Phycologia (1985) Volume 24 (3), 317-337

A taxonomic and nomenclatural reassessment of Tenarea, Titanoderma and Dermatolithon (, Rhodophyta) based on studies of type and other · critical specimens*

WM. J. WOELKERLING', YVONNE M. CHAMBERLAIN2,4 AND PAUL C. SILVA3

'Department 0/ Botany, La Trobe University, Bundoora, Vic. 3083, Australia 2The Marine Laboratory, Portsmouth Polytechnic, Ferry Road, Hayling Island POll ODG, England 3Herbarium, Department 0/ Botany, University o/ California, Berkeley, CA 94720, USA

WM. J. WOELKERLING, Y.M. CHAMBERLAIN AND P.C. SILVA. 1985. A taxonomic and nomenclatural reassessment of Tenarea, Titanodermaand Dermatolithon (Corallinaceae, Rhodophyta) ba��d on studies of type and other critical specimens. Phyc% gia 24: 317 -337.

The taxonomy and nomenclature of genera of the TenarealDermatolithon complex has been reas­ sessed on the basis of a study of generitype and other critical specimens. The type specimen of Millepora tortuosa Esper, which is the type of Tenarea Bory 1832, proves to be conspecific with the specimen on which Bory based his description of the genus. Tenarea undu/osa Bory is an illegitimate name for T.tortuosa (Esper) Lemoine rather than an independent species. Since 1898, M. tortuosa has been incorrectly associated with the alga that is the principal component of the 'trottoirs' (coralline pavements) of the westernMediterranean. This alga is referable to Lithophyllum lichenoides Philippi. TitanodermaNageli 1858 is an earlier homotypic synonym of Dermatolithon Foslie 1898, both being based on Me/obesia pustu/ata Lamouroux. Titanoderma,in view of its clear record, is adopted in preference to proposing Dermatolithon for conservation. Tenarea comprises a single species of in which the thallus is composed of erect or ascending lamellae whose vegetative tissues are organized in an isobilateral manner and include a medulla of two layers of palisade cells. Titanoderma (syn. Dermatolithon) encompasses those Lithophylloideae in which the thallus consists largely or entirely of a prostrate crust whose tissues are organized in a dorsiventral manner and include a unistratose hypothallium composed of palisade-like cells. Titanoderma includes at least 16 species.

INTRODUCTION ically. The controversy was settled in favour of the second alternative(in the absence of an over­ The circumscription and naming of genera in the whelming case for conservation) in 1981 by the TenarealDermatolithon complex (Corallina­ Nomenclature Section of the International Bo­ ceae, Rhodophyta) has been attended by uncer­ tanical Congress at Sydney and is incorporated tainties as is indicated by Johansen (1981, pp. in Articles 10.2 and 10.3 of the current Inter­ 42, 217, 223, 225, 226) in his recent survey of national Code of Botanical Nomenclature . In addition to·diff erences in taxo­ (ICBN-see Voss 1983). Second, no attempt has nomic opinion, two factors are responsible for been made to examine the type specimens of these uncertainties. First, it has been a long­ these generic names and thus the nomenclature standing controversy whether a generic name of this complex has lacked the foundation that must be typified with material in the hands of is essential to stability. the author (that is, material upon which the ge­ During the course of studies on the nongenic­ neric description was primarily based) or, alter­ ulate Corallinaceae of the British Isles (by Y.M.C.) natively, with the type of a cited species, whether and of southern Australia (by W.J.W.), indepen­ or not the citation proves to be correct taxonom- dent visits were made by these authors to CN (Universite de Caen, France) in 1979 and 1980 * Dedicated to the memory of Mme Marie Lemoine to examine Lamouroux's specimens of Melobes­ (1887-1984) in recognition of her generous help over ia pustulata, Der­ 76 years (1909-1984) to all people interested in cor­ on which the generic name allines. matolithon is based. Subsequently (1981), Y.M.C. • Mrs Y.M. Butler. and W.J.W. concurrently studied material on

317 318 Phycologia, Vol. 24 (3),1985

which Bory based his description of Tenarea, abbreviations are taken from Holmgren et al housed at PC (Laboratoire de Cryptogamie, Mu­ (1981). seum National d'Histoire Naturelle, Paris). The whereabouts of Esper's specimen of Millepora tortuosa, on which the concept of Tenarea must HISTORICAL BACKGROUND be based in accordance with the Sydney decision, remained unknown until 1983 when one of us Tenarea (Y.M.C.) discovered that it was at FR (For­ Bory (1832, p. 207, pI. 54, fig. 3) established the schungsinstitut Senckenberg, Frankfurt, Ger­ genus Tenarea to accommodate a 'polypier' many). [Dr M. Grasshoff from FR has indicated (plant-like animal) collected at Cap Tenare (Cape in correspondence (with W.J.W.) that it was Tainaron), Greece. The organisms were de­ transferred from ER (Botanisches Institut der scribed as calcareous, canary yellow when grow­ Universitiit Erlangen, Germany) to FR in 1970.] ing but bleached when dry and composed of Comparative examination of the Lamouroux, lamellae that were undulate-tortuouse, anasto­ Bory, and Esper specimens together with studies mosing, and folded together. Bory was explicit of more recently collected materials enabled us in indicating that it was not new to science at the to resolve various uncertainties in the Tenareal species level and cited Millepora tortuosa Esper Dermatolithon complex at the generic level. Type (1796, p. 118, Millepora pI. xxii) in synonymy. specimens have been designated and relevant This name had been applied by Esper to a more historical background data are provided. or less globular calcareous organism with thin, ascending, variously twisted lamellae which were MATERIALS AND METHODS closely united with one another and which had scattered hemispherical 'pores' (conceptacles). Data were obtained from type and other critical Esper's specimen presumably came from the specimens housed at CN, FR, L (Rijksherba­ Mediterranean, and he compared his organism rium, Leiden, Netherlands) and PC and from with several others, including Corallina offici­ additional collections housed at L TB (Depart­ nalis Linnaeus and Millepora lichenoides (Ellis) ment of Botany, La Trobe University, Bundoora, Ellis & Solander. Unfortunately, in an action that Victoria, Australia). Microtechnique follows complicated subsequent nomenclature, Bory Jones & Woelkerling (1984) and permanent slides (1832) changed the specific epithet to undulosa. have been retained at LTB. Fractures for scan­ The ICBN always has proscribed unnecessary ning electron microscopy were prepared by changing of epithets and if Bory had not estab­ breaking small pieces of thallus, mounting these lished a new genus at the same time that he on aluminium stubs with 'Fotobond' acrylic ad­ changed the epithet, no problem would have aris­ hesive (Agfa-Gevaert Limited), and double-coat­ en. Establishing a new genus, however, allowed ing the specimens with carbon and then with gold for the interpretation of Tenarea undulosa as a prior to viewing in a Siemens ETEC Autoscan new species and the type of its genus, an inter­ microscope. Wherever cell measurements are pretation which has been held by many botanists given, 'L' denotes length, 'D' denotes diameter, but which was ruled out by the Sydney decision and 'LID' denotes the ratio of length to diameter. in 1981 (ICBN, Arts 10.2, 10.3; see Voss 1983). Measurements include the cell walls. Herbarium T'enareawas overlooked until Hariot (1895)

Figs 1-6. Lec totype of Tenarea tortuosa (Esper) Lemoine. Fig. 1. Lec totype specimen (housed at FR). Fig. 2. Surf ace of lectotype showing numerous, tortuouse, intertwined, branched lamellae. Fig. 3. Apex of lamella showing characteristic groove. Fig. 4. Close-up of portion of groove (arrows). Note distinct cell row on opposite sides of groove. Fig. 5. Surf ace of lamella showing early stage of branch development, three co nceptacles, and numerous co n­ cavities denoting the positions of epithallial cells. Fig. 6. Fracture showing one lamella (a) abutting against the surface of another lamella (b). Both lamellae are fractured in a transverse plane. Woelkerling et al: Tenarea, Titanoderma and Dermatolithon 319 320 Phycologia, Vol. 24 (3), 1985

Figs 7-11. Tenarea tortuosa. Lectotype, Figs 7- 10; LTB 14645, Fig. II. Fig. 7. Penetration of one lamel1a (a) by another lamel1a (b); the latter is transversely fractured above the region of penetratio n. Woelkerling et al: Tenarea, Titanodermaand Dermatolithon 321

re-examined Bory's specimen and confirmed that hypothallium that is unistratose and palisade­ it was a nongeniculate coralline alga. He noted like (i.e. with vertically elongate, more or less Bory's citation of Esper's diagnosis of Millepora obliquely arranged cells) and thus generically dis­ tortuosa, but chose to retain the epithet undulosa. tinct from Lithophyllum; and Considering mainly external morphology, Ha­ (2) that plants identifiedby Lemoine and oth­ riot concluded that Tenarea Bory (1832) was ers as Tenarea tortuosa are truly referable to congeneric with and had priority over Litho­ Lithophyllum, belonging to the species L. tor­ phyllum Philippi (1837). He did not examine tuosum (Esper) Foslie. Huve's concept of Lith­ Philippi's collections, however, and the first ophyllum tortuosum, however, was not based on modem account of this critical material was not an examination of Esper's type. published until 88 years later (Woelkerling 1983). Dermatolithon Without examining the type collections, both Foslie (1895, p. 178, footnote; 1898d) and Kuntze This name was applied by Foslie (1898b, p. 11) (1898, p. 433) accepted Hariot's conclusion, and to his emended concept of Melobesia (Foslie, although Kuntze adopted Tenarea over Litho­ 1898a, p. 6). This genus was characterized by phyllum, Foslie rejected Tenarea on the grounds uniporate conceptacles in which sporangia occur " that Lithophyllum . . . has been accepted and around or between a bundle of cylindric-clavate applied for about 60 years ... " (Foslie 1898d, paraphyses. He typified Dermatolithon with Me­ p. 5). Foslie (1898b, p. 9; 1898d, p. 5), however, lobesia pustulata Lamouroux (1816, p. 315, pI. adopted the epithet tortuosa over the epithet un­ 12, fig. 2; see also Lamouroux 1821, p. 46, pI. dulosa, believing that the two taxa were conspe­ 73, figs 17, 18). This 'polypier' was said by La­ cific (as originally indicated by Bory). moroux to grow on along the coast of Subsequently, Lemoine (1910, p. 368; 1911, France, and to take the form of convex orbicular pp. 5, 62, 169) undertook an anatomical exam­ plates with ostioles (of the conceptacles) visible ination of the alga which forms the 'trottoirs' to the naked eye. (coralline pavements) commonly seen on west­ Subsequently, Foslie (1904, p. 3) reduced Der­ ern Mediterranean shores (Lemoine 1911; Feld­ matolithon to a subgenus of Lithophyllum, but mann 1937; Lewalle 1961). She concluded that later (1909, p. 57) again elevated it to generic this alga, which was then generally known as rank, distinguishing it from Lithophyllum on the Lithophyllum tortuosum (Esper) Foslie, differed basis that thalli of Dermatolithon have a uni­ from other members of Lithophyllum (sensu Le­ stratose, palisade-like hypothallium whereas moine) in having a multistratose but noncoaxial those of Lithophyllum have a multistratose, non­ hypothallium and a perithallium composed of palisade-like hypothallium. Foslie, however, ap­ more or less juxtaposed filaments. She adopted parently never examined relevant generitype the generic name Tenarea for the 'trottoir' alga specimens. without examining Bory's specimen anatomi­ Between 1909 and 1957, some authors (e.g. cally. Had she done so, she would have found Rosenvinge 1917; Newton 1931; Feldmann 1939; that the two entities were not congeneric, as did Suneson 1950) treated Dermatolithon as a sub­ Huve (1957). Afterstudy ing Bory's specimen and genus of Lithophyllum while other authors (e.g. recently collected material from the original 10 - Svedelius 1911; Mazza 1917; De Toni 1924; Funk cality, Huve concluded: 1927, 1955; Hamel & Lemoine 1953; Mason (1) that Bory's alga (for which she retained the 1953; Dawson 1955; Kylin 1956; Huve 1957) name Tenarea undulosa) is characterized by a considered Dermatolithon to be a distinct genus,

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Fig. 8. Lon gitudinal fr acture through apex of a lamella. Note minute epithallial cells (small black arrows), elongate medullary ce lls (m), meristem cells at the apex (large black arrows), primary pit connection between cells of the same fil ament (large white arrow), and position of secondary pit connections between cells of contiguous filaments (small white arrows). Fig. 9. Transverse fracture of margin of a lamella. Epithallial ce lls are indi cated with small arrows; remainin g layers are composed of medullary cells viewed 'end on'. Fig. 10. Transverse fracture of central portion of older lamella. All ce lls are medullary cells; epithallial cells have not been preserved. Fig. 11. Fracture showing relationships of tissues as seen in transverse (T) and longitudinal (L) views. Note epithallial cells (small black arrows) and cells of medulla (m). 322 Phycoiogia, Vol. 24 (3), 1985

based on the occurrence of a unistratose hypo­ thallium. Huve (1957), after concluding that Bory's alga was generically distinct fr om Litho­ phyllum, differentiated it fr om Dermatolithon on the presence or absence of erect lamellae-pres­ ent in Tenarea, absent in Dermatolithon. Sub­ sequently, some authors (e.g. Cabioch 1972; Bressan 1974; Johansen 1981) have accepted this distinction while others (Adey 1965, 1970; Lit­ tler 1971; Adey & Adey 1973; Adey & Macintyre 1973; Notoya 1974; Johansen 1976a, 1976b; Townsend 1981) have recognized only one genus (Tenarea), in which they include all nongenicu­ late Corallinaceae with secondary pit connec­ tions and a palisade-like hypothallium. Still oth­ er authors (e.g. Mason 1953; Dawson 1955, 1960; Masaki 1968) place all Lithophylloideae (see Jo­ hansen 1981, p. 11, for subfamily characteristics) with a unistratose, palisade-like hypothallium in one genus, which they call Dermatolithon. Fi­ nally, Adey (1965, p. 78) suggested that Esper's plant represented an unnamed genus. None of these concepts and opinions, however, were based on studies of relevant generitypes.

OBSERVATIONS

Millepora tortuosa Esper

Lectotypification

The Esper collections of Millepora at FRinclude a single unnumbered specimen (Fig. 1) labeled Millepora tortuosa Esper. According to Dr M. Grasshoff (personal communication with W.J.W.), the label is not in Esper's handwriting, but apparently all original labels have been re­ placed. The specimen, whose dimensions and other attributes are concordant with those men­ tioned in the protologue (Esper 1796, p. 118), is chosen here as lectotype.

the lamellar groove, two rows of medullary cell de­ rivatives (m), each with a minute epithallial cell(black arrows). LTB 14645. Fig. 13. Fracture through a uniporate co nceptacle of the lectotype specimen. Note pore (arrow) and inside Figs 12-14. Tenarea tortuosa. of empty chamber. Fig. 12. In terference contrast view of longitudinal Fig. 14. Conceptacle oflectotype specimen with part section through apex of a lamella showing two dense­ of roof removed to show remains of a central col­ ly stained meristem cells (white arrows) situated in umella. Woelkerling et al: Tenarea, Titanoderma and Dermatolithon 323

Morphology and anatomy comments on these ter ms.] Each palisade-like medullary cell bears a small epithallial cell (Figs The lectotype (Fig. I) is a somewhat elongate 8, 9). Collectively these cells form a unistratose conglomerate of intertwined lamellae 120 mm epithallium on each lamellar surface, but this long, 91 mm broad, and 53 mm thick. Esper layer is poorly preserved in the lectotype, and listed the dimensions as approximately 4 inches when lamellae are examined in surface view (Fig. (102 mm) long and broad. The lower surface is 5), numerous concavities denoting the position flat, more or less dull white, and contains a cy­ of epithallial cells are evident. Within a given lindrical hole c. 11.5 mm in diameter and c. 25 filament, medullary cells are joined by primary mm deep. Whether Esper's fig. 3 is based on pit connections (Fig. 8). Medullary cells of lat­ material taken fr om this hole is uncertain. Re­ erally contiguous filaments may be joined by sec­ maining surfaces vary fr om dull white to sooty ondary pit connections (Fig. 8), but pit connec­ grey, the latter colour probably due to long ac­ tions across the midline were not observed. Cell cumulation of dust. fu sions do not occur. In transverse sections of The lectotype lacks any conspicuous holdfast, lamellae (Figs 9, 10), the palisade-like nature of prostrate system, or other evidence of attach­ the oblique medullary cells and the isobilateral ment and is composed almost entirely of erect organization of the thallus are obscured. For or ascending lamellae (Fig. 2) which are mod­ comparison, the relationship between tissue ap­ erately to heavily calcified, flattened, variously pearance in transverse and longitudinal sections twisted, irregularly branched to differing degees, is shown in Fig. II, which was prepared fr om and densely intertwined or interlocked. Most la­ recently collected material (LTB 14645, leg. E. mellae are 1-5 mm broad, and while some apices Coppejans, South Coast, Isle of Sikinos, Cy­ remain intact, many have been damaged. A fairly clades, Greece, August 1972). distinct groove occurs around the lateral and api­ Detailed data on lamellar ontogeny and on cal edges of most lamellae (Fig. 3), and the cell meristems were not obtainable fr om the lecto­ rows on the two sides of the groove tend to be type, and only limited information was gleaned aligned oppositely (Fig. 4). Branches appear to fr om recently collected specimens. It appears that result fr om continued growth of lobes formed at the palisade cells comprising each medullary cell the apex of a lamella (Fig. 5), probably caused row are derived basipetally fr om anticlinal­ by differential activity of meristem cells in a transaxial divisions of a terminal meristem cell manner analogous to that fo und in Mastophora (Figs 8, 12) which lies in the groove (Fig. 3) at (Turner & Woelkerling 1982). Branches com­ the lamellar apex. Cell elongation occurs only in monly abut one another, or in some cases one the basipetal derivative, the meristem cell never branch penetrates another (Figs 6, 7). Where becoming greatly elongated. Soon after fo rma­ abutment occurs, the two lamellae become firmly tion, each medullary cell undergoes a single adherent to one another (Fig. 6). asymmetric, periclinal or somewhat oblique di­ Anatomically, the lamellae are organized in an vision to produce a small epithallial cell acrop­ isobilateral manner, and their appearance in lon­ etally. Medullary cells apparently do not undergo gitudinal and transverse views is quite different further divisions except in association with con­ (Figs 8-10). Except near conceptacles, each la­ ceptacle production, but details on this aspect mella usually consists of fo ur cell layers (two are uncertain. medullary and two epithallial) when seen in lon­ Further studies, based on liquid-preserved gitudinal view (Fig. 8). The central two rows of specimens and on cultured plants, are needed to cells are vertically elongate (L 75-130 JLm;max. determine just how the lamellar meristem is D 12-25 JLm; LID 4-7) and slanted towards the formed. In more mature lamellae, the meristem apex. Some authors (e.g. Huve 1957; Cabioch appears to be composed of two rows of contig­ 1972; Bressan 1974; Johansen 198 1) have used uous cells situated in the lamellar groove (Figs the term 'hypothallium' or 'palisade hypothal­ 3, 4, 12). Each row gives rise to one of the two lium' when referring to the tissue comprising these layers of subtending medullary cells (Figs 8, 12), cells, but it must be remembered that this tissue and it appears that cell division activity within is in the center of an isobilaterally organized la­ the two rows of meristem cells is more or less mella and thus is more properly termed a me­ equal in rate, resulting in the isobilateral orga­ dulla. [See Turner & Woelkerling (l982, pp. 214- nization of tissues (Figs 8-12). What remains 215), and Woelkerling (1980a, p. 221) for further unknown is whether the two rows of meristem 324 Phycologia, Vol. 24 (3), 1985

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Figs 15-20. Bory's specimen of Tenarea undulosa. Fig. 15. Bory's specimen housed at PC. Fig. 16. Surf ace of specimen showing apices of numerous intertwined lamellae. Fig. 17. Apices of two lamellae showing characteristic groove. Fig. 18. Lo ngitudinal fracture through apex of a lamella. Compare with Fig. 8. Fig. 19. Transverse fracture of margin of a lamella. Compare with Fig. 9. Fig. 20. Transverse fracture of central portion of a lamella. Compare with Fig. 10. Woelkerling et al: Tenarea, Titanoderma and Dermatolithon 325

cells originate independently or are derived fr om closely with Lamouroux's rather stylized draw­ a single row during very early stages of lamellar ing (1816, pI. 12, fig. 2). Collection data are lack­ formation. If we consider Cabioch's comments ing. and figure (1972, p. 209, pI. 5, fig.6), the second course seems more probable, a single row of mer­ Morphology and anatomy istem cells at some stage undergoing a synchro­ nous periclinal�oaxial division. Plants in the lectotype collection form a series Uniporate conceptacles are scattered over both of flattened, irregularly shaped crusts less than 1 surfaces of the lamellae in the lectotype (Fig. 5). mm thick on both sides of the compressed sur­ Conceptacles are solitary or in groups, and the fa ce of the host. Adjacent crusts abut (Fig. 30) more or less domoid roofs are up to 350 �m but do not overgrow one another, and most or across and protrude up to 130 �m above the all of the central surface of each plant is firmly lamellar surface. Conceptacles (Figs 13, 14) pre­ attached to the host, as evidenced by cellular sumably arise from derivatives of medullary cells remains on the host surface where portions of and possess more or less ovoid chambers up to the epiphyte have become detached (Fig. 30). 305 �m broad and 90 �m high. The remains of Excrescent and ascending lamellae do not occur, a central columella (Fig. 14) suggest that the lec­ and except for concavities which mark the po­ totype is (bi-) tetrasporangial, but sporangia were sition of epithallial cells (Figs 30, 32), the dorsal not seen. Details of conceptacle roof formation surface of the vegetative thallus is smooth. Uni­ are uncertain; at maturity the roof consists most­ porate conceptacles occur in a scattered fa shion ly of one or two layers of cells which appear to on some crusts; conceptacle roofs protrude only be interconnected by secondary pits. Cell fu sions slightly above the thallus surface and are more between roof cells were not seen. or less domoid or have slightly flattened tops (Figs 37-39). Tenarea undulosa sensu Hory Anatomical examination ofthe lectotype shows that the thallus is organized dorsiventrally; the Morphology and anatomy appearance of perithallial and epithallial cells is The specimen on which Bory based his descrip­ similar in longitudinal and transverse views (Figs tion of Tenarea (Fig. 15), as noted by Hariot 34, 35). In young plants or near the crust margin (1895, p. 113) and Huve (1957, p. 136), is pre­ (Figs 31, 32), the thallus consists of a unistratose served at Pc. It consists of numerous inter­ hypo thallium composed of palisade cells (L 40- twined, erect or ascending lamellae which fo rm 85 �m; D 10-23 �m; LID 2-6). These hypo­ an irregularly shaped conglomerate up to 215 thallial cells arise basipetally fr om a primary mm long and 155 mm broad. The specimen does marginal meristem (Fig. 31) and each hypo­ not have a holdfast or other conspicuous attach­ thallial cell, in turn,gives rise dorsally to a small, ment mechanism. noncalcified epithallial cell (Fig. 36). Epithallial Morphologically and anatomically, Bory's cells are not apparent on much of the lectotype, specimen (Figs 15-26) and Esper's specimen (Figs but their positions are delineated on the dorsal 1-14) are concordant, and there is no doubt that surface by concavities. In older portions of the Bory was correct in concluding that the two en­ lectotype plants, perithallial filaments up to five tities were conspecific. All of the features de­ cells deep also occur (Figs 34, 35). In these por­ scribed in Esper's material are equally apparent tions of the thallus, hypothallial cells appear to in Bory's collection. be obliquely slanted when viewed longitudinally (Fig. 34), but appear more upright when viewed Melobesia pustulata Lamouroux transversely (Fig. 35). The production of peri­ thallial filaments in this species is similar to that Lectotypification in Metamastophora (Woelkerling 1980a) and in Lamouroux (1816, p. 315) did not designate a Lithoporella (Turner & Woelkerling 1982). First, type specimen. His herbarium at CN contains a a secondary meristem is fo rmed as a result of a single unnumbered collection (Fig. 29) labeled series of periclinal, transaxial divisions of hy­ Melobesiapustulata in his handwriting, which is pothallial celis (Figs 32, 33). These meristem cells chosen here as lectotype. The collection consists undergo fu rther periclinal divisions and give rise of a few plants ofthe coralline growing on Chon­ basipetally to additional perithaliial cells (Figs drus (Rhodophyta), but the host does not agree 33-35) which are quadrate to vertically elongate 326 Phyc% gia, Vol. 24 (3), 1985 Woelkerling et al: Tenarea, Titanodermaand Dermatolithon 327

Figs 27, 28. Substrate relations of Tenarea tortuosa (LTB 14646). Fig. 27. Interface between lamellae of T .. tortuosa and a host coralline alga. Note how lamella abut the substrate (arrows). Fig. 28. Erect portions of lamellae (arrows) arising from prostrate portions adherent to substrate.

(L 34-123 !Lm; D 18-29 !Lm; LID 1.1-4.6) and occurred beneath the conceptacle floorand two may be aligned in more or less regular rows. or three layers of cells occurred in the roof; sec­ Within a given hypothallial or perithallial cell ondary pits were evident but cell fu sions were row, adjacent cells are connected by primary pits; not observed. contiguous cell rows may be interconnected by 'Trottoir' alga secondary pits (Fig. 36). Cell fu sions were not observed. Specimens in PC fr om Banyuls (Mediterranean Two conceptacles from lectotype material were France) identified by Lemoine as Lithophyllum examined anatomically (Figs 38, 39). These pos­ tortuosum and a more recent similarly named sessed more or less ovoid chambers which were collection by Coppejans fr om La Dromont up to 400 !Lmacross and 130 !Lm high and were (Mediterranean France) have been examined to devoid of contents. Two to four layers of cells determine the correct name for this alga. We

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Figs 21-26. Bory's specimen of Tenarea undulosa. Fig. 21. Longitudinal fracture through portion of a lamella showing minute epithallial cells (small black arrows), elongate medullary cells (m) with swollen tips (t), primary pit connections between cells of the same filament (large black arrows) and position of secondary pit connections between cells of contiguous filaments (small white arrows). Fig. 22. Fracture showing relationships of tissues as seen in transverse (T) and longitudinal (L) views. Compare with Fig. II. Fig. 23. Surface view of uniporate conceptacle. Fig. 24. Fracture through a uniporate conceptacle showing structure of roof. Pore not shown. Note parts of two intact conceptacles (C) on lower surface of lamella. Fig. 25. Portion of a host (H) Tenarea lamella with an epiphytic plant (E) of Titanoderma. Note size difference in conceptacles of the host (lower surface) and the epiphyte (upper surface). Fig. 26. Longitudinal fracture ofa host (H) Tenarea lamella and a Titanoderma epiphyte (E). Note size difference between hypothallial cells (white arrow) of the epiphyte and the corresponding medullary cells (black arrow) of the host. 328 Phyc% gia, Vol. 24 (3), 1985

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/I<>� ty"'""J� b..t- Ii> 1L-<� [p'�. WoJ.W, ..J( .i'tW>...... � '""" �_ � (.;rrrw\.k � fk(\'1(. h I.. �.2. �. �(h'f'f Woelkerling et al: Tenarea, Titanoderma and Dermatolithon 329

have found that these specimens are conspecific undulosa share the same type (Table 1). The with the type specimen of Llthophyllum lichen­ 'trottoir' alga, to which the name Tenarea tor­ oides Philippi (1837, p. 389) fr om Sicily, a de­ tuosa has generally been misapplied since the tailed account of which has been published else­ time of Lemoine (1910, 1911)*, is ref erable to where (Woelkerling 1983, p. 317, figs 24-32). It Lithophyllum (see Woelkerling 1983, p. 324) is of interest to note that Lemoine (1911, p. 169) rather than to its own genus as thought by Le­ listed L. lichenoides Philippi as a synonym of moine, and moreover is referable to L. lichen­ Tenarea tortuosa, the name that she incorrectly oides Philippi. applied to the 'trottoir' alga. An examination of the type specimen of Me-

DISCUSSION AND CONCLUSIONS * The misapplication of the epithet tortuosa was ini­ tiated by Foslie (1898b, p. 9), who placed Esper's species Nomenclatural implications in the genus Goniolithon [as G. tortuosum (Esper) Fos­ lie] . Foslie circumscribed Esper's species to include The discovery of the type specimen of Millepora Lithophyllum cristatum Meneghini (1840), based on tortuosa Esper together with the fixing of pro­ specimens from the Ligurian Sea, and L. crassum Ro­ cedures fo r typifying generic names permits clar­ sanoff(l866, p. 93, pI. 7, figs 5, 7), based on specimens from Atlantic France. The fo rmer synonym was rec­ ification of the status of Tenarea and thus re­ ognized as forma cristatum, the latter as forma cras­ moves an extremely bothersome uncertainty from sum. Sub sequently, Foslie's (1898c, p. IS) stated that the nomenclature of nongeniculate Corallin­ "M. tortuosa Esp. without any doubt represents an old aceae. Demonstrating the con specificity of the specimen of L. cristatum f. crassa." Lithophyllum cras­ sum L. crassum type of M. tortuosa and the specimen upon which Rosanoff [not to be confused with (Philippi) Heydrich] had been treated previously as a Bory based his description of Tenarea was an fo rm of L. cristatum by Hauck (1883, p. 27 1, pI. 2, fig. important and satisfying result of the present 5, pI. 3, fig . 9) and as a variety of Tenarea undulosa study. Had the two specimens proved to belong Bory by Hariot (1895). Two years after having pro­ to different genera, there would have been the posed the genus Goniolithon, Foslie (1900a) remodeled it in such a way as to exclude Millepora tortuosa, wh ich possibility of conserving the name Tenarea as he removed to Lithophyllum [as L. tortuosum (Esper) typified by Bory's specimen, an exceptional pro­ Foslie]. De Toni (1905, p. 1792) adopted Foslie's treat­ cedure which is allowed by the ICBN (Art. 10.3) ment and was fo llowed by Lemoine (1910, 1911), who, but which might engender controversy. As it is, however, perceived a generic distinction between the 'trottoir' alga and Lithophyllum and therefore applied there is only one possible application of the to it the name Tenarea, making the combination T. name- the one determined by the type specimen tortuosa (Esper) Lemoine. The subsequent accounts of of the type species, Millepora tortuosa Esper. Tenarea tortuosa (or Lithophyllum tortuosum) provid­ Tenarea undulosa Bory must be considered an ed by Mazza (1917), De Toni (1924), Funk (1927, illegitimate name for the type species, sharing its 19 55), Feldmann (1939), Hamel & Lemoine (1 953), Masaki (1968), Ardre (1970, 19 71), Bressan (1974), type specimen in accordance with Arts 63.1, 63.2, and Boudouresque & Perret (1977), among others, are and 7.11 of the ICBN. All names based on Mil­ almost certainly based on plants referable to Litho­ lepora tortuosa and all those based on Tenarea phyllum lichenoides Philippi.

Figs 29-33. Lectotype of Titanoderma pustulata (Lamouroux) Nageli. Fig. 29. Lectotype collection housed at CN. A number of plants cover the host thallus (Chondrus). Data on annotation label of Chamberlain refer to Lamouroux 1816. Other labels are in Lamouroux's script. Fig. 30. Abutment of two adjacent cr usts (a, b) in lectotype collection. Notejuncture line (large arrow), concavities (small arrow) marking position of epithallial cells, and cellular remains on surface of host (Chondrus, H) where portions of the epiphyte have become detached. Fig. 31. Longitudinal fracture of crust margin showing terminal primary meristem cell (m), hypothallial cell derivatives (h) and concavities marking position of epithallial cells on dorsal surf ace. Fig. 32. Longitudinal fracture of crust showing fo rmation of secondary (s) meristem cells as a result of periclinal transaxial divisions of hypothallial cells (h). Fig. 33. Lo ngitudinal fracture showing early development of perithallial cells (p). Note hypothallium (h) and perithallial meristem (m). 330 Phyco!ogia. Vol. 24 (3), 1985

Figs 34-39. Lectotype of Titanoderma pustulata (Lamouroux) Nageli. Fig. 34. Longitudinal fracture of older portion of crust. Note oblique hypothallial cells (h). Fig. 35. Transverse fracture of older portion of crust. Note more or less upright hypothallial cells (h). Woelkerling et al: Tenarea, Titanoderma and Dermatolithon 331

lobesia pustulata Lamouroux, which is the type see also Adey 1965, p. 79, 1970, p. 6; Cabioch of Dermatolithon, has shown that it agrees with 1972, pp. 209, 270; Johansen 1981, p. 43, fig. the concept of that genus held by recent authors 3B) that erect lamellae form when two prostrate (e.g. Cabioch 1972; Bressan 1974; Johansen crusts come into contact and then grow upright 1981). back to back. Indeed, ascending lamellae can arise Having cleared up much of the uncertainty in directly as intact units fr om portions of the thal­ the Tenareal Dermatolithon complex, we now lus which adhere to an underlying substrate (Figs must introduce a slightly discordant note. In a 27, 28). Moreover, when Tenarea lamellae be­ work that has been almost completely over­ come forked (Fig. 5), the resulting branches al­ looked, Nageli (1858, p. 532) established the ge­ ways retain an isobilateral organization, provid­ nus Titanoderma to receive Melobesia pustulata ing further evidence that functionally the lamellae Lamouroux, distinguishing it fr om Melobesia by are integral units. In no case was an intact lamella its production of bispores rather than tetra­ found which lacked an isobilateral organization spores. Nageli included only one species in the or looked like a monostromatic crust. Adey (1970, genus. To our knowledge, the only subsequent p. 6) noted in relation to his concept of Tenarea appearance of this name in literature is its listing that "... the characteristic of thallic upgrowing by Silva in the Index Nominum Genericorum base to base is a common and apparently largely (Plantarum) (Farr et al 1979, p. 1764). Titano­ fo rtuitous fe ature in many crustose corallines ..." derma is clearly an earlier homotypic synonym and that " ...it is not warranted to consider this of Dermatolithon. Although it would be possible a generic fe ature." In view of evidence that the to propose conservation of Dermatolithon, this lamellae of Tenarea can arise and fu nction as name has undergone such vicissitudes that it single, integral units, and do not form by the seems better to adopt Titanoderma, a name with fo rtuitous union of independent crusts, this ar­ a completely clear record. gument of Adey's no longer appears tenable. Secondly, Cabioch (1972) and Johansen (1981) The genus Tenarea incorrectly characterized Tenarea by the occur­ CHA RACTERIZATION: Tenarea, as delineated rence of a unistratose, palisade hypothallium, a here, encompasses those nongeniculate Corall­ characterization probably based on data provid­ inaceae which have uniporate (bi-) tetrasporan­ ed by Huve (1957, p. 134, fig. 8). Huve reported gial conceptacles, secondary pit connections be­ that in "young plants" of Tenarea tortuosa (as tween cells of adjacent filaments, and a thallus T.undulosa) the thallus was prostrate and pos­ composed of erect or ascending lamellae whose sessed a unistratose hypothallium whose cells vegetative tissues are organized in an isobilateral were 40-50 /Lm long and 5-10 /Lm broad. These manner and include a central region of two layers characteristics contrast with those ofthe lamellae of palisade-like cells. Nomenclatural and taxo­ (Huve 1957, fig. 4), which were erect and pos­ nomic data relating to Tenarea are summarized sessed two layers of"hypothallial" cells, each 80- in Table 1. 100 /Lm long. Huve makes no mention of seeing COMMENTS: The above circumscription of Ten­ erect lamellae arising fr om the "young" plants, area is most closely associated with the concept and while the "young" plants appear (Huve 1957, of the genus provided by Huve (1957), Cabioch fig. 3) to be largely affixed to an unidentified sub­ (1972), Bressan (1974) and Johansen (1981), but strate, the older plants (Huve 1957, p. 133) are several points require emphasis. Firstly, the iso­ stated to be attached to the substrate only in bilateral lamellae appear to be integral structures several places and removed very easily. which, as shown by Cabioch (1972, p. 209b, pI. During the present study, plants corresponding Y, fig. 6), can arise as single units fr om one sur­ to those described as "young" thalli by Huve fa ce ofa parent lamella. No evidence was found were encountered as epiphytes on lamellae of in support of Huve's hypothesis (1957, p. 134; Bory's specimen of Tenarea tortuosa (Figs 25,

.-

Fig. 36. Fracture of upper portion of crust showing remains of two epithallial cells (E) and secondary pit connections (arrows) between cells of contiguous filaments. Fig. 37. Surf ace view of uniporate conceptacle. Note ostiole (arrow). Figs 38,39. Fractures of two conceptacles. Note ostiole (black arrows) and remnants of columella (white arro w) . 332 Phycologia, Vol. 24 (3), 1985

Table 1. Nomenclatural and taxonomic data relating 26) as well as on more recently collected speci­ to Tenarea and Titanoderma and their type species mens (LTB 14645). The epiphytes are prostrate and have a unistratose, palisade hypothallium Tenarea Bory 1832, p. 207 whose cells are much smaller than the corre­ Type species: Tenarea tortuosa (Esper) Lemoine 1910, sponding palisade cells of the lamellae of Ten­ p. 368 (only as to binomial). Basionym: Millepora tortuosa Esper 17 96, p. 118, area tortuosa (Fig. 26). Moreover, the epiphyte Millepora Tab. XXII. possesses conceptacles which are considerably Nomenclatural synonyms: smaller than those of T.tortuosa (Fig. 25). The Crodelia incrustans f. tortuosa (Esper) Heydrich occurrence of this epiphyte, combined with the 1911, p. 13 (only as to trinomial) facts that Huve makes no mention of erect la­ Goniolithon tortuosum (Esper) Foslie 1898b, p. 9 (only as to binomial) mellae arising from her "young" plants and that Lithophyl/um tortuosum (Esper) Foslie 1900a, p. the cellular dimensions of Huve's young plants 20 (only as to binomial) are only half those of the lamellae, suggests to us Superfluous substitute names: that what Huve considered to be "young" plants Tenarea undulosa Bory 1832, p. 207 of T.tortuosa may really represent a species of Goniolithon tortuosum f. undulosum (Bory) Fos­ Titanoderma. lie 1898b, p. 9 (,undulosa') Lithophyllum cristatum f. undulosum (Bory) Finally, Bressan (1974, p. 106) noted that Ten­ Heydrich 1901, p. 537 (,undulosa') area could be distinguished from Dermatolithon Lithophyllum tortuosum f. undulosum (Bory) (= Titanoderma) by the occurrence of triangular Foslie 1900a, p. 20 ('undulosa') epithallial cells. We have observed triangular Tenarea tortuosa f. undulosa (Bory) Lemoine 19 10, p. 368 epithallial cells in longitudinal sections of the generitype specimen of Tenarea (Fig. 9) and more Type locality: Mediterranean Sea (Esper 1796, p. 118). Lectotype specimen: FR(Fig. I) or less rounded-rectangular epithallial cells in the generitype specimen of Titanoderma (Fig. 38). Additional references (all as Tenarea undulosa): Further studies are required before the generic Bressan 1974, p. 105, fig. 30; Cabioch 19 72, p. 208, pI. Y, figs6- 11; Coppejans 1974, p. 399; Huve significance of this differencecan be assessed, 19 57, 1963, p. 150. especially since Littler (1971) recorded triangular cells in Titanoderma tessellata [as Tenarea]. Titanoderma Nageli 1858, p. 532 Since 1832, at least 32 specific and infraspecific Nomenclatural synonym: Dermatolithon Foslie 1898b, p. 11 taxa have been referred to Tenarea. Based on Type species: Titanoderma pustulatum (Lamouroux) studies of several type collections and on a survey Nageli 1858, p. 532. of the extant literature relating to these taxa, it Basionym: Melobesia pustulata Lamouroux 1816, would appear that Tenarea as circumscribed here p. 315 includes only one species, namely T.tortuosa. Nomenclatural synonyms: Most other taxa placed in Tenarea by recent au­ Dermatolithon pustulatum (Lamouroux) Foslie 1898b, p. 11 thors (e.g. Adey 1970) belong to Titanoderma Epilithon pustulatum (Lamouroux) Lemoine while most taxa referred to Tenarea by earlier 1921, p. 10* authors (e.g. Kuntze 1898; Lemoine 1911, 1929; Lithophyl/um pustulatum (Lamouroux) Foslie Airoldi 1936; Feldmann 1939) probably repre­ 1904, p. 8 Tenarea pustulata (Lamouroux) Adey 1965, p. sent taxa of Lithophyllum. The proper disposi­ 88 (comb. invalid) tion of these Lithophyllum-like taxa, however, Type locality: France (Lamouroux 1816, p. 315) can be determined only after examination of the Lectotype specimen: CN (Fig. 29) relevant type collections, a task beyond the scope Additional references: Hamel & Lemoine 1953, p. of the present study. 59, text fig. 22, pI. IX, figs 1, 2 (as Dermatolithon); At present, Tenarea tortuosa is known only Kommann & Sahling 1977, p. 206, Abb. 115 (as from the Aegean and parts of the north-eastern Dermatolithon). Mediterranean Sea (Huve 1957, p. 133). * Mme Lemoine (pers. comm.) has indicated that this combination was made by mistake ; elsew here on the same page the intended binomial Lithophyllum The genus Titanoderma pustulatum was used. CHA RACTERIZATION: Titanoderma, as delin­ eated here, encompasses those nongeniculate Corallinaceae which have uniporate (bi-) tetra­ sporangial conceptacles, secondary pit connec­ tions between cells of adjacent filaments, and a Woelkerling et al: Tenarea, Titanoderma and Dermatolithon 333

thallus consisting largely or entirely of a prostrate Table 2. Associated nomenclatural data for taxa trans­ crust whose tissues are organized in a dorsiven­ ferred to Titanoderma Nageli tral manner and include a unistratose hypo­ Taxa whose type specimens were examined by thallium composed of palisade cells. (a) Adey (1970, pp. 6, 7) This characterization is concordant with the Titanoderma ascripticium concept of Dermatolithon presented by Cabioch (Foslie) comb. nov. Basionym: Lithophyllum pustulatum f. ascrip­ (1972), Bressan (1974), and Johansen (1981), ex­ ticium Foslie 1907a, p. 34 cept that our circumscription emphasizes the Titanoderma bermudense (Foslie & Howe) comb. dorsi ventral organization of thallus tissues. No­ nov. menclatural and taxonomic data relating to Ti­ Basionym: Lithophyllum bermudense Foslie & tanoderma are summarized in Table 1. Howe 1906, p. 132 COMMENTS: Only one species (T. pustulatum) Titanoderma canescens (Foslie) comb. nov. Basionym: Melobesia canescens Foslie 1900b, has been referred to Titanoderma. Since 1898, p. 6 however, at least 67 specific and infraspecific taxa Titanoderma conspectum (Foslie) comb. nov. have been placed in Dermatolithon, a homotypic Basionym: Lithophyllum conspectum Foslie synonym of Titanoderma. The proper disposi­ 1907b, p. 29 tion of many of these taxa will have to await re­ Titanoderma dispar (Foslie) comb. nov. examination of the relevant type collections. Basionym: Lithophyllum tumidulum f. dispar Foslie 1907b, p. 29 During the course of studies on the Foslie her­ Titanoderma polycephalum (Foslie) comb. nov. barium, however, Adey (1970) examined a num­ Basionym: Lithophyllum polycephalum Foslie ber of type specimens of taxa which he referred 1905, p. 16 to Tenarea. Because Adey's concept of Tenarea Titanoderma polyclonum (Foslie) comb. nov. is equivalent to Titanoderma as delineated here, Basionym: Lithophyllum polyclonum Foslie . those species that he recognized can be trans­ 1905,p. 18 ferred into Titanoderma. Similarly, published Titanoderma prototypum (Foslie) comb. nov. Basionym: Lithothamnion prototypum Foslie accounts (protologues) of several other taxa pro­ 1897, p. 18 vide unequivocal evidence that they also belong Titanoderma rasile (Foslie) comb. nov. to Titanoderma. The necessary nomenclatural Basionym: Lithophyllum rasile Foslie 1907a, changes are summarized in Table 2. Further p. 34 studies of taxa referred here to Titanoderma are Titanoderma tumidulum (Foslie) comb. nov. needed to clarify taxonomic limits and relation­ Basionym: Lithophyllum tumidulum Foslie 1901, p. 5 ships at the species level; some of these taxa ul­ timately may be demonstrated to be conspecific. (b) Other taxa The genus Titanoderma, as represented by the Titanoderma corallinae (P. Crouan & H. Crouan) taxa listed in Table 2, appears to be widespread, comb. nov. but the geographic distributions of individual Basionym: Melobesia corallinae P. Crouan & H. Crouan 1867, p. ISO species are less certain and require checking of Titanoderma cystoseirae (Hauck) comb. nov. specimens upon which published records are Basionym: Melobesia cystoseirae Hauck 1883, based. p. 266 Titanoderma mutabile (Lemoine) comb. nov. Basionym: Lithophyllum mutabile Lemoine Relationships of Tenarea and Titanoderma 1930, p. 70 to other Corallinaceae Titanoderma stephensonii (Lemoine) comb. nov. Basionym: Dermatolithon stephensonii Le­ In considering the relationships of Tenarea and moine 1971, p. 558 Titanoderma to other Corallinaceae, the classi­ Titanoderma tessellatum (Lemoine) comb. nov. Basionym: Lithophyllum tessellatum Lemoine fication system of Johansen (1981, p. 10, tables 1930, p. 68 3, 4) has been selected to provide a fr amework for discussion. In this system, both Tenarea and Titanoderma are assignable to the Lithophylloi­ deae, which is characterized by the absence of geniculae, the presence of secondary pit connec­ Dermatolithon), Johansen includes fo ur other tions between cells and the occurrence of uni­ genera in the Lithophylloideae: Ezo, Gonioli­ porate tetrasporangial conceptacles. thon, Lithophyllum, and Metamastophora. Ten­ In addition to Tenarea and Titanoderma (syn. area is readily set apart fr om these genera in 334 Phycologia, Vol. 24 (3), 1985

possessing a thallus whose tissues are organized fusions (Denizot & Massieux 1965, p. 388, pI. 7, in an isobilateral manner. In the remaining gen­ fig. 5), and of the relationships of tissues as seen era, the thallus is organized in a dorsiventral in longitudinal and transverse views (compare manner or in a dorsiventraVradial manner (the Denizot & Massieux 1965, text fig. 2B and Pia latter occurring in excrescences). Titanoderma is 1934, fig. 8 with our Figs II, 22). Cell fusions distinctive in possessing a prostrate thallus with are unknown in Tenarea, and the tissues of Ten­ a unistratose hypothallium composed of pali­ area in transverse view show a more complex sade-like cells interconnected by secondary pits. organization than is thought to occur in Disti­ This structure contrasts with that of Ezo (a para­ choplax. Confirmation of the occurrence of con­ sitic genus; Adey et al 19 74) and Lithophyllum ceptacles in Distichoplax also is needed. (Woelkerling 1983) which lack a palisade hy­ pothallium, and with Metamastophora (Woelk­ erling 1980a, 1980b) in which the thallus is an erect, branched, taeniform structure with both ACKNOWLEDGMENTS secondary pit connections and cell fusions. Fur­ ther study is needed to clarify the relationships We are especially grateful to the late Mme Marie between Titanoderma Nageli (1858) and Go­ Lemoine who, through correspondence with niolithon Foslie (1898a). Huve (1962) provided Y.M.C., took great trouble to answer a number an extensive account of Goniolithon papillosum of queries concerning the research reported on (Zanardini ex Hauck) Foslie (as Lithophyllum), here. Sincere thanks also are due Dr E. Coppejans the type species of Goniolithon, and further stud­ (Laboratorium voor Morfologie Systematiek en ies were undertaken by Cabioch (1970, 1972). Ecologie van de Planten, Rijksuniversiteit-Gent, Based on these studies, Cabioch characterized Belgium) for supplying specimens, Dr Fran90ise Goniolithon by the occurrence of numerous lay­ Ardre (Laboratoire de Cryptogamie, Museum ers of palisade-like cells which in excrescences National d'Histoire Naturelle, Paris) for photo­ are aligned in coaxial tiers. The young thallus is graphs of Bory's specimen of Tenarea undulosa considered (Cabioch 1972, pp. 210, 270) to have and for supplying other specimens, Professor P. a Titanoderma-like structure, including a uni­ Gayral for supplying photographic negatives of stratose palisade-like hypothallium. Whether the type specimen of Melobesia pustulata, Pro­ Goniolithon should be treated as a distinct genus fessor Peter Dixon for calling our attention to remains uncertain. the paper by Denizot & Massieux (1 965), P.I. Finally, the relationships between Tenarea and and B.M. Mullock for collecting specimens of the fossil genus Distichoplax require comment. Tenarea tortuosa (LTB 14 646) in Crete espe­ Pia (1934, p. 18) established the genus for D. cially for this study, the curators at CN, FR, L, biserialis (Dietrich) Pia [Basionym: Lithotham­ and PC who permitted examination of relevant nium biserialis Dietrich 1927, p. 46 1, pI. 2, fig. type specimens, Dr M. Grasshofffor information I] and characterized it by the occurrence of cal­ about Esper's collections of Millepora and Dr cified thalli which formed free-standing plates I.A. Staff(Department of Botany, La Trobe Uni­ composed of two rows of cells. Keij (1963, 1964) versity) for discussions relating to anatomical recorded possible conceptacles in fossil speci­ terminology. mens from Borneo, and Denizot & Massieux (1965) concluded, after studying additional ma­ terial, that Distichoplax was most closely related REFERENCES to Tenarea. Although no material has been ex­ amined during the present study, it is obvious AOEY W.H. 1965. The algal tribe Lithophylleae and from the accounts and illustrations of the above its included genera, pp. 67-102. In: Johnson J.H. and Adey W.H., Studies ofLithophyllum and related authors that Distichoplax, like Tenarea, is or­ algal genera. Colo. Sch. Mines Q. 60(2): I-lOS. ganized in an isobilateral manner. Formal judge­ AOEY W.H. 19 70. A revision of the Foslie crustose ment as to whether these two taxa are related or coralline herbarium. K.norske Vidensk. Selsk. Skr. even possibly congeneric, however, must await 1970(1): 1-46. AOEY W.H. & AOEY P.J. 19 73. Studies on the bio­ further study of Distichoplax, especially of the systematics and ecology of the epilithic crustose Cor­ reported occurrence of trichocytes (Denizot & allinaceae of the British Isles. Br. phycol. J. 8: 343- Massieux 1965), of the possible occurrence of cell 407. Woelkerling et al: Tenarea, Titanodermaand Dermatolithon 335

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Index Herbarioru m, Pt.1. The Herbaria of the World. Subfam. 1. Melobesieae. Pages 19-68 in Boergesen, Ed. 7. Bohn, Scheltema, & Holkema, Utrecht. [Reg­ F., Marine algae from the Canary Islands. III. Rho­ num Vegetabile Vol. 106.] dophyceae. Part II. Cryptonemiales, Gigartinales, and HUVE H. 19 57. Sur l'individua1ite gent!rique du Ten­ Rhodymeniales. Bioi. Medd. 8( 1): 1-97. area undulosa Bory 1832 et du Tenarea tortuosa LEMOINE MME P. 1930. Les Corallinacees de I'archi­ (Esper) Lemoine 191 1. Bull. Soc. bot. Fr. 104: 13 2- pel des Galapagos et du Golfe de Panama. Archs 140. Mus. Hist. Na t., Paris. Ser. 64: 37-88, pI. 1-4 (Note: HuvE H. 1962. Taxonomie, eco10gie et distribution Journal is dated 1929, but was published in 1930.) d'une Melobesiee mediterraneenne: Lithophyllum LEMOINE M. 1971. Apparition de la structure mon­ papillosum (Zanardini) comb. nov., non Lithophyl­ ostromatique dans un thalle epais de Dermatolithon lum (Dermatolithon) papillosum (Zanard.) Foslie. Bot. 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Note: SUNESON S. 1950. The cytology of the bispore fo r­ Also issued in thesis form with identical pagination mation in two species of Lithophyllum and the sig­ but a different title page. A summary appears in Bull. nificance of the bispores in the Corallinaceae. Bot. Soc. bot. Fr. 58: 394-397 (P. Hariot, 1911). Notiser 1950: 429-450. LEMOINE MME P. 1921. Note sur les algues calcaires SVEDELIUS N. 1911. Rhodophyceae (Bangia1es und recueillies par MM. A. et L. Joleaud et catalogue des ). In: Die Naturlichen Pj lanzenfamilien (Ed. Melobesiees des cotes fran�aises de la Mediterranee. by A. Engler) Vol. 2, Nachtrage, Lief. 243-244, pp. Bull. Soc. linn. Provence 3: 5-15. 193-284. W. Engelmann, Leipzig. LEMOINE M ME P. 1929. Fam . 6-Corallinaceae. TOWNSEND R.A. 1981. Tetrasporangial conceptacle Woelkerling et al: Tenarea, Titanoderma and Dermatolithon 337

development as a taxonomic character in the Mas­ lata (Sonder) Setchell: morphology and anatomy. Br. tophoroideae and Lithophylloideae (Rhodophyta). phycol. J. IS: 201-225. Phycologia 20: 407-4 14. WOELKERLING WM. J. 19 80b. Studies on Metamas­ TURNER J.A. & WOELKERLING WM. J. 19 82. Studies tophora (Corallinaceae, Rhodophyta). II. Systemat­ on the Mastophora-Lithoporella complex (Coralli­ ics and distribution. Br. phycol. J. IS: 227-245. naceae, Rhodophyta). I. Meristems and thallus struc­ WOELKERLING WM. J. 1983. A taxonomic reassess­ ture and development. Phycologia 21: 20 1-217. ment of Lithophyllum Philippi (Corallinaceae, Rho­ Voss E.G. (CHM.) 1983. International Code of Bo­ dophyta) based on studies of R.A. Philippi's original tanical No menclature adopted by the Thirteenth In­ collections. Br. phycol. J. 18: 299-328. ternational Botanical Congress, Sydney, August 1981. Bohn, Scheltema & Holkema, Utrecht. xv + 472 pp. WOELKERLING WM. J. 19 80a. Studies on Metamas­ Accepted 18 January 19 85 tophora (Corallinaceae, Rhodophyta). I. M. f/abel-