Phycologia (1974) Volume 13 (1), 31-44

The morphology and taxonomy of the red alga Sarconema (: )

GEORGE F. PAPENFUSS* AND TrKVAH EOELSTEINt

* Department of Botany, University of California, Berkeley, California 94720, U.S.A. t Atlantic Regional Laboratory, National Research Council of Canada, Halifax, Nova Scotia, Canada

A study of the structure and reproduction of two species of Sarcollemafrom East Africa has shown that the genus is correctly placed in the Solieriaceae. The genus agrees with Solieria in that the auxiliary cell is indistinguishable before diploidization. A sheath of peri car pic filaments is not produced about the gonimoblast as it is in CallophyclIs, Solieria, Agardhiella and . Only two species of Sarcollemaare recognized hy us, S. filiforme(Sander) Kylin and S. scillaioidesB0rgesen. S. Jurcellatum Zanardini, S. montagnei (Grunow) Kylin, S. indicum (J. Agardh) Kylin, S. Jurcatum B0rgesen and S. filiformef. gracillima Rayss are regarded as representative of S. filiforme. Dicranema setaceum Sonder from Queensland, Australia, and D. setaceum var. "polensis Grunow from Samoa were also found to be S. filiforme.

Introduction (now known as Callophycus Trevisan, 1848; Silva, 1957). The same seven genera were placed in the The history of the family Solieriaceae, to which Solieriaceae by Schmitz & Hauptfleisch (1896). Sarconema belongs, begins with J. Agardh, who in However, in both these accounts the family was 1852 (pp. 718 and 721) erected the tribe Solierieae for treated as a subfamily of the RhodophyJIidaceae. the genus Solieria J. Agardh (1842). Harvey (1853, De Toni (1897, 1924) adopted the classification of & p. 115) elevated the Solierieae to the rank of sub­ Schmitz HauptfJeisch.. family (except that he called it suborder) and included Kylin in 1925 (p . 25) erected the family Rhab­ in it the genus J. Agardh (1847) in addition doniaceae for the reception of Rhabdonia, Agardhiella to Solieria. In 1872 J. Agardh included in the Schmitz (Schmitz & Hauptfleisch, 1896), Anatheca Solierieae his new genus Meristotheca and Rhabdonia Schmitz (Schmitz & Hauptfleisch, 1896), Flahaliltia Harvey (Hooker & Harvey, 1847), in addition to Bornet (1892) and Solieria. (The inclusion of Solieria, Solieria and Eucheuma, and in 1876 he added: the type genus of the Solieriaceae, in the Rhab­ Gelinaria Sonder (with a query; the genus is now doniaceae by Kylin illegitimatizes the family.) In placed in the Cryptonemiaceae), Carpococcus 1932 Kylin removed Solieria, Anatheca, Flahaultia J. Agardh (currently treated as asynonym of Sarcodia and Agardhiella to the Solieriaceae (the existence of J. Agardh of the family Sarcodiaceae), and Catenella which he apparently had overlooked in 1925) and he Greville (1830) (with a query; the genus is now referred Catenella, Erythroclonium and Areschougia, referred to the Rhabdoniaceae). in addition to Rhabdonia, to the Rhabdoniaceae. In Hauck in 1883 (pp. 17 and 186) elevated the sub­ his Gattungen Kylin (1956) added Catenellocolax family Solierieae to the rank of fami Iy and in 1886 he Weber-van Bosse (1928) to the Rhabdoniaceae and placed Sarconema Zanardini (1858) in the family. dates the family as of 1932. Searles (1968, pp. 44 and Schmitz in 1889 (p. 442) placed the following seven 63) has provisionally also assigned Caulacanthus genera in the Solieriaceae (J. Agard h) Hauck: Kiitzing (1843), Taylorophyclls Dawson (1961) and Rhabdonia Harvey, Erythroclonium Sonder (1852), Heringia J. Agardh (1844) to the Rhabdoniaceae. Areschougia Harvey (1855, nomen conservandum), Joly & Alveal (1969) and Augier & Boudouresque Solieria J. Agardh, EllcheumaJ. Agardh, Sarconema (1971) have provisionally also placed the new genera Zanardini, and Thysanocladia (Endlicher) Lindley Montemaria and Feldmannophycus, respectively, in the Rhabdoniaceae. As an appendix to the present Issued as NRCC No. 13699. paper, the family is proposed for conservation.

31 32 Phyc% gia, Vol. 13 (1), 1974

In 1932 Kylin placed the following ten genera in In view of our very meagre knowledge of the the Solieriaceae: Thysanocladia (= Callophycus), development of the female reproductive apparatus Flahaultia, Sarcodiotheca Kylin, gen. nov., Agard­ and the carposporophyte in the Solieriaceae and the hiella, Solieria, Sarconema, Eucheuma, Anatheca, availability of fertile material of Sarconema from Meristotheca, and Euryomma Schmitz (Schmitz & East Africa, the present study was undertaken with Hauptfleisch, 1896). In 1934 Kylin included Opun­ the hope of contributing to a better understanding tiella Kylin (1925) and Turnerella Schmitz (Schmitz of this genus and the family. Two species (the only & Hauptfleisch, 1896) in the family and in 1956 he two in the genus in our opinion, as will be pointed credited the family with two additional genera, out in later pages), S. filiforme(Sonder) Kylin and namely, Predaea G. DeToni (1936) and Gardneriella S. scinaioides B0rgesen were studied. Both liquid Kylin (194 1). preserved and herbarium specimens were examined. Dawson (1949) added the monotypic new genus Reticulobotrys to the family and B0rgesen (1953) added the new genus Tenaciphyllum. Feldmann Structure of the thallus (1942) removed Predaea to the Nemastomaceae, in which family the genus had been placed by Setchell Thalli of Sarconema (Figs. 1-6, 19, 20 and 23-25) are & Gardner (1930) when they described it (as terete, solid, 0,8-2 mm in diameter near the base, up Clarionea gen. nov.). Dawson (1961) also placed to 20 cm long, dichotomously to subdichotomously Predaea in the Nemastomaceae and Kraft & Abbott branched, and attached by a branched, hapteroid (1971) have also convincingly demonstrated that holdfast. The branches taper gradually toward the this genus belongs in the Nemastomaceae rather than tip. Regeneration occurs from branches whose tips in the Solieriaceae. Kraft & Abbott also reduced have been lost. One or several new branches may be Yadranella Ercegovic (1949) to asynonym of Predaea. produced from such tips, which frequently results in The features whereby the Solieriaceae may be a disturbance of the dichotomous habit. characterized are: (1) the thallus is multiaxial in The thallus is muItiaxial in construction and construction and has a filamentous medulla; (2) consists of three clearly definedtissues (Figs. 19 and procarps are lacking; (3) the auxiliary cell initiates 20); an outer cortex of one or two layers of small, only one gonimoblast cell; (4) the gonimoblast at tightly packed pigmented cells (Figs. 8 and 20); an first develops inwardly; (5) the cystocarps are inner cortex of several layers of irregularly shaped, embedded in the thallus or project from the thallus pseudoparenchymatous cells, which in size decrease surface and are usually provided with a distinct toward the outer surface (Figs. 19 and 20); and a ostiole; (6) a pericarp is usually, but not always, prod­ central medulla of interwoven thick-walled filaments, uced; and (7) the tetrasporangia are zonately divided. composed of elongated cells with lateral pit connec­ The family Rhabdoniaceae which has been tions between adjacent filaments (Figs. 19 and 20). confused with the Solieriaceae differs from the latter especially in that (1) the thallus is uniaxial in construction, and (2) the cystocarp of all genera Development of the female reproductive system lacks a special pericarp. Information of a greater or lesser amount on the As in other members of the Solieriaceae, procarps Sarconema. ontogeny of the cystocarp in the Solieriaceae is are not formed in Carpogonial branches available for the following genera: Sarcodiotheca are produced in large numbers in the tips of the * (Kylin, 1925, as Anatheca), Turnerella (Kylin, 1934; thallus branches. The carpogonial branch is com- Tokida & Masaki, 1957),* Sarconema (Kylin, 1932; * Young stages were best observed in material of S. scinaioides Rayss, 1963), Opuntiella (Kylin, 1934), Agardhiella that was collected by Papenfuss & Scagel (PR-YIl-\02) near (Osterhout, 1896; Kylin, 1928), Solieria (Bornet, in Dar-es-Salaam, Tanganyika, in October 1962. Rayss's (1963) cystocarpic material of Sarconema from the eastern Mediter­ Bornet & Thuret, 1880; Kylin, 1932), Meristotheca ranean was collected in July and October. Hauck's (1886) (Kylin, 1932), Gardneriella (Kylin, 1941) and material of this genus, collected by Hildebrandt in Somalia, Callophyclls(Hewitt, 1960). contained cystocarps in March, Newton's (1953) plant from the Hanish Islands, Red Sea, contained tetrasporangia in * Tetrasporophytes have been unknown in Tlirnerella. September, and Aleem (1948), who was the firstto report the Recently South et al. (1972) showed that T. pennyi) of genus from the Mediterranean, obtained plants with tetra­ the north Atlantic Ocean possesses a crustose, Crlloria-like sporangia in October. B0rgesen (1932) had tetrasporic tetrasporic phase. specimens gathered (in Pakistan or India) in January. Papenfuss and Edelstein: Morphology and taxonomy of Sarconema 33

2em � ,' 2 em .2 I....-..J

FIGS. 1-3. Sarcol1ema /ili/orme. FIG. 1. Habit of a liquid preserved cystocarpic specimen collected by Papenfuss & Scagel (PR·YUJ-48) near Dar·es·Salaam, Tanganyika, on 11-13 October 1962. FIG. 2. Habit of herbarium specimen (showing the dichotomously branched nature of the species) collected by Papenfuss & Friedmann (E62/20036) at Entedebir lslet, Dahlak Archipelago, Ethiopia, on 13 March 1962. FIG. 3. Voucher specimen of Borgensen's 1934 record of S. jill'carul1IBorgesen from Pakistan (NY).

c 34 Phyc% gia, Vol. 13 (1), 1974

trichogyne bends as it emerges from the carpogonium and grows toward the surface of the thallus (Fig. 8). An intercalary cell of the inner cortex functions as auxiliary cell (Fig. 10). This cell is spatially removed from the carpogonial branch and is not distinguish­ able from neighbouring vegetative cells before diploidization and the initiation of the gonimoblast. In this latter feature, Sarconema resembles Solieria (Kylin, 1932), whose auxiliary cells also are indistin­ guishable before diploidization. In the other investi­ gated genera of the Solieriaceae, for example, Sarcodiotheca (Kylin, 1925, as Anatheca) and Callophycus (Hewitt, 1960), the auxiliary cell is recognizable at an early stage by its dense contents which stain deeply with dye. After fertilization of thecarpogonium, a connecting filament is produced from the region between the carpogonium and the trichogyne, which structure is often pushed to one side (Fig. 9). Long, branched connecting filamentsare common in fertile branch tips, but their union with an auxiliary cell was not observed. The abundance of carpogonial branches and paucity of gonimoblasts in fertile tips suggest that as in other Solieriaceae (Kylin, 1956, p. 274) the number of auxiliary cells is small in comparison with the number of carpogonial branches. One of the first signs that an auxiliary cell has been diploidized (presumably) is the development toward the surface of the thallus of a small-celled, deeply staining nutritive tissue from the auxiliary cell (Fig. 10). No trace of the nutritive cells was recognized in half-grown cystocarps, which fact leads us to conclude that their contents are used up during the development of the gonimoblast or they become incorporated in the fusion cell. The gonimo­ blast is initiated inwardly (Figs. 11 and 14-18) as in Callophyclls(Hewitt, 1960), Solieria (Kylin, 1932) and other genera of the Solieriaceae. At an early stage, the auxiliary cell, some of the neighbouring

FIGS. 4-6. Sarconema scillaioides. FIG. 4. Habit of a cysto­ vegetative cells, and probably some of the first­ carpic specimen collected by Papenfuss & Scagel (PR-VII [- formed gonimoblast cells fuse and form a large 102) near Dar-es-Salaam, Tanganyika, on 11-13 October irregularly shaped, deeply staining fusion cell 1962. FIG. 5. Habit of a sterile specimen from the same col­ lection as F!g. 4. FIG. 6. Type of the species from Pakistan (Fig. 15). In older stages, arms of the fusion cell (NY). extend into the cortex in the region where the ostiole will be formed (Figs. 12, 16-19 and 21 ). In the mature condition, gonimoblast filaments radiate posed of three or four cells (Figs. 7-9) borne on an from the fusion cell (Fig. 22). Carposporangia are undifferentiated, intercalary cell of the inner cortex. cut off in chains from the tips of the gonimoblast This cell, the supporting cell, is larger than the cells filaments (Figs. 17-19, 21 and 22). [(According to of the carpogonial branch (Figs. 7 and 8). As in Kylin (1932) the carposporangia are formed singly, other Solieriaceae, it produces a single carpogonial an erroneous impression based on the fact, no branch on the side facing the medulla (Fig. 8). The doubt, that he had only herbarium material at his Papenfuss and Edelstein: Morphology and taxonomy of Sarconema 35

-I

[ 20fLm

13

FIGS. 7-12. Sarconema scinaioides (PR-VlII-I02). FIG. 7. Carpogonial branch and supporting cell. FIG. 8. Part of cross section through fertile tip of thalllls, showing a carpogonial branch and its supporting cell,which is an inter­ calary cell of the inner cortex. FIG. 9. Two carpogonial branches, the carpogonium of one of which has produced a connecting filament. FIG. 10. Part of cross section through fertile branch of thallus,showing an auxiliary cell (an intercalary cell of the inner cortex) and a group of small nutritive cells which have been produced by it toward the surface of the thallus. FIG. II. Auxiliary cell and young gonimoblast produced by it toward the interior of the thallus. FIG. 12. Older gonimoblast with a large fusion cell and carposporangia. F,G. 13. S. filiforme (PR-VIlT-48). Part of cross section of tetrasporic plant, showing tetrasporangia embedded in the outer part of the inner cortex (aux, auxiliary cell; ca, carposporangia; cf, connecting filament;cp,carpo­ gonium; cpb, carpogonial branch; fc, fusion cell; go, gonimoblast; ic, inner cortex; nc, nutritive cells; oc, outer cortex; sc, supporting cell; tr, trichogyne).

disposal.] Some of the gonimoblast filaments remain ",m in diameter, more or less embedded in the sterile and seem to have a nutritive function. Early thallus (Figs. 17, 18 and 21), although the cystocarps in the development of the carposporophyte the are readily recognized with the unaided eye as region where the ostiole will develop becomes bulges produced on the thallus (Figs. 1, 18 and 21). recognizable by the formation of files of small In our material cystocarpic plants wer¢ rare-of cells (Fig. 14). However, the pore is formed only over 100 specimens examined only four bore cysto­ when the gonimoblast approaches maturity (Fig. 22). carps. The carposporangia are round to ovoid, measuring No male plants were observed in the material 12-15 x 15-25 ",m. Mature gonimoblasts are 500-750 examined by us. , O.lmm

.\ E E 10 o �]

t8 '"16 17

FIGs. 14-18. Sarconema scinaioides (PR-VIII-102). Parts of cross sections through fertile thallus, showing stages in the development of the gonimoblast (aux, auxiliary cell; ca, carposporangia; fe, fusion cell; go, gonimoblast). E E 10 d

E E E E 10 10 d o

·22

FIG. 19. Sarconema scinaioides (PR-VIlI-102). Part of cross section through fertile thallus, showing three embedded cystocarps, the core of medullary filaments, and the pseudoparenchymatous inner cortex. FIGs. 20-22. S. fili/orme (PR-VllI-48). FIG. 20. Longitudinal section of a vegetative branch. FIG. 21. Cross section through thallus, showing a cystocarp. FIG. 22. Part of cross section through thallus, showing a mature cystocarp (ca, carposporangia; fc, fusion cell; go, gonimoblast; ic, inner cortex; mf, medullary filaments; oc, outer cortex; os, ostiole). ,�t:Jlllllltllllflllf!!tl!IIIIIII!llltlllllllllllllll!11111111111111111111111111111111 , 2 3 • 56?8

,25 23 -/

44489

TYPE

1�1:111"111"111111111111"1"''1IIIII''IIII.I . 2 3 I. Papenfuss and Edelstein: Morphology and taxonomy of Sarconema 39

Development of tetrasporangia Sarcodiaceae Kylin (1932) should be retained as a family distinct from the Solieriaceae. As far as The tetrasporic plants are morphologically similar Sarconema and other better-known genera of the to the female plants. The sporangia are scattered Solieriaceae (see introduction) are concerned, there over the surface of the thallus, except that they are are several features, both anatomical and reproduc­ not present in the lowermost parts of the branches. tive, whereby the members of this family differfrom They develop as lateral outgrowths from cortical the type of the Sarcodiaceae, Sarcodia montagneana cells. When mature, they are zonately divided, appear (J.D. Hooker & Harvey) J. Agardh, as revealed by as dark, round spots, as seen in surface view, are Rasmussen's (1964) study of this species. It would covered by one or two layers of cortical cells (Fig. 13), seem, therefore, that the Sarcodiaceae should be and are relatively large, measuring 20-25 ",m in retained for the genus Sarcodia (and perhaps the width and 35-50 ",m in length. little-known Chondrymenia Zanardini, see Kylin, 1956) and another home sought for the other genus currently placed in the family, namely, Discussion Trematocarpus Kiitzing [see Searles (1969), who has merged Dicllrella Harvey ex J. Agardh in Trema­ In spite of the then scanty knowledge of Sarconema, tocarpus). Hauck (1886) and Kylin (1932) were correct in placing the genus in the Solieriaceae. In some members of this family, for example, Taxonomy Callophycus (Hewitt, 1960, pI. 29), Solieria (Bornet & Thuret, 1880, pI. 50, fig. 5; Kylin, 1932, fig. 3B; When Zanardini (1858) erected the genus Sarconema Thein, personal communication), Agardhiella (Kylin, he credited it with only one species, S. fllrcellatllm, 1928, fig. 45A; 1956, fig.206G), and Meristotheca from the Red Sea. Kylin (1932) placed four species (Kylin, 1932, fig. 6A; 1956, fig. 208B), a sheath of in the genus: S. fllrcellatllm,S. monfagnei (Grunow) pericarpic filaments is formed about the gonimoblast. Kylin (=Dicranema monfagnei Grunow, 1873-74= Such a sheath is not produced in Sarconema (Figs. Plocaria furcellata Montagne, 1850) from the Red

18 and 22; see also Kylin, 1932, fig. 4A; 1956, fig. Sea, S. indicum (J. A-gardh) Kylin ( =Solieria indica 208D; Rayss, 1963, fig. 6). A somewhat weakly J. Agardh, 1852) from India, and S. /i/iforme

developed cellular pericarp is formed, however, by (Sonder) Kylin (=Dicranema /ili/orme Sonder, proliferation of the cortex (Figs. 16-18, 21 and 22). 1845, 1846) from Western Australia. In Agardhiella fenera* (Osterhout, 1896, p. 416, B0rgesen (1932) reported Sarconema furcellatum fig. 1) and Ellchellma procrusteanllm(Kraft, 1970, from Pakistan and India, and in 1934 added two new fig. 4), sterile gonimoblast filaments establish species from Karachi, Pakistan, to the genus: S. secondary pit connections with the pericarpic tissue. furcatum and S. scinaioides. In 1937 he reported In CallophycliS africanus the fusion cell is connected, S. indicllmfrom Cape Comorin in southern India, by means of filaments, to the pericarpic cortex and in ] 939 he reduced S. montagnei and S. indicllm (Hewitt, 1960, fig. 21) but in this taxon the filaments to the synonymy of S. fllrcellafllm. Newton (1953) issue from the inner cortex. Comparable filaments agreed with B0rgesen that S. montagnei and S. were not observed in Sarconema. It is seen, therefore, indicllm were merely variants of S. fllrcellatum. She that the Solieriaceae contains some genera whose thought that S./iliformemay well belong in the genus cystocarps are provided with a filamentous pericarpic Dicranema, where Sonder (1845, 1846) had originally tissue about the gonimoblast and others (Sarconema, placed it, rather than in Sarconema. On the other Turnerella and Opllntiella) in which a filamentous hand, she found that No. 316a of Harvey's Australian pericarpic tissue is lacking. Algae (named D. /i/iformeby Harvey), a duplicate Se:!rles (1968, p. 75) was inclined to doubt that the of which Kylin (1932) had seen in the Agardh Sarconema, • This species is now known as Agardhie/la bailey; (Harvey Herbarium, was indeed a species of viz. ex Ktitzing) Taylor (Taylor & Rhyne, J 970, p. J 3). S. fllrcellatllm.

FIGS. 23-25. Sarconema fili/orme. FIG. 23. Type of DicI"Gnema filiforme Sonder from Western Australia (MEL). FIG. 24. Lectotype of Dicranema selacellm Sonder from Queensland, Australia (MEL). FIG. 25. Lectotype of D. selacellm var. lIpolensis Grunow from Samoa (W). 40 Phyeologia, Vol. 13 (1), 1974

An important point as regards the nomenclature proved to be representative of Sareonema fili/orme. of Sareonema /ureellatum Zanardini (1858) that A comparison of one of the Kotwal specimens needed to be settled was whether Dieranema filt/orme (in Herb. N.Y. Bot. Gard., Fig. 3) of Sareonema Sonder (1845, 1846) is representative of Sareonema /ureatum from Pakistan, cited by B0rgesen (1934) [as Kylin (1 932) and BrJrgesen (1934, 1938, 1943, when he erected this species, with material from East 1950, 1952) believed] or of Dieranema [as Newton Africa and elsewhere has convinced us that this (1953) thought], and if it should prove to be a species species also should be merged in S. fili/orme. of Sareonema, whether it is an autonomous species Rayss in 1963 described two new formae under or the same as S. /ureellatum, a species which Sareonema fili/orme,forma curta (from Bombay, Newton (1953) had shown to occur in Western Karachi, Yemen, Mauritius and the eastern Mediter­ Australia. The second author was privileged to see ranean) and forma graeillima (from the Mediter­ Sonder's type specimen (Fig. 23) while she worked ranean coast of Israel). Papenfuss (1968) concluded in Dr Womersley's laboratory at the University of that forma curta probably is representative of S. Adelaide, South Australia, during part of 1972. fureel/atum. However, Rayss failed to indicate a Examination of this specimen (MEL) showed not type for the taxon, which renders it invalid in only that Sonder's species is a species of Sareonema accordance with Article 37 of the Seattle Code. In but that it indeed is the same as S. /ureel/atum. The the light of present knowledge of the variability of Australian plants of this taxon are somewhat more S. fili/ormeit seems reasonable to conclude that slender and elongate than those from the Red Sea S. fili/ormeforma gracil/ima is the same as S. and seldom exceed 0·8 mm in diameter. B0rgesen fili/orme. (1934) thought that the cells of the parenchymatous It would seem, therefore, that as far as known, tissue were larger in S. fili/ormethan in S. /ureel/atum Sareonema includes only two species, S. fili/orme but we have seen no evidence of this. In writing to the (Sonder) Kylin and S. scinaioides B0rgesen. S. first author about Sonder's specimen, Dr Womersley scinaioides (Figs. 4-6) is similar in general appear­ commented as follows: 'The type specimen of ance to S. fili/orme(Figs. 1-3 and 23-25), but [Dieranema] fili/ormefits the habit description of plants of the former are of greater diameter (up to Sonder. The plant is tetrasporangial and many of 2 mm in diameter), of a much softer texture and the axes are truncated, probably by grazing. New adhere more firmly to paper. The two species were branches have arisen from many of the truncations, obtained at the same locality near Dar-es-Salaam, and thus appear "articulated" and somewhat Tanganyika, by Papenfuss & Scagel. [Schmitz swollen, as referred to by Sonder. They bear tetra­ (1895, p. 139) had previously reported S.fili/orme[as sporangia and might give the impression that the S. /ureel/atum] from Dar-es-Salaam and Zanzi­ sporangia were confined to them if no other parts bar.] were sectioned. Sporangia are general throughout Sareonema scinaioides has no synonyms and the most of the plant.' It is the misconception of Sonder present report of the presence of this species at that the tetrasporangia were confined to the some­ Mambrui, Kenya, and Dar-es-Salaam, Tanganyika, what swollen tips of the branches that caused Newton constitutes the first record of its occurrence outside (1953) to conclude that Dieranema /ili/ormeSonder of Pakistan, whence it was described by B0rgesen probably is a species of Dieranema rather than of (1934). Sareonema. The synonymy of Sareonema fili/ormefollows: Since the binomial Dieranema fili/ormeSonder (1845) has priority over Sareonema /ureel/atum Sareonema fili/orme(Sonder) Kylin Zanardini (1858) the correct name of the species that Kylin, 1932, p. 22. for more than 100 years has been known as S. /ureellatum is S. fili/orme (Sonder) Kylin. Dieranema fili/ormeSonder, 1845, col. 56; 1846, While in Adelaide, the second author also had the p. 173. opportunity of examining the lectotypes of Diera­ Cystoclonium fili/orme(Sonder) Kiitzing, 1849, nema setaeeum Sonder (1 871) (Fig. 24) and D. seta­ p. 757; 1868, p. 6, PI. 18, Figs. a and b. eeum var. upolensis Grunow (1 873-74) (Fig. 25), Sareonema fili/orme f.graeillima Rayss, 1963, which specimens Dr Womersley had borrowed p. 101, Fig. 7. from Melbourne and Vienna, respectively, for Sareonema fili/ormef. curta Rayss, 1963, p. 98, Mr G. Kraft's study of Dieranema. Both these taxa Figs. 1-6 (nomen nudum). Papenfuss and Edelstein: Morphology and taxonomy of Sarconema 41

Sarconema furcellatum Zanardini, 1858, p. 264, Mozambique: Praia Chokas (north of Lumbo), PI. 10, Fig. 1. 15-17.xi .1962, Papenfuss & Scagel PR-XXVIII-92; Plocaria furcellata Montagne, 1850, p. 243. Zavora Reef, 4 .xi .1962, Papenfuss & Scagel Graci/aria furcellata (Montagne) Zanardini, 1858, PR-XX-54. p.266. Pakistan: near Karachi, no date, Kotwal (NY: Trematocarpus furcellatus (Monta gne) Kiitzing, voucher specimen of B0rgesen's 1934, p. 12 record 1869, p. 27, PI. 73, Figs. c and d. of S. furcatum [our Fig. 3)). Dicranema furcellatum (M ontagne) J. Agardh, India: Bombay, xii. 1927 and i.1928, Borgesen 1876, p. 436 (non J. D. Hooker et Harvey). (UC 581368 and 581375, respectively; voucher Dicranema montagnei Grunow, 1873-74, p. 43 material of B0rgesen's 1934 p. 11 record of S. (=Plocaria furcellata Montagne non D. furcel­ fi/iformefrom Bombay). latum J. D. Hooker et Harvey). Australia-Western Australia: no date, Preiss Sarconema montagnei (Grunow) Kylin, 1932, [2557] (MEL, Type of Dicranema filiforme Sonder p. 21, PI. 8, Fig. 16. 1845 col. 56; 1846 p. 173 [our Fig. 23)); no date, Solieria indica J. Agardh, 1852, p. 723. Harvey 316a (MEL, voucher specimen of Harvey's Sarconema indicum (J. Agardh) Kylin, 1932, Australian Algae No. 316a of D. filiforme); Perth p. 22, PI. 8, Fig. 17. (North Beach area), iii .1959, Norris (ADU 22268); Dicranema setaceum Sonder, 1871, p. 58. Fremantle, iii.1951, Royce (ADU 14143); Cockburn Dicranema setaceum var. upolensis Grunow, 1873- Sound, iv.1966, Kraft (G. T. Kraft's personal 74, p. 43. herbarium); Safety Bay, vii .1966, Kraft 1913 and Sarconema furcatum B0rgesen, 1934, p. 12, Fig. 8, 1919 (Kraft's herbarium). PI. 2, upper figure. South Australia: Elliston (inner reef), i. 1951, Womersley (ADU 13695a). New South Wales: Port Stephens, no date (MEL, Material examined Algae Muellerianae curante J. G. Agardh distri­ butae, as D. filiforme). Sarconema filiforme(Sonder) Kylin Queensland: Port Denison, no date [Fitzalan] Red Sea-Ethiopia-Dahlak Archipelago: Ente­ (MEL, Lectotype of D. setaceum Sonder 1871 p. 58 debir Islet (Landing Bay), 11. iii .1962, Papenfuss & [our Fig. 24]). Friedmann E62/2003 (tetrasporic); 13 . iii .1962, Samoa: Upolu, no date, Graeffe fYI, Collectio Papenfuss & Friedmann E62/20036 (Fig. 2) (tetra­ Grunow No. 33896, Lectotype of D. setaceum vaL sporic) and E62/20037; (Goliath Bay), 23.iii.1962, upolensis Grunow 1873-74 p. 43 [our Fig. 25]). Papenfuss E62/20167; Harmil Islet, 28 . iii .1962, Papenfuss E62/20287 (tetrasporic). Massawa, iv. Sarconema scinaioides B0rgesen 1870, Issei (UC 90890; voucher material of Piccone's Kenya: Mambrui (north of Malindi), xii.1967, 1884 p. 320 Massawa record of Dicranema furcel­ Isaac latum). (ADU 40418). Tanganyika: Dar-es-Salaam (point north of French Somaliland: Djibouti (Le Plage), 15 and Oyster Bay), 11-13.x. 1962, Papenfuss & Scagel 16.ix.1962, Papenfuss & Scagel PR-I-65. PR-VIII-102 (female, tetrasporic and sterile [Figs. 4 Somalia: Las Khoreh, iii .1873, Hildebrandt and 5]). (UC 408988; voucher material of Hauck's 1886 Pakistan: near Karachi, no date, Kotwal (NY, p. 167 record of S. furcellatum). 'Type'; right half of the specimen illustrated by Kenya: Mambrui (north of Malindi), xii.1967, Isaac (ADU 40349, 40418); Malindi (Outer Malindi B0rgesen 1934 PI. 2; our Fig. 6). Bank), 5.x.1962, Papenfuss & ScageIPR-VC-55. Tanganyika: Kingombe (35 km south of Tanga Railway Station), 7.x.1962, Papenfuss & Scagel Acknowledgments PR-VII-34; Dar-es-Salaam (point north of Oyster Bay), 11-13 .x.1962, Papenfuss & Scagel PR-VIII-48 This work was done while the second author was on (female, Fig. 1, and tetrasporic). leave of absence from NRCC at the Department of Zanzibar: Bwenju Reef, 18 .x.1962, Papenfuss & Botany of the University of California, Berkeley, Scagel PR-XV-62. and the Department of Botany of the University of 42 Phyc% gia, Vol. 13 (1), 1974

Adelaide, South Australia. The material from the References Red Sea and East Africa was collected under the auspices of the United States Programme in Biology AGARDH, J.G. (1842) Algae Maris Mediterranei et Adriatici. Masson, Paris, x+ 164 pp. for the International Indian Ocean Expedition, which AGARDH, J.G. (1844) In: Systemata Algarum Hodierna Programme received its funds from the National Adversaria. [Thesis.] Lund, 56 pp. [Published by the author.] Science Foundation. As a participant in the United AGARDH, J.G. (1847) Nya alger fran Mexico. DIvers. K. VetenskAkad. Forh., 4, 5-17. States Programme the first author had two oppor­ AGARDH, J.G. (1852) Species Genera et Ordines Aiganlm, tunities in 1962 of collecting marine algae in East Vol. 2 (3 : I) Gleerup, Lund. pp. 701-786. AGARDH, J.G. (1872) Bidrag till florideernes systematik. African waters. During March and April he was a Lunds Univ. Arsskr., 8 (Afd. 2, No.6), 60 pp. participant in the Israel South Red Sea Expedition AGARDH, J.G. (1876) Species Genera et Ordines Alganlm, and from September to December Dr R. F. Scagel Vol. 3 (I). Epicrisis systematisjloridearum. Weigel, Leipzig, vii+724 pp. of the University of British Columbia and he had ALEEM, A.A. (1948) The recent migration of certain Indo­ their own expedition to East Africa. He should like Pacific algae from the Red Sea into the Mediterranean. 47, to express his appreciation to the Israel South Red New Phytol., 88-94, 2 pis. AUGIER, H. & BOUDOURESQUE, C.-F. (197 J) Decouverte des Sea Expedition and especially to its leader, the late cystocarpes de Feldmannophycus rayssiae (1. Feld. et Professor H. Steinitz of the Hebrew University of G. Feld.) nov. gen. (Rhodophycees, Gigartinales). Bull. Soc. phycol. Fr., No. 6, 2-30, 5 figs.,I table. Jerusalem, for inviting him to join the expedition. ] B0RGESEN, F. (1932) Some Indian Rhodophyceae, especially Dr E. Imre Friedmann helped with the collection of from the shores of the Presidency of Bombay: 1 1. KelV some of the Red Sea material. Dr H. B. S. Womersley Bull., 1932, 113-134, 18 figs., pis. 2-5. B0RGESEN, F. (1934) Some Indian Rhodophyceae, especially and Mr G. T. Kraft kindly allowed us to incorporate from the shores of the Presidency of Bombay: [V. K ew Bull., in our study the material of putative taxa of Dicra­ 1934, 1-30, 19 figs., vols. 1--4. nema which they had borrowed from Melbourne B0RGSEN, F. (1937) Contributions to a South Indian marine algal flora-ILJ. fndianbot. Soc., 16, 311-357, 20 figs. (MEL) and Vienna (W) for Mr Kraft's study of this B0RGESEN, F. (1938) Contributions to a South Indian genus and which proved to be representative of marine algal flora-Ill. J. Indian bot. Soc., 17, 205-242, 13 figs., pIs. 7 and 8. Sarconema. Mr Kraft generously also gave us photo­ B0RGESEN, F. (1939) Marine algae from the Iranian Gulf graphs of these taxa (reproduced as Figs. 23-25). especially from the innermost part near Bushire and the We are indebted to Miss Charlotte Mentges for help Island Kharg. Danish Scientific Investigations in Iran, Part I. Munksgaard, Copenhagen, pp. 47-141, 43 figs., with the illustrations. The processing of the Papen­ I map. fuss-Scagel collections was done with the aid of a B0RGESEN, F. (1943) Some marine algae from Mauritius. grant from the National Science Foundation III. Rhodophyceae, Part 2, Gelidiales, Cryptonemiales, Gigartinales, K. danske Vidensk. Selsk. Bioi. Medd., 19 (GB-1656). (I), 85 pp., 42 figs. I pI. B0RGESEN, F. (1950) Some marine algae from Mauritius. Additions to the parts previously published. H. K. danske Vidensk. Selsk. BioI. Medd., 18 (II), 46pp., 22 figs. B0RGESEN, F. (1952) Some marine algae from Mauritius. Additions to the parts previously published. IV. K. danske Vidensk. Selsk. Bioi. Medd., 18 (19),72 pp., 33 figs., 5 pIs. Appendix B0RGESEN, F. (1953) Some marine algae from Mauritius. Additions to the parts previously published. V. K. danske Vidensk. Selsk. Bioi. Medd., 21 (9), 62 pp., 20 figs.,3 pis. Rhabdol1iaceaeKylin (1932) proposed for conserva­ BORNET, E. (1892) Les algues de P.K.A. Schousboe [recoltees tion. au Maroc et dans la Mectiterranee de 1815 it18291. Mem. Soc. natn. Sci. nat. math. Cherbourg, 28, J 65-376 pls.I-3 BORNET, E. & THURET, G. (J 880) Notes Algologiques ... Rhabdoniaceae Kylin, Lunds Univ. Arsskr., N.F., Masson, Paris. Fasc. 2, pp. 73-196, pIs. 26-50. Avd. 2, 28 (8): 32, J932. Nomen /amiliarum conser­ DAWSON, E.Y. (1949) Contributions toward a marine flora of the southern California Channel Islands, 1-111. Allan vandum propositum. Genus typicus: Rhabdonia Hancock Found. Publ., Occas. Pap. No. 8, 57 pp., 15 pIs. W. H. Harvey. Nomen /amiliarum rejiciendum DAWSON, E.Y. (1961) Marine of Pacific Mexico, 2, propositum: Rhabdoniaceae Kylin, Lunds Univ. Part 4, Gigartinales. Pac. Nat., 191-341,63 pIs. DE TONI, G. (1936) Noterelle di Nomenclatura Algologica, Arsskr., N.F., Avd. 2, 21 (9): 38, 1925 [fam. illeg. VI L Primo elenco di Floridee omonime. Brescia. [8] pp. pro Solieriaceae (J. Agardh) Hauck, 1883]. [Published by the author.] DE TONI, J.B. (1897) Sylloge Alganlln .. , Vol. 4. , The Rhabdoniaceae as currently circumscribed . Sect. I. Padua, xx + Ixi + 386 pp. + [index, pp. 387 and 388�. includes five, and possibly ten, genera. The family [Published by the author. I has no synonyms; it would be advantageous, there­ DE TONI, J.B. (1924) Sylloge Aiganim ... , Vol. 6. Padua. xi + 767 pp. [Published by the author.] fore, to conserve the name rather than propose a new ERCEGOVIC, A. (1949) Sur la Yadranella, nouveau genre family name. d'algues de I'Adriatique et sur son developpement. Acta Papenfuss and Edelstein: Morphology and taxonomy of Sarconema 43

Adriatica [Inst. Oceanogr. Ribarstvo Split], 4, 27-39, 4 figs. Expedition 1933-34, Scientific Reports, Vol. 9. British Mus. FELDMANN, J. (1942) Remarques sur les Nemastomacees. (Nat. Hist.), London. pp. 395-420, pis. 1-4. Bull. Soc. bot. Fr., 89,104-113, I fig. OSTERHOUT, W.J.V. (1896) On the life-history of Rhabdonia GREVILLE, R.K. (1830) Algae Britannicae. Maclachlan and tenera J. Ag. Ann. Bot., 10, 403-427, pis. 20 and 21. Stewart, Edinburgh, Ixxxviii+218 pp., 19 pis. PAPENFUSS, G.F. (1968) A history, catalogue, and bibliography GRUNOW, A. (1873-74) Algen der Fidschi-, Tonga- und of Red Sea benthic algae. Israel J. Bot., 17, 1-118, I map. Samoa-Inseln, gesammelt von Dr E. Graeffe. J. Mus. PICCONE, A. (1884) Contribuzioni all'algologia Eritrea. Godeffroy, 3, 23-50. Nuovo Gior. Bot. Ilal., 16, 281-332, pis. 7-9. HARVEY,W.H. (1853) Nereis Boreali-Americana. Smithsonian RASMUSSEN, R.A. (1964) The structure and reproduction of ContI'. Knowledge, 5 [Art. 5], 285 pp., pis. 13-36. Sarcodia montagneana (Rhodophyta). Phycologia, 4, 1-7, HARVEY, W.H. (1855) Some account of the marine botany of 9 figs. the Colony of Western Australia. Trans. R. Irish A cad. , RAYSS, TSCHARNA (1963) Sur la presence dans la Mediter­ 22, 525-566. ranee orientale des algues tropicales de la famille des HAUCK, F. (1883-85) Die Meeresalgen Deutschlands und Solieriacees. Acta Bot. Horti Bucharest, 1961-1962, Oesterreichs. In: L. Rabenhorst's Kryptogamen-Flora von 91-106,10 figs. Deutschland, Oesterreich und del' Schweiz, 2nd Ed., Vol, 2. SCHMITZ, F. (1889) Systematische Ubersicht der bisher E. Kummer, Leipzig, xxiii + [I] + 575 pp., 236 figs., 5 pis. bekannten Gattungen der Florideen. Flora, 72, 435-456, HAUCK, F. (1886) Ueber einige von J. M. Hildebrandt im pI. 21. Rothen Meere und Indischen Ocean gesammelte Algen. SCHMITZ, F. (1895) Marine Florideen von Deutsch-Ostarrika. I. Hedwigia, 25, 165-168. Engler's Bot. Jahrb., 2],137-177. HEWITT, FLORENCE E. (1960) A morphological study of three SCHMITZ, F. & HAUPTFLEISCH, P. (1896-97) Rhodopbyceae. South African Gigartinales. Univ. Calif. Publ. Bot., 32, In: Die natiirlichen Pjlanzell!amilien (By A. Engler 195-233, 35 figs., pis. 26-32. and K. Prantl), W. Engelmann, Leipzig, Teil I, Abt. 2, HOOKER, J.D. & HARVEY, W.H. (1847) Algae tasmanicae: pp. 298-544, figs. 192-288. being a catalogue of the species of algae collected on the SEARLES, R.B. (1968) Morphological studies of red algae of shores of Tasmania by Ronald Gunn, Esq., Dr Jeannerett, the order Gigartinales. Ulliv. Calif. Publ. Bot., 43, 100 pp., Mrs Smith, Dr Lyall, and Dr J.D. Hooker; with characters 33 figs., 12 pis. of the new species. London J. Bot., 6, 397-417. SEARLES, R.B. (1969) Observations on the morphology of JOLY, A.B. & ALVEAL, K. (1969) Notes on Chilean algae-III. Trematocarpus dichotomlls KUtzing and the status of the The identity of Caulacanthus horridulus Montagne. genus Dicurella. Phycologia, 8, 21-25, 7 figs. Rickia, 4, 83-92, 15 figs. SETCHELL, W.A. & GARDNER, N.L. (1930) Marine algae of KRAFT, G.T. (1970) Eucheuma procrusteanum, a new red the Revillagigedo Islands Expedition in 1925. Proc. Calif. algal species from the Philippines. Phycologia, 8, 215-219, A cad. Sc., Ser. 4, 19,109-215, pis. 4-15. 7 figs. SILVA, P.e. (1957) Remarks on algal nomenclature. Taxon, KRAFT, G.T. & ABBOTT, ISABELLA A. (1971) Predaea weldii, 6,141-145. a new species of Rhodophyta from Hawaii, with an SONDER, O.G. (1845) Nova algarum genera et species, quas evaluation of the genus. J. Phycol., 7, 194-202, 15 figs., in itinere ad oras occidentales Novae Hollandiae, collegit 2 tables. L. Preiss, Ph. Dr. Bot. Ztg., 3, 49-57. KUTZING, F.T. (1843) Phycologia Generalis ... Brockhaus, SONDER, O.G. (1846) Algae. In: Plantae Preissianae ... Leipzig, xvi + I-I 44 +xvii-xxxii + 145-485 + [I] pp., 80 pis. quas in Australasia occidentali el meridionali-occidenta!i KUTZING, F.T. (1849) Species Aiganim. Brockhaus, Leipzig, annis 1838-1841 collegit Llldovicus Preiss, Ph. Dr. (By C. vi+922 pp. Lehmann). Meissner, Hamburg. Vol. 2 (I), pp. 148-195. KUTZING, F.T. (1868) Tabulae Phycologicae ... Vol. 18. SONDER, O.G. (1852) Plantae muellerianae. Beitrag zur Flora Nordhausen, [vi] + 35 pp., 100 pis. [Published by the SUdaustraliens, aus den Sammlungen des Dr Ferd. MUlier. author.] Algae. Linnaea, 25, 657-703. KUTZING, F.T. (1869) Tabulae Phycologicae ... Vol. 19. SONDER, O.G. (1871) Die Algen des tropischen Australiens. Nordhausen, iv + 36 pp., 100 pis. [Published by the author.] Abh. Natunviss. Hamburg, 5, 33-74, 6 pIs. KYLIN, H. (1925) The marine red algae in the vicinity of the SOUTH, G.R., HOOPER, R.G. & IRVINE, LINDA M. (1972) The Biological Station at Friday Harbor, Wash. Lunds Univ. life history of Turnerella pennyi (Harv.) Schmitz. Br. Arssk., N.F., Avd. 2,21 (9), 87 pp., 47 figs. phycol. J., 7, 221-233,13 figs., I table. KYLIN, H. (1928) Entwicklungsgeschichtliche Florideen­ TAYLOR, W.R. & RHYNE, e.F. (1970) Marine algae of studien. Lunds Univ. Arssk., N.F., Avd. 2, 24 (4), 127 pp., Dominica. Smithsonian Contrib. Bot., No. 3, 16 pp., 64 figs. 2 figs. KYLIN, H. (1932) Die Florideenordnung Gigartinales. Lunds TOKIDA, J. & MASAKI, T. (1957) On the female organ of Univ. Arssk., N.F., Avd. 2, 28 (8),88 pp., 22 figs., 28 pis. Tllrnerella mertensiana (Post. et Rupr.) Schmitz , Bull. Jap. KYLIN, H. (934) Uber die systematische Stellung der Soc. Phycol., 5, 5-7, 2 figs. [In Japanese with an English Gattungen Opuntiella und Turnerella. K. Fysiogr. Siillskap. summary.] Lund Forhandl., 4 (8), 6 pp., 2 figs. TREVISAN, V.B.A. (1848) Saggio di una Monografia delle KYLIN, H. (1941) Californische Rhodophyceen. Lunds Univ. Aighe Coccotalle. Padua. 112 pp, [Published by the author.] Arssk., N.F., Avd. 2, 37 (2), 51 pp., 7 figs., 13 pis. WEBER-VAN BOSSE, ANNA (1928) Liste des algues du Siboga. KYLIN, H. (1956) Die Galfungen der Rhodophyceen. Gleerup, IV. Rhodophyceae, partie 3, Gigartinales et Rhody­ Lund, xv + 673 pp., 458 figs. meniales. Siboga-Exped. MOllogr. 59d, Brill, Leiden. MONTAGNE, J.F.e. (1850) Pugillus algarum yemensium, quas pp. 393-533,71 figs.,pis. 11-16. collegerunt annis 1847-1849, clarr. Arnaud et Vaysiere. ZANARDINI, G. (1858) Plantarum in mari rubro hucusque Ann. Sc. Nat. Bot. Ser. 3, 13, 236-248. collectarum enumeratio. Mem. 1st. Veneto Sc. Lett. Arti, NEWTON, LINDA M. (1953) Marine algae. In: The John Murray 7,209-309, pIs. 3-14.

(Accepted 5 October 1973)