A Synoptic Key to the Rhodophyta of the Western Mediterranean Part. I Gig

A SYNOPTIC KEY TO THE RHODOPHYTA OF THE WESTERN MEDITERRANEAN PART. I GIG ARTIN ALES, H AL YMENIALES, HILDENBRANDIALES AND PLOCAMIALES C Rodriguez-Prieto, M Sabater, A Vergés

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C Rodriguez-Prieto, M Sabater, A Vergés. A SYNOPTIC KEY TO THE RHODOPHYTA OF THE WESTERN MEDITERRANEAN PART. I GIG ARTIN ALES, H AL YMENIALES, HILDENBRAN- DIALES AND PLOCAMIALES. Vie et Milieu / Life & Environment, Observatoire Océanologique - Laboratoire Arago, 2001, pp.55-65. hal-03192091

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HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. VIE ET MILIEU, 2001, 51 (1-2) : 55-65

A SYNOPTIC KEY TO THE RHODOPHYTA OF THE WESTERN MEDITERRANEAN PART. I GIG ARTIN ALES, H AL YMENIALES, HILDENBRANDIALES AND PLOCAMIALES

C. RODRIGUEZ-PRIETO, M.A. SABATER, A. VERGÉS Universitat de Girona, Facultat de Ciències, Campus de Montilivi, s/n. 17071 Girona, Spain E-mail:cacrp @fc. udg. es

RHODOPHYTES RÉSUMÉ. - Une clé dichotomique pour la détermination des Gigartinales, Halymé- GIG ARTIN ALES niales, Hildenbrandiales et Plocamiales de Méditerranée occidentale est présentée. HALYMENIALES Elle a pour objectif de faciliter la reconnaissance de ces algues rouges. Ce travail HILDENBRANDIALES PLOCAMIALES inclut l'ordre des Gigartinales, avec 69 espèces groupées en 19 familles et 35 gen- CLÉ TAXONOMIQUE res, l'ordre des Halyméniales, avec 13 espèces appartenant à 2 familles et 6 genres, STRUCTURES DE REPRODUCTION l'ordre des Hildenbrandiales, avec 1 famille, 1 genre et 3 espèces, et l'ordre des MÉDITERRANÉE DE L'OUEST Plocamiales, avec 1 famille, 1 genre et 2 espèces. Le genre Wurdermannia Harvey est inclus dans la clé, bien que son appartenance aux Gigartinales reste incertaine. Quelques descriptions morphologiques et anatomiques des structures de reproduction sont aussi inclues, d'après des données de la littérature et de nos propres observations.

RHODOPHYTA ABSTRACT. - A synoptic key for identifying the Gigartinales, Halyméniales, Hil- GIGARTINALES denbrandiales and Plocamiales from the Western Mediterranean has been prepared HALYMENIALES in order to facilitate the récognition of thèse red algae. The orders Gigartinales, HILDENBRANDIALES PLOCAMIALES with some 69 species grouped in 19 families and 35 gênera, Halyméniales, with 13 TAXONOMIC KEY species grouped in 2 families and 6 gênera, Hildenbrandiales, with 1 family and 1 REPRODUCTIVE STRUCTURES gênera of 3 species, and Plocamiales, with 1 family and 1 genus of 2 species are in- WESTERN MEDITERRANEAN cluded. The genus Wurdermannia Harvey is also included in the key, although it has not finally been established whether it belongs to the order Gigartinales or not. Morphological and anatomical descriptions of the reproductive structures, using data from the literature and from personal observations are presented.

INTRODUCTION than a single distinction. Végétative characteristics are used to separate gênera and species where ap- propriate, but as définitive identification can only This is intended to be the first publication in a rarely be obtained from gênerai appearance and séries of keys to the différent orders of benthic ma- form of the Rhodophyta, or from stérile material, rine algal flora from the western Mediterranean reproductive stages have been incorporated in un- (excluding the recently introduced species). The avoidable cases. Many of the families have a présent paper includes the order Gigartinales with highly distinctive female reproductive System from 69 species grouped in 19 families and 35 gênera, which they can be recognized, although such char- the order Halyméniales with 13 species grouped in acters may often be difficult to observe. The main two families and six gênera, the order reproductive characteristics are listed for each ge- Hildenbrandiales with one family and one genus of nus in Table I. The use of dimensions such as the three species, and the order Plocamiales, with one length and width of thalli is only a guide in the key, family and one genus of two species. The genus since such characteristics tend to be very variable Wurdermannia Harvey is also included in the key and unstable. although it has not yet been finally established Following the key is a listing of the gênera and whether it belongs to the order Gigartinales or not. species currently recognized in the studied area. The key should serve as a tool to identify gênera We have included the order Cryptonemiales and species of thèse groups. An effort has been (Kylin, 1956) in the order Gigartinales, following made to base dichotomies in the key upon more Kraft & Robins (1985), because we consider that structure Uniaxial/Multiaxial LIFE HISTORY Triphasic/Biphasic Isonnorphic/Heteromorphic FEMALE GAMETANGIA Monocarpogonial/Polycarpogonial number of cells of the carpogonial branch carpogonial branch Simple/Branched Procarpic/Non-procarpic supporting cell acting as auxiliary cells (Yes/No) auxiliary branch borne on supporting cell (Yes) auxiliary cell a nearby cortical or medullar cell (Yes) auxiliary cells in spécial filaments (Yes) carpogonial and auxiliary cells in ampullae subsidiary cells présent (Yes' carpogonial branch with nutritive auxiliary cells (Yes) auxiliary cell with adjacent nutritive cells (Yes) MALE GAMETANGIA Scattered/in Clusters/in SOri/in CHains in spécial branchlets or bladelets (Yes/No) GONIMOBLAST developing Inwards/Outwards on Auxiliary cell/on Connecting filament with involucre Cellular/Filamentous/Absent with ostiole Présent/Absent in spécial branchlets or bladelets (Yes/No) TETRASPORANGIA Cruciately/Tetrahedrally/Zonately/Irregularly arranged Scattered/in SOri/in Conceptacles in spécial branchlets or bladelets (Yes/No)

Table 1 Type of structure and main reproductive characteristics of the gênera treated (sv = several); (* = usually); (? = unknown); (grey colour = unattainable characteristics). A KEY TO THE RHODOPHYTA OF WESTERN MEDITERRANEAN, PART I 57 the fact that the Cryptonemiales have auxiliary 4. Thallus brownish red. Conceptacle chamber sphe- cells in accessory Systems within the thallus, as op- rical, with a small ostiole. Tetrasporangia with pa- posed to Gigartinales supposedly transforming rallel, transverse or oblique cleavage, 5-10 x 20- végétative cells into auxiliary cells, is insufficient 30 um H. crouanii 4. Thallus purplish red. Conceptacle chamber ovoid, basis for placing the Cryptonemiales in a separate with a large ostiole. Tetrasporangia always with order. The Hildenbrandiales (Pueschel & Cole, transverse cleavage, 10-15 x 25-40 um 1982), Plocamiales (Saunders & Kraft, 1994) and H. occidentalis Halyméniales (Saunders & Kraft, 1996) are consid- 5. Hypothallus with some filaments larger than the érée! as separate orders, following récent molecular others, appearing in surface view as a more or less studies, but since their morphology and reproduc- conspicuous percurrent midrib and as a distinct cen- tive structures are similar to the Gigartinales, the tral axis in cross section. Each hypothallial cell gi- families in thèse orders are not separated in the ving rise to more than one upper and one lower coxal key. Other taxa have been separated as orders and cells. The filaments of the perithallus and subhypo- are not covered in this first part. Thèse taxa in- thallus are erect. Thallus not calcified. Gland cells cluded the Ahnfeltiales (Maggs & Pueschel, 1989), are présent, situated in the upper surface of the blade représentatives of which are not known in Western and apparent in surface view as white microscopic Mediterranean, the Corallinales (Silva & Johansen, rounded spots Family RHIZOPHYLLIDACEAE 1986), which is a clearly distinguishable order, and Genus Contarinia 6 the Gracilariales (Fredericq & Hommersand, 1989) 5. Hypothallus either sparingly branched and more or which requires more investigation in the Mediterra- less parallel (radially arranged), or much branched to nean. The taxonomic arrangement below the level form a polyflabellate layer (polyflabellately arran- of families follows Silva et al. (1996). ged), but not appearing as a distinct central axis in cross section. Each hypothallial cell giving rise to no more than one upper and one lower coxal cells. The CURRENT KEY filaments of the perithallus and subhypothallus are usually inclined towards the margin. Thallus calcified 1. Thallus prostrate, with a dorsiventral symmetry. or not. Gland cells are absent 7 Plants calcified or not. Structure consisting of a 6. Genus Contarinia single basai layer of branched filaments (hypothal- 6. Plants are flattened, forming reniform and encrus- lus), with each cell dividing into one or more upper ting fronds, irregularly branched, with sinuous cells (upper coxal cells) and, occasionally, one or and lobed margins. In longitudinal section, hypo- more lower cells (lower coxal cells). The upper coxal thallial cells up to 35 (-40) um long and 20 um cells give rise to filaments of apical growth directed broad. Erect filaments up to 10-25 um wide at ba- to the upper surface (perithallus), erect or slightly in- sai parts. Gland cells ovoid or pyriform in cross clined towards the margin. In the latter case, they can section. Tetrasporangia cruciately arranged become erect later (secondary perithallus) due to per- C. peyssonneliaeformis sistent growth. The species with lower coxal cells can 6. Plants range from compressed to flattened, for- also develop some filaments directed to the undersur- ming band-shaped segments almost regular in face (subhypothallus) 2 width (1-2 mm), irregularly dichotomously-disti- 1. Thallus erect (at least the gametophyte), without a chously branched, with dentate margins. In longi- dorsiventral symmetry. Plants not calcified. Structure tudinal section, hypothallial cells up to 130 (-150) différent from above 28 um long and 60 um broad. Erect filaments up to 2. Plants crustose, not calcified. Cells very small, usual- 30-45 um at basai parts. Gland cells spherical. Te- ly =S 5 um in diameter. Hypothallus with each cell di- trasporangia irregularly zonate C. squamariae viding into an erect unbranched or sparsely branched 7. Each hypothallial cell cutting off a single upper coxal filament. Secondary perithallus and subhypothallus cell. Subhypothallus absent. Secondary perithallus absent. Rhizoids absent. Tetrasporangia developing absent. Plants fleshy, gelatinous. Carpogonial branch from spherical or ovate conceptacles 2-3 celled. Tetrasporangia zonately arranged Family HILDENBRANDIACEAE Family CRUORIACEAE Genus Hildenbrandia 3 Genus Cruoria 2. Plants crustose or not, calcified or not. Cells not very C. cruoriaeformis small, > 5 um in diameter. Hypothallus with each cell 7. Each hypothallial cell cutting off either one single dividing into one or more coxal, upper cells and, oc- upper coxal cell, or one upper and one lower coxal casionally, one or more lower coxal cells. Perithallus cell, or one indiscriminately upper or lower coxal erect or slightly inclined towards the margin. Secon- cell. Subhypothallus présent or absent. Secondary pe- dary perithallus and subhypothallus sometimes pré- rithallus présent or absent. Plants gelatinous or firm sent. Rhizoids présent or absent. Tetrasporangia not and coriaceous. Carpogonial branch 3-6 celled. Te- developing from conceptacles, scattered or in sori 5 trasporangia (when présent) cruciately or irregularly 3. Genus Hildenbrandia cruciately arranged 3. Thallus rosy to dark red. Tetrasporangia ovoid, Family PEYSSONNELIACEAE.... 8 appearing cruciate or irregular, with non-parallel 8. Thallus lacking rhizoids. Plants encrusting, entirely cleavage H. rubra calcified. Each hypothallial cell cutting off one upper 3. Thallus brownish or purplish red. Tetrasporangia and one lower coxal cell. Secondary perithallus pré- elongated, appearing zonate 4 sent Genus Polystrata .... 9 58 RODPJGUEZ-PRIETO C, SABATER MA, VERGÉS A

8. Thallus adhering to the substratum by rhizoids which the thallus delimits large cavities filled up with arise along the whole of their undersurface. Plants fo- fine sédiments liose or encrusting, calcified or not. Each hypothallial P. rosa-marina f. rosa-marina cell dividing into either one upper and one lower 15. Plants growing on hard bottom, with a planar or coxal cell, or only one single upper coxal cell, or one slightly undulate frond...P. rosa-marina f. saxicola indiscriminately upper or lower coxal cell. Secondary perithallus présent or absent 10 16. Rhizoids one cell long. Cystoliths absent. Hypothal- lial cells radially or polyflabellately arranged... 17 9. Genus Polystrata 9. Subhypothallus not well developed, sometimes 16. Rhizoids more than one cell long. Cystoliths présent absent near the margins, up to 2 cells long (com- or absent. Hypothallial cells radially arranged.. 27 prising the coxal cell). Perithallus cohesive after 17. Hypothallus polyflabellate 18 décalcification P. compacta 17. Hypothallus not distinctly polyflabellate 20 9. Subhypothallus well developed, but a little shorter 18. Perithallial filaments relatively free. Plants loose than perithallus. Perithallus loosely cohesive after and gelatinous. Hypobasal calcification not well de- décalcification P. fosliei veloped P. armorica 10. In the centre of the thallus, the hypothallus gives rise only to a simple perithallus, but, near the mar- 18. Perithallial filaments laterally cohesive. Plants nei- gin, it gives rise also to a subhypothallus, due to the ther loose nor gelatinous. Hypobasal calcification fact that the single coxal cell is situated indiscrimi- well developed 19 nately at the upper or the lower part of the hypothal- 19. Secondary perithallus présent. Thallus 300-500 lial cell. Thallus entirely calcified. Hypothallus (- 800) um thick. Tetrasporangia laterally inserted polyflabellately arranged. Secondary perithallus on the basai cell of the stérile filaments of the sori.... présent. Rhizoids one cell in length P. codana Genus Metapeyssonnelia 19. Secondary perithallus absent. Thallus 70-125 um M. feldmannii thick. Tetrasporangia inserted at the apex of the ba- 10. The hypothallus générâtes only a single upper coxal sa/ cell of the stérile filaments of the sori P. dubyi cell. Subhypothallus usually absent, but, in some 20. Angle of perithallus/hypothallus =s 50° 21 species, a lower coxal cell per hypothallial cell and a few developed subhypothallus can be also présent. 20. Angle of perithallus/hypothallus > 50° 23 Thallus calcified or not. Hypothallus radially or po- 21. Thallus « 190 um thick. Hypothallial cells « 30 um lyflabellately arranged. Secondary perithallus pré- long. Secondary perithallus absent P. crispata sent or absent. Rhizoids one or more cells in length. 21. Thallus up to 300 (-500) um thick. Hypothallial Genus Peyssonnelia 11 cells usually > 30 um long. Secondary perithallus 11. Genus Peyssonnelia présent 22 11. Thallus not calcified. Subhypothallus présent. 22. Hypothallial cells 60-175 um long. Rhizoids long Hypothallus radially arranged. Rhizoids multi- P. atropurpurea cellular L2 22. Hypothallial cells 25-80 um long. Rhizoids short 11. Thallus calcified. Subhypothallus absent. Hypo- and globose P. magna thallus radially or polyflabellately arranged. Rhi- 23. Thallus ^ 425 um thick P. harveyana zoids one or more cells long 13 23. Thallus s; 200 um thick 24 12. Only one subhypothallial cell présent (lower 24. Thallus s 140 um thick. Secondary perithallus pré- coxal cell). Rhizoids branched. Plants between sent 25 95 and 295 um thick. Thallus membranous to 24. Thallus < 140 um thick. Secondary perithallus ab- cartilaginous in texture P. squamaria sent or not well developed 26 12. Two subhypothallial cells présent (including the 25. Tetrasporangia inserted at the apex of the basai cell lower coxal cell). Rhizoids unbranched. Plants of the stérile filaments of the sori P. rara-avis between 155 and 385 um thick. Thallus firm, car- 25. Tetrasporangia laterally inserted on the basai cell of tilaginous to coriaceous in texture...P. coriacea the stérile filaments of the sori P. hongii 13. Thallus entirely calcified. Secondary perithallus 26. Hypobasal calcification well developed, to 70-110 présent 14 um thick. Secondary perithallus absent. Surface 13. Thallus with hypobasal calcification and with or plane. Thallus firm, coriaceous, but fragile and bro- without cystoliths. Secondary perithallus présent ken easily P. stoechas or absent 16 26. Hypobasal calcification not well developed, to 45- 14. Rhizoids unicellular, unbranched. Plants with 70 um thick. Secondary perithallus sometimes pré- concentric growth zones from the marginal thal- sent but not well developed. Surface undulated. lus meristem. Thallus =S 565 um thick. Hypothal- Thallus membranous to coriaceous, not fragile lial cells =S 90 um long P. rosa-marina... 15 P. inamoena 14. Rhizoids more than one cell long, usually bran- 27. Cystoliths présent. Thallus « 145 um thick, mem- ched. Plants without concentric growth zones branous to cartilaginous in texture, whitish seen from the marginal thallus meristem. Thallus «S from below. Secondary perithallus absent. Hypoba- 385 um thick. Hypothallial cells =s 65 um long .... sal calcification up to 70 um thick P. P. polymorpha rubra 15. Plants free-living, forming more or less globu- 27. Cystoliths absent. Thallus « 300 um thick, firm, co- lous rhodoliths due to the occasional overturning riaceous in texture, not whitish seen from below. Se- of the plant on itself and the conséquent change condary perithallus présent. Hypobasal calcification of its growing direction. The loose wrapping of up to 240 um thick P. bornetii A KEY TO THE RHODOPHYTA OF WESTERN MEDITERRANEAN, PART I 59

28. Structure uniaxial. Growth by a single apical cell or 34. Carpogonial branches > 3 cells long. Auxiliary cells by a small group of them 29 in spécial filaments, easily distinguishable before 28. Structure multiaxial. Growth by a marginal meris- fertilization. Whorls of 4 branched filaments. Outer tem of many apical cells 50 cells moniliform or radially elongated 37 29. Growth maintained by transverse division of a 35. Gonimoblast arising from the auxiliary cell. Outer single apical cell. Thallus erect and bushy (at least cells always lacking terminal hyaline hairs the gametophyte). Tetrasporangia cruciately or zo- Genus Calosiphonia 36 nately arranged 30 35. Gonimoblast arising from the Connecting filament, 29. Growth by oblique division of a single apical cell or near the auxiliary cell. Outer cells sometimes provi- by a small group of them. Thallus erect, decumbent ded with hyaline hairs Genus Schmitzia or prostrate. Tetrasporangia zonately arranged ... 41 S. neapolitana 30. Thallus is terete to flat. Plants rose pink in colour, 36. Genus Calosiphonia mucilaginous and soft in texture. Structure consis- 36. Main axes usually simple, occasionally with ting of a very apparent axial filament of elongated some spine-like alternate or unilatéral branches. cells producing whorls of 3-4 repeatedly branched Plants up to 2 (-4) cm high and 1-2 mm broad filaments composed of cells diminishing in size to- C. dalmatica ward the periphery, where they are embedded in mu- 36. Main axes many times branched alternately. cilage to form a loose cortex. In younger parts the Plants up to 1-8 (-12) cm high and 2 mm broad.... whorls are distinct in surface view as annulations, C. vermicularis but, in adult parts, the axial filament is usually sur- 37. Outer cortical cells moniliform, occasionally provi- rounded by a compact layer of branched rhizoidal fi- ded with terminal hyaline hairs. Carpogonial bran- laments produced by the periaxial cells and the ches initially unbranched, latter producing opposite bands are not distinct. Procarpic or non-procarpic. laterals to one or two orders, except for the carpogo- Carpogonial branch 3 or more cells long. Carpospo- nium and its two subtending cells. Auxiliary cell rophytes lacking involucre and ostiole. Spermatan- branch 5-16 cells long, with auxiliary cell terminal... gia grouped on terminais or on subterminal cells of Family ACROSYMPHYTACEAE whorled branches. Tetrasporangia cruciately or zo- Genus Acrosymphyton nately arranged 31 A. purpuriferum 30. Thallus flattened. Plants dark red in colour, membra- 37. Outer cortical cells radially elongated, not provided nous to cartilaginous in texture. Structure différent with hyaline hairs. Carpogonial branches usually from above. Procarpic. Carpogonial branch 3-celled. simple. Auxiliary cell branch 10-20-cells long, with Carposporophytes with a cellular pericarp, non os- auxiliary cell larger than the others, very distinct, tiolate. Spermatangia in depressed sori. Tetrasporan- intercalary, with the cells situated under and over it gia zonately arranged 38 acting as nutritive cells 31. Plants are terete or flat.Whorls of 4 branched fila- Family DUMONTIACEAE ments. Procarpic. Auxiliary cell situated on a branch Genus Dudresnaya arising from the supporting cell. Outer cells monili- D. verticillata form Family GLOIOSIPHONACEAE 32 38. Plants dichotomously or laterally branched. Axial 31. Plants are terete or slightly compressed.Whorls of 3-4 filament distinct throughout, surrounded by rhi- branched filaments. Non-procarpic. Auxiliary cell not zoids, and the whole immersed in some layers of situated on a branch arising from the supporting cell. large rounded or elongated thick-walled and com- Outer cells moniliform or radially elongated 34 pacter! cells. Cortex consisting of an outer part of several layers of moniliform cells arranged in radial 32. Plants flattened. Axes branching somewhat pinnate. rows, and a subcortex of slightly larger ovoid cells... Carpogonial branch 4-celled Family SPHAEROCOCCACEAE Genus Schimmelmania Genus Sphaerococcus 3.9 S. ornata 38. Plants repeatedly pinnate branched, with alternating 32. Plants terete. Axes branching alternately or irregu- groups of 2-5 branchlets. Axial filament sometimes larly. Carpogonial branch 3-celled 33 distinct throughout, immersed in some layers of 33. Outer cells not provided with terminal hair cells. large rounded or elongated thick-walled and com- Monocarpogonial. Auxiliary cell branch 2-celled, pacted cells. Cortex of cells not arranged in radial unbranched, with auxiliary cell intercalated rows Family PLOCAMIACEAE Genus Thuretella Genus Plocamium 4.0 T. schousbœi 39. Genus Sphaerococcus 33. Outer cells occasionally provided with terminal hair 39. Plants up to 4-5 cm broad, with a non conspi- cells. Mono- or polycarpogonial. Auxiliary branch cuous percurrent midrib. Larger medullary cells usually 5-celled, occasionally branched from subten- up to 60-70 um in diameter. Outer cortical cells ding cells, with auxiliary cell terminal 10-15 um in diameter in surface view Genus Gloiosiphonia S. rhizophylloides G. capillaris 39. Plants up to 10-15 (-25) cm, lacking a midrib. 34. Carpogonial branch 3-celled. Auxiliary cells nume- Larger medullary cells up to 100-130 um in dia- rous in cortex, indistinguishable before fertilization. meter. Outer cortical cells 5-10 um in diameter in Whorls of 3-4 branched filaments. Outer cells moni- surface view S. coronopifolius liform Family CALOSIPHONACEAE 35 40. Genus Plocamium 60 RODRIGUEZ-PRIETO C, SABATER MA, VERGÉS A

40. Main axis « 2000 um broad. Ramuli and secon- 45. Medullary cells lacking secondary pit-connections. dary branchlets and arrangement of ramuli along Non-procarpic. Auxiliary cell is a nearby cortical axes not always unilatéral P. cartilagineum cell. Gonimoblast not developing adjacent nutritive- 40. Main axis « 250 um broad. Ramuli, secondary cells. Carposporophytes without pericarp, usually branchlets and arrangement of ramuli along axes ostiolate. Spermatangia in clusters or rosettes on unilatéral P. secundatum thallus surface. Tetrasporangia scattered or in sori 41. Thallus flattened, usually complanately branched on thallus surface from the margins, lacking spine-like branchlets. Family CAULACANTHACEAE 48 Procarpic. Nutritive cells présent 46. Genus Hypnea Family CYSTOCLONIACEAE 42 46. Thallus ^ 3 cm high. Main axes =s 800 um in 41. Thallus terete or only slightly compressed, not com- diameter H, furnariana planately branched, sometimes bearing numerous 46. Thallus > 3 cm high. Main axes s 1000 um in spine-like branchlets. Procarpic or non-procarpic. diameter 47 Nutritive cells présent or absent 45 47. Plants without definite main axes, 2-6 (-15) cm 42. Thallus erect, growing from a branched holdfast in high and to (110-) 250-660 (-1000) um in dia- the Mediterranean species. Plants cartilaginous in meter. Main branches usually straight. In cross texture. Fronds not anastomosing. Growth by a section, axial cells 20-40 (-65) um in diameter, single apical cell. Thallus transversed by a less- surrounded by cells ^160 um wide. In longitu- conspicuous axial filament, surrounded by several dinal section, medullary cells =£ 250 um long. lacunose layers of longitudinally elongated cells.... H. cervicornis Genus Calliblepharis 43 47. Plants with main axis usually remaining conspi- 42. Thallus erect or somewhat prostrate, growing from a cuous, 10-16 (-20) cm high and 1000 (-2000) discoid holdfast. Plants thin and délicate, membra- um in diameter. Main branches usually swollen nous when young and more firm in older spécimens. and crozier-hooked. In cross section, axial cells Fronds occasionally anastomosed or fixed to other 25-30 um thick, surrounded by cells =£ 90 um species by rhizoids produced secondarily from the wide. In longitudinal section, medullary cells =£ frond surface. Growth by a small group of apical 320 um long H. musciformis cells. Thallus appearing transverse by a diffuse axial filament in longitudinal section, and scarcely and ir- 48. Branches segmented with elongated-ovoid interno- regularly celled in transverse section des and constricted nodes. Segments terete to com- Genus Rhodophyllis 44 pressed. Cortex 2-5 cells thick. Growth by a single 43. Genus Calliblepharis apical cell or by a small group of apical cells 43. Thallus lanceolate, > 10 mm wide. Blade is Genus Catenella simple or irregularly branched. Marginal proli- C. caespitosa férations usually =s 5 mm long, simple. Thallus 48. Branches not segmented, axes of constant diameter =£ 650 um thick. Axial filament surrounded by but pointed apices. Plant terete. Cortex one cell cells up to 50-200 um longitudinally elongated thick. Growth by a single apical cell 49 up to 2 times their diameter. Cortex of 2-3 layers of pigmented cells C. ciliata 49. Medulla lacunose. Two periaxial cells per axial cell. 43. Thallus linear-lanceolate, usually 2-6 mm Tetrasporangia scattered Genus Caulacanthus broad. Blade dichotomous or irregularly bran- C. ustulatus ched. Marginal proliférations up to 15-30 mm 49. Medulla not lacunose. Only one periaxial cell per long, sometimes hooked, especially those ari- axial cell. Tetrasporangia grouped in sori sing from the apex. Thallus up to 350 (-400) um Genus Feldmannophycus thick. Axial filament surrounded by cells from F. rayssiae 50 um to over 100 um in diameter, longitudinally elongated up to 5 times their diameter. Cor- 50. Plants with a large, much-lobed supporting cell, that tex of a single layer of pigmented cells gives rise to several stérile (subsidiary) cells, simi- C. jubata lar in shape to the supporting cell. Carpogonial branch 2-celled, arising from the subsidiary cells. 44. Genus Rhodophyllis Plants flattened, foliose. Cortex of cells decreasing 44. Thallus up to 1-5 (-7) cm high and 1 cm wide, in size from the medulla outwards, not arranged in erect or somewhat prostrate R. divaricata radial rows Family KALLYMENIACEAE... 51 44. Thallus < 1 cm long and 2 mm wide, prostrate.... R. strafforellii 50. Plants without subsidiary cells. Carpogonial branch s 2-celled, arising from the supporting cell. Plants 45. Medullary cells with numerous secondary pit- flattened, compressed or terete. Cortex of cells so- connections. Procarpic. Auxiliary cell branch origi- metimes arranged in radial rows 58 nating from the supporting cell and adjacent outwar- dly to the carpogonial branch, with intercalary 51. Medulla compact, composed of large, rounded, auxiliary cell. Gonimoblast developing adjacent nu- thick-walled cells through which a délicate network tritive cells. Carposporophytes enclosed in a hemis- of cobwebby filaments permeates, denser at the ba- pherical poreless pericarp. Spermatangia formed in sai parts. Inner cortical cells ovoid to angular; outer- chains in outer cortical cells of short latéral bran- most rounded. Procarpic. Supporting cell acting as chlets. Tetrasporangia grouped in the swollen parts auxiliary cell. Carposporophytes surrounded by sté- of spécial short latéral branchlets rile strands from the gametophytes, usually ostiolate Family HYPNEACEAE Genus Callophyllis Genus Hypnea .46 C. laciniata A KEY TO THE RHODOPHYTA OF WESTERN MEDITERRANEAN, PART I 61

51. Medulla lax, filamentous, interwoven with very ob- stellate or ganglionic in shape, with radiating vious stellate or ganglionic cells. Inner cortical cells arms =£ 320 um in length. Polycarpogonial stellate, the following large and angular or slightly K. reniformis stellate, and the outermost small, ovoid or irregular- 57. Outer cortical cells usually polyhedral in surface ly shaped. Non-procarpic. Auxiliary cell Systems view, occasionally irregularly shaped. Medullar with several subspherical to ovoid subsidiary cells cells ganglionic, with radiating arms « 1000 um on an ovoid auxiliary cell. Carposporophytes lac- in length. Monocarpogonial K. requienii king filamentous involucre and ostiole 52 58. Medulla compact, of hyaline cells 59 52. Gametophytes erect, with a simple or branched stipe from which arise a séries of more or less alternately 58. Medulla lax, filamentous, occasionally with stellate arranged blades. Plants strongly concave and auricu- or X-shaped cells (small ganglionic cells with body late at the base, the older blades enveloping the cell about twice the medullary filament diameter) younger. Thallus dark red in colour; gelatinous and remaining 65 thin when young, and cartilaginous and thicker in 59. Medulla of large cells with a core of small cells wi- older spécimens. Medullar ganglionic cells présent, thin. Plants membranous, terete, consisting of a yellow staining. Gametophytes and tetrasporophytes prostrate and erect System of entangled and irregu- heteromorphic. Tetrasporangia zonately arranged ... larly divided branches. Tetrasporangia in small ra- Genus Meredithia muli, zonately arranged Genus Wurdermannia M. microphylla W. miniata 52. Thallus erect or somewhat decumbent, sessile or 59. Medulla lacking a core of small cells. Plants cartila- shortly stalked, broadly expanded above to form a ginous, compressed or flattened, erect, usually dicho- single blade. Plants initially undivided, but adults tomously or irregularly dichotomously branched. can be lobed, laciniate or also irregularly dichoto- Tetrasporangia not in small ramuli, cruciately arran- mously branched, sometimes perforated (not in Me- ged Family PHYLLOPHORACEAE 60 diterranean species). Thallus rose-pink to dark red 60. Cortex of several layers of small isodiametric cells in colour, fleshy-membranous in texture. Medullar mostly arranged in radial rows. Plants compressed, cells présent, either ganglionic and yellow staining, shortly stipitate, dividing in one plane. Life history or stellate and hyaline. Gametophytes and tetraspo- biphasic, with tetrasporophytes hemiparasitic, as rophytes isomorphic. Tetrasporangia cruciately or pustules on the surface of female gametophytes irregularly arranged Genus Kallymenia 53 Genus Gymnogongrus 61 53. Genus Kallymenia 53. Inner cortical cells S (40-)60 um in diameter. 60. Cortex of small cells not arranged in radial rows. Outer cortical cells rounded in surface view, Plants flattened, with a long stipe, not complanately and developing rosettes around large inner branched. Life cycle triphasic with isomorphic ga- cells. Medullar cells stellate, usually colourless, metophytes and tetrasporophytes 62 50 um large 54 61. Genus Gymnogongrus 53. Inner cortical cells < 60 um in diameter. Outer 61. Thallus up to 5-7 (-15) cm high, and 2-4 mm cortical cells ovoid, polyhedral or irregularly broad. Apices obtuse. Medullar cells 100-250 shaped in surface view, not forming rosettes um and 2-3 times longer. Pustule small in pro- around larger inner cells. Medullar cells ganglio- portion to branches, scattered over both surfa- nic or stellate, yellow staining, usually < 50 um ces G. crenulatus large 55 61. Thallus up to 2.5 (-5) cm high and 0.5-0.6 mm 54. Thallus lobed. Inner cortical cells between 40- broad, except in bifurcations, which are 0.8-1.1 100 um in greatest dimension. Medullar stellate mm wide. Apices acute. Medullar cells 10-20 cells with a rounded to ovoid body, (25-) 50-100 um, and 4-8 (10) times longer. Pustule large in um in diameter, and radiating arms «s 300 um proportion to branches, sometimes completely long encircling them G. griffithsiae K. feldmannii 62. Thallus arising from a branched cylindrical rhizoma- 54. Thallus laciniate and sometimes slightly pinnate tous holdfast. Midrib absent. Carpogonial branches at the ends. Inner cortical cells up to 70-185 um situated in the blade surface. Tetrasporangia aggre- in size. Medullar stellate cells with a very elongated in a séries of sequentially developing sori that gated body, to 70-230 um long and 20-120 um protrude as bands on both surfaces of frond wide, and arms 20-115 um in length originating Genus Schottera mainly at distal parts of the cell K. lacerata S. nicaeensis 55. Frond irregularly, di- or trichotomously branched, reniform when very young. Outer cortical cells 62. Plants arising from a discoid holdfast. Midrib-like irregularly shaped K. patens thickening présent in some species. Carpogonial 55. Frond simple, lobed or laciniate, sometimes with branches situated in spécial bladelets. Tetrasporan- marginal proliférations. Outer cortical cells roun- gia in elevated sori, not sequentially developed ded, polyhedral or irregularly shaped 56 Genus Phyllophora.... 63 56. Inner cortical cells up to 25-30 um in diameter 63. Genus Phyllophora K. spathulata 63. Stipe short in relation to total length of the 56. Inner cortical cells up to 40-50 um in diameter plant. Blade more or less parallel-sided. Margin 57 undulated. Thallus with a basai or médium mi- 57. Outer cortical cells rounded to ovoid or irregu- drib-like thickening formed by secondary larly shaped in surface view. Medullary cells growth of the cortical layer P. crispa 62 RODRiGUEZ-PRIETO C, SABATER MA, VERGÉS A

63. Stipe long in relation to total length of the plant. groups of 1-5 per bearing cell, in close related Blade more or less fan-shaped. Margin plane. cells, but not always in the upper and the lower Thallus lacking midrib-like thickening 64 part of the auxiliary cell. Gonimoblast initially 64. Blade very divided, with acute apices. Stipe formed on the auxiliary cell, and originating op- usually representing up to half of the total posite to the Connecting filament. Gonimoblast length of the frond. Medullary cells up to 40-50 ovoid to lobed P. pusilla um in diameter P. heredia 70. Carpogonial and auxiliary branches in a spécial 64. Blade usually with a single dichotomy and ob- compact branch System (ampulla). Non-procarpic. tuse apices. Stipe usually less than one third of Tetrasporangia cruciately arranged total length of the frond. Medullary cells 90 Family H AL YMENI ACE AE.. ..71 um in diameter P. sicula 70. Carpogonial and auxiliary branches not in a spécial 65. Outer cortex arranged in filamentous fascicles. Non- compact branch System (ampulla). Procarpic or non- procarpic 66 procarpic. Tetrasporangia cruciately or zonately ar- 65. Outer cortex not arranged in filamentous fascicles. ranged 80 Procarpic or non-procarpic 70 71. Cortex of cells arranged in radial rows. Inner corti- 66. Carpogonial branch with adjacent nutritive auxiliary cal cells ovoid to ameboid. Medulla filamentous, cells (nutritive cells originating from the same with numerous X-shaped cells remaining 72 branch System of the carpogonial branch) 71. Cortex not in radial rows. Inner cortical cells stellate Family SCHIZYMENIACEAE.. 67 and lying in the plane of the flattened surface. Me- 66. Carpogonial branch without adjacent nutritive auxi- dulla filamentous but lacking X-shaped cells... 76 liary cells Family NEMASTOMATACEAE . 68 72. Margin rolled over one of the sides giving a 67. Gland cells rounded to ovoid in shape formed in an concave-convex appearance. Plants segmented, ri- intercalary position in cortical fascicles. Gameto- gid, sometimes spirally twisted..Genus Acrodiscus phytes erect or somewhat decumbent, the branching A. vidovichii is dichotomous or irregularly lobed or palmate to pinnate. Branches often anastomosing. Carposporo- 72. Margin not rolled over one of the sides. Plants not phytes lacking a filamentous involucre segmented, not rigid, not spirally twisted 73 Genus Platoma 73. Thallus with a canaliculated margin in the Mediter- P. cyclocolpum ranean species, sometimes distinct only with a raa- 67. Gland cells ellipsoid to ovoid in shape, situated at gnifying glass. Plants simple, flattened the end of the cortical filaments. Gametophytes Genus Aeodes erect, simple or irregularly lobed, sometimes laci- A. marginata niate. Branches not anastomosing. Carposporophy- 73. Margin not canaliculated. Plants simple or variously tes with a filamentous involucre divided, flattened or compressed Genus Schizymenia Genus Grateloupia.... 74 S. dubyi 74. Genus Grateloupia 68. Gland cells présent in gametophytes of Mediterra- 74. Thallus flattened, foliose, simple or deeply di- nean species. Fronds gelatinous but cartilaginous at vided once to several times, often proliferous the base. Auxiliary cell without nutritive cells. Goni- from damaged or truncated parts. Plants 2-6 cm moblasts developing from the Connecting filament. wide G. doryphora Genus Nemastoma 74. Thallus compressed, not foliose, dichotomously N. dichotomum or pinnately branched. Plants s; 5 mm broad 68. Gland cells absent in gametophytes of Mediterra- 75 nean species. Fronds soft and mucilaginous, very dé- 75. Frond branching repeatedly dichotomous. licate. Auxiliary cell bearing short simple or Plants up to 1.5-3.0 mm wide, attenuated to- branched chains of subspherical nutritive cells. Go- wards the outer surface G. dichotoma nimoblast developing either on auxiliary cells or on 75. Frond initially dichotomous and becoming lar- Connecting filaments near the point of union with gely complanately branched, bi- or tripinnate auxiliary cell Genus Predaea 69 when old. Plants up to 3 (-5) mm wide 69. Genus Predaea G. filicina 69. Plants up to 4-5 (-9) cm high, dichotomously or 76. Medulla lax in young parts, with filaments forming irregularly branched. Outer cortical cells ovoid bridges from cortex to cortex. Thallus simple or va- to cylindrical. Carpogonial branch 2-celled. Nu- riously branched. Stipe filamentous. Thallus usually tritive cells originating in groups of (3-) 5-12 (- rose pink in colour, very soft and gelatinous in tex- 15) cells, situated on the cells site under and ture. Carposporophyte and tetrasporangia scattered over the auxiliary cell. Gonimoblast developing and immersed in outer cortex on a latéral swelling of the junction filament, Genus Halymenia...77 the gonimoblast and the Connecting filament re- 76. Medulla usually fairly compact and strong, wi- maining side by side. Gonimoblast rounded or thout filaments forming bridges from cortex to sometimes pyriform, radially orientated cortex. Thallus leaf-like, with proliférations of P. ollivieri the same shape of the main blade, with the thal- 69. Plants up to 1-4 (-7) cm high, simple or sparsely lus appearing catenate. In cross section, the stipe branched dichotomously or irregularly at the api- is celled, with a secondary developed cortex ar- ces. Outer central cells cylindrical. Carpogonial ranged in concentric circles. Thallus dark-red in branch 3-celled. Nutritive cells originating in colour, membranous to cartilaginous in texture. A KEY TO THE RHODOPHYTA OF WESTERN MEDITERRANEAN, PART I 63

Carposporophyte and tetrasporangia scattered or 84. Medulla lax, in younger parts transversed by a net- occurring in secondary bladelets work of filaments mostly at right angles to plane of Genus Cryptonemia.... 79 the flattened surface, and dense and transversely 77. Genus Halymenia oriented in older ones. Cortex thin. Outer cortical 77. Plants terete or slightly compressed, dichoto- cells radially elongated. Gland cells absent mously or subdichotomously branched, someti- Family FURCELLARIACEAE...85 mes whorled. Medulla lax, with occasionally 84. Medulla appearing empty in younger part, and trans- transversely orientated filaments. When the thal- versed by a network of filaments mostly at right an- lus is folded, creases are formed H. trigona gles to plane of the flattened surface in older ones, 77. Thallus flattened, simple, pinnate or dichoto- usually with numerous stellate cells remaining. Cor- mously branched. Medulla lax in young parts tex thick. Outer cortical cells not radially arranged. with many to most filaments transversely orien- Gland cells sometimes présent tated, dense and irregular in older parts. When Family SEBDENIACEAE the thallus is folded, creases are not formed. 78 Genus Sebdenia...&6 78. Thallus lanceolate, simple or, rarely, divided once dichotomously. Plants up to 180 (-300) um 85. Fronds bearing a long, terete and caulescent stipe, thick. Medullar filaments unbranched, 8-14 um usually branched in mature plants, with an inner part thick. Outer cortical cells polyhedral in surface filamentous, and an outer part of little and radially view, compact, 15 um in greatest dimension elongated cells. Plants flattened, with blades broa- H. latifolia der than high and peltate. Plants cartilaginous 78. Thallus complanately much-branched to 4-5 or- Genus Neurocaulon ders, pinnate or occasionally subdichotomous. N. foliosum Plants =£ 800 um thick. Medullar filaments bran- 85. Fronds lacking a caulescent branched stipe. Plants ched, of 2 sizes, 10-15 and 15-20 um. Outer cor- compressed or flattened (subcylindrical when nar- tical cells ovoid in surface view, « 10 um in row), entire or irregularly dichotomously branched, diameter, and radially elongated (up to 10-15 um often furnished with ligulate proliférations from the high) H. floresia margins and blade surface. Thin plants soft and dia- 79. Genus Cryptonemia phanous, gelatinous; thicker plants subcartilaginous 79. Blades =s 3 cm broad, into which the stipe ex- Genus Halarachnion tends as an evanescent midrib-like thickening in H. ligulatum mature plants C. lomation 86. Genus Sebdenia 79. Blades «s 0.5 cm broad. Midrib absent 86. Réfringent gland cells présent. Medullary stel- C. tunaeformis late cells abondant 87 80. Cortex of cells arranged in radial rows. Inner corti- 86. Gland cells absent. Medullary stellate cells rare. cal cells ovoid to ameboid. Medulla of a network of S. monardiana irregularly oriented filaments and numerous X-sha- 87. Gland cells very obvious situated in the center ped cells remaining. Thallus erect or decumbent. of the stellate cells and 20-30 um in diameter, Procarpic or non-procarpic 81 or over the medullar filaments, and then 15-20 80. Cortex of cells not arranged in radial rows. Inner um thick. Thallus 400-500 um thick, cuneiform, cortical cells stellate and lying in the plane of the thicker at the base and margins, first simple and flattened surface. Medulla filamentous, lacking X- lanceolate, later divided in large subdichoto- shaped cells, but occasionally with some stellate mous lobes, with deep sinus. Margin whole. cells remaining. Thallus erect. Non-procarpic ....84 Stellate cortical cells 20-50 (-70) um in diame- 81. Medulla consisting of primary filaments, usually ter S. rodrigueziana longitudinally orientated, interlined with some se- 87. Gland cells not obvious, situated in the center condary short filaments, 1-3 cells long, thinner, of the stellate cells and 10-15 um in diameter. usually transversely oriented. Procarpic Thallus =s 1500 um thick, more or less flatte- Family GIGARTINACEAE ned, but usually cylindrical at the apices, dicho- Genus Chondracanthus 82 tomously or subdichotomously branched, 81. Medulla lacking secondary filaments. Procarpic or usually with marginal proliférations. Stellate non-procarpic 83 cortical cells up to 15-20 (-25) um in diameter... 82. Genus Chondracanthus S. dichotoma 82. Fronds compressed or terete, irregularly branched. Plants erect or decumbent, often arching TAXONOMIC ARRANGEMENT and reattaching on contact C. acicularis OF THE GENERA TREATED 82. Fronds flattened, regularly pinnately branched. Plants erect, not reattaching C. teedii 83. Margin ribbed and dentate. Thallus leafy and simple Order GIGARTINALES Schmitz in Engler or dichotomously branched. Procarpic. Plants midlit- Family ACROSYMPHYTACEAE Lindstrom toral Family RISSOELLACEAE Genus ACROSYMPHYTON Sjôstedt; "Acrosymphytone- Genus Rissoëlla ma" stadium Boudouresque, Perret-Boudouresque & R. verruculosa Knoepffler-Péguy 83. Margins smooth or ribbed, but not dentate. Thallus A. purpuriferum (J. Agardh) Sjôstedt almost as long as it is broad, lobed. Non-procarpic. Tetrasporophyte: "Acrosymphytonema breemaniae " Plants sublittoral Genus Chondrymenia Boudouresque, Perret-Boudouresque & Knoepf- C. lobata fler-Péguy 64 RODRIGUEZ-PRIETO C, SABATER MA, VERGÉS A

Family CALOSIPHONfrCEAE Kylin K. spathulata (J. Agardh) Parkinson Genus CALOSIPHONIA P.L. Crouan & H.M. Crouan Genus MEREDITHIA J. Agardh C. dalmatica (Kiitzing) G.B. De Toni M. microphylla (J. Agardh) J. Agardh C. vermicularis (J. Agardh) Schmitz Tetrasporophyte: Rhodochorton hauckii (Schiff- Genus SCHMITZIA Lagerheim ex P.C. Silva ner) Hamel S. neapolitana (Berthold) Lagerheim ex P.C. Silva Family NEMASTOMATACEAE Schmitz in Engler Family CAULACANTHACEAE Kiitzing Genus NEMASTOMA J. Agardh Genus CATENELLA Greville N. dichotomum J. Agardh C. caespitosa (Withering) Irvine in Parke & P.S. Genus PREDAEA G. De Toni Dixon P. ollivieri J. Feldmann Genus CAULACANTHUS Kiitzing P. pusilla (Berthold) J. Feldmann C. ustulatus (Turner) Kiitzing Family PEYSSONNELIACEAE Denizot Genus FELDMANNOPHYCUS Augier & Boudouresque Genus METAPEYSSONNELIA Boudouresque, Coppe- F. rayssiae (J. Feldmann & G. Feldmann) Augier & jans & Marcot Boudouresque M. feldmannii Boudouresque, Coppejans & Marcot Family CRUORIACEAE Kylin Genus PEYSSONNELIA Decaisne Genus CRUORIA Fries P. armorica (P.L. Crouan & H.M. Crouan) Bôrgesen C. cruoriaeformis (P.L. Crouan & H.M. Crouan) De- P. atropurpurea P.L. Crouan & H.M. Crouan nizot P. bornetii Boudouresque & Denizot Family CYSTOCLONIACEAE Kiitzing P. codana (Rosenvinge) Denizot Genus CALLIBLEPHARIS Kiitzing P. coriacea J. Feldmann C. ciliata (Hudson) Kiitzing P. crispata Boudouresque & Denizot C. jubata (Goodenough & Woodward) Kiitzing P. dubyi P.L. Crouan & H.M. Crouan Genus RHODOPHYLLIS Kiitzing P. harveyana P.L. Crouan & H.M. Crouan R. divaricata (Stackhouse) Papenfuss P. hongii Marcot-Coqueugniot R. strafforellii Ardissone P. inamoena Pilger P. magna Ercegovic Family DUMONTIACEAE Bory de Saint-Vincent P. polymorpha (Zanardini) Schmitz in Falkenberg Genus DUDRESNAYA P.L. Crouan & H.M. Crouan P. rara-avis Marcot & Boudouresque D. verticillata (Withering) Le Jolis P. rosa-marina Boudouresque & Denizot Family FURCELLARIACEAE Greville f. rosa-marina Boudouresque & Denizot Genus HALARACHNION Kiitzing f. saxicola Boudouresque & Denizot H. ligulatum (Woodward) Kiitzing P. rubra (Greville) J. Agardh Tetrasporophyte: Cruoria rosea P.L. Crouan & P. squamaria (Gmelin) Decaisne H.M. Crouan P. stoechas Boudouresque & Denizot Genus NEUROCAULON Zanardini ex Kiitzing Genus POLYSTRATA Heydrich N. foliosum (Meneghini) Zanardini P. compacta (Foslie) Denizot Family GIG ARTIN ACEAE Kiitzing P. fosliei (Weber-van-Bosse) Denizot Genus CHONDRACANTHUS Kiitzing Family PHYLLOPHORACEAE Nageli C. acicularis (Roth) Fredericq in Hommersand, Gui- Genus GYMNOGONGRUS Martius ry, Fredericq & Leister G. crenulatus (Turner) J. Agardh C. teedii (Mertens ex Roth) Kiitzing G. griffithsiae (Turner) Martius Family GLOIOSIPHONACEAE Schmitz Genus PHYLLOPHORA Greville Genus GLOIOSIPHONIA Carmichael in Berkeley P. crispa (Hudson) Dixon G. capillaris (Hudson) Carmichael in Berkeley P. heredia (Clémente y Rubio) J. Agardh Tetrasporophyte: Rhododiscus pulcherrimus P.L. P. sicula (Kiitzing) Guiry & Irvine Genus SCHOTTERA Guiry & Hollenberg Crouan & H.M. Crouan Genus THURETELLA Schmitz S. nicaëensis (Lamouroux ex Duby) Guiry & Hollen- berg T. schousbœi (Thuret) Schmitz Genus SCHIMMELMANNIA Schousboe ex Kiitzing Family RHYZOPHYLLIDACEAE Schmitz S. ornata Schousboe ex Kiitzing Genus CONTARINIA Zanardini C. peyssonneliaeformis Zanardini Family HYPNEACEAE J. Agardh C. squamariae (Meneghini) Denizot Genus HYPNEA Lamouroux H. cervicornis J. Agardh Family RIS SOËLLACEAE Kylin H. furnariana Cormaci, Alongi & Dinaro Genus RISSOËLLA J. Agardh H. musciformis (Wulfen) Lamouroux R. verruculosa (Bertoloni) J. Agardh Family KALLYMENIACEAE (J. Agardh) Kylin Family SARCODIACEAE Kylin Genus CALLOPHYLLIS Kiitzing Genus CHONDRYMENIA Zanardini C. laciniata (Hudson) Kiitzing C. lobata (Meneghini) Zanardini Genus KALLYMENIA J. Agardh Family SCHIZYMENI ACEAE (Schmitz & Haupt- K. feldmannii Codomier fleisch) Masuda & Guiry K. lacerata S. Feldmann Genus PLATOMA Schousboe ex Schmitz K. patens (J. Agardh) Parkinson P. cyclocolpum (Montagne) Schmitz K. reniformis (Turner) J. Agardh Genus SCHIZYMENIA (J. Agardh) J. Agardh K. requienii J. Agardh 5. dubyi (Chauvin ex Duby) J. Agardh A KEY TO THE RHODOPHYTA OF WESTERN MEDITERRANEAN, PART I 65

Tetrasporophyte: Haematocelis rubens J. Agardh University of Barcelona, the Laboratoire de Biologie Family SPHAEROCOCCACEAE Dumortier emend. Marine et d'Écologie du Benthos from the University of Searles Marseille (France) and the Observatoire Océanologique Genus SPHAEROCOCCUS Stackhouse de Banyuls-sur-Mer from the University of Paris VI 5. coronopifolius (Goodenough & Woodward) Stack- (France) for providing indispensable bibliography, and house to M Perret-Boudouresque, MT Panouse and S Maria for Tetrasporophyte: Haematocelis fissurata P.L. their indispensable help. Valuable material was provided Crouan & H.M. Crouan by K Ballesteros, N Sant and L Polo. Thanks also for as- S. rhizophylloides Rodriguez Fememas sistance in the laboratory to N Niell and G Vila, and to X Quintana for his hepful translations from German to Order HALYMENIALES Saunders & Kraft Spanish. This project was supported by the Ministerio de Family HALYMENIACEAE Bory de Saint-Vincent Educaciôn y Cultura (Spain), PB95-0385-C06-06. Genus ACRODISCUS Zanardini A. vidovichii (Meneghini) Zanardini Genus AEODES J. Agardh A. marginata (Roussel ex Montagne) Schmitz REFERENCES Genus CRYPTONEMIA J. Agardh C. lomation (Bertoloni) J. Agardh C. tunaeformis (Bertoloni) Zanardini FREDERICQ S, HOMMERSAND MH 1989. Proposai Genus GRATELOUPIA C. Agardh of the Gracilariales ord. nov. (Rhodophyta) based on G. dichotoma J. Agardh an analysis of the reproductive development of Graci- G. doryphora (Montagne) Howe laria verrucosa. J Phycol 25: 213-227. G. filicina (Lamouroux) C. Agardh Genus HALYMENIA C. Agardh KRAFT GT, ROBINS PA 1985. Is the order Cryptone- miales (Rhodophyta) defensible ? Phycologia 24: 67- H. floresia (Clémente y Rubio) C. Agardh H. latifolia P.L. Crouan & H.M. Crouan 77. H. trigona (Clémente y Rubio) C. Agardh KYLIN H 1956. Die Gattungen der Rhodophyceen. Lund: C W K Gleerups Fôrlag. XV + 673 p., 458 figs. Family SEBDENIACEAE Kylin MAGGS CA, PUESCHEL CM 1989. Morphology and Genus SEBDENIA (J. Agardh) Berthold development of Ahnfeltia plicata (Rhodophyta): pro- 5. dichotoma (J. Agardh) Berthold posai of Ahnfeltiales ord. nov. J Phycol 25: 233-351. S. monardiana (Montagne) Berthold PUESCHEL CM, COLE H 1982. Rhodophycean pit S. rodrigueziana (J. Feldmann) Codomier plugs: an ultrastructural survey with taxonomic impli- Order HILDENBRANDIALES Pueschel & Cole cations. Ann J Bot 69: 703-720. Family HILDENBRANDIACEAE Rabenhorst SAUNDERS GW, KRAFT GT 1994. Small subunit Genus HILDENBRANDIA Nardo rRNA gene séquences from représentatives of selec- H. crouanii J. Agardh ted families of the Gigartinales and Rhodymeniales H. occidentalis Setchell in Gardner (Rhodophyta). 1. Evidence for the Plocamiales ord. H. rubra (Sommerfelt) Meneghini nov. Can J Bot 72: 1250-1263. SAUNDERS GW, KRAFT GT 1996. Small subunit Order PLOCAMIALES Saunders & Kraft rRNA gene séquences from représentatives of selec- Family PLOC AMI ACEAE Kiitzing ted families of the Gigartinales and Rhodymeniales Genus PLOCAMIUM Lamouroux (Rhodophyta). 2. Récognition of the Halyméniales P. cartilagineum (Linnaeus) P.S. Dixon ord. nov. Can J Bot 74: 694-707. P. secundatum (Kiitzing) Kiitzing SILVA PC, BASSON PW, MOE RL 1996. Catalogue of INCERTAE SED1S the benthic marine algae of the Indian Océan. Univ Genus WURDERMANNIA Harvey Calif Pubis Bot 1259 p. W. miniata (Sprengel) J. Feldmann & Hamel SILVA PC, JOHANSEN HW 1986. A reappraisal of the order Corallinales (Rhodophyta). Br Phycol J 21: ACKNOWLEDGEMENTS. - We would like to thank K 245-254. Ballesteros and M Verlaque for their very helpful suggestions and comments. We are very grateful to the La- Reçu le 3 décembre 1999; receivedDecember 3,1999 boratori de Botànica (Facultat de Farmàcia) from the Accepté le 4 juillet 2000 ; accepted July 4, 2000 66 ANALYSE D'OUVRAGE/BOOtf REVIEW

ANALYSE D'OUVRAGE/500£ REVIEW

M. NORMAN, 2000. Cephalopods - a world guide. Con- scientifique de renom international, a su créer, avec le chBooks (E-mail : [email protected]), D- concours d'une maison d'édition spécialisée dans le do- 55546 Hackenheim (Allemagne), p., 708 fig., 16 x 24 maine du livre naturaliste, un guide « haut de gamme ». cm. Couverture : carton rigide plastifié. 41.00 Euros M. NORMAN & A. REID, 2000. A Guide to Squid, (+ frais d'expédition en Europe : 5.50 Euros). [Deman- Cuttlefish and Octopuses of Australasia. CSIRO Publis- der les conditions de vente pour l'édition en Allemand]. hing (E-mail : [email protected]), 150 Oxford Depuis la parution, en 1987, du grand livre des Cé- Street, Collingwood, Victoria 3066 (Australie). 96 p., phalopodes rédigé et illustré par Kir. N. Nesis (« Cepha- 242 figs, 16.5 x 24 cm. Couverture : carton souple plasti- lopods of the World »), les teuthologistes et les fié. 32.00 $AU. naturalistes amateurs s'intéressant aux Poulpes, Seiches, Bien que ce guide soit le « petit frère » de Sépioles et Calmars, attendaient un ouvrage faisant CEPHALOPODS n A WORLD GUIDE, il peut être uti- « systématiquement » appel aux photos en couleur. lisé de manière parfaitement indépendante, pour identi- L'excellent livre de Norman répond pleinement à cette fier la soixantaine d'espèces de Céphalopodes recensés attente, en présentant près de 230 espèces par une véri- sur les côtes Australiennes et dans les régions avoisinan- table galerie de portraits d'animaux vivants - pour la tes comprises entre 10°N et 50°S, et entre 95° et 190°E. plupart photographiés dans leur environnement naturel. Dans un avant-propos enthousiaste, Kir Nesis insiste sur L'introduction est suivie de brefs rappels sur (1) le l'importance de cette approche : « [les céphalopodes] rapport systématique entre les Céphalopodes et les au- sont difficiles à identifier quand ils sont morts, voire tres Mollusques, (2) l'évolution des Céphalopodes, (3) fixés, alors qu'ils sont très différents entre eux dans leur leur importance dans la faune marine actuelle, (4) les environnement naturel, grâce aux couleurs et comporte- grands groupes de Céphalopodes vivants et l'essentiel de ments si diversifiés ». leur biologie. Afin de permettre au lecteur non initié de se familiari- A quelques exceptions près, la présentation de chaque ser avec ces animaux, Norman commence par une intro- espèce occupe une page entière : une grande photo en duction se terminant par un glossaire, continue par une couleur montrant l'adulte, une ou deux petites photos brève description des caractéristiques de la classe des consacrées à un détail morphologique ou de comporte- Céphalopodes, avec un survol de la systématique - d'ail- ment, toutes avec de brèves légendes, un texte succinct leurs bien visualisée dès le sommaire - couvrant les dif- indiquant le mode de vie, la taille adulte, quelques carac- férents groupes, en ajoutant un bref rappel sur les formes téristiques de la reproduction, puis un cartouche conte- fossiles. Pour chaque groupe taxonomique, une brève in- nant les caractères distinctifs de l'espèce et une petite troduction présente l'essentiel de sa répartition géogra- carte montrant sa répartition géographique. phique et de sa biologie/écologie, avec des renvois à une ou plusieurs des 35 « histoires illustrées [picture sto- Pour une trentaine d'espèces de Seiches, l'os (sépion) ries] ». Ces histoires offrent des observations inédites ou est présenté en vues dorsale et ventrale ; ces « portraits peu connues, qui sont autant de suggestions pour des de pièces détachées » sont réunis dans une série de plan- études futures. ches qui seront particulièrement appréciées par les col- Contrairement aux compilations plus ou moins réus- lectionneurs fréquentant les plages. En effet, les sépions, sies de certains « éditeurs d'images », Norman offre un détachés du corps après la mort de l'animal, peuvent véritable précis de teuthologie, richement illustré, avec s'échouer sur une plage, à l'issue d'une dérive plus ou un index des noms scientifiques et vernaculaires ; en moins longue ; ils sont alors des indicateurs intéressants, même temps il attire l'attention du lecteur (et en particu- quoique « approximatifs », de la présence d'une ou de lier du plongeur scientifique) sur les nombreuses ques- plusieurs espèces de seiches dans une zone plus ou tions non encore résolues. Ainsi 20 espèces de Poulpes moins vaste. non encore décrites « lege artis » (ce qui exigerait le dé- Notons enfin l'excellent glossaire ainsi que l'index pôt d'un ou de plusieurs spécimens types) sont représen- réunissant les noms scientifiques et vernaculaires. Tout tées par d'excellentes photos accompagnées de brèves comme son « grand frère » le « Guide mondial des Cé- descriptions biologiques -voilà un exemple de ce que phalopodes », ce fascicule est un excellent outil de tra- Nesis dit dans son avant-propos ! vail doté d'une grande qualité esthétique. Ce très beau livre témoigne d'un souci permanent de soigner le contenu autant que l'esthétique. L'auteur, un S. v. Boletzky