JLm wide, increased in diameter at the rate of about 1-2 with stalks 10-15 JLm high and 10-15 JLm in diam. mm per month. After I year in culture with one Each contained 15-30 spores and possessed a to two changes of medium per month the max­ thick refringent plug that separated the sporan­ imum diameter was 15 mm. The pores of the gium from the filament (Figs 9, 17). Spores were disc were circular to obovoid (Figs 5, 6). In sur­ released only after the sporangium wall ruptured. face view, white translucent patches (pillars) were Spores were motile for a brief time (ca. 1 min), also observed close to the pores (Fig. 6). resulting in random settlement on the bottom, The continued growth of the rugose filaments or sometimes at the air-water interface. Zoo­ and continued formation and growth of new discs spores were stephanokontic and spherical, 30- resulted in a jumbled tuft. The upward-growing, 35 JLm in diam. (Fig. 18). free filaments attained a maximum height of 10 In some cultures non-motile spores were re­ mm after 6 months, and in all situations the leased in groups and germinated a short distance ultimate branches again grew downward giving from the parent sporangium or in situ (Fig. 10). a stoloniferous appearance. After attachment to the substrate the zoospores Tufts of sporangium-bearing branches were emitted one germ tube or two at opposite sides produced in five replicates, 15-30 days after PES (Fig. 19). The germ tubes produced rugose fila­ enrichment was increased from '1;0 to 112 strength ments or discs depending on culture conditions. at 30-45 JLmolm-2 S-I. These filamentsdiffered Very infrequently and especially in old cul­ from the rugose filaments mainly in their more tures the sporangial development patterns varied linear and smooth appearance (Figs 7, 8). Before substantially even in the same culture. Three pat­ the production of this stage both discs and rugose terns occurred: filaments became dark green and accumulated (1) A young, non-septate sporangium (Fig. starch. This adult stage can be propagated veg­ 16) formed one or two germ tubes that grew etatively in PES ('1;0 strength) and in low PFD over the mother plant. When isolated into a (10-20 JLmolm -2 S-I). In all conditions tested it new culture this 'germling' reproduced the same did not revert to the other stages. When apices stage, i.e. mature, sporangium-bearing fila­ were used as inoculum they grew and branched ments. either free from the substrate or fixed on the (2) A septate sporangium enlarged (172-264 bottom of flasks. When they grew attached to the JLm long, 144-192 JLm wide), failed to produce substrate many unbranched, erect filaments were zoospores, and detached to form one to two produced (Fig. 8). More commonly these adult germ tubes (Fig. 20) producing discs and/or filaments grew upright and ramified irregularly rugose filaments. (Fig. 11). The maximum length attained by the (3) A septate, normal-size sporangium be­ adult filaments was 25-30 mm in a culture 40 came dark green, accumulated large quantities days old. The adult filaments developed at many of starch and then extruded a single protoplast points along the rugose filaments showing spher­ that usually died after release. A few of these ical enlargements in the transition zone (Fig. 15). protoplasts enlarged somewhat on the bottom No discs produced adult filaments directly. of the flasks and formed two germ tubes at Two to 7 days after transfer to fresh medium opposite ends, and produced discs and/or ru­ (1/4 to 112strength, PFD 30-45 JLmol m-2 S-I) gose filaments. hundreds of sporangia appeared on the upright adult filaments. They were numerous along the In Table 1 the main characters of the adult main branches and on long lateral branches in sporophytic thallus for the known Pedobesia the median portions of the filament tuft (Figs 7, species are listed. The Brazilian specimens of 11, 16). On filaments attached to the substrate Pedobesia show the basic characters of the genus: 486 Phycologia, Vol. 25 (4), 1986

IOOWM -

Figs 7-13. Pedobesia ryukyuensis. Fig. 7. Sporophytic thallus with sporangia. Fig. 8. Sporophytic thallus attached to the substrate, showing unbranched erect portions. Fig. 9. Sporangia. Fig. 10. In situ germinating spores. Fig. 11. General view of the sporophytic thallus. Fig. 12. Chloroplasts from rugose filaments. Fig. 13. Chloroplasts from sporophytic thallus. de Paula and West: Culture studies on Pedobesia 487

a direct sporophytic recycling life history and besia sporophytic thalli, presenting no special unique calcified discoid stage. The adult sporan­ diagnostic characters for the generic circum­ gium-bearing thallus is composed of many erect, scription. branched, siphonous filaments originating from (2) The life history of many species of Der­ a stolon-like system of rugose filaments (Fig. IS ). besia is unknown. There are about 20 species Discs and rugose filaments are interconvertible of Derbesia and only six species of Halicystis by changing the culture conditions whereas the (Ziegler & Kingsbury 1964; Sears & Wilce sporangium-bearing filaments are not. The adult 1970). sporangium-bearing erect thalli are branched once (3) Because of the small size of some species or several times, (10) 30-50 (70) /.tm in diam. and because of their habitats in deep water and almost the same diameter throughout (Figs along rough coasts they are rarely collected 11, 15, 16). Rugose filaments are negatively pho­ (MacRaild & Womersley 1974). totropic, much branched, a maximum of 10 In nature the absence of the adult sporangium­ mm high, measuring (10) 20-30 (60) /.tm in diam. bearing filaments from Brazilian specimens could Basal discs possess circular to obovate pores, 7- be explained by intense grazing by invertebrates 12 /.tm in diam., arranged in rows on the surface. such as molluscs, echinoderms and fishes that The discs growing in nature showed similar mor­ characterize the sites studied. Littler et al (1983) phology and pore size. indicate that crustose forms are favoured under The chloroplasts are lenticular, 2-5 /.tm long exposure to high grazing pressures. by 1-2 /.tm wide, and without a pyrenoid in all Although the life history of Pedobesia is in­ developmental stages of the plant. They are pa­ terpreted in this paper as being direct and spo­ rietal with long axes parallel to the axis of the rophytic, some intriguing questions arise. The filament. In the basal discs or in the rugose fil­ adult thallus differs from the rugose filaments in aments they are always seen in face view ap­ being the only stage that produces sporangia. It pearing circular to obovoid (Fig. 12). In the adult differs also in the more linear and smooth ap­ filaments they appear elongated in side view (Fig. pearance of the filamentwalls, the upright po­ 13). Starch is present most commonly in the sition, and the faster growth when compared with chloroplasts of older filaments and on the dam­ the rugose filaments. The chloroplasts are len­ aged portions. ticular in the two stages, but in surface view are seen as circular to obovoid in the rugose fila­ ments and appear elongate in the adult sporo­ DISCUSSION phytic thallus. The rugose filamentsshow a sharp negative phototropism suggesting different ad­ The growth of the Brazilian specimens of Pe­ aptations to the irradiance. The rugose filament! dobesia is slower than that of Derbesia tenuis­ disc stages are interconvertible depending upon sima (Moris et De Notaris) Crouan and Derbesia culture conditions, whereas the sporangium­ sp. (pers. obs.), especially considering the basal bearing filaments are not, except through the spo­ disc/rugose filament stage. This prevented easy rangium development. Some of these differences isolation from Ochrosphaera sp., a coccolitho­ have a parallel with Derbesia-Halicystis stages. phorid contaminant. The isolation technique de­ The growth of the Derbesia phase is fast when veloped by adding a diatom competitor was use­ compared with that of the Halicystis phase (pers. ful in this case. obs.). According to Ziegler & Kingsbury (1964) The assurgency of new information for Pedo­ the lack of interconversion in culture between besia and the new record in Brazil give rise to the two morphological phases of Derbesia ten­ interesting questions. The genus is known with uissima, except through the normal filament­ assurance in Australia, New Zealand, the Med­ zoospores-vesicle-gamete-zygote sequence or iterranean, the Galapagos Islands, Japan, and now possibly through the parthenogenetic develop­ Brazil. This seemingly irregular distribution sug­ ment of female gametes, indicates that the con­ gests that the genus is more widespread than the trolling factors are not environmental. previous records indicate. The causes for this The developmental patterns of sporangia in apparent rarity may be: Derbesia and Pedobesia exhibit a reduction series (1) The adult sporangium-bearing filaments portrayed in Fig. 21: known for Pedobesia are quite similar to Der- (A) Derbesia and Pedobesia sporangia re- 488 Phycologia, Vol. 25 (4), 1986

• ' ' ' . . . ••. •.(1 " . . ' :.' : •" - · 'il•0 ... ,,, . \'-.0 ' • . ., . .. � . . . . . lmm '.: - Q

lmm

( c

•• ..um - -

Figs 14-20. Pedobesia ryukyuensis, Fig. 14. Discs in different developmental stages, Fig. 15. Adult sporophytic thallus arising from rugose filaments. de Paula and West: Culture studies on Pedobesia 489

Table 1. Primary characters for species delimitation in Pedobesia

Zoospores

Num- Sporangia ber Filaments Size Dia- per Diameter Length Length x meter sporan- Taxon (}tm) (mm) Shape width (}tm) (/Lm) gium References

P. clavaeformis (1. 2000-3000 30-60 Spherical 350-450 30-40 up to MacRaild & Agardh) MacRaild & to ovoid 500 Womersley Womersley 1974 (1974) P. lamourouxii (1. 400-600 Spherical 45-50 Feldmann & Agardh) Feldmann et Codomier al 1975 (1974) 200-500 30-70 Spherical 310-450 23-30 Kobara & Chi- 220-400* 220-450* hara (1984) P. feldmannii Abelard 20-30 3-10 Obovoid 120-210 x 18-35 14-30 Abelard (1982) 1982 50-85 P. ryukyuensis(Yamada 35�50 5-15 Obovoid 115-180 x 24-28 Kobara & Chi- & Tanaka) Kobara & 35-60 60-80 hara (1984) Chihara 1984 (or P. 130-200 x gracilis Chihara & 65-90* Kobara 1982) P. ryukyuensis 30-50 25-30 Obovoid 140-190 x 30-35 15-30 This paper 60-100

* Measurements from laboratory cultured plants.

lease motile spores that develop into new septate bases develop directly into cal­ filamentous sporophytes. careous basal discs. (B) Derbesia sporangia release motile spores (F) Derbesia and Pedobesia sporangia with that develop into a Halicystis-type ga­ non-septate bases form germ tubes that metophyte. Pedobesia sporangia release become sporophyte filaments. motile or non-motile spores that devel­ Some of these patterns were described as ab­ op into a calcareous basal disc. normal (Ziegler & Kingsbury 1964) and their im­ (C) Pedobesia sporangia produce non-mo­ portance in nature is difficult to assess (MacRaild tile spores that germinate in situ to form & Womersley 1974). Although meiosis is known a calcareous basal disc. to occur within the sporangium of Derbesia ma­ (D) Derbesia sporangia release single pro­ rina (Neumann 1967, 1969) there is no cytolog­ toplasts that develop into a sporophyte ical evidence to explain the unusual types of re­ filament. Pedobesia sporangia release production. The absence of sexual reproduction single protoplasts that develop into cal­ (Pattern A, Fig. 21) was demonstrated for a hap­ careous basal discs. loid mutant isolate of D. marina (Kornmann (E) Derbesia sporangia with septate bases 1966, 1970; Neumann 1967) and also referred develop directly into Halicystis-type ga­ to in both D. marina (Sears & Wilce 1970) and metophytes. Pedobesia sporangia with D. tenuissima (Feldmann 1950). Sears & Wilce

�

Fig. 16. Detail ofsporophytic thallus with sporangia. Note a young non-septate in situ germinating sporangium (arrow). Fig. 17. Mature sporangium with spores. Fig. 18. Stephanokontic zoospores. Fig. 19. Germinating spores. Fig. 20. Germinating sporangia. 490 Phycologia, Vol. 25 (4), 1986

D. marina (2, 4, 5) D. marina (1, 10) D. tenuissima (3) D. marina (5) D. marina (5) D. tenuissima D. tenuissima D. tenuissima (3) (2, 11, 15) (2, 3, 10, 11, 15) D. sp. (6) D. sp. (9) D. sp. (15)

1 Y - J p � - '4/ t t A - f - - - Q - � A B � � -- - � c� E !f- , '* / - / � � � � "� {t a a �

P. clavaeformis (7) P. clavaeformis (7) P. ryukyuensis P. ryukyuensis p. ryukyuensis P . clavaeformi s

P. lcunourouxii (8, 13) (14) (14) (14) P. feldmannii P. feldmannii (12) P. ryukyu�nsis (1 4) P. ryukyuensis (13, 14)

Fig. 21. Reduction series of sporangial development; see Discussion for an explanation of each pattern. The numbers in parentheses cite publications that are relevant to each taxon listed. (I) Kornmann 1938; (2) Feldmann 1950; (3) Ziegler & Kingsbury 1964; (4) Kornmann 1966, 1970 and Neumann 1969; (5) Sears & Wilce 1970; (6) Page 1970; (7) MacRaild & Womersley 1974; (8) Feldmann & Codomier 1974; (9) Mayhoub 1976; (10) Kobara & Chihara 1981; (II) Schnetter et a/1981; (12) Abelard 1982; (13) Kobara & Chihara 1984; (14) this paper; (15) unpublished data.

(1970) made the supposition that in the above P. clavaeformis isolated from Australia (Mac­ case the sporophyte is haploid. Feldmann (1950) Raild & Womersley 1974), P. lamourouxii from explained this tentatively as a lack of meiosis. Japan (Kobara & Chihara 1984) and P. ryu­ Pattern E (Fig. 21), observed for D. marina (Sears kyuensis(present paper) produced sporangium­ & Wilce 1970), in which sporangia of the spo­ bearing thalli in culture. Pedobesia lamourouxii rophytic phase failed to cleave into spores but isolated from the Mediterranean (Feldmann & rather developed directly into gametophytes, can Codomier 1974), P. feldmannii from the Gala­ be interpreted as a loss of spore formation with pagos I. (Abelard 1982) and P. ryukyuensisfrom meiosis occurring somatically within the sporan­ Japan (Kobara & Chihara 1984) remained as ru­ gia (see Tanner 1981). gose/discoid sterile stage in laboratory culture. Considering the strong parallels, except for Feldmann & Codomier (1974) and Abelard Patterns C and D, found in the subsequent de­ (1982) suggested that perhaps in some undefined velopment of the sporangia of Derbesia and Pe­ circumstance the rugose/discoid stage could pro­ dobesia (Fig. 21), it is possible that meiosis also duce sexual reproductive structures. No such occurs in Pedobesia (Patterns B, C, D, E) or is structures were observed in the Brazilian speci­ absent (Patterns A, F). From all life history in­ mens. In some instances only a spherical enlarge­ vestigations listed in Fig. 21 for Pedobesia, only ment was observed in the transition zone be- de Paula and West: Culture studies on Pedobesia 491

tween rugose/adult filamentous stage. No special as sporangial stalk and plug and the mode of­ culture conditions were mentioned by MacRaild spore liberation do not differsignificantly. Also, & Womersley (1974) and Kobara & Chihara the shape and size of chloroplasts and absence (1984) for induction of adult filamentsfrom ru­ of pyrenoids are not useful to distinguish them. gose ones. The Brazilian isolates remained sterile The structure of the discs was not studied in for about a year. The adult filaments were pro­ detail for the two species but the size and shape duced only in high PFD and high level of en­ of the pores in the Brazilian specimens coincided richment and were preceded by starch accumu­ with that ascribed to Pedobesia ryukyuensis (Ko­ lation. If the speculation that meiosis occurs in bara & Chihara 1984). The discs found in nature the sporangium of Pedobesia is correct it is pos­ also do not differfrom those in the laboratory. sible that gametic nuclei and diploidization are The 'pillars' seen in surface view as refringent involved in this transition zone. patches are located near the pores in the two Some problems arise in determining the Bra­ entities. zilian Pedobesia isolates. Mature sporangium­ Based on the present information both from bearing filaments were not found in the field. our own results and those presented for Pedo­ Consequently all these descriptions were based besia feldmannii and P. ryukyuensis it is difficult on cultured plants. According to Page (1970) the to distinguish the three species. Pedobesia feld­ morphological characters of Derbesia are pleo­ mannii is recorded from the Galapagos Islands morphic in culture conditions, thus causing ad­ (Ecuador) growing on coral reef in deep water ditional taxonomic problems. (Abelard 1982). Pedobesia ryukyuensisis re­ In filament diameter and length of sporophytic ported from Japan (Kobara & Chihara 1984) thallus, Pedobesiafeldmannii and P. ryukyuensis growing on polychaetes in shaded places near low (Kobara & Chihara 1984; present paper) are very tide level and, according to these authors, also similar whereas P. clavaeformis and P. lamou­ in South Africa (as Derbesia ryukyuensisYa­ rouxii are very different (Table 1). mada & Tanaka, in Papenfuss & Egerod 1957) Pedobesia clavaeformis occurs in Australia and and Ecuador (as D. longifructain Taylor 1945). New Zealand (MacRaild & Womersley 1974; For the Brazilian coasts only Derbesia marina Womersley 1981; Hawkes 1983). Pedobesia la­ (Lyngbye) Solier is recorded (Joly 1965; Oliveira mourouxii occurs in the Mediterranean and in PO. 1977). Our research did not confirm the pres­ Japan (Feldmann & Codomier 1974; Feldmann ence of this species but has shown the occurrence et al 1975; Kobara & Chihara 1984)! According of D .. tenuissima (Moris et De Notaris) Crouan to Kobara & Chihara (1984) this species is also at Praia do Segredo, Sao Sebastiao, SP. This recorded in southern California (as D. lamour­ species differsfrom D. marina mainly in the ouxii in Collins 1928), Guadeloupe I. and Ber­ presence of pyrenoids and shows the character­ muda [as Derbesia lamourouxii in Taylor 1960 istic life history of the genus (unpubl. data). (see also Abelard 1982)]. The occurrence of P. The correct name of the Brazilian specimens lamourouxii for southernCalifornia and Guad­ is uncertain because of incomplete information eloupe and Bermuda needs to be confirmed by and was based on the fact that Pedobesia ryu­ further collections and life history studies. kyuensis Kobara & Chihara (1984) (= Derbesia The sporangium-bearing filaments of Pedo­ ryukyuensisin Yamada & Tanaka 1938; = P. besia feldmannii are described (Abelard 1982) as gracilis in Chihara & Kobara 1982) has priority being attached to the substrate, horizontally over the other names. Thus, the Brazilian spec­ branched, and producing unbranched erect si­ imens are tentatively referred to as P. ryukyuen­ phonous filaments.This pattern of development sis, but further investigation on D. longifructa seems taxonomically to lack importance since it and the three closely related entities of Pedobesia also occurs in P. ryukyuensis(Fig. 8) and Der­ is needed to solve the nomenclatural problem. besia tenuissima and D. sp. (pers. obs.). These same isolates in other cultures grew and ramified irrregularly in the erect position. ACKNOWLEDGMENTS In sporangial shape and size, as well as number and diameter of zoospores, Pedobesiafeldmannii Many thanks are due to Mitsuo Chihara for valu­ and P. ryukyuensisfrom Japan and Brazil also able exchange of information on the systematics show similarities (Table 1). Other characters such of Pedobesia; to Marilza Cordeiro-Marino for 492 Phyc% gia, Vol. 25 (4), 1986

initiating the algal culture technique class at the and taxonomy of two species of Pedobesia (Bryop­ Instituto de Botanica de Sao Paulo, where this sidales, Chlorophyceae) in Japan. Bot. Mag. Tokyo 97: work began, and providing preliminary culture 151-162. A KORNMANN P. 1938. Zur Entwicklungsgeschichte von space; to Claudio Gonr;alves Tiago, lvaro Es­ Derbesia und Halicystis. Planta 28: 464-470. teves Migotto and Aime Rachel Magenta Ma­ KORNMANN P. 1966. Eine erbliche Variante von Der­ galhaes of CEBIMAR for identification of the besia marina. Naturwissenschaften6: 161. invertebrates and other valuable information; and KORNMANN P. 1970. Eine Mutation bei der sipho­ nalen Griinalge Derbesia marina. Helgolander Wiss. to Nanuza Luiza de Menezes, Ana Maria Giu­ Meeresunters. 21: 1-8. lietti and Pablo Garcia Carrasco of the Depar­ LITTLER M.M., LITTLER D.S. & TAYLOR P.R. 1983. tamento de Botanica at USP for photographic Evolutionary strategies in a tropical barrier reef sys­ assistance. This research was supported by CNPq tem: functional-form groups of marine macroalgae. J. Phycol. 19: grant 150-301130/83. Further assistance was 229-237. MACRAILD G.N. & WOMERSLEY H.B.S. 1974. The provided in typing by Barbara Joyce and in pho­ morphology and reproduction of Derbesia clavae­ tographic work by Chris O'Brien (School of Bot­ formis (J. Agardh) De Toni (Chlorophyta). Phyco­ any, University of Melbourne, Australia) where logia 13: 83-93. the second author was on sabbatical leave in MAYHOUB H. 1976. Sur I'existence d'une Derbesia dan Ie cycle de I'Halicystis boergesenii Iyengar et 1984-5. Raman (Chlorophycees, Derbesiales). C. R. Acad. Sci. Paris ser. D 282: 707-710. McLACHLAN J. 1973. Growth media-marine. In: Handbook of Phycological Methods. Culture Meth­ REFERENCES ods and Growth Measurements (Ed. by J. Stein) pp. 25-51. Cambridge University Press, Cambridge. NEUMANN K. 1967. Der Ort der Meiosis und die ABELARD C. 1982. 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