Bulletin British Museu (Natural History) BRITISH MUSfcU (NATURAL WSTORYi PRE8CNTBO GENERAL LIBRA Botany Series VOLUME 22 NUMBER 2 26 NOVEMBER 1992 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Dr S. Blackmore Editor of Bulletin: Dr R. Huxley Assistant Editor: Mrs M.J. West Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. A volume contains about 160 pages, made up by two numbers: published Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 716 Andover Hampshire SP10 1YG Telephone: (0264) 334748 Fax: (0264) 334058 World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.) © British Museum (Natural History), 1992 Botany Series ISSN 0068-2292 Vol. 22, No. 2, pp. 109-170 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 26 November 1992 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by The Alden Press, Oxford Bull. Br. Mas. not. Hist. (Bot.) 22(2): 109-121 Issued 26 November 1992 Palynological evidence for the generic delimitation of Sechium (Cucurbitaceae) and its allies BFHTISH MUSEUM (NAflffiAL MSTOtti JOSE LUIS ALVARADO Institute National de Antropologia e Historia, Mexico RAFAEL LIRA-SAADE Herbaria National de Mexico, Institute de Biologia, U.N.A.M. Mexico JAVIER CABALLERO Jar din Botdnico, Instituto de Biologia, U.N.A.M. Mexico ,,:", ,:,...., PRESENTED GFNERAL LIBRARY CONTENTS Introduction 109 Material and Methods 110 Results and Discussion 112 1 . Pollen Morphology 112 2. Numerical Taxonomic Treatment 119 Conclusions 120 Acknowledgements 121 References . . 121 SYNOPSIS. In order to gather new data about the taxonomic limits of Sechium, a palynological study of 15 taxa belonging to the genera Sechium sensu lato, Frantzia, Parasicyos, Michrosechium, Pterosicyos, Sechiopsis and Sicyos, all in subtribe Sicyinae, was carried out. A numerical taxonomic analysis was conducted using the palynological characters observed. The results obtained showed that pollen morphology is highly significant in the delimitation of genera within the subtribe Sicyinae, and give some support to the conclusions of Jeffrey (1978) that genera Ahzolia Polakowskia and Frantzia, as well as Microsechium hintonii are congeneric with Sechium. INTRODUCTION Table 1 Summary of the two generic conceptions of Sechium and its closest allies. According to Jeffrey (1990), the genus Sechium P.Br. belongs to the subtribe Sicyinae C. Jeffrey, tribe Sicyeae Schrad. For a long time this genus was considered as monospecific, consisting only of 5. edule (Jacq.) Swartz, a highly variable crop species native to Mexico and Central America. Based upon the comparison of macromorphological characters, Jef- frey (1978) enlarged the limits of Sechium by including in two sections a series of Mexican and Central American species formerly pertaining to the genera Ahzolia, Microsechium, Polakowskia and Frantzia (Dieterle, -F7 1976; Pittier, 1910; Wilson, 1958; Wunderlin, 1976; Table 1). However, this circumscription of the genus has not been widely accepted. In a study on the origin and evolution of Sechium edule, Newstrom (1986, 1990) proposed the segregation of Sechium sensu stricto (constituted by three species) and the reinstate- ment of the genera Frantzia and Polakowskia (Table 1), mainly on the basis of the comparison of flower and fruit characters. The controversy described above remains unsolved and Jeffrey (1978, 1990) Newstrom (1986, 1990) Sechium P.Br. (s. /.) Section Sechium ('pouch-like' nectaries) 5. edule (Jacq.) Swartz 5. compositum (Donn. Sm.) C. Jeffrey 5. hintonii (P.O. Wilson) C. Jeffrey 5. tacaco (Pitt.) C. Jeffrey S. talamancense (Wunderlin) C. Jeffrey Section Frantzia ('cushion-like' nectaries 5. pittieri (Cogn.) C. Jeffrey S. villosum (Wunderlin) C. Jeffrey Sechium P.Br. (s. str.) No sections 5. edule (Jacq.) Swartz 5. compositum (Donn. Sm.) C. Jeffrey 5. hintonii (P.O. Wilson) C. Jeffrey Polakowskia Pittier P. tacaco Pitt. P. talamancensis (Wunderlin) Newstrom Frantzia Pittier F. pittieri (Cogn.) Pitt. F. villosa Wunderlin F. venosa L.D. Gomez F. panamensis Wunderlin 110 more comparative evidence is still needed in order to assess the generic limits of Sechium. Related to this, several palyno- logical studies (Ayala-Nieto et al ., 1988; Jeffrey, 1964, 1980; Page & Jeffrey, 1975; Singh & Singh, 1990) have shown that pollen morphology varies considerably between taxa within the Cucurbitaceae and that this kind of character could be highly significant in the delimitation of tribes, subtribes and even genera. Following this approach, the authors conducted a palynological study of the species of Sechium sensu lato (Cucurbitaceae) and its allies in order to provide some empirical evidence for the taxonomic delimitation of the genus Sechium. This study included the description of the morphological characters of the pollen grains of 15 different taxa of Sechium and their relatives, as well as the utilization of such characters to assess the generic limits of Sechium by means of a numerical taxonomic analysis. The 15 taxa studied include: the seven species recognized by Jeffrey (1978, 1990) as constituting Sechium sensu lato (wild and cultivated forms of 5. edule were considered as different taxa for the purpose of the study); two species of Frantzia not yet transferred to Sechium (F. panamensis Wunderlin and F. venosa L.D. Gomez) (Gomez & Gomez, 1984; Wunderlin, 1977); and five other species each one belonging to a distinct genus of the subtribe Sicyinae (Microsechium helleri (Peyr.) Cogn., Parasicyos maculatus Dieterle, Sechiopsis triquetra (Ser.) Naudin, Pterosicyos laciniatus Brandegee, and Sicyos motozintlensis Lott & Fryx- ell). Of the 15 taxa included in this study only Microsechium helleri (Rodriguez- Jimenez & Palacios-Chavez, 1991), Sech- ium compositum (Dieterle, 1976; Marticorena, 1963 as Ahzo- lia composita), S. pittieri and 5. tacaco (Marticorena, 1963 as Frantzia pittieri and Polakowskia tacaco respectively) have been palynologically studied before. As for Sechium edule, only the pollen grains of cultivated plants have been previ- ously described (Aleshina, 1964; Ayala-Nieto et al., 1988; Erdtman, 1952; Marticorena, 1963; Rangaswami, 1976; Singh & Singh, 1990), therefore this study included the examination of pollen from wild populations. MATERIALS AND METHODS Pollen samples of the 15 taxa of Sechium and their relatives were taken from herbarium specimens (Table 2) available at the following institutions: Colegio de Postgraduados, Chap- ingo, Mexico (CHAPA); Herbario Nacional de Mexico, U.N.A.M.(MEXU); Institute de Ecologfa, Xalapa, Mexico (XAL); British Museum (Natural History) (BM); Gray Her- barium, Harvard University (GH); Royal Botanic Gardens, Kew (K); University of Michigan (MICH); and Missouri Botanical Garden (MO). The samples were examined by light microscope (LM). Anthers, flowers and small buds were removed from the specimens and then acetolized according to the method described by Erdtman (1943). Permanent slides were made using glycerine jelly. Two slides were made from each specimen examined (Table 2). Measurements of the 12 palynological characters considered in this study (Table 3) were made from 10 pollen grains in polar view and 10 pollen grains in equatorial view from each slide. Microphotographs of the pollen grains were made using a LEITZ-DIALUX ALVARADO, LIRA-SAADE & CABALLERO microscope under 40 x objective, and 100 x oil immersion objective. These microphotographs were made using Kodak Plus-X film. In order to obtain additional information on the pollen morphology of the taxa under study, samples of some of them were also examined under Scanning Electron Micro- scope (SEM; JEOL-SM) and photographed (3000 X). The slides are deposited in the Pollen Collections of the Laborato- rio de Paleobotanica of the Instituto Nacional de Antropo- logfa e Historia, Mexico, and the Herbario Nacional de Mexico, U.N.A.M. (MEXU). The terminology followed in the description of pollen grains is that of Erdtman (1952), Faegri & Iversen (1964) and Kremp (1965). The numerical taxonomic analysis of Sechium sensu lato and its relatives made use of two complementary methods: cluster analysis and ordination techniques. These techniques were used to investigate the grouping of taxa within the genus, on the basis of their overall palynological similarity. Both cluster analysis and ordination techniques were applied using the 'Numerical Taxonomy and Multivariate Analysis System' (NTSYS-PC version 1.50; Rohlf 1989). Twelve palynological characters were included in the analysis. In addition, two floral characters (nectary and stamen structure) were used, both being features of great significance to the classification of the genera of the subtribe Sicyinae, and to the separation of the sections of Sechium sensu lato (Jeffrey, 1978, 1990). The characters used included both metric and non-metric characters (Table 3). A basic data matrix (BDM) was formed with the different character average values for each species. Since characters 8 to 14 are non-metric (see Table 3), their values in the BDM do not constitute averages, but represent codes for the character states observed in each species. In all the species studied, the states of the seven qualitative characters were uniform for all the speci- mens examined. The result was a 15 x 14 matrix in which rows were Operational Taxonomic Units (OTUs) and columns were characters. This BDM is shown in Table 4 in which OTUs are presented as rows and characters are presented as columns. To reduce the effects of different measure scales in different characters and to make the characters contribute more equally to similarity, the BDM was standardized by rows using the linear transformation algorithm provided by NTSYS (Rohlf, 1989: 6.89). The subtract option used in the standardization was the mean of each variable and the divide option was the standard deviation. For the cluster analysis, a dissimilarity matrix of distances between OTUs was calculated from the BDM standardized using the Average Taxonomic Distance Coefficient (see Rohlf, 1989: 6.13). This coefficient is a variation of the widely known Euclidian Distance Coefficient (see Dunn and Everitt, 1982: 32; Rohlf, :6.13; Romesburg, 1984: 12). A cluster analysis using the unweighted pair-group method (UPGMA), as described by Romesburg (1984: 15), was carried out using the above dissimilarity matrix. The ordination of the 15 OTUs in a multidimensional space of characters was made by means of a Principal Coordinate Analysis (PCO). This method was used instead of Principal Components Analysis or other similar metric technique the characters used included qualitative characters. PCO is an extension of Principal Component Analysis which allows the utilization of qualita- tive or mixed data (see Rohlf, 1989: 6.25 and 8.3). PCO was run in two directions: by columns to obtain the representation of the OTUs in a multidimensional space of characters; and by rows, in order to identify those characters which contrib- ute the most to clustering. In the case of the analysis by PALYNOLOGICAL EVIDENCE FOR TAXONOMIC LIMIT OF SECHIUM Table 2 List of voucher specimens of the 15 taxa included in the palynological study. Ill SECHIUM S. L. 1. S. compositum Chiapas Breedlove 31625 MEXU Chiapas Breedlove 55725 MEXU Chiapas Heath & Long 577 MEXU Chiapas Lira 950 MEXU Chiapas Lira 952 MEXU, K Chiapas Lira 960 MEXU, K Chiaqpas Matuda 2151 MEXU Chiapas Matuda 17047 MEXU Guatemala Lira 1004 MEXU, K 2. S. edule (wild=w) Veracruz Beristain 2247 (w) CHAPA Veracruz Beristain 2248 (w) CHAPA 3. S. edule (cultivated =c) Veracruz Beristain 2250 (w) CHAPA Veracruz Beristain 225 l(w) CHAPA Veracruz Beristain 2252 (w) CHAPA Veracruz Beristain 2253 (w) CHAPA Veracruz Beristain 2254 (w) CHAPA Veracruz Beristain 2257 (w) CHAPA Veracruz Cruz-Leon 2211 (w) CHAPA Veracruz Lira 1029 (w) MEXU Yucatan Lira 507 (c) MEXU Campeche Lira 807 (c) MEXU Hidalgo Villa 72 (c) XAL 4. 5. hintonii Edo. de Mexico Hinton 4808 GH Edo. de Mexico Lira & Soto 1082 MEXU, K 5. S. pittieri Costa Rica Chacon 397 MEXU Costa Rica Garwood 423 BM Costa Rica Grayum 8103 BM Costa Rica Khan 1020 BM Panama Hampshire 676 BM Panama Mori 5636 MEXU Panama Schmalzel 1942 BM 6. S. tacaco Costa Rica Lira 1062 MEXU, K Costa Rica Lira 1064 MEXU, K Costa Rica Raven 21793 MO 7. 5. talamancense Costa Rica Grayum 8159 BM Costa Rica Lira 1055 MEXU, K 8. 5. villosum Costa Rica Skutch 3163 GH FRANTZIA 9. F. panamensis Panama Antonio 2687 MO 10. F. venosa Costa Rica Gomez 20483 MO Costa Rica Lira 1036 MEXU, K PARASICYOS 11. P. maculatus Guatemala Williams 40205 MIDH MICROSECHIUM 12. M. helleri Edo. de Mexico Lira 1077 MEXU, K PTEROSICYOS 13. P. lacinlatus Chiapas Lira 956-A MEXU, K SECHIOPSIS 14. S. triquetra Guerrero Lira & Soto 1087 MEXU, K SICYOS 15. 5. motozintlensis Chiapas Lira 951 MEXU, K 112 ALVARADO, LIRA-SAADE & CABALLERO Table 3 List of the palynological characters (1-12) and the floral ones (13, 14) used in the numerical taxonomic study of Sechium s. I. and its allies. 1. Polar axis (um) 2. Equatorial axis (urn) 3. Spine length (urn) 4. Exine thickness (urn) 5. Number of colpi o. Polar/Equatorial Ratio 7. Polar area Index 8. Ornamentation type 0 = spiniferous, microreticulate 1 = spiniferous, clavate, microreticulate 2 = spiniferous, reticulate 9. Shape 0 = oblate 1 = suboblate 2 = oblate-spheroidal 10. Spine type 0 = conic, short (x length < 4.5 um) 1 = conic, long (x length > 4.5 um) 2 = cylindric, very variable length 11. Spines apex 0 = without a small cavity 1 = with a small cavity 12. Colpus shape 0 = regular or continuous 1 = irregular or discontinuous 13. Floral nectaries 0 = without nectaries 1 = pore shape nectaries 2 = cushion shape nectaries 14. Stamen structure 0 = free anthers on branches of filaments 1 = free anthers sessile on filaments column 2 = anthers and filaments fused into a column columns, PCO was performed using the same dissimilarity matrix as in the cluster analysis. For the PCO by rows, a new dissimilarity matrix with the distances between variables was calculated. In both cases the dissimilarity matrices were transformed into their scalar products by means of a double centring procedure (see Rohlf, 1989: 6.25), so that their eigenvalues and eigenvectors could be computed. RESULTS AND DISCUSSION The results and discussion are presented in two sections: the first includes descriptions and illustrations of the pollen morphology, while the second encompasses the analysis and discussion of the results of the numerical taxonomic study. 1. Pollen Morphology A. General Features of the Pollen Grains Comparative illustrations of the taxa studied are included in Plates I (LM) and II (SEM). Plate I includes complete pollen grains (in polar view) of the 15 taxa considered in the study, showing the most typical palynological features of the sub- tribe Sicyinae (spiniferous, colporate) recognized by Jeffrey (1990). However, general interspecific differences in size, shape and number of colpi of the grains, as well as in size and shape of the spines, can be clearly observed. Under the SEM (Plate II), differences in ornamentation (e.g. reticulation in the interspinal area) are more evident and confirm the observations made under light microscope. B. Palynological Descriptions Sechium compositum (Donn.-Sm.) C.Jeffrey (Plate III: 1-6) Eumonad, isopolar, radiosymmetric; oblate (60.8 x 81.0 um), stephanocolpate (8-9 colpi). Tectate; spiniferous, conical spines 4.3-6.3 um long, with a small cavity near the top. Interspinal area microreticulate. Exine 3.0 um thick; sexine thicker than nexine; amb circular. Sechium edule (Jacq.) Swartz S. edule (wild) (Plate III: 7-13) Eumonad, isopolar, radiosymmetric; suboblate (64.0 x 82.7 um), stephanocolpate (8-9 colpi). Tectate; spiniferous, conical spines 4.9-7.2 um long, with a small cavity near the Table 4 Data matrix showing the values found for the 14 characters (columns) in the 15 taxa (rows) of Sechium sensu lato and its allies included in this study. For the purpose of the numerical taxonomic study, OTUs were used as columns and characters were used as rows (N = number of pollen grains examined). 10 11 12 13 14 Sechium compositum (N=240) 60.8 81.0 5.1 3.0 9 0.74 0.26 0 0 1 1 0 1 0 S. edule 'wild' (N=240) 64.0 82.7 6.0 2.5 9 0.77 0.55 0 1 1 1 0 1 0 S. edule 'cultivated' (N=240) 66.8 90.4 3.4 2.3 11 0.73 0.60 1 0 2 1 0 1 0 5. hintonii (N=160) 52.6 68.8 5.8 2.5 10 0.76 0.36 0 1 1 1 0 1 0 S. pittieri (N=240) 48.5 64.4 6.4 2.8 9 0.75 0.36 0 1 1 1 0 2 2 S. tacaco (N=240) 78.3 87.6 6.5 3.0 9 0.89 0.34 0 2 1 1 0 1 1 S. talamancense (N=160) 46.3 60.8 5.5 3.5 9 -.76 0.49 0 1 1 1 0 1 1 S. villosum (N= 80) 54.3 73.7 8.4 2.5 9 0.73 0.40 2 0 1 1 0 2 2 Frantzia panamensis (N= 80) 73.0 81.2 5.4 2.7 9 0.89 0.32 0 2 1 1 0 2 2 F. venosa (N=160) 69.3 76.1 4.9 2.8 9 0.91 0.34 0 2 1 1 0 2 2 Parasicyos maculatus (N= 80) 40.1 41.7 4.4 2.2 9 0.96 0.21 2 2 0 0 0 0 1 Microsechium helleri (N= 80) 68.9 72.8 5.1 2.3 7 0.94 0.19 0 2 1 0 1 0 0 Pterosicyos laciniatus (N= 80) 58.1 60.3 4.3 2.5 9 0.96 0.36 0 2 0 0 0 1 2 Sechiopsis triquetra (N= 80) 52.2 60.5 4.0 2.5 10 0.86 0.39 0 1 0 1 0 1 2 Sicyos motozintlensis (N= 80) 59.7 61.7 3.8 2.1 9 0.96 0.30 0 2 0 0 0 0 2 PALYNOLOGICAL EVIDENCE FOR TAXONOMIC LIMIT OF SECHIUM BRITISH MUSEUM (NATURAL HISTORY) 113 -3DEC1992 PRESENTED L LIBRARY >,•••;-• vv, 1 «*J\ ^* Plate I Comparative illustrations of the 15 taxa studied; polar view. 1, Sechium composition; 2, S. edule (wild); 3, 4, 5. edule (cultivated); 5, 5. hintonii; 6, 5. pittieri; 7, 5. tacaco; 8, 5. talamancense; 9, 5. villosum; 10, Frantzia panamensis; 11, F. veno^o; 12, Parasicyos maculatus; 13, Microsechium helleri; 14, Pteroslcyos laciniatus; 15, Sechiopsis triquetra; 16, Sicyos motozintlensis . (bar scale - 10 nm). top. Interspinal area microreticulate. Exine 2.5 sexine thicker than nexine; amb circular. thick; S. edule (cultivated) (Plate III: 14-19) Eumonad, isopolar, radiosymmetric; oblate (66.8 x 90.4 urn), stephanocolpate (9-11 colpi). Tectate; spiniferous, cylindrical spines very variable in size (1.3-6.7 ^m), with blunt end, with a small cavity near the top; also with clavate and verrucate ornamentation. Interspinal area faintly microreticulate. Exine 2.3 [Am thick; sexine thinner than nexine toward the colpi margin; amb circular. Sechium hintonii (P.O. Wilson) C. Jeffrey (Plate IV: 1-6) Eumonad, isopolar, radiosymmetric; suboblate (52.6 x 68.8 [im), stephanocolpate (9-10 colpi). Tectate; spiniferous, conical spines 5.4-6.3 [im long, with a small cavity near the top. Interspinal area faintly microreticulate. Exine 2.5 um thick; sexine slightly thicker than nexine; amb circular. 114 ALVARADO, LIRA-SAADE & CABALLERO Plate II Comparative SEM illustrations of the pollen grains of nine of the taxa included in the study. 1, Sechium edule (wild); 2, 5. pittieri; 3, 5. tacaco; 4, S. talamancense; 5, S. villosum; 6, Frantzia panamensis; 7, F. venosa; 8, Parasicyos maculatus; 9, Microsechium helleri. (bar scale = 5 m. Sechium pittieri (Cogn.) C. Jeffrey (Plate IV: 7-10) Sechium tacaco (Pittier) C. Jeffrey (Plate IV: 11-14) Eumonad, isopolar, radiosymmetric; suboblate (48.5 x 64.4 urn), stephanocolpate (8-9 colpi). Tectate; spiniferous, conical spines 5.6-7.1 um long, with a small cavity near the top. Interspinal area faintly microreticulate. Exine 2.8 urn thick; sexine slightly thicker than nexine; amb circular. Eumonad, isopolar, radiosymmetric; oblate-spheroidal (78.3 x 87.6 um), stephanocolpate (8-9 colpi). Tectate; spinifer- ous, conical spines 6.1-7.2 um long, with a small cavity near the top. Interspinal area microreticulate. Exine 3.0 um thick; sexine slightly thicker than nexine; amb circular. PALYNOLOGICAL EVIDENCE FOR TAXONOMIC LIMIT OF SECHIUM 115 Plate III Sechium composition. 1, equatorial view, high focus; 2, equatorial view, cross section; 3, polar view, cross section; 4, polar view, high focus; 5, apocolpium; 6, polar view, cross section. S. edule (wild). 7, equatorial view, high focus; 8, 9, polar view, high focus; 10, apocolpium; 11-13, polar view, cross section. S. edule (cultivated). 14, equatorial view, high focus; 15, 16, 19, polar view, cross section; 17, 18, apocolpium. (bar scale = 10 urn). Sechium talamancense (Wunderlin) C.Jeffrey (Plate IV: 15-17) Eumonad, isopolar, radiosymmetric; suboblate (46.3 x 60.8 p,m), stephanocolpate (8-9 colpi). Tectate; spiniferous, conical spines very variable in size (3.6-8.1 um), with a small cavity near the top. Interspinal area micro reticulate. Exine 3.5 um thick; sexine thicker than nexine; amb circular. Sechium villosum (Wunderlin) C. Jeffrey (Plate V: 1-4) Eumonad, isopolar, radiosymmetric; oblate (54.3 x 73.7 um), stephanocolpate (8-9 colpi). Tectate; spiniferous, conical spines very long (7.2-9.9 [Am), with a small cavity near the top. Interspinal area reticulate. Exine 2.5 um thick; sexine as thick as nexine; amb circular. 116 ALVARADO, LIRA-SAADE & CABALLERO Plate IV Sechium hintonii. 1, equatorial view, high focus; 2, equatorial view, cross section; 3, 5, polar view, high focus; 4, apocolpium; 6, polar view, cross section. 5. pittieri. 7, equatorial view, high focus; 8, equatorial view; cross section; 9, polar view, high focus; 10, polar view, cross section. 5. tacaco. 11, equatorial view, high focus; 12, equatorial view; cross section; 13, polar view, cross section; 14, polar view, high focus. 5. talamencense. 15, equatorial view, high focus; 16, polar view, apocolpium; 17, polar view, cross section, (bar scale = lOum). Frantzia panamensis Wunderlin (Plate V: 5-8) Eumonad, isopolar, radiosymmetric; oblate-Spheroidal (73.0 x 81.2 urn), stephanocolpate (8-9 colpi). Tectate; spinifer- ous, conical spines 4.9-5.8 um long, with a small cavity near the top. Interspinal area faintly microreticulate. Exine 2.7 um thick; sexine slightly thicker than nexine; amb circular. Frantzia venosa L.D. Gomez (Plate V: 9-12) Eumonad, isopolar, radiosymmetric; oblate-spheroidal (69.3 x 76.1 um), stephanocolpate (8-9 colpi). Tectate; spinifer- ous, conical spines 4.4-5.7 um long, with a small cavity near the top. Interspinal area faintly microreticulate. Exine 2.8 um thick; sexine as thick as nexine; amb circular. PALYNOLOGICAL EVIDENCE FOR TAXONOMIC LIMIT OF SECHIUM 117 Plate V Sechium villosum. 1, equatorial view, high focus; 2, polar view, cross section; 3, polar view, high focus; 4, polar view, cross section. Frantziapanamensis. 5, equatorial view; cross section; 6, mesocolpium detail; 7, polar view, high focus; 8, polar view, detail of spines, cross section. F. venosa. 9, 10, equatorial view; 11, polar view, high focus; 12, polar view, cross section. Parasicyos maculatus. 13, equatorial view, high focus; 14, equatorial view; cross section; 15, 16, polar view, cross section, detail of spines, (bar scale = lOum). Parasicyos maculatus Dieterle (Plate V: 13-16) Eumonad, isopolar, radiosymmetric; oblate-spheroidal (40.1 x 41.7 um), stephanocolpate (8-9 colpi). Tectate; spinifer- ous, conical spines 4.1^4.9 mm. Interspinal area reticulate. Exine 2.2 urn thick; sexine thicker than nexine; amb circular. Microsechium helleri (Peyr.) Cogn. (Plate VI: Eumonad, pseudoisopolar, asymmetric; oblate-spheroidal (68.9 x 72.8 urn); heterocolpate (colpi 7 of different sizes). Tectate; spiniferous, conical spines 4.5-5.7 um long. Interspi- nal area microreticulate. Exine 2.3 um thick; sexine thicker than nexine; amb circular. 118 ALVARADO, LIRA-SAADE & CABALLERO Plate VI Microsechium helleri. I, equatorial view, high focus; 2, equatorial view, cross section; 3, polar view, high focus; 4, polar view, cross section. Pterosicysos laciniatus. 5, equatorial view, high focus; 6, equatorial view; cross section, spines detail; 7, polar view, high focus; 8, polar view, cross section. Sechiopsis triquetra. 9, equatorial view, high focus; 10, equatorial view; cross section; II, polar view, high focus; 12, polar view, cross section. Sicyos motozintlensis . 13, equatorial view, high focus; 14, equatorial view; cross section; 15, polar view, high focus; 16, polar view, cross section, (bar scale = lOum). PALYNOLOGICAL EVIDENCE FOR TAXONOMIC LIMIT OF SECHIUM 119 Pterosicyos laciniatus Brandegee (Plate VI: 5-8) Eumonad, isopolar, radiosymmetric; oblate-spheroidal (58.1 x 60.3 urn), stephanocolpate (8-9 colpi). Tectate; spinifer- ous, conical spines 3.6-5.1 um long. Interspinal area microre- ticulate. Exine 2.5 um thick; sexine thicker than nexine; amb circular. Sechiopsis triquetra (Ser.) Naudin (Plate VI: 9-12) Eumonad, isopolar, radiosymmetric; suboblate (52.2 x 60.5 urn), stephanocolpate (9-10 colpi). Tectate; spiniferous, conical spines 3.6-4.7 um long, with a small cavity near the top. Interspinal area microreticulate. Exine 2.5 um thick; sexine thicker than nexine; amb circular. Sicyos motozintlensis Lott & Fryxell (Plate VI: 13-16) Eumonad, isopolar, radiosymmetric; oblate-spheroidal (59.7 x 61.7 jim), stephanocolpate (8-9 colpi). Tectate; spinifer- ous, conical spines 3.1-4.5 um long. Interspinal area microre- ticulate. Exine 2.1 um thick; sexine as thick as nexine; amb circular. 2. Numerical Taxonomic Study A. Cluster Analysis As can be seen from the phenogram resulting from the cluster analysis (Fig. 1), two main clusters can be observed. The first is made up of eight taxa, all belonging to Sechium and Frantzia; they are Sechium compositum, the wild forms of S. edule, S. hintonii, S. pittieri, S. tacaco, S. talamancense, Frantzia panamensis and F. venosa. The second cluster includes only three species, two of which (Sechiopsis triquetra and Pterosicyos laciniatus) belong to the two closely related genera, which Kearns (in press) has proposed should be united under Sechiopsis. The other species in this cluster is Sicyos motozintlensis, a species pertaining to a genus which has been considered as not closely related to either Sechiopsis or Pterosicyos. Nevertheless, the phenogram shows the above-mentioned species as palynologically similar to 5. triquetra. The four remaining taxa (cultivated Sechium edule, S. villosum, Microsechium helleri and Parasicyos maculatus), are clearly apart from the two previously described groups. In the first cluster, two subgroups can be observed. The first places taxa belonging to the two sections of Sechium sensu lato close to each other. This subgroup is formed by Sechium compositum, wild 5. edule and 5. hintonii (all of section Sechium) and S. pittieri (of section Frantzia), which is placed next to 5. talamancense (of section Sechium). The second subgroup is formed by S. tacaco (of section Sechium), and Frantzia panamensis and F. venosa, both of which, if transfered to Sechium, would fall under section Frantzia, for they present 'cushion like' nectaries. Of the 15 taxa studied, Frantzia panamensis and F. venosa are the two most closely related species, although some macromorphological features (e.g. inflorescence type), readily set them apart. B. Ordination Techniques As can be seen from Fig. 2, the results of PCO are in general consistent with those of the Cluster Analysis. Furthermore they provide a sharper definition of the different clusters described above. The results of PCO indicate that the charac- ters which contribute the most to cluster formation are the polar/equatorial ratio and the type of spine apex in the first principal coordinate, while in the second principal coordi- nate, the most important characters are the length of the polar axis, and the number of colpi (Table 5). The character with higher loadings in the third principal coordinate is the type of grain ornamentation. In contrast with the palynologi- cal characters, none of the floral characters contributed significantly to the variation. The first three principal coordi- nates explain 70.7% of the variation (Table 5). The representation of the OTUs in a bidimensional space of characters (Fig. 2.) shows that all the species of Sechium and Frantzia are closer to one another than to the majority of the species of the other genera. In the cluster formed by these species it is possible to identify subgroups that are in agree- ment with the interspecific relationships proposed elsewhere. Thus, wild Sechium edule and S. compositum are very close to each other. This is consistent with the chemical evidence of kinship between these two species found by Newstrom (1986, 1989). Likewise Frantzia venosa and F. panamensis are the 2.0 1.5 1.0 0.5 0.0 SCO SEW SHI SPI STL STA FPA FVE PLA SIM SET SVI SEC PMA MHE Fig. 1 Phenogram showing the similarity between the taxa of Sechium s. I. and its allies (Cophenetic Correlation Coefficient = 0.849). Taxa acronyms = SCO (Sechium compositum); SEC (cultivated S. edule); SEW (wild S. edule); SHI (5. hintonii); SPI (5. pittieri); STA (S. tacaco); STL (S. talamancense); SVI (5. villosum); FPA (Frantzia panamensis); FVE (F. venosa); MHE (Microsechium helleri); PMA (Parasicyos maculatus); PLA (Pterosicyos laciniatus); SET (Sechiopsis triquetra); SIM (Sicyos motozintlensis). 120 ALVARADO, LIRA-SAADE & CABALLERO 1.2 0.6 » o.o- ex o. •o 8 -0.6 -1.2 J MHE STA FPA FVE SEW SCO SEC SIM PLA SPI SHI STL SVI SET PMA -1.2 -0.6 0.0 First principle coordinate 0.6 1.2 Fig. 2 Representation of; the taxa of Sechium s. I. and its allies in a bidimensional space of characters resulting form a Principal Coordinate Analysis. Taxa acronyms = SCO (Sechium composition); SEC (cultivated 5. edule); SEW (wild 5. edule); SHI (5. hintonii); SPI (5. pittieri); STA (5. tacaco); STL (5. talamancense); SVI (5. villosum); FPA (Front zia panamensis) ; FVE (F. venosa); MHE (Microsechium helleri); PMA (Parasicyos maculatus); PLA (Pterosicyos laciniatus); SET (Sechiopsis triquetra); SIM (Sicyos motozintlensis). Table 5 Variation explained by the three first principal coordinates and the palynological characters with higher loading in each principal coordinate. Variation explained (%) Characters 1st principal coordinate 39.46 2nd principal coordinate 18.05 3rd principal coordinate 13.24 Polar/equatorial ratio and apex of spines Polar axis and number of colpi Grain ornamentation two most closely related species and make up a very well- defined group, though, in this case, they lie somewhat apart from 5. tacaco. The grouping of the species of Sechium according to the two Sechium sections defined by Jeffrey (1978), could not be observed with this type of analysis, since species belonging to different sections appear very close together, as in the group formed by Sechium hintonii, S. talamancense, S. villosum (all of section Sechium) and S. pittieri (of section Frantzid). It is important to notice the position of the two cultivated taxa (5. edule and 5. tacaco) in the graph resulting from PCO (Fig. 2). These two taxa, particularly 5. edule, lie far apart from the rest of the species of the Sechium-Frantzia group. We do not have a clear explanation for this. However, some answers may be found in the extreme average values that these species have for characters which contribute the most to the variation, such as the polar/equatorial axis ratio and the ornamentation in S.edule, as well as the length of the polar axis in 5. tacaco. In the case of S. tacaco, the most notable difference in its pollen grains is the length of the polar axis (78.3 urn), which was the longest recorded in this study (Table 4). As for cultivated Sechium edule, the polar/ equatorial ratio of its pollen grains (0.73) was one of the two lowest recorded. Furthermore, the ornamentation observed in the grains of this taxon (spiniferous- clavate- microreticulate) was unique among all the taxa studied. These kinds of palynological differences have also been found between cultivated and wild species of Cucurbita by Ayala- Nieto et al. (1988). This suggests that pollen of Cucurbitaceae is a structure which has been unconsciously modified through the process of selection under domestication by humans. Related to the species of Sechiopsis and Pterosicyos, the results obtained from both Cluster Analysis and PCO show that these two genera are relatively close to one another, occupying an intermediate position between the Sechium- Frantzia group and the rest of the genera (Microsechium, Parasicyos and Sicyos), though now clearly separated from Sicyos motozintlensis. On the other hand, Microsechium, Parasicyos and Sicyos, are definitely distant both from one another and from the two main groupings described above (Sechium-Frantzia and Pterosicyos-Sechiopsis). CONCLUSIONS The segregation of the subtribe Sicyinae proposed by Jeffrey (1990) was partially based on a series of palynological attributes such as the 7-10 colporate and the spiniferous pollen grain characters. The pollen morphology observed from the species studied here is consistent with Jeffrey's criterion. However, as seen from this study, the pollen morphology is highly variable at the level of genera and species within the above-mentioned subtribe. The more vari- PALYNOLOGICAL EVIDENCE FOR TAXONOMIC LIMIT OF SECHIUM 121 able pollen characters are the polar/equatorial index, the form of the apex of the spines, the length of the polar axis, the number of colpi and the grain ornamentation. The results from the numeric taxonomic study give some support to the idea that pollen morphology is significant in the delimitation of genera within the Cucurbitaceae. These results also indicate a strong palynological relationship between the species of Sechium sensu lato and Frantzia, since both cluster analysis and ordination techniques showed these species grouping together and separated from the other species and genera studied here. This palynological relationship supports Jeffrey's (1978) proposal of including Frantzia, Polakowskia and Ahzolia within the limits of Sechium. However, the splitting of this genus into sections based on floral characters by that author was not evident. The numerical taxonomic analysis has shown that the clustering of the species does not reflect such sections, even though the analysis included important floral characters. Nevertheless, it is clear that the final assessment of the generic limits of Sechium and its relatives should be the result of an integrative study involving a larger number of charac- ters as well as a variety of systematic approaches. ACKNOWLEDGEMENTS. This paper forms part of the project Taxo- nomic and Ecogeographic Studies in Cucurbitaceae of Latin Amer- ica', sponsored by the International Board for Plant Genetic Resources (IBPGR), and conducted by the second author in the Herbario Nacional de Mexico, Instituto de Biologia, U.N.A.M. We wish to thank the authorities of Herbario Nacional de Mexico, especially Alfonso Delgado and Patricia Davila, and Instituto Nacio- nal de Antropologfa e Historia de Mexico, especially Mario Aliphat, for the facilities provided to carry out this study. To Maria del Rosario Garcia for the procurement of specimen loans, and to the curators of BM, CHAPA, GH, K, MICH, MO and XAL for the loans of the specimens entrusted to their care. Thanks to the Laboratorio de Micromecanica, Instituto de Ingenieria, U.N.A.M., especially Ricardo Peralta, Armando Peralta and Orlando Palacios for their help to use the Scanning Electron Microscope. Robert Bye (Jardin Botanico del Instituto de Biologia, U.N.A.M.), Fernando Chiang, Patricia Davila, Oswaldo Tellez (MEXU), Daniel Debouck (IBPGR-Roma), Stephen Blackmore, Rob Huxley and David Sutton (BM) read and commented on the manuscript. The latter two also assisted in the English of the manuscript. The second author thanks Jose Carmen Soto (Colegio de Ciencias y Humanidades, U.N.A.M.), Alberto Reyes (MEXU), Rafael Ocampo (CATIE), Carlos Astorga (IBPGR-CATIE) and Abdenago Brenes (Universidad Nacional Heredia Costa Rica), for assistance in the field. REFERENCES Aleshina, L.A. 1964. On the pollen grains of the Cucurbitaceae (in Russian). Bot. Zhurn. SSSR 49(12): 1773-1776. Ayala-Nieto, M., Lira-Saade, R. & Alvarado, J.L. 1988. Morfologia polinica de las Cucurbitaceae de la Peninsula de Yucatan. Pollen et Spores 30(1): 5-28. Dieterle, J.V.A. 1976. Cucurbitaceae. In: Flora of Guatemala. Fieldiana Bot. 24(11): 306-395. Dunn, G. & Everitt, B.S. 1982. An Introduction to Mathematical Taxonomy. Cambridge. Erdtman, G. 1943. An Introduction to Pollen Analysis. New York. Erdtman, G. 1952. Pollen Morphology and Plant Taxonomy. Angiosperms. New York. Faegri, K & Iversen, J. 1964. Text Book of Pollen Analysis. Copenhagen. G6mez, L.D. & Gome/ J. 1984. Plantae Mesoamericanae Novae. IX. Phytolo- gia 53(6): 447-448. Jeffrey, C. 1964. A note on pollen morphology in Cucurbitaceae. Kew Bull. 17: 473-477. 1978. Further notes on Cucurbitaceae. IV. Some New World taxa. Kew Bull. 33: 347-380. 1980. A review of the Cucurbitaceae. /. Linn. Soc. Bot. 81: 233-247. 1990. An outline classification of the Cucurbitaceae. In: D.M. Bates, R.W. Robinson & C. Jeffrey (Eds). Biology and Utilization of the Cucurbi- taceae: 449-463. Ithaca. Kearns, D.M. A review of the genus Sechiopsis. Syst. Bot. in press. Kremp, G.O.W. 1965. Morphologic Encyclopedia of Palynology . Tucson. Marticorena, C. 1963. Material para una monograffa del polen de Cucurbita- ceae. Grana Palynologica 4(1): 78-91. Newstrom, L. 1986. Studies in the origin and evolution of Sechium edule (Jacq.) Sw. Ph.D. Thesis. University of California at Berkeley. 1989. Reproductive biology and evolution of the cultivated chayote Sechium edule: Cucurbitaceae. In: J.H. Bock & Y.B. Linhart (Eds). Evolutionary Ecology of Plants: 491-509. Boulder. 1990. Origin and evolution of chayote, Sechium edule. In: D.M. Bates, R.W. Robinson & C. Jeffrey (Eds). Biology and Utilization of the Cucurbi- taceae: 141-149. Ithaca. Page, J.S. & Jeffrey, C. 1975. A palynotaxonomic study of African Peponium. Kew Bull. 30: 495-502. Pittier, H. 1910. New or noteworthy plants from Colombia and Central America. Contr. U.S. Natl. Herb. 13: 118-131. Rangaswami, K. 1976. Palynological highlights of Cucurbitaceae. In: P.K.K. Nair (Ed.). Advances in Pollen-Spore Research. Vol.11 54-59. New Delhi. Rodriguez- Jimenez, C. & PaIacios-Chavez,R. 1991. Nota sobre la variation morfo!6gica en las flores masculinas de Microsechium helleri (Peyr.) Cogn. (Cucurbitaceae). Palynologica et Palaeobotanica. 3 (1): 99-106. Rohlf, F.J. 1989. NTSYS-pc. Numerical Taxonomy and Multivariate Analysis System. Version 1.50. New York. Romesburg, H. C. 1984. Cluster Analysis for Researchers. Belmont, California. Singh, S & Singh,!). 1990. Palynology of the Indian Cucurbitaceae. In: D.M. Bates, R.W. Robinson & C. Jeffrey (Eds.) Biology and Utilization of the Cucurbitaceae: Ithaca. Wilson, P.G. 1958. Microsechium hintonii. In: Contributions to the Flora of Tropical America: LXIII. Kew Bull. 13: 161. Wunderlin, R.P. 1976. Two new species and a new combination in Frantzia (Cucurbitaceae). Brittonia 28(2): 239-244. 1977. A new species of Frantzia (Cucurbitaceae) from Panama. Bull. Torrey Bot. Club 104(2): 102-104. Bull. Br. Mus. nat. Hist. (Bot.) 22(2): 123-146 Issued 26 November 1992 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 3. Genera H-K JAMES H. PRICE, DAVID M. JOHN AND GEORGE W. LAWSON Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS Synopsis 123 Introduction 1 23 Species list 124 Acknowledgements 137 References in numerical order 137 References . 140 SYNOPSIS. This paper assembles and, so far as is possible without extended field and herbarium studies, examines critically the validity of records of marine and brackish-water Rhodophyta (Florideae) for the western coast of tropical Africa. The whole mainland coastline from the northern boundary of Western Sahara southwards to the southern boundary of Namibia, the oceanic islands from the Salvage Islands southwards to Ascension and St Helena, and all islands close to the African mainland coast are included in the area covered. Each species entry includes all traced records for the species, the names which have previously been applied to it for the area, and additional comments or evaluation, as necessary. Comments are also provided at generic or generic group levels in very complex cases. INTRODUCTION The area dealt with in this part of the work is identical with that covered in parts published previously (Lawson & Price, 1969; Price, John & Lawson, 1978, 1986, 1988; John, Price, Maggs & Lawson, 1979). Country names employed and their earlier equivalents, and the names of island groups, are either listed in the legend or both listed and shown on the map in Fig. 1. Genera with the initial letter H-K and constituent species are listed in alphabetical order. Each main entry consists of: (i) The major bold heading, representing the currently- accepted name and authorities. (ii) Subsidiary italicised headings at intervals within the entry. These are in square brackets and essentially subdivide the overall entry. They represent the differ- ent ways in which the species has been referred to throughout in past publications for the area. Incorrect citations from past literature have been maintained in these subsidiary heads so that there shall be no doubt as to which record we attribute to which species or lower taxon level; only when clarification was required for comprehension have changes been made in subhead citation, in which case explanation is given in interme- diary or terminal notes. (iii) The distributional data, with countries and island groups arranged in a single alphabetical order. More generalised but relevant statements of distribution follow the specific country list. Complete distribution patterns require a scan of records established under all names by which a species is known for this or adjacent areas. Hence, generalised distribution statements are included verbatim since it is not always clear for precisely which countries within the area they establish records. References are presented in two ways: (a) as a numbered list (p. 137) to allow inclusion of manuscript and expedition sources, and (b) as a list of full references in alphabetical order. Numbers within parentheses after the geographical names refer to cor- responding numbers in the references. References rele- vant only in previous parts of this series are omitted here and numbers are therefore not fully standardised between the present and past parts. (iv) Additional qualifying notes appear below whole entries or individual parts of entries to which they specifically refer. In these notes, references containing species records consist of authors' names, followed by the reference number in the terminal list and, where appro- priate, the relevant page numbers after a colon. Other references consist of authors' names, date of publica- tion and sometimes page numbers after a colon. 124 PRICE, JOHN & LAWSON Fig. 1. The coastline of tropical west Africa and the offshore islands. 1, Salvage Islands; 2, Canary Islands; 3,*Western Sahara [=former Spanish Sahara, Spanish West Africa] (includes the often quoted Rio de Oro, the southern region of the country, but excludes Ifni); 4, Mauritanie; 5, Senegal; 6, Gambia; 7, Guinea-Bissau [-Portuguese Guinea]; 8, Guinee; 9, Sierra Leone; 10, Liberia; 11, Cote d'lvoire; 12, Ghana; 13, Togo; 14, Benin [=Dahomey]; 15, Nigeria; 16, Cameroun; 17,** Bioko [=Macias Nguema Biyogo, Fernando Poo]; 18, Principe; 19, Sao Tome; 20,** Equatorial Guinea [^Spanish Guinea]; 21, Gabon; 22,*** Republic of the Congo; 23, Cabinda; 24, Zaire [=Congo Republic]; 25, Angola; 26, Namibia [=South West Africa]; 27, Ascension Island; 28, Saint Helena; 29, Pagalu [=Annobon]. The Cape Verde Islands, which lie immediately to the west of Dakar (Senegal), have been omitted from this map but are included in the species list that follows. * The former colony of Spanish Sahara no longer officially exists, the territory it once covered being divided, by agreement between Morocco and Mauritanie. The effective date of the division, Spain concurring, was 28 February 1976, although guerilla opposition delayed matters until a formal agreement on 14 April 1976. The attempt to maintain the territory as the Democratic Sahara [Saharan] Arab Republic has apparently entered the 'realm of myth' (Gretton, 1976). The authors' citation terminology is maintained throughout the records. ** Nos 17 (Bioko) and 20 (Spanish Guinea, = Rio Muni) on the original map (part I) are now jointly administered as Equatorial Guinea. Bioko is entered separately, where appropriate, in the species list. *** Loango, a name much used by earlier collectors such as Welwitsch, was formerly a coastal region of West Africa. Its application appears to have included much of the coastline of the Republic of the Congo (22), as well as of Cabinda (23) and Zaire (24). Because by far the longest and rockiest part of the Loango coast lies now within the Republic of the Congo we have attributed all marine algal records from Loango to the Congo. Species nomenclature has been revised as far as possible and the complete author citation is given for each currently- accepted combination. The subsidiary italicised headings and any other discarded combinations that require reference are included as cross-referencing entries to the currently- accepted names in the overall list. Without extended field and herbarium studies, the treatments presented here are essen- tially preliminary. Critical updating of the overall text is kept firmly in mind for the whole series and we would appreciate notification of any detected errors and omissions from any of the parts. SPECIES LIST Halarachnion ligulatum (Woodward) Kiitzing Canaries (38D; 70; 191; 226; 227; 232B; 392; 584; 598). RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 'From Great Britain southwards to the Canary Islands' (70). [As Halymenia (Halarachnion) ligulata (Woodward) Kiitzing] Canaries (439). Note. For interrelationship in life history between Halarachnion ligulatum and the crustose tetrasporophyte Cruoria rosea, see Boillot 1965, 1972). Data established by Boillot have subsequently been confirmed by Kornmann & Sahling (1977) and Maggs (1983). For a detailed study in Halarachnion ligulatum of development, vegetative and reproductive, see Knauss & Hommersand (1989), where also are discussed its grouping in Furcellariaceae and characterization/ analysis of this family. Halarachnion sp. For possible Canaries and Senegal records, see Halymenia spp. Halichrysis depressa (J. Agardh) Schousboe in Bornet Bioko (346; 350; 586). Ghana (350; 377; 586). '. . . in the Mediterranean and tropical parts of the eastern Atlantic' (350; 586). Tropical Africa (N. Gambia — Congo river)'(598). [As Halichrysis depressa (J. Agardh) Schmitz] '. . . Atlantischer Ozean: Nordwestafrikanische ktiste. . .' (499). [As Halichrysis depressa (Schousboe) Schmitz] '. . . in warmeren Teile des atlantischen Oceans. . .'(507). Note. Plants with similar coloration, habit and vegetative struc- ture have been found in Cameroun and Gabon. Gabon material has zonate tetraspores and was tentatively referred to Meristotheca (295). Sterile Ghanaian plants resemble M. coacta Okamura (299). Recent collections of cystocarpic plants have all been found to be Halichrysis depressa (350; 586), save for the record (Heincke Expedition, 1991) cited under Halichrysis peltata (q.v.). Halichrysis peltata (W.R. Taylor) P. Huve & H. Huve Canaries (635). Haliptilon Johansen & Womersley (1986: 562) commented that: 'The systematics of Haliptilon on a world-wide basis requires further study'. Haliptilon and Jania are more closely related than either is to Corallina — see the generic note to Jania. See also Johansen & Womersley (1986: 563) who presented a table of the characteristics distinguishing Haliptilon from Corallina in southern Australia. Haliptilon cubense (Montagne ex Kiitzing) Garbary & Johansen [As Corallina cubensis (Montagne) Kiitzing] Canaries (13; 128 A; 227; 306B; 582). Cape Verde Islands (191; 500). [As Jania cubensis Montagne in Kiitzing] Cape Verde Islands (259). [As Jania cubensis Kiitzing] Cape Verde Islands (145). [As Jania cubensis Montagne] Cape Verde Islands (260). [As Corallina (Jania) cubensis Montagne in Kiitzing] Cape Verde Islands (38). Haliptilon squamatum (Linnaeus) Johansen, L. Irvine & Webster Canaries (Herb. BM; 541) Western Sahara (349). [As Corallina squamata Ellis et Solander] Canaries (25; 134; 239; 283; 497; 499; 582). Senegal (50). 125 Western Sahara (33; 222). '. . . a Britannia meridionali usque ad Insulas Canarienses. . .' (25; 134). '. . . de PAngleterre au Rio de Oro. . .' (33; 222). '. . . depuis le partie sud de 1'Angleterre jusqu'aux Canaries. . .' (283). '. . .In Atlantischen Ozean von der sudenglischen Kiiste bis zu den Kanaren. . .' (497; 499). '. . . Nordwestafrika. . .' (499). [As Corallina squamata Park.] Canaries (254; 305). Note. Bodard (50: 85) stated regarding his Senegal material 'C. squamata. . . n'est d'ailleurs a notre avis qu'une forme de C. mediterranea . . '. According to Afonso-Carrillo et al. (582), the presence of this species in the Canaries is doubtful. Haliptilon subulatum (Ellis & Solander) Johansen Ghana (350; 586). Liberia (350; 586). Mauritanie (349; 598; 624). 'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598). Tropical Africa (N. Gambia — Congo river)' (598). [As Haliptylon subulata (Ellis & Solander) Johansen] Liberia (129). Note. Johansen & Womersley (1986: 562) stated: 'The diagnostic characteristics of H. subulatum are unknown, and as the type specimen is lost this species must be considered a nomen nudum. . .'. It seems highly likely then that if this is a good taxon it is more widespread throughout the list area than the few records would suggest. Haliptilon virgatum (Zanardini) Garbary & Johansen Canaries (387; 439; 448; 541; 598; 604; 605). Salvage Islands (598). Tropical Africa (N. Gambia — Congo river)' (598). Note. The epithet is rendered virgata by Garbary & Johansen (1982). They examined the ?type of Corallina virgata Zanardini and found it to possess surface morphology and trichocytes of a type similar to Jania rubens. See also notes to Corallina granifera Ellis & Solander. Haliptilon cubense (Montagne ex Kiitzing) Garbary & Johansen may also be implicated in the background to H. virgatum; see Johansen & Womersley (1986: 562). According to Prud'homme van Reine (pen. comm., 19/11/91), from discussions with W. Johansen and J. Afonso-Carrillo it appears probable that material of Corallina lobata Lamouroux (q.v., Price et al. 1986: 75) from the Canaries actually represented Haliptilon virgatum. Haliptilon spp. West Africa (290). Halodictyon mirabile Zanardini Canaries (13; 38D; 71; 177; 190; 191; 227; 232B; 350; 556; 584; 586; 598; 604; 605). Cape Verde Islands (598). Salvage Islands (38B; 38D; 556; 598). Sierra Leone (30; 350; 586). Note. Aleem (1978) reported this plant growing on hydroids in the lower littoral at one location on the Sierra Leone peninsula. Although known from the islands, this is the first mention of the species from the mainland of West Africa and, until confirmed, the record must remain doubtful. Weisscher (1983: 66) indicated, with doubt, a record, presumably secondary, for Morocco. Halophytis incurvus See Halopitys incurvus (Hudson) Batters. Halopithys incurvans (Hudson) Batters See Halopitys incurvus (Hudson) Batters. 126 PRICE, JOHN & LAWSON Halopitys incurvus (Hudson) Batters Canaries (13; 38D; 128A; 191; 227; 232B; 306B; 375; 392; 517; 584; 598; 621). Mauritanie (38D; 349). '. . . Atlantique (de 1'Angleterre a la Mauritanie). . .' (33). '. . .Atlantique nord, du sud de 1'Angleterre aux Canaries. . .' (190). '. . . Atlantique nord jusqu'en Mauritanie. . .' (222). '. . . British Isles and southward to W. Africa. . .'(375). '. . . mediterraneo desde el sur de Inglaterra a Canarias' (517). 'Subtropical Africa [Senegal (N. of Gambia), Mauritania, Former W. Sahara]' (598). [As Halophytis incurvus sine auctorum] Canaries (128 A) [As Halopithys incurvans (Hudson) Batters] Canaries (227; 238). [As Halopithys pinastroides Kiitzing] Canaries (89; 547). '. . . Du sud de la Grande-Bretagne aux Canaries. . .' (89). [As Halopithys pinastroide] Canaries (237). [As Halopitys pinastroides (Gmelin) Kiitzing] Canaries (70; 71; 133; 236; 252; 322). Mauritanie (252). '. . . von Siidengland bis zu den Canarischen Inseln. . .' (179). '. . . From south of England down to the Canary islands. . .' (71). '. . . warmeren Teile des atlantischen Oceans. . .' (501). [As Halopithys pinastroides] Canaries (2). '. . . Atlantique nord jusqu'en Mauritanie. . .' (222). [As Lophura episcopalis (Montagne) Kiitzing] Canaries (318; 323). [As Rhodomela episcopalis Montagne] Canaries (402; 407). [As Rhodomela pinastroides Agardh] Canaries (44). [As Rhodomela pinastroides (Gmelin) C. Agardh var. episco- palis Montagne] Canaries (401; 402). [As Rytiphlaea episcopalia (Montagne) Endlicher] Canaries (26). [As Ryt iphloea pinastroides C. Agardh] Canaries (37; 401; 439). [As Rytiphlaea pinastroides (Gmelin) C. Agardh] '. . . in oceano atlantico calidiore ab insulis Canariensibus usque ad littora meridianalia Angliae. . .' (26). '. . . Atlantischen Ocean (von den Kanaren bis England). . .' (37). [As Rytiphlaea pinastroides (Turner) J. Agardh] Canaries (258). [As Fucus lycopodioides Flor. Dan.] Canaries (90). Note. There have been so many variations, accidental and pur- poseful, of spelling in generic and specific names that the majority have been ignored unless they were likely to lead to misunderstand- ing or nomenclatural confusion. For details of the attribution of Fucus lycopodioides here, see the earlier entry (Price et al., 1986: 92-93) under that name. Kiitzing himself remarked (323: 15, no. 3535) below the figure legends for Lophura episcopalis that the alga belonged to the genus Halopithys [sic!], very close to and perhaps not specifically distinct from H. pinastroides. Essentially similar com- ments had earlier (1841) been made by Montagne (401: 153-154) in considering his var. episcopalis of Rhodomela pinastroides, based on material sent under the name Rhodomela lycopodioides by Despreaux to Webb. Halopithys pinastroides sine auctorum See Halopitys incurvus (Hudson) Batters. Haloplegma duperreyi Montagne '. . . ad oras Africae australis et austro-occidentales (sec. Sender). . .' (24; 27; 133). Note. It is not clear whether the south west coast here relates to the west coast of South Africa or the coast of South West Africa (now Namibia). In view of the lack of other records and general Indo- Pacific distribution probably the former is true and the entry there- fore not relevant. Included for completeness only. Halurus equisetifolius (Lightfoot) Kiitzing Canaries (565 A; 598; 605). Mauritanie (33; 349; 529). 'de Inglaterra a Mauritania' (619; 622). '. . . Atlantique (de 1'Angleterre a la Mauritanie). . .' (33) '. . . Atlantique nord: des cotes anglaises jusqu'au cap Blanc (Mauritanie). . .' (222). [As Halurus sp.] Mauritanie (245B). Halymenia This genus is imperfectly known and apparently highly poly- morphic. Despite work by many authors over the last three decades there is still an insufficiently broad and critical database for the establishment of a satisfactory working taxonomy. The entries given here are therefore essentially preliminary and biogeographic conclusions are unwarranted as yet. Bodard (55: 14) commented on the difficulty of comparing herbarium materials in which resuscitation of structure is imperfect because of inseparable adjacent wall layers and faulty preservation in liquids, especially of the secretory cells. The wide distribution of forms referred to the genus in warm and temperate seas further complicates assign- ment of records. Recent treatment by Schneider & Searles (642: 276-282) should be consulted for details of some of the difficulties encountered, especially between the genera Haly- menia and Sebdenia (642: 269-270). Halymenia actinophysa Howe Ghana (290; 292; 299; 300; 350; 376; 567; 586). Mauritanie (624). '. . .in tropical parts of the Pacific and Atlantic Oceans' (350; 586). Tropical Africa (N. Gambia — Congo river)' (598). Halymenia agardhii De Toni Ghana (292; 299; 350; 376; 586). Senegal (50; 55; 56; 59; 399; 598). Tropical Africa (N. Gambia — Congo river)' (598). '. . . widespread in warm temperate and tropical seas' (350; 586). Note. See Cordeiro-Marino (108: 59) on the possible conspecific- ity of this species with Halymenia polydactyla B0rgesen (from India) [= Sebdenia polydactyla (B0rgesen) Balakrishnan]. See also the generic note above regarding data in Schneider & Searles (642) on these two potentially confusing genera. They indicated (p. 282) that Parkinson (1980) showed Halymenia agardhii to be Sebdenia flabel- lata, causing terete plants of Halymenia from the USA, Bermuda and the Caribbean to be referred thereafter to Halymenia trigona (Clem- ente) G. Agardh. Terete Halymenia spp. world-wide have been found (Codomier, 1974; Millar, 1990) to be conspecific with H. RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 127 trigona, described from Atlantic Spain. Cremades & Perez-Cirera (1990: 490) indicated the correct name of H. trigona to be Halymenia elongata C. Agardh. Although Schneider & Searles (642: 282) stated that H. elongata (as H. trigona) can be easily distinguished from the similar S. flabellata (the latter bears obvious gland-like cells on inner medullary stellate ganglia; H. elongata has no gland-like cells), both taxa could have been involved in the establishment of these earlier western African records under H. agardhii and we have not attempted definitive allocation to either. Restrictions on time have prevented us from examining actual material involved. Halymenia bermudensis Collins & Howe Salvage Islands (38B; 556; 598). Note. See Wynne (1986) on the authorities for this taxon. Schneider (1975) and Schneider & Searles (642: 277) cite the correct form as in this heading. Halymenia capensis Montagne Canaries (44; 401; 404). Note. For possible attributions of these records, see entry for Gymnophlaea canariensis Kiitzing in Price et al., 1988: 254. Halymenia clavaeformis Suhr Canaries (24; 44; 70; 134; 318; 401; 553; 616). Note. B0rgesen (70), De Toni (134) and Weber-van Bosse (553) all comment that specimens from Suhr in Herb. Montagne, Paris, are possibly representative of Rhodymenia [now Palmaria] palmata rather than H. clavaeformis. Halymenia cyclocolpa Montagne See Platoma cyclocolpa (Montagne) Schmitz. Halymenia dichotoma (J. Agardh) J. Agardh Canaries (70; 134; 188; 191; 227; 390). [As Halymenia dichotoma J. Agardh] '. . . mittellandischen Meer und in den angrenzenden Teilen des atlantischen Oceans' (510). [As Chrysymenia dichotoma J.G. Agardh] Canaries (259). [As Chrysymenia (Halichrysis) dichotoma J. Agardh] Canaries (24). Note. Recorded with doubt by Hemsley (259). This taxon has been reallocated to the genus Sebdenia of Berthold; see Codomier (1973). In view of the confusions in previous records between and within both these genera, we have not attempted reallocation in the absence of opportunity to examine original material in confirmation of generic/specific placement. Halymenia duchassaingii (J. Agardh) Kylin Cameroun (350; 586). Gabon (350;586). '. . . Tropical Africa (N. Gambia — Congo river)' (598). '. . . in tropical parts of the Atlantic Ocean' (350; 586). Halymenia elongata C. Agardh Canaries (635). Senegal (403; 408). [As Halymenia elongata L. et Crouan] Senegal (38). ?Cape Verde Islands (38). Note. Montagne (403: 125-126) commented: '. . . J'ai trouve des te~traspores dans des exemplaires de VH. elongata rapportes du Cap-Vert par M. Leprieur.'. Askenasy (38) later extended this by stating of H. elongata: '. . . n'ont pas 6te rapporte'es des iles du Cap Vert; il est tres probable qu'elles y croissent aussi et qu'on les trouvera plus tard. . .'. The question mark above derives from Askenasy 's statement. Conspecificity world- wide of terete Halyme- nia spp. (Millar, 1990) could result in records being reallocated here from various taxa. See the note to Halymenia agardhii De Toni. Halymenia fasciata Bory de St. Vincent Senegal (282). Halymenia floresia (Clemente) C. Agardh Canaries (24; 33; 60; 70; 97; 102; 134; 177; 188; 191; 227; 350; 375; 390; 401; 517; 584; 586; 598; 635; 642). Mauritanie (624). Sierra Leone (30; 350; 586). '. . . warmer parts of the Atlantic. . .' (60; 70; 375). '. . . im warmeren Teile des atlantischen Oceans, an den Kusten . . . Afrikas. . .' (510). '. . . Ad oras . . . Africae borealis. . .' (318). [As Halymenia floresia (Clemente) J. Agardh] '. . . warmeren Theilen des Atlantischen Oceans. . .' (266). '. . . in oceano atlantico calidiore ad litora . . . Africae. . .' (27). [As Halymenia floresia Agardh] Canaries (44; 89; 256; 257). '. . . De I'embouchure du Guadalquivir aux Canaries. . .' (89). '. . . Atlantic (. . . Africa. . .)' (410). '. . . Atlantic Ocean (. . . African . . . coasts. . .)' (177). Note. Aleem (30) reported this species dredged from 30 m from the shelf of the Sierra Leone peninsula; Lawson & John (350; 586) repeated the record with doubt, since it then constituted the only precise West African mainland report. Borgesen's (60; 70) Canaries report was based on '. . . few small bleached fragments of this plant [in Montagne's Herb.] . . . most probably belong to this species'. Marcot-Coquengniot (624: 198) recently unequivocally reported H. floresia from subtidal wrecks at two locations in the Bane d'Arguin area of Mauritanie. Halymenia floridana J. Agardh Canaries (18; 598). Note. For details of this difficult taxon, see Schneider & Searles (642: 279-280; comments on p. 339 are also useful). Halymenia hancockii W. Taylor Senegal (38D; 59). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). Note. Also recently reported (38D) from Madeira. Halymenia latifolia P. & H. Crouan in Lloyd ex Kiitzing Canaries (70; 191; 226; 227; 584; 598). '. . . Atlantique (de la Manche aux Canaries). . .' (188). [As Halymenia (Halarachnion) latifolia Crouan] Canaries (439). Note. Piccone himself (439), the first to record this species, stated that he had a single sterile specimen and recorded it with grave doubt. Prud'homme van Reine (pers. comm.), on studying the original specimen, concluded that it represented Halymenia vinacea Howe & W.R. Taylor (q.v.). The entry is maintained here for completeness only, since the nature of the later Canaries records has not been established by re-examination of material. Halymenia latifolia P. & H. Crouan f. trabeculata (Ercegovic) Codomier Canaries (643). [As Halymenia (Halarachnion!) spp.] Canaries (439). Note. The reallocation of the record from 439 follows re-examination of material by Prud'homme van Reine (pers. comm.). Halymenia ligulata (Woodward) C. Agardh See Halarachnion ligulatum (Woodward) Kutzing. 128 PRICE, JOHN & LAWSON Mai vine nia pinnulata J. Agardh See the note to Chrysymenia dichotoma J. Agardh. Halymenia polydactyla B0rgesen See the note to Halymenia agardhii De Toni. Halymenia ramosissima Suhr See the note on Agardhiella tenera (J. Agardh) Schmitz [=Solieria filiformis (Kiitzing) Gabrielson]. Halymenia rosea Howe & W.R. Taylor Canaries (635). Halymenia senegalensis Bodard Senegal (55; 56; 59; 290; 399; 598). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). Halymenia trigona (Clemente) C. Agardh See the note to Halymenia agardhii De Toni. Halymenia vinacea Howe & W.R. Taylor Canaries (635; 643). Ghana (350; 586; 590). '. . .in tropical parts of the Atlantic Ocean' (350; 586). Tropical Africa (N. Gambia — Congo river)' (598). Note. See remarks in 350, 586 and 642: 280. Possibly recorded as Halymenia sp. b. from Senegal by Bodard (50); see below. See also the notes to Halymenia latifolia P. & H. Crouan for the early additional record given as 643. Halymenia (Halarachnion?) spp. Canaries (439). Ghana (487). Senegal (50; 182). Note. Bodard (50) reported two undetermined species of Halyme- nia, a and b. Regarding species b he remarked '. . . Par certains caracteres elle rassemble a Halymenia vinacea Howe et Taylor'. The record in 439 from the Canaries is based on material recently re-examined by Prud'homme van Reine (643); he concluded that it represents Halymenia latifolia f. trabeculata (q.v.). Hapalidium phyllactidium Kiitzing See Melobesia confervicola (Kiitzing) Foslie. Haraldia lenormandii (Derbes & Solier) J. Feldmann Canaries (636). Cape Verde Islands (637). Haraldiophyllum bonnemaisonii (Greville) A. Zinova Namibia (36B; as 'prox'). Note. Wynne's (36B: 321) use of 'prox.' indicates wide morpho- logical variation between specimens that are apparently referable to this European Atlantic species; Wynne also indicated a tentative identification from Cape Peninsula (South Africa). Helminthiopsis purpurifera (J. Agardh) Papenfuss [As Acrosymphyton purpuriferum (J. Agardh) Sjostedt] Canaries (38C; 38D; 227; 232; 584; 628). Helminthocladia calvadosii (Lamouroux ex Dufy) Setchell Canaries (4; 8; 16; 128A; 172; 227; 232B; 379; 598). Cape Verde Islands (598). '. . . Atlantique (de 1'Angleterre aux Canaries)' (33). 'de Inglaterra a Canarias' (619). Note. See also Helminthocladia senegalensis Bodard. Helminthocladia senegalensis Bodard Senegal (55; 57; 58; 59; 399; 598). Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara] (598). [As Helminthocladia calvadosii (Lamouroux) Setchell] Senegal (49; 50). Helminthora divaricata (C. Agardh) J. Agardh See Helminthora stackhousei (Clemente) Cremades & Perez- Cirera. Helminthora stackhousei (Clemente) Cremades & Perez- Cirera [As Helminthora divaricata (C. Agardh) J. Agardh] Canaries (638). Heringia mirabilis (C. Agardh) J. Agardh Namibia (36B; 163; 348; 500; 515). Herposiphonia For general background to some aspects of specific level taxonomy and nomenclature in this widespread genus, see Wynne (36). Comments in Lawson & John (586) provide the necessary leads into much of the earlier literature, whilst the present listing updates subsequent taxonomic practice and publications for the area. Herposiphonia brachyclados Pilger Annobon (139; 350; 456; 457; 496; 586). Ascension (475). Note. De Toni (139: 420) considered this species and H. tenella [H. secunda f. tenella, q.v.] to be close. Herposiphonia densa Pilger See Herposiphonia secunda (C. Agardh) Falkenberg forma densa (Pilger) Wynne and Herposiphonia guineensis G. Law- son & D. John. Herposiphonia guineensis G. Lawson & D. John Cote d'lvoire (350; 586). Ghana (350; 586). Liberia (350; 586). Togo (350; 586). Tropical Africa (N. Gambia — Congo river)' (598). '. . . western Africa' (36). [As Herposiphonia densa Pilger] Cote d'lvoire (287; 295). Ghana (295; 297; 338; 537). Liberia (287; 295). Togo (293;295). Herposiphonia heringii (Harvey) Falkenberg Namibia (348). Herposiphonia? parvula (Suhr ex Kiitzing) De Toni Canaries (133). [As Polysiphonia parvula (Suhr in herb. Sender — P. irregu- laris Zanardini. ms]. Canaries (322). Note. De Toni (133) placed this record under 'Species Polysipho- niae inquirendae, forsan ad genus Herposiphoniani pertinentes'. Herposiphonia secunda (C. Agardh) Falkenberg f. secunda (C. Agardh) Wynne Canaries (604; 605). Mauritanie (624). [As Herposiphonia secunda (C. Agardh) Falkenberg] Canaries (108; 604; 605). Mauritanie (624) 'Ampliamente distribuida por los mares calidos y tropicales' (604). [As Herposiphonia secunda (C. Agardh) Nageli] RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 129 Cameroon (337; 537) Canaries (16; 71; 174; 191; 225; 235; 375; 489; 517; 546; 547; 556; 610). Cape Verde Islands (191). Gambia (292). Ghana (292). Salvage Islands (215; 231; 375; 556). Senegal (556). Sierra Leone (30). 'Mediterranean and adjacent warmer parts of the Atlantic . . . Probably widely spread in tropical and subtropical seas' (375). '. . . pantropical. . .' (610). [As Herposiphonia secunda (Polysiphonia secundd) Mon- tagne] Canaries (547). [As Herposiphonia secunda (C. Agardh) Ambronn] Canaries (13; 128A; 190; 226; 227; 229; 253; 306B; 379; 584). Senegal (59). '. . . muy extendida por los mares tropicales y subtropicales' (253). 'Semble repandu dans toutes les mers tropicales et subtropi- cales' (190). [As Herposiphonia tenella var. secunda Hollenberg] Angola (352). Cameroun (350). Canaries (38B; 38C; 38D). Ghana (350). Mauritanie (349). Salvage Islands (38B; 38C; 38D). Senegal (38B; 38C; 38D). Sierra Leone (295; 349; 350). [As Herposiphonia tenella forma secunda (C. Agardh) Hol- lenberg] Canaries (392). [As Herposiphonia tenella var. subsecunda Hollenberg] Mauritanie (349). [As Polysiphonia secunda Montagne] Canaries (38; 44; 89). Cape Verde Islands (38). Senegal (38). [As Polysiphonia secunda (Agardh) Montagne] Canaries (401; 439; 445). Senegal (408). [As Polysiphonia secunda (Agardh) Montagne var. Adunca (Agardh) Montagne] Canaries (401). [As Polysiphonia secunda (Agardh) Zanardini] Canaries (441; 444). f. tenella (C. Agardh) Wynne Canaries (602A; 604). Mauritanie (624). 'Ampliamente distribuida por los mares calidos y tropi- cales'(604). [As Herposiphonia tenella (C. Agardh) Ambronn] Angola (298; 352). Ascension (474; 475). Cameroun (350). Canaries (13; 38B; 38C; 38D; 128A; 226; 227; 253; 306B; 379; 517; 557A; 584). Gambia (296; 350). Ghana (295; 338; 350; 377; 491). Liberia (129; 295; 350). Mauritanie (38B; 38C; 38D; 349). Salvage Islands (38B; 38C; 38D)' Sao Tome (93; 350). Senegal (38B; 38C; 38D; 50; 122; 542). Sierra Leone (30; 295; 350). '. . . Atlantique tropicale et subtropicale. . .' (542). '. . . Muy extendida por los mares tropicales y subtropicales. . .' (253). '. . . probably widespread in warm temperate and tropical seas' (350). 'Semble repandu dans toutes les mers tropicales et subtro- icales' (190). '. . . world-wide ... on tropical to warm temperate coasts. . .' (267 A). '. . .widespread in tropical and subtropical seas' (642). [As Herposiphonia tenella (C. Agardh) Falkenberg] Canaries (598). Cape Verde Islands (598). Salvage Islands (598). '. . . atlantischen kusten von England bis zu den Canarischen Inseln. . .' (501). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). '. . . Tropical Africa (N. Gambia — Congo river)' (598). [As Herposiphonia tenella (C. Agardh) Schmitz] Canaries (392). [As Herposiphonia tenella (C. Agardh) Nageli] Canaries (16; 71; 108; 140; 174; 191; 225; 375; 390; 489; 555; 556). Mauritanie (555; 556). Salvage Islands (231; 555; 556). Sao Tome (93). Senegal (555; 556). '. . . Atlantique (du golfe de Gascogne aux Canaries) . . . repandu dans les mers tropicales et sub-tropicales' (33). '. . . Probably spread in tropical and subtropical waters' (375). [As Herposiphonia tenella — Schmitz (Polysiphonia tenella J. Agardh)] Canaries (547). [As Polysiphonia tenella J. Agardh] Canaries (89). Note. On the status of this taxon as forma or variety of H. secunda (C. Agardh) Falkenberg, or as separate species, see Schneider & Searles (642). f. densa (Pilger) Wynne [As Herposiphonia densa Pilger] Cameroun (122; 139; 288; 337; 350; 454; 500; 537). Cote d'lvoire (287; 288). Ghana (288; 297). Liberia (287; 288). Senegal (47; 55; 59; 122; 529). Togo (288; 293). '. . . 1'aire se limite au golfe du Benin et a la Mauritanie. . .' (59). '. . . espece ouest africaine' (59). '. . . Espece tropicale africaine' (529). [As Herposiphonia tenella (C. Agardh) Ambronn var. densa (Pilger) G. Lawson & D. John] Cameroun (350). Herposiphonia spp. Bioko (346; 350). Canaries (71; 226). 130 PRICE, JOHN & LAWSON Ghana (377; 487; 491). Senegal (59). 'West Africa' (290; 344). Heteroderma lejolisii (Rosanoff) Foslie See Pneophyllum lejolisii (Rosanoff) Chamberlain. Heterosiphonia Many red algal genera show little overlap in species names utilised in different but adjacent areas of the western Atlan- tic. Heterosphonia is such a genus, South Africa (Seagrief, 1984) showing records for five species of which none is recorded for the present list area, although the records to genus only for Namibia could perhaps represent one or more common taxa. Heterosiphonia crispella (C. Agardh) Wynne Canaries (36A; 598; 604; 605; 621). Cape Verde Islands (598). Ghana (586). Mauritanie (624). Salvage Islands (36 A; 598). '. . . ampliamente distribuida por los mares calidos y tropi- cales' (604). '. . . northwestern Africa . . . southward to tropical West Africa . . . and Angola' (36 A). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]. . .' (598). 'Tropical Africa (N. Gambia — Congo River). . .' (598). '. . . widespread from boreal-antiboreal to tropical seas' (586). Note. For detailed reasoning behind the name change from wurdemannii, see Wynne (36A: 85-87). See also the entry (Price et al., 1986: 93) for Fucus plumosus L. [As Heterosiphonia crispella (C. Agardh) Wynne var. laxa (B0rgesen) Wynne] Canaries (36 A). Salvage Islands (36 A). '. . .northwestern Africa . . . southward to tropical West Africa . . . and Angola. . .' (36 A). 'West Africa' (642). Note. Records above are based on those in 71 (Canaries), 535 (Salvage Islands) and 118, 119, 122, 298 and 350 (the generalized statement). [As Heterosiphonia wurdemannii (Bailey ex Harvey) Falken- berg] Angola (298; 352). Ascension Island (475). Canaries (13; 36A; 108; 226; 227; 292*; 392). Ghana (292; 299; 350; 376; 377; 491). Mauritanie (349). Senegal (292). 'Atlantic Ocean (Africa and American coasts, Canary Islands. . .)' (177). '. . . widespread from boreal-antiboreal to tropical seas. . .' (350). '. . . widespread occurrence in warm temperate and tropical seas. . .' (292). Note. No attempt is made separately to distinguish records where the epithet wurdemannii is incorrectly rendered with a single T or 'n'. The record indicated by an asterisk states the describing authority to be Bailey in Harvey. Material reported from Cap Blanc (Mauritania) was detected (349) only as drift. [As Heterosiphonia wurdemannii (Bailey) Falkenberg] Canaries (71; 128A; 174; 191; 306B; 375; 542; 555; 556). Mauritanie (555; 556). Salvage Islands (36A; 231; 375; 555; 556). Senegal (55; 59; 122; 186; 399; 542; 555; 556). '. . . Atlantique (de Cadix aux Canaries. . .)' (190). '. . . warm Atlantic' (78). Note. No attempt is made separately to distinguish records where the epithet wurdemannii is incorrectly rendered with a single 'n' or T. [As Heterosiphonia wurdemannii (Bailey ex Harvey) Falken- berg var. laxa B0rgesen] Canaries (13; 38B; 71). Mauritanie (38B; 349). Salvage Islands (38B). Senegal (38B). [As Heterosiphonia wurdemannii (Bailey ex Harvey) Falken- berg var. typica B0rgesen] Canaries (38B; 38C; 38D; 71; 226). Mauritanie (38B*; 38C; 38D). Salvage Islands (38B; 38C; 38D). Senegal (38B*; 38C; 38D). Note. No attempt is made separately to distinguish records where the specific (not varietal) epithet wurdemannii is incorrectly rendered with a single 'i' or 'n'. Records indicated by an asterisk state 'var. typica Piccone', thereby differing from the usual authority statement. [As Dasya wurdemanni Bailey] Canaries (439; 547). [As Dasya wurdemanni Bailey in Harvey] Canaries (89). Heterosiphonia dubia (Suhr) Falkenberg Namibia (36B). Heterosiphonia pellucida (Harvey) Falkenberg Canaries (303). Note. No authority is given by Johnston (303); conspecificity of material is therefore conjectural. Heterosiphonia plumosa (Ellis) Batters Western Sahara (33; 222; 349). Note. Overall distribution statements in 33 and 222 give the southern limit as 'Rio de Oro'. Heterosiphonia wurdemannii (Bailey ex Harvey) Falkenberg and other variants See Heterosiphonia crispella (C. Agardh) Wynne. Heterosiphonia sp. Namibia (348). Note. See the generic note for possible specific relationships. Hildenbrandia canariensis B0rgesen Canaries (33; 39; 70; 191; 227; 598; 627). Mauritanie (33; 349; 598; 627). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). Hildenbrandia prototypus Nardo See Hildenbrandia rubra (Sommerfelt) Meneghini. Hildenbrandia rosea Kiitzing See Hildenbrandia rubra (Sommerfelt) Meneghini. Hildenbrandia rubra (Sommerfelt) Meneghini Ascension (475). Canaries (227; 598). Cote d'lvoire (350; 586). RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 131 Gambia (296; 350; 586). Ghana (350; 586). Liberia (350; 586). Sao Tome (350; 586). Sierra Leone (350; 586). Tropical Africa (N. Gambia — Congo river)' (598). [As Hildenbrandia prototypus Nardo] Angola (298; 352). Canaries (70; 191; 493; 517). Cote d'lvoire (288). Gambia (288). Ghana (288; 340; 344). Liberia (129; 288). Sao Tome (288). Sierra Leone (30; 288; 295). '. . . Atlantique (du Spitzberg aux Canaries. . .)' (33). '. . . most probably ubiquitous. . .' (61). 'Probably cosmopolitan' (375). [As Hildenbrandia rosea Kiitzing] Namibia (348). Sao Tome (263; 264; 265; 535). [As Hildenbrandia sp.] Sao Tome (535). Note. Although Henriques (264) reported the material from Sao Tome as doubtfully determined since sterile, he still named it H. rosea (263; 264; 265). Steentoft (535), faced with similar problems, preferred to avoid allocation of epithet. Hildenbrandia spp. Ghana (342). Namibia (348). Sao Tome (535). Note. For the probable allocation of records in 348 and 535, see the note to H. rubra (Sommerfelt) Meneghini. Hutchinsia filamentosa (Wulfen) C. Agardh See Spyridia filamentosa (Wulfen) Harvey. Hydrolithon We are indebted to Wm. J. Woelkerling for the following comments. Studies of Penrose & Woelkerling (1992) on Hydrolithon reinboldii, the type species of Hydrolithon, have resulted in a revised circumscription for the genus in which attributes associated with tetrasporangial conceptacle roof anatomy are crucial. Thus Hydrolithon is delimited from other genera of the Corallinaceae by the following combination of features: 1. thallus nongeniculate, nonendophytic and lacking hausto- ria; 2. thallus lacking a basal layer of palisade cells throughout; 3. cells of adjacent filaments joined by cell fusions, secondary pit-connections absent; 4. tetrasporangial conceptacles uniporate and tetrasporangia lacking apical plugs; and 5. pore canals of tetrasporangial conceptacles lined by a ring of conspicuous, elongated cells which arise from filaments interspersed amongst sporangial initials, which do not pro- trude into the pore canal and which are orientated more or less perpendicular to the roof surface. Penrose & Woelkerling (1992) also have determined that Porolithon is a heterotypic synonym of Hydrolithon. Distinc- tions between Hydrolithon and Spongites are outlined by Penrose & Woelkerling (1992), and a detailed study of the type species of Spongites is provided by Penrose (1991). Hydrolithon boergesenii (Foslie) Foslie Sao Tome (350; 586). '. . . in warm temperate and tropical parts of the Atlantic Ocean' (350; 586). '. . . Tropical Africa (N. Gambia — Congo river)' (598). [As Goniolithon boergesenii] Sao Tome (6). [As Goniolithon Boergesenii Foslie forma africana Foslie] Sao Tome (206). [As Goniolithon Boergesenii Foslie var. africana Foslie] Sao Tome (135; 136; 485). [As Goniolithon (Hydrolithon) Boergesenii Foslie forma afri- cana Foslie] Sao Tome (135, 206). 'Coste occid. d' Africa, S. Thomas (Gravier)' (135). [As Goniolithon (Hydrolithon) Boergesenii Foslie var. afri- cana Foslie] Sao Tome (136; 137; 206; 251) [As Porolithon Boergesenii (Foslie) Lemoine forma africana Foslie] Sao Tome (139). [As Porolithon Boergesenii (Foslie) Lemoine var. africana] Sao Tome (357). [As Porolithon Boergesenii (Foslie) Lemoine] Sao Tome (139; 357; 358; 535). Note. Foslie (206) based his St. Thomas (Africa) record on Gravier collection no. 6, communicated by P. Hariot. Similarly, the determination given in Hariot (251) relates to the same Gravier material and is stated to be by Foslie. Lemoine (357: 149) indicated that she could add nothing to the picture of this variety as envisaged by Foslie. The specimen she examined in Paris of 'type' material of the var. africana was so small that to study it was to destroy it — she did not know if Foslie had a better specimen. In their entry on Hydrolithon reinboldii (Weber-van Bosse & Foslie) Foslie, Adey et al. (1982: 26) commented of the two species: '. . . The Caribbean Hydrolithon borgesenii (Foslie) Foslie is quite similar to H. reinboldii in color, surface texture, and anatomy, though it is not usually so strongly mammillate as the Hawaian plant. Also, H. borgesenii has a very wide depth range and, in the Caribbean, appears to be the ecological equivalent of all of the Pacific Hydrolithon species'. Woelkerling (in litt.) has kindly confirmed that the type of H. boergesenii (Foslie) Foslie belongs to Hydrolithon sensu Penrose & Woelkerling (1992). The taxonomic placement of the forma africana is not yet certain. Hydropuntia dent at a (J. Agardh) Wynne Cape Verde Islands (639). Mauritanie (624). Note. Wynne (1989) has reinstated the genus Hydropuntia Mon- tagne to include species of Polycavernosa; see Gracilaria [ = Polycav- ernosa] dentata for comments appropriate at the time (Price et al. 1988). Since then, and subsequent to Wynne (1989), Abbott et al. (1991: 24) have reassessed assignment to Hydropuntia of both H. dentata and H. henriquesiana in the process of arriving at their conclusion that there is little to warrant continued recognition of Hydropuntia (= Polycavernosa) and Gracilariopsis. If their reason- ing, and consequent reassignments, are accepted, both taxa (H. dentata, H. henriquesiana) become synonyms of Gracilaria rangifer- ina (Kiitzing) Piccone. We make no definitive moves in any direction at this time since restoration to Gracilaria by Abbott et al. (op. cit. ) was purely based on previous assignment, not on re-examination of appropriate material. Hydropuntia henriquesiana (Hariot) Wynne Note. See (a) the comments and records under Gracilaria [-Poly- cavernosa] dentata in Price et al. (1988), and (particularly) (b) the detailed note to Hydropuntia dentata (J. Agardh) Wynne, above. 132 PRICE, JOHN & LAWSON Hymenena fissa Greville See Hymenena venosa (L.) Kylin. Hymenena venosa (L.) Kylin Namibia (348). [As Hymenena fissa Greville] Senegal (38; 408). Note. Both citations derive from the island of Gor6e. Askenasy (38) also indicated that plants representative of this taxon '. . . n'ont pas etc" rapporte"es des lies du Cap Vert; il est tres probable qu'elles y croissent aussi et qu'on les trouvera plus tard'. [As Nitophyllum venosum Harvey] Namibia (165; 500). Note. Dinter (165: 315) recorded the taxon from Walfischbay, collected by Cleverley, and the Schmidt & Gerloff (500) statement is merely based on that. [As Nitophyllum fissum (Greville) J. Agardh] Namibia (165; 453). Note. Dinter (165: 315) recorded the taxon from Liideritzbucht, collected by L. Scholtz, a direct derivation from the Pilger (453) statement. Overall Note. The epithets fissa and venosa are now generally accepted to apply to the same plant in this area of the Atlantic (see, for example, Seagrief, 1984). It remains to be established that the determinations in all the records quoted were correct. A record quoted for the Canaries (598) under the name Nitophyllum venulo- sum may have been a mis-statement of epithet for the same taxon; no authorities are listed and the determination needs confirmation since this represents the only record for that island group. The relationship of the records under Hymenena sp. (q.v.) to H. venosa is not clear. Hymenena sp. Ghana (290; 299; 300; 350; 376; 377; 586). Note. See the terminal note to Hymenena venosa (Linnaeus) Kylin. Hypnea This complex genus has a confused nomenclatural history. Additionally, both biology and relation of form-variation to environment are still poorly understood. All this has resulted in a taxonomic and recording mosaic, the resolution of which really requires monographic treatment on a world scale. Since the genus is of substantial commercial (phycocolloid) importance, there is considerable literature, much of it con- cerned with aspects other than taxonomy and biology; the general effect has been further to complicate the taxonomic situation. Commercial data and literature sources appear in Hoppe (269: 253-255) and in Rama Rao (481), both of which also include primary and secondary distributional data for parts of the list area. More recent papers concerned directly with part (Senegal) of the area are those of Mollion (398; 399; 623). An earlier taxonomic coverage by Bodard (53), although valuable, includes many errors of taxonomy and faulty interpretation of distributional and other data. Hypnea arbuscula P. Dangeard Canaries (635; 640). Cape Verde Islands (640). Gambia (296; 350; 586). Guinea-Bissau (529). Mauritanie (624). Senegal (53; 55; 56; 59; 122; 529). '. . . in warm temperate and tropical parts of West Africa' (350; 586). '. . . Tropical Africa ( N.Gambia — Congo river)' (598). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). Hypnea benguelensis Palminha Angola (241; 352; 393; 394; 424; 428). Hypnea cenomyce J. Agardh Cape Verde Islands (637; 640). Sierra Leone (53; 295; 350; 586). Senegal (53; 122; 350; 586). '. . . doit etre largement diffusee sur les cotes du golfe du Benin. . .' (53). '. . . in warm temperate and tropical parts of West Africa. . .' (350; 586). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). Tropical Africa (N. Gambia — Congo river)' (598). [As Hypnea cenomyces J. Agardh] Senegal (59). [As Hypnea ctenamyces J. Agardh] Senegal (55). [As Hypnea ctenomyces J. Agardh] Senegal (53; 122; 535). Note. The record in 122 is given with '?'. Several of the authors (e.g. Bodard, 53; Steentoft, 535) have suggested this to be the same as a small Hypnea spinella (C. Agardh) Kiitzing (q.v.). Hypnea ceramioides Kiitzing Senegal (59; 348; 399). Mauritanie (53; 349). Namibia (348; 438). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). [As Hypnea sp. aff. ceramioides] Senegal (53; 398). [As Hypnea aff. ceramioides Bod. (nomen)] Senegal (55) [As Hypnea ceramoides Kiitzing] Senegal (59; 480). [As Hypnea aff. ceramoides Bodard (nomen)] Senegal (59; 399). [As Hypnea veramoides] Senegal (481). Hypnea cervicornis J. Agardh See Hypnea spinella (J. Agardh) J. Agardh. Hypnea cor nut a (Lamouroux) J. Agardh See Hypnea valentiae (Turner) Montagne. Hypnea ctenomyces J. Agardh Note. See Hypnea cenomyce J. Agardh, for which this name represents a mis-rendering of the specific epithet. Hypnea divaricata Greville Cape Verde Islands (38; 145; 450; 451; 500; 598; 625). Hypnea eckloni Suhr See Hypnea musciformis (Wulfen) Lamouroux. Hypnea esperi Bory, (Bory) Kiitzing, or Kiitzing See Hypnea musciformis (Wulfen) Lamouroux. Hypnea flagelliformis Greville ex J. Agardh [As Hypnea flagelliformis (Greville) J. Agardh] Canaries (640). Cape Verde Islands (640). Cote d'lvoire (59; 586). RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA Ghana (59; 291; 350; 586). Guinea-Bissau (529). Liberia (129). Mauritanie (53; 296). Senegal (53; 59; 122?; 296; 529). '. . . golfe du Benin. . .' (59). Tantropical' (529). Tropical Africa (N. Gambia — Congo river)' (598). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). Hypnea muscif ormis (Wulfen) Lamouroux Angola (213; 269; 352; 500; 535). Cameroun (213; 269; 295; 337; 350; 454; 484; 500; 535; 537; 586). Canaries (5; 13; 16; 25; 38B; 38C; 38D; 70; 71; 89; 108; 128A; 132; 147?; 148?; 191; 213; 226; 227; 230; 235; 236; 238; 252; 267A; 302; 306B; 351; 379; 392; 401; 404; 439; 489; 490; 499; 517; 535; 556; 584; 598; 604; 605; 610; 621). Cape Verde Islands (38B; 38C; 38D; 100; 183; 191; 213; 252; 267A; 408; 423; 442; 443; 445; 446; 449; 483; 499; 535; 556; 598). Congo (535). Cote d'lvoire (287; 288; 350; 586). Gabon (294; 350; 586). Gambia (296; 350; 586). Ghana (213; 288; 291; 335; 338; 340; 344; 350; 393; 394; 409; 481; 487; 491; 535; 537; 586; 594). Guinea-Bissau (529?). Liberia (129; 287; 295; 350; 586). Mauritanie (38B; 38C; 38D; 53; 252; 267A; 269; 349; 481; 516; 535; 556; 624). Namibia (36B; 348). Nigeria (213; 285; 350; 535; 586). Principe (93; 350; 535; 586). Salvage Islands (38B; 375; 555). Sao Tome (213; 251; 261; 265; 295; 350; 535; 586). Senegal (38B; 38C; 38D; 50; 51; 53; 55; 56; 59; 99; 122; 182; 213; 252; 267A; 291; 350; 394; 398; 399; 411; 480; 481; 529; 542; 556; 586; 618; 623). Sierra Leone (30; 213; 295; 336; 350; 535; 586). Togo (293; 295; 350; 586). Western Sahara (38B; 38C; 38D). '. . . alien warmeren Meeren. . .' (482; 483). 'Ampliamente distribuida por los mares calidos y tropicales' (604). '. . . Atlantique tropical et Subtropical. . .'(184; 189). '. . . im warmeren atlantischen . . . Ocean. . .' (506). '. . . Du Golf de Gascogne aux Canaries' (38; 89). 'Gulf of Guinea' (480). 'GulfofGivea[sic!]'(481). Tantropical' (529; 610). 'seems to occur in all warmer seas. . .' (64; 70). '. . . Tropical and subtropical Atlantic. . .' (375). '. . . toutes les mers tropicales et subtropicales' (542). Tropical Africa (N. Gambia — Congo river)' (598). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). 'Warm Atlantic' (410). 'Widespread in warmer waters' (535). 'Widespread in warm temperate and tropical seas' (350; 586). [As Hypnea eckloni (Suhr)] Namibia (128; 164). [As Hypnea esperi (Bory) Kutzing] 133 Cape Verde Islands (38). [As Hypnea musciformis Greville] Senegal (282). [As Hypnea musciformis J. Agardh] Angola (41; 42). Canaries (44; 547). Cape Verde Islands (38; 41; 42; 446; 596). Congo (249;250). Gabon (250). Salvage Islands (231). Sao Tome (263; 264). [As Hypnea musciformis Lamouroux] Canaries (385). [As Hypnea musciformis Lamouroux var. esperi (Bory) Mon- tagne] Cape Verde Islands (38; 408). [As Hypnea musciformis (Wulfen in Jacquin) J. Agardh] '. . .In oceano calidiore atlantico. . .' (27). Note. See Schneider & Searles (642: 308-9) on the potentiality for confusion between this taxon and H. valentiae (Turner) Montagne. Hypnea pannosa J. Agardh Cape Verde Islands (38; 145; 450; 451; 500; 598). Senegal (53; 122; 535). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). Note. The comment (see note to H. spinella (C. Agardh) Kutzing) on genomic distinction possibly applies equally between H. pannosa and H. cervicornis. Hypnea setacea Kutzing See Hypnea unilateralis P. Dangeard. Hypnea spicifera (Suhr) Harvey Namibia (312A; 348; 437; 438; 523). Hypnea spinella (C. Agardh) J. Agardh St. Helena (644) [As Hypnea spinella (C. Agardh) Kutzing] Angola (352). Annobon (456; 457; 535). Ascension (474; 475). Canaries (13; 118; 128A; 226; 227; 232B; 253; 306B; 535; 602 A; 604; 605). Cape Verde Islands (38; 100; 183; 191; 264?; 440; 442; 443; 445; 446; 528; 535?). Gabon (253; 295; 350; 586). Gambia (296; 350). Ghana (350; 491; 586). Sao Tome (251; 263?; 264?; 265; 350; 535; 586). Senegal (53; 535?). Sierra Leone (30; 295; 336; 339; 350; 378; 535; 586). Western Sahara (349). 'Ampliamente distribuida por los mares calidos y tropicales' (604). '. . . Atlantico (Canarias-Gabon . . .)' (253). '. . . widespread in warm temperate and tropical seas' (350; 586). '. . . widely distributed on the Atlantic coast of Africa' (535). [As Hypnea spinella Kutzing] Sao Tome (263; 264). [As Hypnea cervicornis J. Agardh] Angola (352). Canaries (8; 13; 16; 38B; 38C; 38D; 70; 108; 128A; 191; 226; 227; 229; 233; 236; 237; 253A; 306B; 375; 535; 556; 584; 598; 604; 605; 610; 621). 134 PRICE, JOHN & LAWSON Cape Verde Islands (1; 38; 38B; 38D; 150; 500; 556; 598). Gabon (295; 350; 586). Gambia (296; 350; 586). Ghana (288; 290; 299; 300; 335; 338; 350; 376; 377; 491; 537; 586). Liberia (129; 288; 350; 586). Mauritanie (38B; 38C; 38D; 53; 349; 556; 624). Principe (93; 288; 350; 535; 586). Salvage Islands (38B; 38C; 38D; 375; 556; 598). Sao Tome (288; 350; 535; 586). Senegal (38B; 38C; 38D; 49; 53; 56; 59; 122; 290; 350; 398; 399; 480; 481; 535; 542; 556; 586). Sierra Leone (295; 350; 586). Western Sahara (38B). 'Ampliamente distribuida por los mares calidos y tropicales' (604). '. . . most warm seas' (64; 70). '. . . pantropical. . .' (610). 'Gulf of Guinea' (269). Tropical Atlantic' (544). '. . . largement connu de 1'Atlantique tropical. . .' (53). '. . . largement connue en milieu tropicale. . .' (56). 'Tropical Africa (N. Gambia — Congo river)' (598). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). 'West Africa' (615). '. . . widespread in tropical and subtropical seas' (642). [As Hypnea cervicoris (Turner) J. Agardh] Senegal (55). [As Hypnea cervicornis C. Agardh] Canaries (602 A). [As Hypnea musciformis] Sao Tome (251; 261; 263; 264; 265). [As Hypnea spinella J. Agardh] Cape Verde Islands (528). Note. See also Hypnea ctenomyces J. Agardh. Analysis of the respective genomes may be the surest way to resolve the question of whether or not H. spinella and H. cervicornis (q.v.) are conspecific. Haroun & Prud'homme van Reine (640 and in press) have presented detailed discussion on both nomenclatural background and distribu- tion, concluding that H. spinella and H. cervicornis are transitional forms resulting from the effects of habitat on the same genome. Pending genomic analysis, we have accepted their discussion and typification. Hypnea .'tennis Kylin Namibia (348). Note. Recorded from the Cunene River mouth, Angola/Namibia border; otherwise known from east coast of southern Africa. Hypnea unilateralis P. Dangeard Mauritanie (53; 349; 598). Senegal (53; 55; 56; 59; 122; 598). Western Sahara (53; 349; 598). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). '. . . vraisemblable que cette espece existe dans tout 1'Atlan- tique tropical' (53). Note. Bodard (53: 812) commenting on the Hypnea spp. given by Dangeard (122) for 'presqu'ile du Cap- Vert (Senegal)', said of H. unilateralis 'sp. nov.' 'Toutefois nous doutons de 1'originalite de cette espece, il pourrait bien s'agit en fait de YHypnea setacea Kiitzing, quoique cette derniere espece soit tres mal connue et toujours accompagnee . . . d'un?. . .'. Hypnea ustulata (Mertens ex Turner) Montagne See Caulacanthus ustulatus (Turner) Kiitzing. Hypnea valentiae (Turner) Montagne Canaries (25; 132; 401; 404; 407; 598). Cape Verde Islands (640). Ghana (350; 586). Guinee (350; 586). '. . . widespread in warm temperate and tropical seas' (350; 586). '. . . widespread in tropical and subtropical seas' (642). [As Hypnea valentinae] Canaries (598). 'Tropical Africa (N. Gambia — Congo river)' (598). [As Hypnea cornuta (Lamouroux) J. Agardh] Canaries (128A; 306B; 582; 598). 'Guinea' [probably Ghana — see note below] (25; 64; 97; 132). Senegal (542). Sao Tome (132). '. . .In atlantico calidiore. . .' (27). [As Hypnea cornuta var. cornuta J. Agardh] '. . .in oceano atlantico calidiore, ad oras Guineae. . .' (25). Note. B0rgesen (64) stated: 'St. Thomas: in the harbour of Charlotte Amalia . . . Orsted has gathered it at St. Thomas'. Therefore all records of this plant from St. Thomas refer to the West Indian island, not that in the Gulf of Guinea. Nevertheless, B0rgesen mentioned it from Guinea, probably direct attribution of 'ad oras Guineae' in J. Agardh (1852) and De Toni (1900). In the time of Lamouroux, whose herbarium specimen is the source of these records, 'Guineae' referred to what is now Ghana. The references to H. cornuta are secondary records in Lawson & John's (586) entry for H. valentiae; they come from the same publications and should all be localised to Ghana, not to Guinee as stated. See also the note to Hypnea musciformis (Wulfen) Lamouroux. Hypnea spp. Angola (352). Canaries (2; 5; 351). Cote d'lvoire (287). Gambia (296). Ghana (92; 291; 335; 344; 491; 567; 626). Liberia (287). Mauritanie (53; 349). Namibia (348). Senegal (50; 52; 53; 55; 56; 59; 182; 290; 396; 398; 399; 411; 481; 531; 615). '. . . African west coast. . .' (374). 'West Africa' (344; 615). Hypneocolax stellaris B0rgesen Canaries (640). Cape Verde Islands (640). Hypoglossum For an analysis of morphological distinctions between the sometimes confused forms in the genera Caloglossa and Hypoglossum, see Wynne & Kraft (561A). There is growing literature on the family Delesseriaceae in southern Africa to which effective entry can be made through the recent paper by Wynne & Norris (1991). Hypoglossum abyssicolum W.R. Taylor Angola (352). Hypoglossum guineense G. Lawson & D. John Ghana (350; 377; 586). 'Tropical Africa (N. Gambia — Congo river)' (598). RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA 135 [As Hypoglossum sp.] Ghana (300; 376). Hypoglossum hypoglossoides (Stackhouse) Collins & Hervey Canaries (128A; 306B; 598; 604; 605; 621; 642). Cape Verde Islands (598). Salvage Islands (598). 'Gran Bretana — Rio de Oro' (604). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). [As Delesseria hypoglossum Agardh] Canaries (254; 257; 305). [As Delesseria Hypoglossum Goodenough & Woodward) Lamouroux] '. . . D'Angleterre aux Canaries. . .' (89). [As Delesseria hypoglossum (Woodward) Lamouroux] Canaries (25; 44; 401; 547). [As Hypoglossum woodwardii Kiitzing] Canaries (13; 38B; 38D; 71; 132; 191; 221; 226; 227; 232B; 238; 304; 375; 379; 392; 489; 517; 565A; 584; 617; 619). Mauritanie (38B; 38D). Salvage Islands (38B; 38D). Senegal (38B; 38D; 55; 59; 122; 399; 529). Western Sahara (349; 565 A; 618). [As Hypoglossum woodwardii Kylin] Mauritanie (516). 'Atlantischen Inseln' (239). '. . . Atlantique nord et subtropical' (190). '. . . Atlantique: depuis les cotes anglaises jusqu'au Rio de Oro. . .' (222). '. . . Atlantic ocean (from England to the Canary Islands, . . .)' (177). '. . . Atlantique (de 1'Angleterre au Rio de Oro). . .' (33). '. . . Atlantique: cotes d'Europe jusqu'aux Canaries. . .' (221). '. . . Atlantico de Inglaterra a Canarias. . .' (517). '. . . de Inglaterra a Rio de Oro' (619). '. . . English coast southwards to the Canary Islands. . .' (71). '. . . Im Atlantischen Ozean von den englischen kiisten bis zu den kanaren. . .' (495). '. . . in oceano Atlantico ab oris Angliae usque ad Tingin Africae et insulas Canarienses. . .' (132). Hypoglossum woodwardii Kiitzing See Hypoglossum hypoglossoides (Stackhouse) Collins & Hervey. Hypoglossum spp. Ghana (299; 300; 376; 491). Namibia (348). Senegal (50; 55; 59; 542). '. . . ouest africaines. . .' (59). Note. Bodard & Mollion (59: 201) referred to this Hypoglossum sp. as '. . . est a decrire. . .'. Iridaea capensis J. Agardh Angola (352). Namibia (36B; 348; 352; 437; 523; 595). Iridaea elongata Suhr Table Bay to Walfisch Bay' (128). Note. Possibly a synonym for /. capensis. Janczewskia verrucaeformis Solms-Laubach Canaries (71; 190; 191; 227; 232B; 375; 379; 598). Cape Verde Islands (598). Salvage Islands (38B; 231; 375; 598). '. . . Atlantique (. . . Canaries). . .' (33). Note. On Janczewskia in general, see Apt (1987). Jania The genus is more closely related to Haliptilon than it is to Corallina (Johansen, 1981; Johansen & Silva, 1978; Johansen & Womersley, 1986: 562). The taxonomic boundaries between Haliptilon and Jania are of different clarity in different parts of the world (Johansen & Womersley, 1986: 562); in southern Australia, they are very clear, whereas elsewhere (e.g. Europe) there are problems. There, Jania corniculata (Linnaeus) Lamouroux (q.v.) has lateral branches reminiscent of Haliptilon. Jania adhaerens Lamouroux Canaries (9; 13; 38B; 38C; 38D; 128 A; 227; 306B; 379; 556; 582; 584; 598; 602; 604; 605). Cameroun (122; 350; 500; 586). Cape Verde Islands (598). Gambia (296; 350; 586). Ghana (350; 376; 377; 586). Liberia (129). Mauritanie (38B; 38C; 38D; 349; 556; 624). Salvage Islands (38B; 38C; 38D; 556; 598). Sao Tome (93; 350; 535; 586). Senegal (38B; 38C; 38D; 122; 556). Sierra Leone (30; 295; 349; 350; 586). St Helena (644). 'Gulf of Guinea' (582; 604). '. . . more or less pantropical. . .' (535). Tropical Africa (N. Gambia — Congo river' (598). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W.Sahara]' (598). '. . . widespread in warm temperate and tropical seas' (350; 586). '. . . widespread in tropical and subtropical seas' (642). [As Jania sp. (/. adhaerens Lamouroux?)] Senegal (529). 'Atlantique tropicale' (529). [As Corallina adhaerens (Lamouroux) Kiitzing] Cameroun (454; 500). Jania capillacea Harvey Ascension (475). Canaries (38B; 38D; 38C; 555; 556; 598; 602; 621). Cape Verde Islands (38B; 38C; 38D; 100; 183; 191; 555; 556). Gabon (294; 350; 586). Ghana (377; 586). Mauritanie (38C; 624). Salvage Islands (38B; 38C; 38D; 555; 556; 598). '. . .in tropical parts of the Atlantic and Pacific Oceans' (350; 586). '. . . widespread in tropical and subtropical seas' (642). [As Jania capillacea Lamouroux] Canaries (18; 582). Gulf of Guinea (582). Jania corniculata (Linnaeus) Lamouroux Canaries (38D; 227; 582; 584; 602). Salvage Islands (598). Note. For an additional Canaries record possibly attributable here, see Jania rubens (Linnaeus) Lamouroux f. corniculata. 136 PRICE, JOHN & LAWSON Jania crassa Lamouroux Angola (352; 541). Cape Verde Islands (641). Cote d'lvoire (295; 350; 586). Gabon (295; 350; 586). Ghana (295; 350; 586). Tropical Africa (N. Gambia — Congo river)' (598). 'widespread in warm temperate and tropical seas' (350; 586). [As Jania natalensis Harvey] Gabon (498; 535). Ghana (338; 537; 541). '. . . Westafrika (Gabun) bis zum Kap der Guten Hoffung' (498; 499; 535). Jania cubensis Montagne ex Kutzing See Haliptilon cubensis (Montagne ex Kutzing) Garbary & Johansen. Jania fastigiata Harvey See Jania verrucosa Lamouroux. Jania granifera Decaisne See Corallina granifera Ellis & Solander and Haliptilon virgatum (Zanardini) Garbary & Johansen. Jania longifurca Zanardini Congo (249; 250; 350; 586). Mauritanie (252; 349; 359). Senegal (252). 'Atlantico (desde la costa vasca a Mauritania)' (517). '. . . Atlantique (de la cote basque a la Mauritanie)' (188). '. . . Subtropical Africa [Senegal (N. of Gambia); Maurita- nia, Former W. Sahara]' (598). [As Corallina (Jania) longifurca] Congo (535). Jania micrarthrodia Lamouroux Canaries (541). Note. It has not been possible to locate authenticating material for this record. Jania natalensis Harvey See Jania crassa Lamouroux. Jania pumila Lamouroux Ascension (474). Canaries (5; 16; 70; 71; 191; 227; 235; 238; 375; 582; 598; 602; 610). St Helena (541). Salvage Islands (38B; 231; 375). '. . . pantropical. . .' (610). Note. No authenticating material can be traced for the record presented in 541 (St Helena). Determination of Canaries material reported in 602 was provisional since the reduced-size plants lacked sufficiently definitive and well-documented characteristics, a situa- tion common to all species in the genus (see 602: 20). Jania rubens (Linnaeus) Lamouroux Ascension (37; 474; 475). Cameroun (213; 350; 454; 500; 586). Canaries (2; 5; 8; 9; 13; 16; 38D; 70; 108; 128A; 175; 191; 213; 225; 226; 227; 228; 229; 230; 236; 237; 238; 252; 302; 304; 306B; 351; 356; 359; 379; 386; 392; 414; 439; 489; 490; 517; 546; 548; 556; 582; 584; 594; 598; 602; 604; 605; 610; 621). Cape Verde Islands (37; 38D; 213; 252; 441; 556; 598; 620). Gambia (296; 350; 586). Ghana (299; 300; 338; 340; 344; 350; 376; 377; 491; 567; 586). Mauritanie (38D; 252; 349; 359; 556; 624). Nigeria (213; 350; 586). Salvage Islands (38B; 38D; 231; 375; 556; 598). Sao Tome (350; 535; 586;). Senegal (38D; 59; 213; 556). 'Subtropical Africa [Senegal (N. of Gambia); Mauritania; Former W. Sahara]' (598). Tropical Africa (N. Gambia — Congo river)' (598). 'Atlantique depuis la Norvege jusqu'au littoral de 1'Afrique du Nord et les Canaries; sud de 1'Afrique' (356). '. . . Atlantique: des cotes anglaises a la Mauritanie. . .' (222). '. . . Atlantic Ocean (European, African and American coasts, Canary Islands. . .)' (177). '. . . Atlantic coast from Norway to the Canary Islands. . .' (70). '. . . Atlantico de Noruega a las islas Canarias. . .' (517). '. . . Atlantique (de la Norvege a la Mauritanie, Canaries. . .)' (33). 'Atlantico (de Noruega a Mauritania. . .)' (620). 'Gulf of Guinea' (582). '. . . widespread in warm temperate and tropical seas' (350; 586). '. . . widespread in warm temperate to subtropical seas' (642). '. . . cosmopolite. . .' (59). [As Corallina rubens Ellis & Solander] Canaries (236). Note. The juxtaposition of records under both Jania rubens and Corallina rubens Ellis & Solander in 236 may imply that female material recorded as C. rubens represented a different taxon. Nomenclatural attribution suggests both are the same species. [As Corallina rubens Linnaeus] Cameroun (239; 454). Cape Verde Islands (483). '. . . alien warmaren Meeren' (483). 'An der afrikanischen Kiiste um Marokko und den Atlantis- chen inseln bis Kamerun. . .' (239). '. . . De Norvege aux Canaries. . .' (89). [As Corallina (Jania) rubens Linnaeus] Cape Verde Islands (38). '. . . De la Norvege aux Canaries. . .' (38). [As Jania rubens Lamouroux] Canaries (439; 499; 547). Cape Verde Islands (445; 446; 499). '. . .In den warmeren Teilen aller Ozeane. . .' (499). '. . . Nordwestafrika' (499). [As Jania rubens (Linnaeus) Lamouroux forma corniculata] Canaries (70). [As Jania rubens (L.)] Canaries (3). Overall Note. See also the notes to Jania verrucosa Lam- ouroux. Jania tenella Kutzing Cape Verde Islands (150; 598; 541). [As Corallina (Jania) tenella Kutzing] Cape Verde Islands (38). Jania verrucosa Lamouroux Cape Verde Islands (641). Cote d'lvoire (350; 586). Ghana (350; 376; 377; 586; 590). Liberia (350; 586). Mauritanie (624). RHODOPHYTA (FLORIDEAE) OF TROPICAL AFRICA Sierra Leone (295; 350; 586). Tropical Africa (N. Gambia — Congo river)' (598). '. . . widespread in warm temperate and tropical seas' (350- 586). [As Jania fastigiata Harvey] Angola (352). Cote d'lvoire (287; 295). Ghana (295; 590). Liberia (129; 287; 295). [As Jania rubens (Linnaeus) Lamouroux] Nigeria (Herb. BM). Sao Tome (535). Note. The Mauritania record in Marcot-Coguengniot (624: 198) expressed doubt as to the specific determination. It may well be that material from Nigeria collected by Steentoft and examined by Johansen contains both /. verrucosa (as 'fastigiata prox.') and /. rubens (specimen clump in BM). See Steentoft (535: 127-8) and Lawson & John (350: 230) for additional data on the taxa and distinctions between them. Jania spp. Angola (352). Canaries (5; 175; 229; 232B; 301; 302; 303; 393; 394). Cape Verde Islands (423). Cote d'lvoire (287). Ghana (290; 297; 340; 344; 376; 377; 487; 567). Liberia (287). Senegal (50; 399). 'West Africa' (290; 344). Kallymenia dentata (Suhr) J. Agardh See Anatheca montagnei Schmitz. Kaliv iiienia feldmannii Codomier Canaries (635). Kallymenia microphylla J. Agardh Canaries (191; 273; 392; 584). [As Callymenia microphylla J. Agardh] Canaries (70; 177). [As Callymenia (Meredithia) microphylla J. Agardh] Canaries (493). Note. Both Sauvageau (493: 184) and B0rgesen (70) expressed doubt on the exact nature of their material due to its juvenile state, poor development, scarcity or sterility. If correctly determined, all records should appear under Meredithia microphylla (J. Agardh) J. Agardh. Kallymenia perforata J. Agardh Canaries (598). Note. According to Schneider & Searles (642: 285-6), the mate- rial from North Carolina named thus has subsequently proved to relate to K. westii Ganesan, described from Venezuela. Ganesan (1976) suggested, when describing the latter, that all Caribbean vegetative material might really be K. westii, K. perforata being restricted to the Indian and Pacific Oceans. This possibility applies to eastern as well as to western Atlantic material — Prud'homme van Reine (pers. comm.), whose records in 598 are the basis for this entry, certainly considers so. Kallymenia reniformis (Turner) J. Agardh Canaries (191; 227; 273; 547; 598). Cape Verde Islands (598) [As Callymenia reniformis Agardh] Canaries (70; 191; 547). Note. B0rgesen (77: 230) confirmed the determination of Vick- ers's specimens. Much of the material detected (70; 77) has been small and sterile. 137 [As Callymenia reniformis (Turner) J. Agardh] Canaries (77; 499). '. . . in oceano Atlantico caldiore. . .' (131). 'Nordwestafrika' (499). Kallymenia requienii J. Agardh Canaries (635). Kallymenia schizophylla J. Agardh Cape Verde Islands (36B; 191; 598). Namibia (36B; 348). [As Callymenia schizophylla (Harvey) J. Agardh] Cape Verde Islands (38; 117; 118; 119; 123). Senegal (123). '. . . communes aux iles du Cap Vert et a 1'Afrique meridionale. . .' (38) [As Kallymenia schizophylla Harvey] Cape Verde Islands (145). Kallymenia westii Ganesan See the note to Kallymenia perforata J. Agardh. Kallymenia spp. Namibia (348). [As Callymenia sp.] Senegal (529). ACKNOWLEDGEMENTS. We acknowledge with thanks assistance from many sources, of which the principal are: (i) The Natural History Museum, Department of Botany, London, for provision of the necessary research facilities to continue the series of papers of which the present is part; (ii) Dr W. J. Woelkerling for helping us with critical aspects of genera of crustose algae (especially Hydrolithon) ; (iii) the editorial expertise of Dr. R. Huxley for detecting inaccura- cies in the submitted text; (iv) a grant (for GWL) towards this work from the Systematics Association; (v) Dr W.F. Prud'homme van Reine for additional unpublished data in the form of records and comments, as well as for partial revision of the manuscript. 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Gracilaria mixta, sp. nov. and other western Pacific species of the genus (Rhodophyta: Gracilariaceae). Pacific Sci. 45: 12-27. Acuna Gonzales, A. 1970. Algunos aspectos de la vegetaci6n submarina de las Islas Canarias. Vieraea [I]: 2-5. 1972 ['1968-70']. Cinco nuevas citas de algae Rhodophyceae en la Isla de Tenerife. Anales Univ. La Laguna 7: 3-6. Acuna G[onzales,], A. Santos G[uerra], A. & Wildpret [de la Torre], W. 1970. Algunos aspectos de la vegetacidn algal de la Playa de San Marcos, Icod, Tenerife. Cuad. Bot. canaria 9: 30-36. Adey, W.H. & Lebednik, P.A. 1967. Catalog of the Foslie Herbarium. Trondheim, Norway: Det Kongelige Norske Videnskabers Selskab Museet. pp. [l]+92. , Townsend, R.A. & Boykins, W.T. 1982. The crustose coralline algae (Rhodophyta: Corallinaceae) of the Hawaian Islands. Smiths. Contrib. mar. Sci. 15: i-iv+1-74. Afonso-Carrillo, J. 1980a. Algunas observaciones sobre la distri buci6n vertical de las algas en la Isla del Hierro (Canarias). Vieraea 10(1-2): 3-16. 1980b. Nota sobra algunas Corallinaceae (Rhodophyta) nuevas para la flora ficologica de las Islas Canarias. Vieraea 10(1-2): 53-58. 1985. Conexiones intercelulares entre diferentes talos de Neogoniolithon absimile (Foslie et Howe) Cabioch (Corallinaceae, Rhodophyta). Vieraea 15: 139-142. & Gil-Rodriguez, M.C. 1980. Datos para la flora marina de la Isla de Fuerte ventura. Vieraea 10(1-2): 147-170. & Gil-Rodriguez, M.C. 1981. Sobre el limite meridional de Sauvageau- gloia chordariaeformis (Crouan) Kylin (Chordariaceae, Phaeophyta). Inves- tigacidn pesq. 45: 297-300. , Gil-Rodriguez, M.C. & Wildpret de la Torre, W. 1979 ['1978']. Estudio de la vegetaci6n algal de la costa del future poligono industrial de Granadilla (Tenerife). Vieraea 8: 201-242. , GU-Rodrfguez, M. C. & Wildpret de la Torre, W. 1985 ['1984']. Algunas consideraciones floristicas, coro!6gicas y ecoldgicas sobre las algas Coralli- naceae (Rhodophyta) de las Islas Canarias. An. Biol. Fac. Biol. Univ. Murcia 2 (Secc. Esp. 2): 23-37. , Gil-Rodriguez, M. C., Haroun Tabraue, R., Villena Balsa, M. & Wildpret de la Torre, W. 1984 ['1983']. Adiciones y correcciones al catalogo de algas marinas bentonicas para el Archipielago Canario. Vieraea 13: 27-49. Agardh, J.G. 1851. Species genera et ordines algarum, . . . floridearum. . . 2(1). Lund. Note. Facsimile reprint, J. Cramer, 1977. 1852. Species genera et ordines algarum, . . . floridearum, . . . 2(2). Lundae. Notes. 1. The Corallineae (Ordo XII. pp. 506-576) was by I.E. Areschoug — so stated in volume. 2. Some versions of this second volume, Pars II, were issued as two separate texts — Pars 11:1 being dated 1851 and Pars 11:2 1852. 3. There are also internal differences of numbering of pages between copies. BM copy is numbered straight through from 337-720, including therein the Addenda [701-706] and Index [707-720]. In the copy from which the Cramer (1977) reprint was prepared, the Index [unnum- bered, of 14 sides] is placed immediately after p. 700 and is followed by six sides [also unnumbered] of Addenda. Content of these unnumbered sides is exactly as the numbered BM pages. The BM copy of Vol. 2 Part 3 (1863) commences with pages headed 'Ordo XIV. Wrangelieae', and numbered 701-715; these are followed by 'Ordo XV. Chondrieae' [pp. 716-786]. All these latter pages [701-786], also so numbered, are in the Cramer (1977) version placed immediately after the unnumbered Addenda pages (see above) and before the title page to Vol. 2, Part 3, of 1863, thereby implying that the copy facsimiled was also so arranged. Pages from 787 to 1291 are in both cases in Vol. 2, Part 3, 1863. Despite this, the Index in the end of the Cramer (1977) facsimile of Vol. 2, Part 3 (pp. 1279-1291) indicates the same page numbers as does the BM version. Both dated texts are indicated as the sources of data where the records occur in pp. 701-786 [textual pages]. The implication behind all this is that there may be yet other differently paged versions elsewhere. 1863. Species genera et ordines algarum. .. .floridearum . . . 2(3). Lundae. 1876. Species genera et ordines algarum ... 3: De Florideis curae posteriores. Part 1. Epicrisis systematic floridearum . Lipsiae. Aleem, A. A. 1978. A preliminary list of marine algae fromSierra Leone. Botanica mar. 21: 397-399. Apt, K.E. 1987. A new species of Janczewskia (Rhodomelaceae, Rhodophyta) from the Hawaiian Islands. Phycologia 26: 328- 333. Ardre, F. 1959. Un interessant Hildenbrandtia du Portugal. Revue algol. ser., 4: 227-237. 1970 ['1969-70']. Contribution a 1'etude des algues marines du Portugal I— La Flore. Port. Acta biol. B, 10 137-555 + [56]. Note. The reprint is paged l-423+[56]. Askenasy, E. 1888 ['1889']. Algen, mit Unterstutzung der Herren E. Bornet, A. Grunow, P. Hariot, M. 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Die Meeresalgen der deutschen Tiefsee — Expedition 1898-1899 [pp. 549-586+[8], pis LV-LVIII]. In C.Chun, Wissenschaftliche Ergebnisse der deutschen Tiefsee — Expedition auf dem Dampfer 'Valdivia' 1898-1899. Bd. H, Teil II, Lieferung IV. pp. 549-586+[8]+pls LV-LVIII. Jena. 1908 ['1928']. Die Meeresalgen der Deutschen Sudpolar-Expedition 1901-1903 [Heft II, 1. pp. [2] + 179 +202]. In E. von Drygalski, Deutsche Sudpolar-Expedition 1901-1903 im Auftrage des Reichsministeriums des Innern 8 Botanik. pp. [4] + 178+[2] + 179-372+[12]+373-715+[27]. Berlin & Leipzig. Note. 1928 is the publication date of the whole volume and of the last part; the earlier parts were published separately at various dates as completed (1906; 1908; 1911; 1924; 1928). Rengasamy, R. & Ilanchelian, K. 1988. Seasonal studies on Hypnea valentiae (Turn.) Mont. (Gigartinales, Rhodophyta) at Muttukadu near Madras. Seaweed Res. Utiln 11: 21-26. Ribera Sigirfn, M. A., Gomez Garreta, A. & Seoane-Camba, J. A. 1985 ['1984']. 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Rhodomelaceae [pp. 421-480; Lief. 149/150]. In A. Engler & K. Prantl, Die naturlichen Planzenfamilien. . ., I Teil. 2. Abteilung. pp. XII+580. Leipzig. 1890-1897. & Hauptfleisch, P. 1896-1897a. Sphaerococcaceae [pp. 382-396]. In A. Engler & K. Prantl, Die naturlichen Pflanzenfamilien. . ., I Teil. 2. Abteilung. pp. XH+580. Leipzig. 1890-1897. & Hauptfleisch, P. 1897b. Rhodymeniaceae [pp. 396-405; Liefs. 149 & 150]. In A. Engler & K. Prantl, Die naturlichen Pflanzenfamilien . . ., I Teil. 2. Abteilung. pp. XII + 580. Leipzig, 1890-1897. & Hauptfleisch, P. 1897c. Grateloupiaceae. [pp.508-514]. In A. Engler & K. Prantl, Die naturlichen Pflanzenfamilien. . . I. Teil. 2. Abteilung. pp. XII + 580. Leipzig, 1890-1897. Schneider, C.W. 1975. North Caroline marine algae. V. Additions to the flora of Onslow Bay, including the reassignment of Fauchea peltata Taylor to Weberella Schmitz. Br. phycol. J. 10: 129-138. & Searles, R.B. 1991. Seaweeds of the Southeastern United States Cape Hatteras to Cape Canaveral, pp. xi+[2]+553+[l]. Durham, North Carolina & London. Searles, R.B. 1968. Morphological studies of red algae of the order Gigartina- les. Univ. Calif. Publns. Bot. 43: vi+[2]+ 1-100. Seagrief, S.C. 1984. A catalogue of South African green, brown and red marine algae. Mem. hot. Soc. S. Africa 47: i-iv+1-72. Seoane-Camba, J. 1960. Nota sobre algunas especies de algas de la costa occidental africana (sur de Cabo Blanco). Investigacion pesq. 16: 91-103. 1965. Estudios sobre las algas bent6nicas en la costa sur de la Peninsula Iberica (litoral de Cadiz). Investigacion pesq. 29: 3-216. Simons, R.[H]. 1974. Algae, (including diatoms and seaweeds), [pp. 239-261]. In J.H. Day, N.A.H. Millard, & M.-L. Penrith (eds), A Guide to marine life on South African shores. Second (revised) edition, pp. [4]+i-iii+[l]+300. Cape Town & Rotterdam. Note. The first edition (not seen) was probably 1969, since that is the copyright date and the preface is dated April 1968. Overall tripartite editorship is obvious from both copyright allocation and statement in preface. Latter also clearly indicates responsibility of R.H. Simons for Algae section. Presumably algal content of first edition was as in second, since preface statement unchanged from past. Sender, [O.W.] 1852. Algae [pp. 125-127]. In J.A. Schmidt, Beitrage zur Flora der Cap Verdischen Inseln. Mil Berucksichtigung aller bis jetzt daselbst bekannten wildwachsenden und kultivirten Pflanzen. Nach eigenen Untersu- chungen und mil Benutzung der gewonnenen Resultate anderer Reisenden. [2]+viii + l-356+[l] pp. Heidelberg. Note. The algal material was sent to Sender in Hamburg; he worked on the plants and provided both the determinative data and text as printed. Schmidt (the overall editor) added a small terminal section with Cape Verde Islands records. Sourie, R. 1954a. Contribution a 1'etude ecologique des cotes rocheuses du Senegal. Mem. Inst. fr. Afr. noire 38: l-342+[l]. Note. From the note on p. 117, it is clear that the algae were worked on mainly by J. Feldmann, but that Sourie took account of some of the views of Dangeard as expressed in the latter's memoir on the Cap Vert (Dakar) peninsula algae. Since the exact contribution of the various people involved is in doubt, we have left the reference in the name of Sourie, who seems to have exercised overall authorship. 1954b. Etude ecologique sommaire des fond sableux en Baie de Dakar. Annls EC. sup. Sci., Dakar 1: 141-155. Note. Sourie stated (p. 141) that many of the specific determinations of algae were by J. Feldmann. Steentoft, M. 1967. A revision of the marine algae of Sao Tome and Principe (Gulf of Guinea). J. Linn. Soc. (Bot.) 60: 99-146. Stephenson, T.A. & Stephenson, A. 1972. Life between tidemarks on rocky shores. xii+[2]+425 pp. San Francisco. Suhr, J.N. von 1831. 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The background to this work is explained in detail by Martin Aguado (1957) who outlined (p. 8) the career of Viera y Clavijo and the progress of his work. The MS was completed in 1799, with the title Diccionario de Historia Natural de las Canarias, but was not published until the indicated dates. See also references 549 and 386. A further version of the work appeared in 1942 under the 'Publicaciones de la Biblioteca Canaria' series (Tenerife). A more recent (1982) new edition was edited by M. Alvar and included an introduction and appendix, with purely historical literary data. Viera-Rodriguez, M.A. 1987. Contribuci6n al estudio de la florula bent6nica de la isla de La Graciosa. Canarias. Vieraea 17: 237-259. Collections from Canary Islands, determined by collector and pers. comm. by W.F. Prud'homme van Reine to D.M. John, 19/11/91. , Gil-Rodriguez, M.C., Audiffred, P.A.J., Prud'Homme Van Reine, W.F., Haroun-Tabraue, R. & Wildpret de la Torre, W. 1987. Contribuci6n al estudio de la fl6rula bent6nica del islote de Montana Clara. Canarias. Vieraea 17: 271-279. Villena-Balsa, M., Afonso-Carrillo, J. & Wildpret de la Torre, W. 1987. Morfologia, estructura y reproducci6n de una especie epffita del g6nero Janiii (Corallinaceae, Rhodophyta) de las Islas Canarias. Vieraea 17: 19—42. Weber-van Bosse, A. 1921. Liste des algues du Siboga II Rhodophyceae Premiere Partie Protoflorideae, Nemalionales, Cryptonemiales. In M. Weber, Siboga-Expeditie . . . Monographic LIXb. Uitkomsten op zoolo- gisch, botanisch, oceanographisch en geologisch Gebied . . ., Livr. LXXX1X. pp. [6] + 187-392+[4]. Weisscher, F.C.M. 1982. Marine algae from Ilheu de Fora (Salvage Islands). Bol. Mus. munlc. Funchal 34: 23-34. 1983. Marine algae from Selvagem Peguena (Salvage Islands). Bol. Mus. munic. Funchal 35: 41-80. Woelkerling, W.J. 1985. A taxonomic reassessment of Spongites (Coralli- naceae, Rhodophyta) based on studies of Kiitzing's original collections. Br. phycol. J. 20: 123-153. Wynne, M.J. 1985. ['1984') Notes on Herposiphonia (Rhodomelaceae, Rhodophyta) in South Africa, with a description of a new species. Cryptoga- mie: algologie 5:167-177. 1985. Concerning the names Scagelia corallina and Heterosiphonia wurdemannii (Ceramiales, Rhodophyta). Cryptogamie: algologie. 6: 81-90. 1986. A checklist of benthic marine algae of the tropical and subtropical western Atlantic. Can. J. Bot. 64: 2239-2281. 1986. Report on a collection of benthic marine algae from the Namibian coast (southwestern Africa). Nova Hedwigia 43: 311-355. 1989. The re-instatement of Hydropuntia Montagne (Gracilariaceae, Rhodophyta). Taxon 38: 476-479. & Kraft, G.T. 1985. Hypoglossum caloglossoides sp. nov. (Delesseri- aceae, Rhodophyta) from Lord Howe Island, South Pacific. Br. phycol. J. 20: 9-19. & Morris, R.E. 1991. Branchioglossum pygmaeum sp. nov. and new records of other delesseriaceous algae (Rhodophyta) from Natal, South Africa. Phycologia 30: 262-271. Yarish, C., Breeman, A.M. & Hoek, C. van den 1985? ['1984']. Temperature, light, and photoperiod responses of some Northeast American and West European endemic rhodophytes in relation to their geographic distribution. Helgoldnder Meeresunters. 38: 273-308. Bull. Br. Mus. not. Hist. (Bot.) 22(2): 147-152 Issued 26 November 1992 Two new species of Solanum section Geminata (Solanaceae) from Cerro del Torra in western Colombia SANDRA KNAPP Department of Botany, British Museum (Natural History), Cromwell Road, London SW7 5BD SYNOPSIS. Two new species of Solanum section Geminata (G. Don) Walp. are described from the Choco floristic province in western Colombia. Solanum cyclophyllum and 5. nematorhachis are both found on the peaks of the Serrania de los Paraguas on the Choc6/Valle de Cauca border. Relationships of each of the species are briefly discussed. Recent collecting in the Serrania de los Paraguas on the Choco/Valle de Cauca border has brought to light two new and interesting species of the large and diverse Solanum section Geminata (G.Don) Walp. These are described here so the names can be used in floristic and monographic works. NEW SPECIES DESCRIPTIONS 1. Solanum cyclophyllum S. Knapp, sp. nov. Species Sol. unifoliato S. Knapp af finis, sed foliis late ovatis, venatione camptodroma nervis ad basin 3-5 jugis differt. Type. Colom- bia. Choco: San Jose del Palmar, Cerro del Torra, base of E. slope, right edge of heliport, 2000 m, 31 Aug 1988, Ramos et al. 1663 (CUVC-holotype; IBE, MO, NY-isotypes). (Fig. 1) Shrubs 1-3 m tall; young stems and leaves glabrous; bark of older stems pale yellowish-grey. Sympodial units unifoliate. Leaves broadly elliptic to orbicular, widest at or just below the middle, glabrous and shining adaxially, glabrous and drying somewhat golden abaxially, with 3-9 pairs of primary veins, the secondary venation conspicuously parallel, lamina 11-21 cm long, 6-19 cm wide, the apex acute, the base acute to rounded; petioles 1-2 cm long. Inflorescences opposite the leaves or sometimes internodal, simple, 3-10-flowered, gla- brous except for a few reddish papillose trichomes on the tips of the axes; pedicel scars closely packed, not overlapping. Buds ovoid with pointed tips, c. 1/2 exserted from the calyx tube. Pedicels at anthesis deflexed, glabrous, 5-7 mm long, c. 0.5 mm in diameter at the s deltate to long-triangular, 1-1.5 mm long, glabrous; corolla greenish-white to bright green (fide Ramos et al.), 0.7-1 cm in diameter, lobed c. 3/4 of the way to the base, the lobes more or less planar at anthesis, the tips and margins of the lobes minutely papillose; anthers 2-2.5 mm long, c. 1 mm wide, poridical at the tips, the pores lengthening to slits with age; free portion of the filaments 0.5-1 mm long, the filament tube 0.25-0.5 mm long, glabrous; ovary glabrous; style straight, 4-5 mm long, glabrous; stigma capitate, the surface minutely papillose. Fruit a globose, green to greenish-white berry, 1-1.5 cm in diameter; fruiting pedicels woody, erect, 1-1.5 cm long, 1.5-2 mm in diameter at the base. Seeds dark brown, ovoid- reniform, 4-4.5 mm long, 2-2.5 mm wide, the surfaces minutely pitted. Chromosome number not known. DISTRIBUTION. Western Colombia in the departments of Choco and adjacent Valle de Cauca, 800 to 2000 m, in wet cloud forest (Fig. 2). SPECIMENS EXAMINED. Colombia. Choco: Novita, N. slope of Cerro del Torra, ridge to W. of Rio Surama, trail to Alto de Oso, 600-900 m, 22 Feb 1977, Forero et al. 3175 (COL, MO); San Jose del Palmar, Cerro del Torra, E slope, near edge of heliport, 1600 m, 26 Aug 1988, Ramos et al. 1603 (CUVC, MO, NY). Valle: Bosque de San Antonio, W. of Cali, near television tower, 1950-2050 m, 15 Jul 1984, Gentry et al. 48121 (MO, NY); old road to Buenaventura kms 14-18, above Saladito, c. 1800 m, 21 Oct 1987, Ramos 803 (CUVC, MO, NY); El Cairo, Cerro del Ingles, Serrania de los Paraguas, Cordillera Occidental, Valle-Choco border 1 hour by jeep from El Cairo, 2070-2400 m, 1 Apr 1988, Silverstone- Sopkin et al. 3946 (CUVC, MO, NY). Solanum cyclophyllum is a member of the unifoliatum species group, a small group of 6 species confined to the Choco floristic province. It is closely related to both S. unifoliatum S. Knapp of the Rio Atrato drainage and 5. triplinervium Morton of Isla Gorgona and in some ways is almost intermediate between them. Solanum cyclophyllum has leaves with three to five main veins from the base, like S. triplinervium, but lacks the distinctive almost Melastomata- ceous secondary venation of that species. The broadly elliptic to orbicular leaves of 5. cyclophyllum (from which the epithet is derived) immediately distinguish it from 5. unifoliatum. 2. Solanum nematorhachis S. Knapp, sp. nov. Species Sol. leptorhachidi Bitter similis, sed foliis pubescentibus, inflores- centiis et floribus maioribus differt. Type. Colombia. Choco: San Jose del Palmar, Cerro del Torra, E. slope between the heliport and the summit, 2000-2400 m, 28 Aug 1988, Silverstone-Sopkin et al. 4846 (CUVC-holotype; CHOCO, MO, NY-isotypes). (Fig. 3) Shrubs, 1-3 m tall; young stems and leaves densely pubescent 148 S. KNAPP HOLOTYPE Systematic Studies in Solanum Solanum cyclophyllum S. Knapp Sandra Knapp (I BE) 1991 FIIAWIADO BOS HA* PIANTAS DE COLOMB HERBARIO (CUVCJ, CAil Fanritte; Cerro del forr^t vertleats ert del agua qo« toaja d*l Oeryo, .'• t puerto. Arbolito ca. 3 a «lturaj emrSi sSpaloo verde claros, hordes i lias* fruto inaaduro glotoso. : noo pfllidas ajnarillentaa; garganta oon tricomas verdo- SOB; ovario, eatilo y estiRma blancosf fruto Imaduro verde claro, blanouzcof sendllas COB un lado d« 2 coeti- llas. Ail.. 1600 a Col.. Jorge E. Samoa, No.: 1603 P.Silveratone, et al. 150 S. KNAPP S. cyclophyllum • S. nematorhachis D r» I \ Fig. 2 Distribution of 5. cyclophyllum and 5. nematorhachis. with golden curl-tipped trichomes c. 0.5 mm long; bark of older stems dark brown, glabrous or only sparsely pubescent. Sympodial units unifoliate. Leaves elliptic, with 8-9 pairs of brochiodromus primary veins, glabrous but with a few scat- tered golden uniseriate trichomes along the midrib adaxially, sparsely to densely pubescent with golden trichomes along the main veins abaxially, 8.5-17 cm long, 2.5-8 cm wide, the apex acuminate, the base acute, not winged onto the petiole; petiole c. 0.5 mm long. Inflorescences opposite the leaves or internodal, simple, glabrous to sparsely puberulent at the tip, 6-17 cm long, 30-60-flowered, but bearing only two or three flowers at a time; pedicel scars widely and unevenly spaced 4-5 mm apart. Buds globose, the corolla exserted from the calyx tube. Pedicels at anthesis deflexed, glabrous, filiform, 5-8 mm long, c. 0.25 mm in diameter at the base. Flowers with the calyx tube conical to somewhat cyathiform and swollen (Cerro Ingles), 0.5-1.5 mm long, the lobes deltoid, c. 0.5 mm long, glabrous or sparsely pubescent with uniseriate trichomes like those of the stems and leaves; corolla white, 7-8 mm in diameter, lobed nearly to the base, the lobes more or less reflexed at anthesis, minutely papillose at the tips and margins; anthers 1.5-2.5 mm long, c. 1 mm wide, poricidal at the tips, the pores tear-drop-shaped; free portion of the filaments c. 0.5 mm long, the filament tube c. 0.25 mm long; ovary glabrous; style straight, glabrous, c. 4 mm long; stigma capitate, the surface minutely papillose. Fruit a globose, green berry, c. 1.5 cm in diameter when fresh, 1-1.2 cm in diameter dried; fruiting pedicels woody, deflexed, 1.5-2 cm long, c. 1.5 mm in dimaeter at the base, 4 mm in diameter at the apex. Seeds dark brown, ovoid-reniform, 3.5-4 mm long, 2.5-3 mm wide, the surfaces smooth to minutely pitted. Chromosome number not known. DISTRIBUTION. In the Serrania de los Paraguas on Cerros Torra and Ingles, on the Choco/Valle border from 2000 to 2400 m (Fig. 2). SPECIMENS EXAMINED. Colombia. Choco: San Jose del Pal- mar, Cerro del Torra, W. slope, below the heliport at base of cerro, Rio Negro drainage, 1680-1940 m, 10 Aug 1988, Ramos et al. 1111 (CUVC, MO, NY). Valle de Cauca: S. slope of Cerro Ingles (Cordillera Occidental, Serrania de los Paraguas), 1 hour in jeep from El Cairo, 2260 m, 5 Jan 1987, Silverstone-Sopkin et al. 2975 (CUVC, MO, NY). Solanum nematorhachis is very similar to and probably closely related to 5. leptorhachis Bitter of low to middle elevation western Ecuador. It differs from S. leptorhachis in its pubescent leaf surfaces and inflorescences, white flowers, and much shorter fruiting pedicels. Solanum tenuiflagellatum Bitter ex S. Knapp of montane coastal Venezuela also has elongate inflorescences and pubescent leaves, but its flowers are much smaller and the fruiting pedicels are more elongate than those of either 5. leptorhachis or 5. nematorhachis. The extreme length of the inflorescence in S. nematorha- chis may be related to its fruit dispersal. The hard green fruits of many species of section Geminata are dispersed by bats and are taken on the wing. The long inflorescence effectively extends the fruiting pedicel length, perhaps making the fruit itself easier to find in the dense rainforest understory. Only one or at most two fruits seem to develop on each inflores- cence in 5. nematorhachis (also in 5. leptorhachis and 5. tenuiflagellatum), perhaps suggesting an andromonoecious breeding system (Whalen & Costich, 1986). This however, as in many other species of section Geminata (Knapp, 1986b; 1991), needs to be tested in the lab and in the field. ACKNOWLEDGEMENTS. I thank the curators of CUVC for their kind loan and gifts of material of these species and the curators of other herbaria mentioned in the text for loan of material. Specimens from Cerro del Torra were collected by staff of CUVC on an expedition financed by National Geographic Grant no. 388-88. REFERENCES Knapp, S. 1986a. A revision of Solanum section Geminata (G.Don) Walpers. Ph.D. thesis, Cornell University. 1986b. Reproductive biology of Solanum section Geminata in a Costa Rican cloud forest. In: W.G. D'Arcy (Ed.). Solanaceae: Systematics and Biology: 253-263. New York. 1991 . A revision of the Solanum sessile species group (section Geminata proparte: Solanaceae). Bot. J. Linn. Soc. 105: 179-210. Whalen, M.D. & Costich, D.E. 1986. Andromonoecy in Solanum. In: W.G. D'Arcy (Ed.). Solanaceae: Systematics and Biology: 284-302. New York. TWO NEW SPECIES OF SOLANUM SECTION GEM IN ATA 151 Systematic Studies in Solanum Solatium nematorhachis S. Knapp Sandra Knapp (IBE) 1991 'ADO FOR RATIONAL GEOGRAPHIC SOCIETY GHANT FLANTAS DE COLOMBIA # 3884-88 Depto.i Ctaocdj Municlpio San JosS del Palmar Hombre « Solanum 30LAKACEAE Arbusto oa. 3 m alturaj hojaa aubcorlaceas, haz verde oscuro, euvfis venle ciaro; flores pgndulasj corola blunca, 16buios reflexosj anteraa axarillas; eatilo co; frutos pendulos, verdea, globosos, el mas gran- de 1. '> etc longitud X 1,5 cm ancho, Fechai ^ ue ii;.:iiio de 19t-i8 Felipe A. Silverstone-Sopkin Mo. 4846 • f . • • • .-intjOo, L. Hatnos, A. Henao Fig. 3 Holotype of Solanum nematorhachis S. Knapp (Silverstone-Sopkin el al. 4846, CUVC); (b) isotype of Solanum nematorhachis S. Knapp (NY) 152 S. KNAPP PM&NCJIABO POlt NATIONAL GJSOCritAl'HIC FLANTAS m COLOMBIA G^f #5884-88 Depto. i Jnoeo; I«unicipio aan Jose del Palmar J Torra, vertiente oriental, entre to y siesa, bosque nublado primario, ;:,-,:. v»;i'.:o Odcuro, envea verde pendulas; oorola blanoe.» lobu. ... i ut, a;:,arillas j estilo blai penduloa, verdes, globosoe, ei 1.5 cm longitud. x 1.5 cm anohi as, ISOTYPE Ho. 4! Systematic Studies in Solanum Solan urn nematorhachis S. Knapp Sandra Knapp 1 1 BE) 1991 Fechat 28 de agosto de 1988 Felipe A. Silvwrstone-Sopkin Fig. 3 (b) isotype of Solanum nematorhachis S. Knapp (NY) Bull. Br. Mus. not. Hist. (Hot.) 22(2): 153-156 Issued 26 November 1992 Fissidens ceylonensis Dozy & Molkenb. (Musci: Fissidentaceae) and some allied taxa from southern India L.T. ELLIS Department of Botany, The Natural History Museum, Cromwell Road, London SW75BD SYNOPSIS. Fissidens ceylonensis Dozy & Molkenb. is described and compared with some related taxa. F. ceylonensis var. acutifolius Dixon & P. Varde is considered to be a distinct taxon possibly warranting the rank of species, whereas F. pennatulus Thwaites & Mitten and F. perpusillus Wilson ex Mitten are regarded as forms of F. ceylonensis. The taxonomic status of F. karwarensis Dixon remains uncertain, its resolution will require the examination of a wider range of specimens. The late A.H. Norkett, an acknowledged authority on the moss genus Fissidens, undertook two expeditions to India (1966-1967, 1970-1971) during his career at the Natural History Museum (BM). Material of Fissidens ceylonensis Dozy & Molkenb. and closely related taxa, which he col- lected in southern India on the first of these expeditions, has formed the basis for a preliminary investigation into the interrelationships of these plants. Fissidens ceylonensis Dozy & Molkenb., Annls Sci. nat. Bot. ser. 3, 2: 304 (1844). Type: Sri Lanka [Ceylon], [KoenigJ s.n. (BM!,NY!— isotypes). Fissidens bicolor Thwaites & Mitten, /. Linn. Soc. Bot. 13: 322 (1873). Type: Sri Lanka [Ceylon], G. Thwaites [137b in Hb. Dixon] (BM!— isotype). Fissidens perpusillus Wils. ex Mitten, /. Linn. Soc. Bot. Suppl. 1: 141 (1859). Type: Sri Lanka [Ceylon], Gardner 630 (NY!— holotype; BM!— isotype). Fissidens thwaitesii Par., Index Bryol. 488 (1896). [F. papillosus Thwaites & Mitt., /. Linn. Soc. Bot. 13: 323 (1873), horn, illeg.] Type: Sri Lanka [Ceylon], G. Thwaites [137 in Hb. Dixon] BM!— isotype). Fissidens pennatulus Thwaites & Mitten, /. Linn. Soc. Bot. 13: 325 (1873). Type: Sri Lanka [Ceylon], G. Thwaites [136] (NY!— holotype; BM!— isotype). Plants forming pale yellowish-green to dark green, sparse to dense patches; shoots 2-6 (- 9) mm long, straight to curved, sometimes branched; stems in cross section elliptical, central strand barely perceptible, cells of cortex small and thick walled, 1-2 layers (those of the outer layer often protruding slightly from the stem surface). Leaves in about 6-18 (- 30) pairs, often slightly overlapping, straight when moist, hardly altered to curving ventrally when dry, mostly 0.8-1.2 x (0.15 -) 0.2-0.25 (- 0.3) mm, oblong-lanceolate with mucronate apices (Fig. la). Costa usually excurrent, occasionally percur- rent. Vaginant lamina about 3/5-4/5 of leaf length, unequal at apex (point of apex usually lying near to leaf margin), margins of vaginant lamina often limbate from base to apex or to near apex; limbidium sometimes poorly developed, or lacking (especially in lower leaves of the shoot), marginal for most of its length, composed of linear to sublinear cells, the outermost row frequently give rise to small, distant teeth; most leaves with a single row of small, quadrate to shortly rectangular (sometimes papillose) cells forming part of the margin along the mid to basal vaginant lamina and giving the limbidium an intramarginal appearance (Fig. Ik). Dorsal lamina narrowing gradually from above mid-leaf, usually tapering into costa at, or slightly above, the leaf base (some- times curving abruptly into the base of the costa). Margins of dorsal and apical laminae papillose-crenulate. Laminal cells mostly isodiametric with 4-6 sides, 5-10 (- 12.5) x 5-10 urn, multi-papillose, in the leaf base becoming shortly subrectan- gular and smoother. Autoecious (Dioecious in 'perpusillus' form): perichaetia terminal; male branches axillary on female shoots, normally diminutive with underdeveloped leaves or bud-like (Fig. 2a, b) , sometimes larger with leaves well developed (in 'perpusillus' form apparently independent of female shoots)(Fig. 2c, d), perigonia terminal. Seta 2-3 mm long, yellow to red; capsule erect to inclined, theca shortly cylindrical to ellipsoidal, symmetrical, 0.4-0.6 mm long, walls of epithecial cells thickened at angles; operculum with subu- late beak. Spores spherical, 15-17.5 mm diam., finely papil- lose. ECOLOGY AND DISTRIBUTION. Fissidens ceylonensis Dozy & Molk. occurs on sandy and clay soils, and has been collected on bare ground, soil overlying rocks, amongst the roots of trees and on the banks of streams. Shaded situations are favoured. F. ceylonensis (sensu lato) is well represented in southern India and Sri Lanka and the type variety, at least, appears to be generally distributed throughout tropical Asia. DISCUSSION. Fleischer (1904) and Norris and Koponen (1987) place Fissidens perpusillus in synonymy with F. ceylon- ensis, while Eddy (1988) retains it as a distinct species. The majority of specimens from the study area are easily recogn- isable as one or the other, but there are a significant number which possess ambiguous features and are difficult to place. A key difference used to differentiate the two taxa is that most leaves in specimens of F. ceylonensis possess vaginant laminae with limbidia; whereas in specimens of F. perpusillus similar limbidia occur only in the perichaetial and subpericha- etial leaves. Specimens with limbidia distributed as in F. ceylonensis often possess excurrent costae and dorsal laminae 154 L.T. ELLIS 50gm Fig. 1 Fissidens ceylonensis Dozy & Molkenb. a-c: leaves; d-f: limbidium in vaginant lamina (d: near apex, e,f: near base. Fissidens ceylonensis var. acutifolius Dixon & P. Varde. g: leaf; h: limbidium near base of vaginant lamina. Fissidens karwarensis Dixon. i: leaf from mid-shoot; j, k: limbidium near base of vaginant lamina (j: as frequently occurring, k: as very rarely occurring), a, d, e, Drawn from Sri Lanka, Alston 1769 (BM); b, from Sri Lanka. Thwaites 136 (type of F. pennatulus) (BM); g, h, from southern India, Foreau 216 (BM); i-k, from southern India, Sedgwick 6368 (lectotype of F. karwarensis) (BM). which gradually taper into the base of the costa. Many specimens in which the limbidia occur as in F. perpusillus , (the 'perpusillus' form), possess leaves with percurrent costae and dorsal laminae which curve abruptly into the base of the costa. However, all of these features can be rather variable and do not always occur together. In most material from the study area, identified as F. ceylonensis or F. perpusillus, the form of the male shoots is correlated to some extent with the distribution of limbidia. Plants exhibiting the features associ- ated with F. perpusillus tend to produce perigonia at the apex of small leafy shoots (occasionally comparable in size to female shoots); their attachment to the female shoots is difficult to establish. In specimens apparently referable to F. ceylonensis the perigonia occur as, or on, diminutive axillary branches. These are unequivocally attached to the female shoots and rarely produce fully developed leaves. Unfortu- nately, some specimens do not show this apparent correlation of features — for example, southern Indian specimens other- wise resembling F. perpusillus, such as Norkett 11369 (BM), which has large, leafy shoots with terminal (rarely lateral) groups of bud-like perigonia and solitary archegonia sparsely scattered on the upper stem (Fig. 2e). SELECTION OF HERBARIUM SPECIMENS EXAMINED. F. CEYLONENSIS (TYPICAL FORM). Southern India. Kerala State: Kerala, near Kellar, 1967, A.H. Norkett 12645 (BM!); 12 miles from Trivandrum, 1967, A.H. Norkett 12650 (BM!). Tamil Nadu, Courtallam, 1967, A.H. Norkett 13133 (BM!). Sri Lanka. Morowak Korale, Campein Hill Estate, 1890, A. Waddells.n. (BM!); 1928, A. Alston 1769 (BM!); Henerat- goda Gardens, A. Alston 1983. Malay Peninsula, Singapore, 1898, M. Fleischer exs. no. 74 (NY!). F. CEYLONENSIS ('PERPUSILLUS FORM'). Southern India, Tamil Nadu, Courtallam: 1966, A. H. Norkett 11340/11338 (BM!); near Old Falls, 4 miles south of Courtallam, 1966, A.H. Norkett 11369 (BM!); above Tourist Bungalow, 1967, A.H. Norkett 13105 (BM!); jungle area, 1967, A.H. Norkett 13118 (BM!); Nilgiri Mountains, Beddome 707 (NY!). Sri Lanka. Uva Province, Badulla District, between Ella and Wellawaya, south of Ravanala Falls, 1973, C.C. Townsend 1125, 1126 (BM!). Fissidens ceylonensis var. acutifolius Dix. & P. Varde, Rev. bryol. 52: 38 (1925). Type. Southern India, Shembaganur, 1912, G. Foreau 235 ['1824' in Hb. Dixon} (BM!— isotype?) FISSIDENS CEYLONENSIS AND ALLIED TAXA 155 0-5mm Fig. 2 Fissidens ceylonensis Dozy & Molkenb. (typical form) a: outline of a female shoot bearing axillary male branch; b: a detached axillary male branch. Fissidens ceylonensis ('perpusillus' form), c-e: shoots bearing gametangia (c: independent leafy male shoot (with groups of antheridia and perigonial leaf shown detached): outlines of male shoots showing positions of perigonia; e: apical region of shoot (leaves removed) showing terminal perigonia and scattered, solitary archegonia). Fissidens karwarensis Dixon. f: part of old, degenerating shoot supporting new leafy male and female shoots (male shoot with group of antheridia and inner-perigonial leaf shown detached), a, Drawn from Sri Lanka, Alston 1769 (BM); b, from Sri Lanka, Thwaites 137b (BM) (type of F. bicolor); c, from Sri Lanka, Gardner 630 (type of F. perpusillus) (BM); d, from southern India, Norkett 11328 p.p. (BM); e, from southern India Norkett 11369; f, from southern India Sedgwick 6368 (type of F. karwarensis) (BM). Eddy (1988) found Fissidens pennatulus to be conspecific with Fissidens ceylonensis. In the type specimens of these taxa the limbidium in at least some vaginant laminae appears to be intramarginal for a short part of its length where a row of subquadrate cells form the leaf margin (Fig. If). The major- ity of specimens from southern India (in NY and BM) originally identified as F. pennatulus possess a broad limbid- ium which usually extends the full length of the vaginant lamina and completely lacks a margin of subquadrate cells (Fig. Ih). In this and other features these specimens closely resemble type and authentic material of F. ceylonensis var. acutifolius Dixon & P. Varde. Several specimens referable to F. ceylonensis var. acutifolius possess slender leaves with acute apices (Fig. Ig). However, it is the structure of the limbidium, and not the form of the leaf, which is the distinctive and consistent feature. The type (Foreau [235] '1824', BM) is unusual in including female shoots with small, bud-like, axillary male branches as well as apparently inde- pendent, leafy male shoots with terminal perigonia. Foreau [235] '1824', along with authentic material (det. P. Varde) from the type locality (NY) and the other specimens listed below appear to form a discrete group, and thus provide justification for retaining F. ceylonensis var. acutifo- lius as a distinct taxon. (It may even warrant the rank of species.) Four collections from Burma, identified as Fissidens cey- lonensis (M oiler s.n. NY!, BM!), include shoots in which the limbidia are arranged on the leaves as in those found in 'var. acutifolius'. Some of the sporophytes in this material are terminal on well-developed shoots (as would be expected in F. ceylonensis), while others appear to emerge from the axils of basal leaves, or, at least, are terminal on very short axillary branches. The setae attain more than twice the length com- monly observed in southern Indian collections of F. ceylonen- sis. These Burmese collections can not yet be comfortably grouped with Fissidens ceylonensis var. acutifolius or the type variety. SELECTION OF HERBARIUM SPECIMENS EXAMINED. F. CEYLONENSIS VAR. ACUTIFOLIUS. Southern India, Tamil Nadu, Palni Hills: Kodai-Kanal, 1913 F. Marler 35 (BM!); Perumalmali Shola, 1925, G. Foreau 216 (BM!, NY!); Shem- baganur, G. Foreau 110, 368, 614, 1312, s.n.(NY!); Travan- core, Kumili, G. Foreau 589 (NY!). Tamil Nadu, Nilgiri Hills: Beddome 499, 547 (NY!); Kotagiri Road, 1966, A.M. Norkett 11411 (BM!). Tamil Nadu/Kerala border, Mangoli, High Wavy Mountains, 1967, A.M. Norkett 13269 (BM!). Sri Lanka. Nuwara Eliya, 1913, C.H. Binstead 252 (BM!); Udapassellawa, 1913, C.H. Binstead 402 (BM!). Fissidens kawarensis Dixon', /. Indian Bot. 2: 179 (1921). Type: Southern India, Karwar, October 1919, L.J. Sedgwick 6368 (BM! — lectotype proposed here). 156 L.T. ELLIS This species resembles the 'perpusillus' form of Fissidens ceylonensis except that the leaves and the sporophytes attain relatively larger dimensions. In the type specimen, the long- est leaves reach 1.5-1.8 x 0.2- 0.4 mm. Limbidia are best developed in the upper leaves of fertile shoots (Fig. lj,k). They are often obscure in leaves lower on the stem and obscure or absent in the leaves of sterile shoots. The dorsal lamina usually curves abruptly into the base of the costa, but in some leaves (especially those toward the apex of the shoot) it tapers gradually into the leaf base. Laminal cells show a similar range of size to those of F. ceylonensis, >5-12.5 x >5-12.5 um. Autoecious (and possibly dioecious), the male and female branches may originate from different leaf-axils on the same degenerating shoot of the previous generation and are similar in size and proportions (Fig. 2f). Perichaetia and perigonia are terminal. Sporophytes are similar to those in F. ceylonensis except that the seta reaches 3-5 mm in length and the longest theca examined is 0.75 mm. The lectotype, Sedgwick 6368 was collected on stones and a paratype, Sedgwick 3485 (Shiggaon, Dharwar District) (BM!) from the bole of a coconut palm. On the available evidence it would be unsafe to consider Fissidens karwarensis to be conspecific with F. ceylonensis. ACKNOWLEDGEMENTS. I am grateful to Dr A.J. Harrington for his valuable comments on the manuscript and also thank the Directors and staff at New York (NY) and Helsinki (H) for the loan of specimens. REFERENCES Eddy, A. 1988. A Handbook of Males ian Mosses. 1. Sphagnales to Dicranales. London. Fleischer, M. 1904. Die Musci der Flora von Buitenzorg 1. Leiden. Norris, D. & Koponen, T. 1987. Bryophyte flora of the Huon Peninsula, Papua New Guinea XX. Fissidentaceae, Mitteniaceae, Phyllodrepaniaceae, Phyll- ogoniaceae and Sorapillaceae :(Musci). Ann. hot. fenn. 24: 177-219. Bull. Br. Mus. not. Hist. (Bot.) 22(2): 157-158 Issued 26 November 1992 New species of Piper (Piperaceae) from Central America MARGARET TEBBS do Royal Botanic Gardens, Kew, London SYNOPSIS. During the preparation of the genus Piper (Piperaceae) for Flora Mesoamericana, two new species were discovered, both from Panama. The first, Piper canaliculum, is a fleshy-stemmed shrub with large, obovate leaves and stout inflorescences, collected from the provinces of Code, Panama, and San Bias. The second, P. premnospicum, is a low shrub or subshrub, with yellowish-pubescent stems and short, subglobose inflorescences, collected only from the province of Code. Piper canaliculum Tebbs sp. nov. (Fig. la,b,c) Caules carnosus, canaliculatus. Folia obovata, glandulosa, pili petioli vaginans. Holotype: Panama, Cerro Obu, Comarca de San Bias, 400-500 m, Nevers, Herrera & Gernada 8079, (BM!); isotype (MO!). Shrub 1-2.5 m; stems fleshy, longitudinally deeply grooved, minutely puberulent with white or yellow hairs. Leaves 24-32 x 10-14 cm, obovate, upper surface covered with white hairs 0.2-0.5 mm long, underside paler, with black or brown glands, minutely white puberulent especially along the veins; apex obtuse; base narrowly shallowly lobed. Vena- tion with 4-6 prominent secondary veins arising from the lower to middle part of the midrib, curving steeply to the apex. Petioles 10-40 mm, puberulent, sheathing the stem. Inflorescences 4-7 cm long, stout, erect; peduncles 10-15 mm long, minutely puberulent. Anthers 0.2-0.3 mm. Floral bracts 1-1.5 mm wide, triangular, minutely pale-ciliate, with dark centre. Fruits 1-2 mm, round to trigonous from above, glandular, glabrous; stigmas 3. DISTRIBUTION: Panama, tropical wet forest, low cloud for- est; 300-1200 m. REPRESENTATIVE SPECIMENS: Code, Sytsma & Andersson 4545 (BM!, MO!); Panama, Dressier 3763; San Bias, Nevers 4042 (MO!); Nevers & Herrera 6658 (MO!); Sytsma, Antonio & Dressier 2747 (BM!, MO!). The thick, fleshy stems with deep, ridge-like grooves, and the large, obovate leaves with narrow basal lobes distinguish this species. It has only been collected from three areas of Panama. Piper canaliculum is in Section Peltobryon Klotsch ex Miquel, which contains a group of species with densely glandular leaves, the glands yellow, orange, brown or black, and fleshy, often glandular fruits with minute to elongated styles. Piper premnospicum Tebbs sp. nov. (Fig. ld,e) Folia obovata, supra scabrosa, subter pili densi. Spica erecta, 1.2-2 cm, subglobosa; bacca 1-2 mm, stylus 2-3 mm, rectus, elongatus, stigmata 2. Holotype: Panama, El Valle Mesa, Code, 800 m, Folsom, Hartman & Butcher 4646 (MO!). Low shrub or subshrub 0.3-1 (-2) m; stems densely crisp- pubescent with yellowish hairs. Leaves 10-18 x 6-10 cm, obovate to elliptic-obovate, upper surface dark green, sca- brous, underside paler, densely hirsute, especially on veins; apex acute to obtuse; base unequally obtuse to rounded. Venation with 4-5 pairs of secondary veins arising from the lower to middle part of the midrib, curving steeply to the apex. Petioles 12-25 mm, densely pubescent. Prophylls 10-20 mm, pale-pubescent, drying to dark brown or black. Inflorescences 1.2-2 cm long, erect, subglobose, narrowing to crisp-pubescent tips 1-5 mm long; peduncles 4-12 mm, densely pubescent. Anthers 0.8-1 mm. Floral bracts 1-2 mm, triangular, mostly glabrous, with pale hairs at the base. Fruits 1-2 mm, round to trigonous from above, glabrous, with straight, prominent styles 2-3 mm long; stigmas 2, recurved. DISTRIBUTION: Panama, disturbed areas, dense rain forest, premontane or montane forest, mature forest understory and cloud forest; 400-1000 m. REPRESENTATIVE SPECIMENS: Code, Croat 14279 (MO!); Dwyer 8921 (MO!); Knapp & Dressier 3428 (MO!), 4916 (BM!, MO!); Lewis et al 1784 (MO!); Sytsma et al. 4385 (MO!). Piper premnospicum has only been collected from the prov- ince of Code, Panama. It is closely related to P. cuspidispi- cum Trel., with similar inflorescences, and long, straight styles with two recurved stigmas. However, Piper cuspidispi- cum is a glabrous shrub up to 3 m in height, with glossy, ovate to ovate-elliptic leaves, and glabrous prophylls. Piper prem- nospicum is in Section Isophyllon Miquel, the species of which are distinguished by their short inflorescences, with distinct sterile tips. 158 MARGARET TEBBS mm cm Fig. 1 Piper canaliculum, a: leaf; b: inflorescence; c: fruit and bract. P. premnospicum, d: stem; e: fruit and bract. Bull. Br. Mus. nat. Hist. (Bot.)22(2): 159-164 Issued 26 November 1992 Studies on the Cretan flora 1. Floristic notes NICHOLAS J. TURLAND Ludlow House, 12 Kingsley Avenue, Kettering, Northamptonshire NN16 9EU SYNOPSIS. Distributional, ecological and historical notes are provided for flowering plants occurring in the Cretan area (the islands of Crete, Kasos, Karpathos and Saria) in the South Aegean region. Uncommon, localized and endemic taxa are covered, including four new records from Crete: Arum cyrenaicum Hruby (also new for Europe), Kochia scoparia (L.) Schrader, Phlomis \ cytherea Rech.fil., and Taraxacum hellenicum Dahlst. The occurrence of Abutilon theophrasti Medicus, Carex illegitima Cesati, Colchicum cupanii Guss., Cymbalaria microcalyx subsp. dodekanesi Greuter, and Gageafibrosa (Desf.) Schultes & Schultes fil. in Crete is confirmed. Atractylis gummifera is recorded for the first time from Karpathos. INTRODUCTION The Cretan area, in the south Aegean region, is the most south-easterly floristic territory defined in Flora Europaea. The area includes not only the main island of Crete but the island group of Kasos, Karpathos and Saria to the north-east. All belong to the central part of a partially submerged arc of mountains which once linked the Greek Peloponnisos with the Taurus mountains of southern Turkey. Elements within the flora reveal these former links with Europe and Asia Minor. Occurring together with widespread Euro-Siberian and Mediterranean plants are taxa which otherwise exist only in Greece and the Balkan peninsula, or in south-western Asia. Two examples of Asiatic plants with their only Euro- pean stations in the Cretan area are Datisca cannabina (Datiscaceae) and Lecokia cretica (Umbelliferae). The islands of the Cretan area have remained separate from the continents for several million years and, not surpris- ingly, there is a high level of endemism in the flora, with roughly ten per-cent of the c. 1700 native species of vascular plants occurring nowhere else. Many of these are vicariously represented by relatives in other parts of the Aegean region and adjacent areas, although a few are without close affini- ties, e.g. Petromarula pinnata (Campanulaceae). A full historical account of botanical exploration of the area will not be given here, although a brief outline of some of the more recent and important floristic studies seems appropriate. Karl Heinz Rechinger (Vienna) collected exten- sively in the Aegean region during the 1930s and his Flora Aegaea (Rechinger, 1943a) covers the Cretan flora on the basis of earlier botanists' work. He visited Crete himself in 1942 and published a full account of his itinerary and collec- tions, as well as descriptions of new taxa (Rechinger, 1943b). Subsequent floristic studies are widely dispersed in botani- cal literature and include numerous papers by Werner Greu- ter (Berlin, formerly Geneva), who intensively explored Crete during the 1960s. Among them is a paper correcting the first two volumes of Flora Europaea (as far as the Cretan area is concerned) and providing lists of species doubtfully or erroneously recorded (Greuter, 1974). There are also four papers detailing floristic additions since Flora Aegaea (Greu- ter, 1973; Greuter, Malthas & Risse, 1984a, b, 1985). The most up-to-date and comprehensive work on the Cretan flora (excluding the Karpathos island group) is the checklist com- piled by Sir Colville Barclay (1986). The only recent and comprehensive work on the Karpathos island group is the basic checklist by Greuter, Pleger and Raus (1983). Botanical activity in the Cretan area during the years since 1986 has been considerable. A publication by Jacques Zaf- fran (University of Provence) gives details of his collections together with phytosociological studies (Zaffran, 1990). The author's own field-work in Crete began in 1984, much of it jointly carried out with Mr Lance Chilton, who first visited the island in 1982 and resided there for the greater part of five years from 1987. Their initial floristic findings have been compiled by Turland (1992). The quantity and importance of data which have accumu- lated since 1986 led to the initiation of the project to compile an up-dated and annotated checklist of the Cretan flora. This will be published in a single volume as a product of the Natural History Museum's European Plant Information Cen- tre. The scope of the new checklist will include the Karpathos island group, thereby making it directly comparable with the treatment of the Cretan area provided in Flora Europaea. Annotations will include habitats, altitudinal ranges and flowering times, where known, as well as distributions within and outside the area. A distribution map will be provided for each species, prepared by Chilton, based on his own field observations and those of the author, as well as records from literature sources. In order that the concise presentation planned for the checklist may be maintained, various significant new data are presented here as floristic notes. The plant records are mostly made by Chilton and the author; those by others are included only when they could be confirmed. The sequence of taxa follows that of the checklist, i.e. alphabetical, by families, genera, and species. Localities are headed by their eparchy (administrative district) or nomos (prefecture). Where more than one locality is cited, the order runs from west to east. The transliteration of Greek place-names follows Greuter (1973) and Greuter, Malthas & Risse (1984a, b, 1985), with Ihe following exceplions: Ihe Greek leller gamma always becomes 'g', never 'j', and della becomes 'd', nol 'dh'. This is Ihe syslem currenlly used on Greek road-signs bearing Roman spelling as well as Greek. 160 N.J. TURLAND FLORISTIC NOTES DICOTYLEDONES CHENOPODIACEAE Kochia scoparia (L.) Schrader (new record) Kidonia: Maleme 15 km W of Hania, 10 m, roadside, 1991, Chilian obs.; Hania town, 10 m, roadside, 1990, Chilton obs. Apokoronos: Georgioupoli, 10 m, trackside, 1990, Chilton obs. Agios Vasilios: Plakias, sea-level, stream-gravel, 1990, Chilton obs. Rethimni: Venetian fortress in Rethimno town, 20 m, paths and waste ground, 1990, Chilton obs. ; Misiria just E of Rethimno, 5 m, waste ground at back of beach, 1990, Chilton obs. Pirgiotissi: Matala, 10 m, waste ground and roadside, 1991, Chilton obs.; Agios loannis 5.5 km SE of Timbaki, 30 m, roadside, 1990, Chilton obs. Malevizi/ Temenos: Ammoudara just W of Iraklio, 5 m, roadside, 1990, Chilton obs. Temenos: Venerato 18 km SSW of Iraklio, 250 m, roadside, 1990, Chilton obs. Pedias: Kokkini Hani 10 km E of Iraklio, 5 m, waste ground, 1990, Chilton obs. Lasithi: Tzermiado, 850 m, waste ground, 1990, Chilton obs. Miram- bello: Sissi 9 km NW of Neapoli, 15 m, waste ground, 1990, Chilton obs. This species is cultivated for ornament and is recorded in Flora Europaea as naturalized in central, eastern and south- ern Europe, but not in Crete (Ball, 1964a: 99). The records cited here appear to be the first correct ones from Crete and Chilton's observations suggest that the species is becoming naturalized in at least some of the localities. Kochia scoparia is cultivated in Crete, but none of the plants observed was growing near such material, although some were near gar- dens. An old Cretan record by Gandoger was rejected as erroneous by Greuter (1974: 155). Noaea mucronata (Forskal) Ascherson & Schweinf. Kidonia: Akrotiri, in saxosis maritimis, 10.1899, Mancuso in herb. Baldacci (BM!). This collection from the Akrotiri peninsula in north-western Crete appears to have been over- looked, as the Cretan area is omitted from the range given in Flora Europaea (Ball, 1964b). In the apparent absence of any more recent collections, it is best to consider the species as doubtfully present in Crete until confirmation becomes avail- able. The species occurs in the wider Cretan area, however, having been recorded from Kasos, in the Karpathos island group, by Greuter (1974: 146). A distribution map is given by Jalas and Suominen (1980: 84, map 607). Taraxacum Wiggers Flora Europaea cites Taraxacum megalorhizon (Forskal) Hand.-Mazz. and the T. officinale group from the Cretan area (Richards & Sell, 1976: 334, 341). The presence of the latter is rejected by Greuter, Matthas & Risse (1984b: 274), who state 'The old record by Heldreich quoted by Rechinger (1943a: 692) has never been disclaimed, although it obviously refers to the very broad aggregate comprising the whole genus.' They accept, however, the presence of Taraxacum sect. Scariosa Hand.-Mazz. emend. Dahlst., to which T. megalorhizon belongs. According to Richards (pers. comm., 1988), T. megalorhizon is best treated as a nomen confusum. The following taxa in sect. Scariosa are known to occur in Crete. They tend to be small plants of rock-crevices and bare soil, often in the mountains, flowering in autumn (sometimes before the leaves develop) and in spring. Taraxacum aleppicum Dahlst. This species occurs also in Greece and south-western Asia and is recorded from Crete by van Soest (1975: 799). Rich- ards (1991: 547) gives it as occurring on the Omalos plain, in the Samaria gorge and the Dikti massif. Taraxacum bithynicum DC. Sfakia: above SE edge of Omalos plain, path from Xiloskala pass to Kallergi mountain refuge, 1200 m, under Zelkova abelicea (Lam.) Boiss., 3.8.1989, A.J.Richards (herb. A.J.Ri- chards). Taraxacum bithynicum DC. is a mostly Turkish alpine species, which has its only European stations in Crete. It is recorded from the island by Rechinger (1943b: 159) and by van Soest (1975: 799), according to whom it possibly does not belong within sect. Scariosa at all. Taraxacum hellenicum Dahlst. (new record) Agios Vasilios: hill 0.5 km E of Aktounda (S of Spili), 600 m, open phrygana over serpentine, with Taraxacum minimum, 30.3.1991, Turland 409 (BM, herb. A.J.Richards)— det. A.J.Richards (Newcastle-Upon-Tyne). Sitia: immediately W of Toplou monastery, 200 m, roadside, one large clump growing with T. minimum, 6.4.1990, A.J.Richards (herb. A.J.Richards). According to Richards (pers. comm. 1990, 1991), these are the first Cretan records of this Mediterranean species. He strongly suspects that the single plant at Toplou is an adven- tive from the Greek mainland, where the species also occurs. It seems reasonable to assume, however, that the population at Aktounda is native. The locality is not near any ports or in an area subjected to heavy tourism, and the habitat is undisturbed, so it is unlikely that an introduced Taraxacum would become established there. COMPOSITAE Atractylis gummifera L. Karpathos: 9.1988, Barclay obs. The occurrence of this Mediterranean species in Crete is well documented but it has not been recorded previously in the Karpathos island group. Taraxacum minimum (Briganti ex Guss.) N. Terrace. According to Richards (pers. comm., 1988-1991), and from the author's own collections and observations to date, it appears that this is the commonest segregate taxon in sect. Scariosa in Crete. It occurs throughout much of the island, as well as more widely in southern Europe and south-western Asia. CRETAN FLORA 1. FLORISTIC NOTES CONVOLVULACEAE Ipomoea imperati (Vahl) Griseb. (/. stolonifera (Cyr.) J.F.Gmelin) Kissamos: coast at Drapanias 4.5 km E of Kastelli Kissamou, near sea-level, sand dunes, 10.1987, C.C.Wilson colour photo! (herb. Turland); same locality, 10.1991, C.C.Wilson seed collection! (hort. Turland). This largely tropical and warm-temperate species was first recorded in Crete by Rechinger, on the island of Elafonisi in the south-west, as 'Calystegia soldanella' (Rechinger, 1943b: 107). He later corrected this mis-identification and cited a second collection by Barneby & Davis from Malia on the northern coast, east of Iraklio (Rechinger, 1949: 205). Greu- ter (1973: 48, 49) confirms the identity of the Elafonisi collection as Ipomoea but suggests that the Malia record is probably referable to Calystegia soldanella (L.) R.Br., which he cites from a point approximately 25 km west along the same stretch of coast. C. soldanella is fairly widespread in Crete, also occurring on the Akrotiri peninsula in the north- west (Greuter, op. cit.: 49) and in the same areas where Ipomoea occurs: on Elafonisi island (27.4.1989, Turland obs.) and on the coast immediately east of Kastelli Kissamou (9.8.1991, Turland 546 (BM)). In the vegetative state, /. imperati may be distiguished by the shape of its leaves, which are oblong or slightly three-lobed, unlike those of C. sol- danella, which are reniform. Until confirmation is available, it seems best to accept Greuter's opinion that the Malia population is probably Calystegia and not Ipomoea, which is known with certainty only from the extreme western part of Crete. In Europe Ipomoea imperati has only a few populations outside Crete: in the Azores, on Cabrera island south of Mallorca (Bonafe Barcelo, 1979: 346), and in southern Italy. These lie on the northern limit of its extensive world distribu- tion, which includes the Americas as well as the Old World. Another Greek occurrence was reported by Goulimis (1956), from the east Aegean island of Rodos, outside the European floristic area. Goulimis cited the record again in Goulandris et al. (1968: 88), where he stated that the species also occurs on the Greek island of Elafonisi, between the eastern penin- sula of the southern Peloponnisos and the island of Kithira. This is erroneous, however, owing to confusion with the Cretan island of the same name (Yannitsaros, 1971: 149). The illustration given by Goulandris et al. (op. cit.: 89) shows the distinctive, oblong leaf-shape referred to above. DIPSACACEAE Lomelosia albocincta (Greuter) Greuter & Burdet (Scabiosa albocincta Greuter) Milopotamos: hill between the Nida plain and Mt Skinakas, 1500 m, W- and NW-facing limestone cliffs, 11.8.1991, Chil- ton & Turland 224 (BM). This is yet another population of this endemic taxon, which was originally known only from a single valley in the Lefka Ori in western Crete but has since been recorded from several new localities (Turland, 1992: 349). This extra locality is worthy of mention because it is in the immediate area where 161 Baldacci collected the species in 1899 (BM!). For fuller details, and a distribution map for the species, see Turland (loc. cit.). LABIATAE Phlomis x cytherea Rech.fil. (new record) = P. cretica C.Presl x P. fruticosa L. Agios Vasilios: between Sellia and Kotsifos gorge N of Plakias, 300 m, phrygana at side of road, 10.4.1989, Turland 79 (BM). This natural hybrid was originally described from the island of Kithira (Rechinger in Greuter & Rechinger, 1967: 115). The record cited here appears to be the first from Crete. The collection is morphologically intermediate between the par- ents, which are both common at the locality. LEGUMINOSAE Astragalus idaeus Bunge According to Strid (1986: 472), this little-known species is apparently endemic to the high mountains of central and eastern Crete. It has been collected only twice, once by Fischer in the Psiloritis massif and once by Heldreich in the Dikti massif of eastern Crete (the former being the type collection). It is apparently similar to Astragalus agraniotii Orph., endemic to Mt Parnon in the south-eastern Pelopon- nisos. It would be surprising if A. idaeus has become extinct, since its dwarf habit should provide some protection from overgrazing, which is the only likely threat. However, the species is not recorded by Zaffran (1990), who has collected extensively in the high mountains of Crete. Confirmation of its continued existence would be welcome. MALVACEAE Abutilon theophrasti Medicus (confirmation) Apokoronos: Kalives, 25 m, waste ground, 10.1990, Chilton obs. Agios Vasilios: Plakias, 3 m, waste ground in the village, 8.1991,C/z//torts.n.!(BM). Greuter (1974: 156) considers old records by Sieber to be doubtful. Chilton's records confirm the presence of this species in Crete, and his collection from Plakias matches material at BM from Nafplio in the Peloponnisos. It seems likely that the plant has been fairly recently introduced to Crete, in view of its weedy habitat, occurrence in touristic areas, and the lack of records since Sieber's times. Its European distribution includes the Mediterranean region, although its native status is doubtful west of the Balkan peninsula (Webb, 1968). 162 RANUNCULACEAE N.J. TURLAND MONOCOTYLEDONES Nigella arvensis subsp. brevifolia Strid Pirgiotissi: 0.75 km NE of Kamilari, by the track from the village to the main Timbaki-Matala road, 50 m, cultivated soil in open olive grove, 7.7.1991, Chilian s.n. (BM!). According to Strid (1970: 47), this taxon has been collected only twice in Crete, on both occasions in the north-west, but not within the last 100 years and is, therefore, mapped as presumably extinct by Jalas & Suominen (1989: 33, map 1541). The new locality confirms its continued existence on the island, the specimen clearly agreeing with one of the old Cretan collections cited by Strid (Kissamos, 2.7.1884, Rever- chon (BM!)). According to Chilton, the habitat is not, at present, threatened by intensive agriculture of the type practised on the nearby Messara plain, with its extensive polythene greenhouses and copious use of fertilizers and herbicides. The only other known localities for this obscure subspecies are on the east Aegean island of Rodos, where it is apparently common, and the nearby small islands of Halki and Alimia (Strid, op. cit.: 45). SCROPHULARIACEAE Cymbalaria microcalyx subsp. dodekanesi Greuter (confirmation) Sitia: gorge NW of Toplou monastery, 50 m, crevices of calcareous rock, 30.3.1990, Turland 114 p.p. (BM) & living material (hort. Turland). The Balkan and south Aegean Cymbalaria microcalyx (Boiss.) Wettst. has a very disjunct distribution in Crete, occurring on the far western coast as subsp. microcalyx, which is otherwise found in the Peloponnisos and on Kithira, and at the far eastern end of the island, where the precise identity of the plants was unknown (Greuter, Matthas & Risse, 1984b: 290). Greuter et al. (loc. cit.) suggested that the eastern population might belong to subsp. dodekanesi, which was already known within the Cretan area on Karpathos and Saria and, outside Europe, on the east Aegean island of Rodos (Greuter in Greuter & Rechinger, 1967: 108). At the Toplou locality the population is a mixture of glabrous plants with small, fleshy leaves and pubescent plants with slightly larger, less fleshy leaves. The two grow together, often in the same crevices but retain their individual characters when separated and grown in cultivation. The pubescent plant agrees with subsp. dodekanesi, thereby confirming the pres- ence of this taxon in Crete. Fruits have been produced in cultivation to enable this determination. The glabrous plants have also fruited and proved not to be subsp. dodekanesi. They probably belong to the south Aegean and eastern Mediterranean C. longipes (Boiss. & Heldr.) A.Cheval., which is recorded by Zaffran (1990: 290) from the same two areas of Crete in which C. microcalyx occurs. ARACEAE Arum cyrenaicum Hruby (new record) Selinos: near Vlithias on Paleohora-Kandanos road, on E side of road, 200 m, 9.4.1984, M.Briggs obs.; same locality, 28.3.1988, M.Briggs living material (hort. P.Boyce); same area but 0.25km to S, on W side of road, 100 m, 28.3.1988, M.Briggs s.n. (K) — det. P.Boyce (Kew); same area but 1 km to N, on E side of road, 250 m, 28.3.1988, M.Briggs obs.; small ravine/valley 1.5 km NE of Vlithias, 300 m, 28.3.1991, Turland 402 (K) & living material (hort. P.Boyce) — det. P.Boyce; near Epanohori N of Sougia, 570 m, 20.3.1989, E.Markus colour transparency (K!) — det. P.Boyce. This species was formerly considered endemic to Cyrenaica in north-eastern Libya. These records from Crete are the first for Europe. Mrs Mary Briggs, who first discovered the plant in Crete, has kindly supplied data, which are quoted here: The first sighting at Vlithias, on 9th April 1984, was of seven flowering plants, which had very dark brownish-purple spathes and pale yellow spadices. They were growing in natural vegetation at the edge of a cultivated plot. Photo- graphs were taken and later shown to staff at Kew, but no determination was made. A return visit to the same locality was made on 28th March 1988, in order to collect material, at the request of Peter Boyce at Kew. Two plants were col- lected, with greenish spathes, purplish at the margins. They were later successfully grown by Peter Boyce. On the western side of the road, 0.25 km to the south, some individuals with very dark spathes were growing out of rock, with tubers firmly embedded. It was impossible to collect living material here but one herbarium specimen was taken. Approximately 1 km to the north, on the eastern side of the road, more individuals were seen growing out of and between rocks and boulders, and out of a low cliff above a stream. In 1991 the plants were identified by Peter Boyce as Arum cyrenaicum Hruby.' At the author's locality north-east of Vlithias, the plants were growing in woodland of Platanus orientalis L., associ- ated with Arum concinnatum Schott, Cyclamen creticum Hildebr., Hedera helix L. and Tamus communis L. The spathes were either unfurled or almost so, whereas none of the plants of Arum concinnatum (distinguishable by the different leaves) had even an immature inflorescence present. This early flowering-period and the superficial resemblance to the common and widespread A. concinnatum might account for A. cyrenaicum having remained undiscovered in Crete for so long. At present, the species appears restricted to the eparhia of Selinos in the far south-western part of the island. A fuller account of the Cretan occurrence is being prepared by Briggs. Biarum tenuifolium Schott Greuter (1973: 78) cites four Cretan localities for Biarum zelebori Schott. Three of these, however, are referable to B. tenuifolium subsp. idomenaeum Boyce & Athanasiou, and are included among the specimen citations accompanying the description of this new, apparently endemic taxon (Boyce & Athanasiou, 1991). The collection from the fourth locality cited by Greuter (Pedias: N slopes of Mt Kouna above CRETAN FLORA 1. FLORISTIC NOTES 163 Milliarado, 21.10.1966, Greuter 7777 (B, G)) is confirmed by Boyce (pers. comm., 1991) as belonging to B. tenuifolium subsp. tenuifolium var. zelebori (Schott) Engler (taxonomy: Boyce, in prep.). Boyce knows of no other localities in Crete for this eastern Aegean and south-western Anatolian taxon. CYPERACEAE Carex illegitima Cesati (confirmation) lerapetra: 1 km SW of Thripti, 650 m, woodland of Pinus brutia Ten., 24.4.1991, Turland 427 (BM). The only Cretan record for this species, by Gandoger, was considered doubtful and in need of confirmation by Greuter (1974: 162), and the Cretan area is omitted from the distribu- tion given in Flora Europaea (Chater, 1980: 313), although it is listed as doubtfully present in Crete by Barclay (1986: 104). The plants are frequent in the field-layer of the pine wood- land south-east of Thripti and the material collected agrees with specimens at BM from the Greek mainland. Outside Crete, the species is found on the island of Pantelleria west of Sicily, along the western coasts of Yugoslavia and Greece, on some of the East Aegean Islands, in western Turkey, and in Cyprus (Chater, loc. cit.; Nilson, 1985: 88). GRAMINEAE Sesleria doerfleri Hayek Rethimni: upper part of Prassiano gorge E of Prassies, 100 m, E-facing limestone cliff, 6.8.1991, Turland obs. Amari: N side of Mt Kedros above Gerakari, 1300 m, N-facing limestone cliff, 10.8.1991, Turland obs. This uncommon, endemic grass is an obligate chasmophyte with only five previously known localities all in western Crete (Greuter, Malthas & Risse, 1985: 38). The two new popula- tions extend the known distribution into central Crete and expand the altitudinal range (formerly considered to be 350-1000 m). The author has seen the plant on several occasions at one of the localities cited by Greuter et al. (Agios Vasilios: Kotsifos gorge N of Plakias). It is a very distinctive species and cannot be mistaken for other Cretan grasses. LILIACEAE Colchicum cupanii Guss. (confirmation) Pedias: near Hersonissos, 300 feet, low limestone hills in company with Ranunculus bullatus L. and Crocus laevigatus Bory & Chaub., 15.12.1939, Davis 1096 (K!). Mirambello: between Milatos and Kounali NNW of Neapoli, 3 km after Milatos, 300 m, stony limestone hillside, 6.11.1980, Brickell & Mathew 10149 (retained by Brickell); between Vrahassi and Voulismeni WNW of Neapoli, 200 m, olive groves, plants abundant with very wide leaves and sometime white- flowered, 12.11.1980, Brickell & Mathew 10260 (retained by Brickell); E of Latsida on road to Neapoli, 320 m, stony limestone hillside with Calicotome villosa (Poiret) Link, Phlomis & Pistachio, mixed with Colchicum pusillum Sieber, 6.11.1980, Brickell & Mathew 10145 (retained by Brickell). Greuter (1974: 161) rejects Cretan records of this Mediter- ranean species as either very doubtful or definitely errone- ous. Among the very doubtful ones, he mentions a record by Turrill (1941: 266) which is, in fact, a citation of the Davis collection detailed above. The specimen at Kew has been determined as Colchicum cupanii Guss. by K.Persson (Gote- borg) and, according to Mathew (pers. comm., 1991), there is no doubt that C. cupanii occurs in Crete. Gageafibrosa (Desf.) Schultes & Schultes fil. (confirmation) Agios Vasilios: 3 km NE of Plakias, col immediately E of the summit of Mt Kirimianou, 600 m, dry stony hillside, approx. 20 plants seen, 4.4.1991, M.J.Y.Foley s.n. (LANC)— conf. E.M.Rix. Mr Michael Foley has kindly supplied data, which is quoted here: 'Their overall height was rarely more than 5 cm, and they invariably possessed only solitary flowers borne on very short stems such that the cauline leaves were distinguished only with difficulty from the basal. Most characteristic was the shape (elongate-acuminate) of the perianth-segments which were of typical colour: bright yellow above and greenish beneath. The single basal leaf was c. 3 mm wide and more or less canaliculate, rigid and tortuous. Two or three bulbs examined showed a dense, fibrous network of roots but no prominent fibrous collar. There was no evidence of stolons bearing bulbils, and the plants occurred in isolation from each other.' Foley's is perhaps the first Cretan record of this species, which is otherwise found elsewhere in the Aegean region, south-western Asia, North Africa and Arabia. The plant was recorded by Sieber over 170 years ago (as Ornithogalum fibrosum Desf.), but Greuter (1973: 66) suggested that this might be referable to Gagea commutata K.Koch, which he gives as new to Crete, citing a Davis collection from the east of the island and one of his own from the Psiloritis massif. In Flora Europaea, Richardson (1980: 27) does not include the Cretan area in the distribution of G. fibrosa. ACKNOWLEDGEMENTS. I am firstly grateful to Lance Chilton for jointly carrying out some of the field-work with me and supplying voucher specimens and field-data for his own records. I should like to thank Dr John Richards (University of Newcastle-Upon-Tyne) for determining my Cretan collections of Taraxacum, supplying data concerning his own collections, and for other valuable advice. I am also most grateful to Mary Briggs for her notes on Arum cyrenaicum, Michael Foley for his notes on Gagea fibrosa, Dr Brian Mathew (Kew) for records of Colchicum cupanii, Sir Colville Barclay for his record of Atractylis gummifera, Peter Boyce (Kew) for advice on Biarum tenuifolium, and Chris Wilson for reporting Ipomoea imper- ati. REFERENCES Ball, P.W. 1964a. Kochia Roth In: T.G. Tutin et al. (Eds.). Flora Europaea 1: 98-99. Cambridge. 1964b. Noaea Moq. In: T.G. Tutin et al. (Eds.). Flora Europaea 1: 107. Cambridge. Barclay, Sir C. 1986. Crete, checklist of the vascular plants. Englera 6. Bonafe Barcelo, F. 1979. Flora de Mallorca 3. Editorial Moll, Palma de Mallorca. Boyce, P. (in prep.) A revision of the genus Biarum Schott. 164 N.J.TURLAND & Athanasiou, K. 1991. A new subspecies of Biarum tenuifolium (Araceae) from Crete. Fl. Medit. 1: 5-13. Chater, A.O. 1980. Carex L. In: T.G. Tutin et al. (Eds.). Flora Europaea 5: 290-323. Cambridge. Coulandris, N.A., Goulimis, C.N. & Steam, W.T. 1968. Wild flowers of Greece. Goulandris Botanical Museum, Kifissia. Goulimis, C.N. 1956. New additions to the Greek flora. Athens. Greater, W. 1973. Additions to the flora of Crete, 1938-1972. Annales Musei Goulandris 1: 15-83. 1974. Floristic report on the Cretan area. Mem. Soc. Brot. 24: 131-171. Greuter, W., Malthas. U., & Risse, H. 1984a. Additions to the flora of Crete, 1973-1983—1. Willdenowia 14: 27-36. 1984b. Additions to the flora of Crete, 1973-1983 (1984)— II. Willdenowia 14: 269-297. 1985. Additions to the flora of Crete, 1973-1983 (1984)— III. Willdenowia 15: 23-60. Pleger, R. & Raus, T. 1983. The vascular flora of the Karpathos island group (Dodecanesos, Greece). A preliminary checklist. Willdenowia 13: 43-78. & Rechinger, K.H. 1967. Flora der Insel Kythera, gleichzeitig Beginn einer nomenklatorischen Uberpriifung der griechischen Gefa|3pflanzenarten. Boissiera 13. Jalas, J. & Suominen, J. 1980; 1989. Atlas Florae Europaeae. Distribution of vascular plants in Europe. 5; 8. Helsinki. Nilsson, 0. 1985. Carex L. In: P.H. Davis (Ed.). Flora of Turkey and the East Aegean Islands 9: 73-158. Edinburgh. Rechinger, K.H. 1943a. Flora Aegaea. Flora der Inseln und Halbinseln der agaischen Meeres. Denkschr. Akad. Wiss. Math.-Nat. Kl. (Wien) 105(1). 1943b. Neue Beitrage zur Flora von Kreta. Denkschr. Akad. Wiss. Math.-Nat. Kl. (Wien) 105(2/1). 1949. Florae aegaeae supplementum. Phyton (Horn) 1: 194-228. Richards, A.J. 1991. Taraxacum Wiggers In: A. Strid & Kit Tan (Eds.). Mountain Flora of Greece 2: 541-572. Edinburgh. & Sell, P.O. 1976. Taraxacum Weber In: T.G. Tutin et al. (Eds.). Flora Europaea 4: 332-343. Cambridge. Richardson, I.B.K. 1980. Gagea Salisb. In: T.G. Tutin et al. (Eds.). Flora .Europaea 5: 26-28. Cambridge. Soest, J.L. van 1975. Taraxacum Wiggers In: P.H. Davis (Ed.). Flora of Turkey and the East Aegean Islands 5: 788-812. Edinburgh. Strid, A. 1970. Studies in the Aegean flora XVI. Biosystematics of the Nigella arvensis complex. With special reference to the problem of non-adaptive radiation. Opera Bot. 28. 1986. Astragalus L. In: A. Strid (Ed.). Mountain Flora of Greece 1: 460-478. Cambridge. Turland, N.J. 1992. Floristic notes from Crete. Bot. J. Linn. Soc. 108: 345-357. Turrill, W.B. 1941. On the flora of the Nearer Esat XXII. New records and new species from Greece and the greek Islands. Kew Bull. 1940: 262-266. Webb, D.A. 1968. Abutilon Miller In: T.G. Tutin et al. (Eds.). Flora Europaea 2: 254-255. Cambridge. Yannitsaros, A.G. 1971. Notes on the flora of Elaphonesos island (Laconia, Greece). Biol. Gallo-Hellen. 3: 149-162. Zaffran, J. 1990. Contributions a la flore et a la vegetation de la Crete. Universite de Provence. Bull. Br. Mus. nat. Hist. (Bot.) 22(2): 165-169 Issued 26 November 1992 Studies on the Cretan flora 2. The Dianthus juniperinus complex (Caryophyllaceae) NICHOLAS J. TURLAND Ludlow House, 12 Kingsley Avenue, Kettering, Northamptonshire NN16 9EU SYNOPSIS. The Cretan endemic Dianthus juniperinus Sm. is treated as a complex of seven subpecies: subsp. pulviniformis (Greuter) N.J.Turland comb, et stat. nov., subsp. juniperinus, subsp. heldreichii Greuter, subsp. idaeus N.J.Turland subsp. nov., subsp. kavusicus N.J.Turland subsp. nov., subsp. aciphyllus (Sieber ex Ser.) N.J.Turland comb, et stat. nov., and subsp. bauhinorum (Greuter) N.J.Turland comb, et stat. nov. Morphological and distributional data are provided, including a key to the taxa recognized. INTRODUCTION This paper is the second in a series resulting from ongoing research connected with the Natural History Museum's project to compile an annotated checklist, with distribution maps, of the vascular flora of the Cretan area in the south Aegean region. The 'Cretan area' is the floristic territory defined by Flora Europaea, and includes the islands of Kasos, Karpathos and Saria to the north-east of Crete itself. The Dianthus juniperinus complex is endemic to the main island of Crete, where the plants grow as obligate chasmo- phytes on vertical, calcareous cliffs. The extremely rugged nature of the landscape, with abundant mountains and deep gorges, limits the extent to which such specialized plants can migrate. Genetic drift in isolated population-groups of a species can eventually result in a complex of closely related taxa. This process probably explains the particularly high degree of differentiation within D. juniperinus, in which seven taxa can be recognized altogether. In this respect, D. juniperinus can be likened to another chasmophyte, D. fruticosus L., within which Runemark (1980) recognized eight allopatric subspecies distributed in the Ionian islands, the southern Peloponnisos and the south Aegean islands. The Erysimum cheiri complex (Crudferae) is also chasmophytic, and behaves similarly, with various species and subspecies endemic to islands and island-groups in the Aegean region (Snogerup, 1967). Tutin (1964) treated the complex as a single species, Dianthus juniperinus Sm., with no infraspecific variation, and included D. aciphyllus Sieber ex Ser. in synonymy. Greuter (1965), however, recognized five taxa, based upon material available to him, describing three as new. D. juniperinus Sm. subsp. juniperinus from the western half of the district of Sfakia in western Crete. D. juniperinus subsp. heldreichii Greuter from the eastern half of the district of Sfakia. D. pulviniformis Greuter from Mount Kedros in central Crete. D. aciphyllus var. bauhinorum Greuter from the area east of the Psiloritis mountain massif, again in central Crete but further east than Kedros. ' d g , f / e Fig. 1 Distribution of Dianthus juniperinus in Crete: a. subsp. pulviniformis; b. subsp. juniperinus; c. subsp. heldreichii; d. subsp. idaeus; e. subsp. kavusicus; f. subsp. aciphyllus; g. subsp. bauhinorum. Open circle indicates indeterminate subspecies. Question-mark indicates doubtful locality. D. aciphyllus Sieber ex Ser. var. aciphyllus from the Dikti mountain massif (Lasithiotika Ori) in eastern Crete. This arrangement can no longer be justified, in the light of subsequent collections augmenting the rather scanty material available to botanists during the early 1960s, and especially because of new information provided by a recent gathering by Chilton and Turland, from central Crete, and one made in 1938 by Barneby and Davis in eastern Crete and apparently not seen by Greuter. If these two are disregarded, then there are clear-cut discontinuities between Dianthus juniperinus and D. aciphyllus. The former has pale pink or bright pink flowers opening together, while those of the latter are pinkish-purple with darker markings and do not open together (i.e. immature buds at various stages of develop- ment are present during the greater part of anthesis). Fur- thermore, D. juniperinus has calyces 10-13 mm long and flowers small, the petal-limb 3-4.5 mm, whereas those of D. aciphyllus measure 15-23 mm and 7-14 mm respectively. However, the new collection from central Crete is similar to D. juniperinus but has a petal-limb measuring 6-7 mm, while Barneby and Davis's collection has flowers matching D. aciphyllus in colour but all opening together, with calyces (13-)14 mm long and petal-limb 6 mm. It is necessary, there- fore, to combine D. aciphyllus within D. juniperinus (the earlier name) because there now exists a continuous range of variants, from the small-flowered D. juniperinus subsp. juni- 166 N.J. TURLAND perinus to the large-flowered D. aciphyllus var. bauhinorum. The two varieties of D. aciphyllus are better placed at at subspecific rank, since they differ from each other in more than one character and have discrete distributions. The maintenance of Dianthus pulviniformis at specific rank is inappropriate because the plant is clearly very close to typical D. juniperinus, differing only in usually being smaller. The single clear-cut discontinuity is the size of the calyx, which is 7-9 mm long (other taxa 10-23 mm). It is better, therefore, to treat this plant, too, as a subspecies of D. juniperinus. The description of Dianthus pulviniformis is somewhat misleading. The plant is said to have flowers not opening together, although more recent observations show this not to be the case. The illustration of the type specimen, accompa- nying the description, shows a plant which is just beginning to flower, with several buds present, all apparently in an advanced stage of development. This gives a wrong impres- sion of the taxon. Had the specimen been collected one week later, all flowers probably would have been open and the affinities with D. juniperinus made less obscure. It is also stated that the flowers are white, although the colour has been pale pink in all the plants seen in the field by the author. In April 1989 sterile material of a Dianthus obviously belonging to the D. juniperinus complex was collected in a gorge near Kamares on the southern side of the Psiloritis mountain massif in central Crete. It subsequently flowered in cultivation in Britain and has proved to be close to D. juniperinus subsp. juniperinus, on account of its calyces, which are 10-11 mm long, and its pale pink flowers. It differs strikingly, however, in its large petals, which have a limb 6-7 mm long. This feature is unlikely to be attributable to the effects of cultivation under glass in Britain, since other taxa have been grown alongside the Kamares plant and their flowers have remained identical in size to those of wild individuals. The geographical position of the Kamares popu- lation is significant, being discrete from the ranges of all other taxa and considerably disjunct from that of D. juniperinus subsp. juniperinus to the west (two other taxa present in the area between). Upon this evidence the Kamares plant is described here as a new subspecies within the complex. The epithet idaeus has been chosen, after Mount Ida, the ancient name for the Psiloritis massif where the new plant is found. One collection in the Kew herbarium, labelled as Dianthus juniperinus var. sieberi Boiss., has a combination of features which make it impossible to assign to any of the six taxa recognized so far. It was gathered by Barneby and Davis in 1938, on Mount Skliros north of Mount Afendis Kavousi (Afendis Stavromenos), on the border of the eparchies of lerapatra and Sitia in eastern Crete. This extends the distri- bution of the complex eastward from the Dikti massif. The inflorescences have 9-10 crowded flowers opening together, the calyces are (13-)14 mm long and the petals are pinkish- purple with darker markings, the limb 6 mm long. In view of all this, it seems appropriate to treat the plant as another, new, seventh subspecies in the complex. The epithet kavusi- cus has been chosen, referring to Mt Afendis Kavousi, the highest peak in the range of mountains where the plant was collected. SYSTEMATIC TREATMENT Key to the subspecies of Dianthus juniperinus la. Leaves 0.4-1.5 mm wide; epicalyx scales c. 1/2 as long as calyx; calyx 7-13 mm; petal-limb 3-7 mm, pale or bright pink: 2a. Calyx (10-)12-13 mm; flowers bright pink c. subsp. heldreichii 2b. Calyx 7-ll(-12) mm; flowers pale pink: 3a. Leaves 0.4-0.8 mm wide; calyx 7-9 mm a. subsp. pulviniformis 3b. Leaves 1-1.5 mm wide; calyx 10-11(-12) mm: 4a. Petal-limb 3-4.5 mm b. subsp. juniperinus 4b. Petal-limb 6-7 mm d. subsp. idaeus lb.:Leaves 1-3.1 mm wide; epicalyx scales c. 1/3 as long as calyx; calyx (13-)14-23 mm; petal-limb 6-14 mm, pinkish-purple with darker stripes: 5a. Calyx (13-)14 mm; petal-limb 6 mm e. subsp. kavusicus 5b. Calyx 15-23 mm; petal-limb 7-14 mm: 6a.: Leaves 1-1.9 mm wide, linear; petal-limb 7-11 mm f. subsp. aciphyllus 6b. Leaves 1.6-3.1 mm wide, linear to linear-lanceolate; petal-limb 8-14 mm g. subsp. bauhinorum 1. Dianthus juniperinus Sm. in Trans. Linn. Soc. London (Bot.) ser.l, 2: 303 (1794). Type: Crete, Gundelsheimer & Tournefort (LINN-holotype). Shrubby, glabrous perennial with densely leafy vegetative shoots normally forming cushions. Leaves 10-40 x 0.4-3.1 mm, flat or slightly involute, coriaceous, linear to linear-lanceolate; apex acute. Flowering stems numerous, herbaceous, 3-30 cm with 3-4 pairs of leaves. Inflorescence with 1-10(-12) flowers opening together or gradually (i.e. with immature buds at various stages of development present during greater part of anthesis). Epicalyx-scales 4-8, c. 1/3 to 1/2 as long as calyx, obovate or sometimes obcordate; apex subulate. Calyx 7-23 mm, nearly cylindrical; teeth 3-6 mm, acute. Petal-limb 3-14 mm, orbicular-obtriangular, dentate, bearded, pale or bright pink, or pinkish-purple with darker stripes. FLOWERING TIME. June to September. ECOLOGY. Crevices and ledges of vertical, calcareous cliffs. DISTRIBUTION. Endemic to Crete, where widespread, occur- ring at altitudes between 600 and 1750 m. a. subsp. pulviniformis (Greuter) N.J.Turland comb, et stat. nov. Basionym: D. pulviniformis Greuter in Candol- lea 20: 189 (1965). Type: Crete, Agios Vasilios, ad parietes deruptorum meridionalium mentis Kedros, prope pagum Kria Vrisi, Snogerup, Strid & von Bothmer 20955 (LD-holotype). Icon. Greuter (1965: 190). Leaves 10-20 x 0.4-0.8 mm, linear. Flowering stems 3-8.5 cm. Inflorescence with 1-4 flowers opening together. Epicalyx-scales c. 1/2 as long as calyx. Calyx 7-9 mm; teeth 3-3.5 mm. Petal-limb 4 mm, pale pink. DISTRIBUTION. Mount Kedros in central Crete, 600-1300 m. SPECIMENS EXAMINED. Crete: Agios Vasilios: in faucibus supra Palea Kria Vrisi ad latus mentis Kedros, 750 m, CRETAN FLORA 2. THE D1ANTHUS JUNIPERINUS COMPLEX 167 2.11.1966, Greuter 7863 (K). Amari: north side of Mt Kedros above Gerakari village, 1300 m, 10.8.1991, Turland 548 (BM). b. subsp. juniperinus Icon. Greuter (1965: 188). Leaves 15-30 x 1-1.5 mm, linear. Flowering stems 8-20 cm. Inflorescnce with 2-6(-7) flowers opening together. Epicalyx- scales c. 1/2 as long as calyx. Calyx 10-11(-12) mm; teeth 3-4 mm. Petal-limb 3-4.5 mm, pale pink. DISTRIBUTION. Western half of district of Sfakia in western Crete, 1400-1750 m; doubtfully on Akrotiri peninsula north- east of Hania (see Supplementary Notes). SPECIMENS EXAMINED. Crete: Sfakia: in rupestribus l.d. Linoseli m. Volakia, 8.7.1893, Baldacci 54 (BM, K); between Mt Strifomadi & the Xiloskala pass, 1450-1500 m, 8.8.1991, Turland obs. ?Kidonia: pr. Acrotiri, Jul? 1932, Guiol 2129 (BM). c. subsp. heldreichii Greuter in Candollea 20: 187 (1965). Type: Crete, Sfakia, fauces Sfakiano, in rupe Asprokremnos, 600 m, Heldreich 1640 (G-Bs-holotype; scheda gallica, BM!, FI, G, G-Bs, G-Bu, K!, W-Hal- isotypes). Icon. Greuter (1965: 188). Leaves (13-)15-25 x 0.5-1.4 mm, linear. Flowering stems 6-13 cm. Inflorescence with 1-4 flowers opening together. Epicalyx-scales c. 1/2 as long as calyx. Calyx (10-)12-13 mm; teeth 3-4 mm. Petal-limb 4 mm, bright pink. DISTRIBUTION. Eastern half of district of Sfakia in western Crete, 600-650 m. SPECIMENS EXAMINED. Crete: 1846, Heldreich (BM, K). Sfakia: in rupe Asprokremno dicta Sphakia, 7.1846, Heldre- ich (BM, K); in faucibus montium Sphacioticorum Cretae, 7.1846, Heldreich (BM, K); gorge south of Asfendos, 500 m, 21.4.1989, Chilton & Turland 10 (BM). 10 cr Fig. 2 Dianthus juniperinus subsp. kavusicus: a. habit; b. flower. D. juniperinus subsp. idaeus: c. habit; d. flower. 168 N.J. TURLAND Table 1 Comparison of features of subspecies of Dianthus juniperinus. Subspecies Size of Length of leaves fig stems Flowers per infl. Flowers opening Epicalyx: calyx ratio Length of calyx Length of calyx-teeth Length of petal-limb Colour of petal-limb pulviniformis 10-20 3-8. 5cm 1-4 together c. 1:2 7-9mm 3-3. 5mm 4mm pale pink xO.4-0.8mm juniperinus 15-30 8-20cm 2-6(-7) together c. 1:2 10-ll(-12)mm 3-4mm 3-4. 5mm pale pink x 1-1. 5mm heldreichii (13-) 15-25 6-13cm 1-4 together c. 1:2 (10-)12-13mm 3-4mm 4mm bright pink xO.5-l.4mm idaeus 20-30 9.5-13.5cm 2-5 together c. 1:2 10-1 1mm 3.5mm 6-7mm pale pink x 1-1. 3mm kavusicus 17-26 8.5-10cm 9-10 together c. 1:3 (13-)14mm 4mm 6mm pink-purple, x 1-1. 3mm with stripes aciphyllus 16-40 ll-30cm (l-)2-6 not c. 1.3 15-22mm 4-6mm 7-1 1m pink-purple, x 1-1. 9mm together with stripes bauhinorum 21-40 13-30cm 2-6(-12) not c. 1:3 16-23mm 4-6mm 8-14mm pink-purple, x 1.6-3. 1mm together with stripes d. subsp. idaeus N.J.Turland subsp. nov. Subsp. juniperino Sm. affinis, sed floribus maioribus differt. Type: Crete, Pirgiotissi, gorge 0.75 km NW of Kamares village, 600 m, 16.4.1989, Turland & Chilian 8 (BM-holotype); same locality, ex hort. Turland (with flowers), 10.7.1991 (BM-paratype). Leaves 20-30 x 1-1.3 mm, linear. Flowering stems 9.5-13.5 cm. Inflorescence with 2-5 flowers opening together. Epicalyx-scales c. 1/2 as long as calyx. Calyx 10-11 mm; teeth 3.5 mm. Petal-limb 6-7 mm, pale pink. DISTRIBUTION. Gorge north-west of Kamares village on southern edge of Psiloritis massif in central Crete, 600 m. NOTE. The flowering stem of the paratype specimen mea- sures 23 cm. This is probably abnormally tall, a result of the light-levels prevalent during its development in cultivation in Britain being lower than those at the type locality in Crete. e. subsp. kavusicus N.J.Turland subsp. nov. Subsp. juni- perino Sm. affinis, sed floribus numerosioribus in inflo- rescentiis 9-10-floris dispositis, calyce longiore, limbo petali longiore striis subroseis-purpureis ornato, differt. Type: Crete, lerapetra-Sitia, Skliros E. [sic] of Mt Kavutsi, 3000 ft, 9.1938, Barneby & Davis (K-holotype). Leaves 17-26 x 1-1.3 mm, linear. Flowering stems 8.5-10 cm. Inflorescences with 9-10 flowers opening together. Epicalyx-scales c. 1/3 as long as calyx. Calyx (13-)14 mm; teeth 4 mm. Petal-limb 6 mm, pinkish-purple with darker stripes. DISTRIBUTION. Mt Skliros north of Mt Afendis Kavousi (Afendis Stavromenos) between lerapetra and Sitia in east- ern Crete, 900 m. f. subsp. aciphyllus (Sieber ex Ser.) N.J.Turland comb, et stat. nov. Basionym: D. aciphyllus Sieber ex Ser. in DC., Prodr. 1: 358 (1824). Type: Crete, Lasithi, ad rupes prope pagum Tzermiado, Sieber (G-DC-holotype; G, G-Bs, K!-isotypes). D. juniperinus var. sieberi Boiss., Fl. Or. 1: 498 (1867). Type: same as that of D. aciphyllus Sieber ex Ser. D. juniperinus var. aciphyllus (Sieber ex Ser.) Halacsy, Consp. Fl. Graec. 1: 202 (1900), nom. illeg. (article 11.3). Icon. Greuter (1965: 191). Leaves 16-40 x 1-1.9 mm, linear. Flowering stems 11-30 cm. Inflorescence with (l-)2-6 flowers not opening together. Epicalyx-scales c. 1/3 as long as calyx. Calyx 15-22 mm; teeth 4-6 mm. Petal-limb 7-11 mm, pinkish-purple with darker stripes. DISTRIBUTION. Dikti mountain massif (Lasithiotika Ori) in eastern Crete, 800-1700 m. SPECIMENS EXAMINED. Crete: 24.5.1948, Davis — seed grown at Kew and specimens prepared 10.10.1951, 25.8.1952, 31.7.1953 & 8.8.1955, K. 2779 (K). Pedias: in latere septen- trionali mentis Kuna supra Miliarado, 750-850 m, 21.10.1966, Greuter 7775 (K); in rupestr. reg. sup. et mediae m. Aphendis Sarakeno, 15.6.1899, Baldacci 108 (BM); Mt Aphendi Sarakino, Atchley 1256 (K). Lasithi: ad rupes prope pagum Tzermiado, Sieber (K); Tzermiado, 3000 feet, 11.7.1938, Davis 284 (BM, K); in rupestribus Katavo[thra] Omalo Lassithi, 2.7.1899, Baldacci 108 bis (BM). g. subsp. bauhinorum (Greuter) N.J.Turland comb, et stat. nov. Basionym: D. aciphyllus var. bauhinorum Greuter in Candollea 20: 190 (1965). Type: Crete, Temenos, in rupestribus m. Joukta, 6.6.1899, Baldacci 57 (G-holotype; BM!, Fl, G-Bu, W-Hal-isotypes). Leaves 21-40 x 1.6-3.1 mm, linear to linear-lanceolate. Flowering stems 13-30 cm. Inflorescence with 2-6(-12) flow- ers not opening together. Epicalyx-scales c. 1/3 as long as calyx. Calyx 16-23 mm; teeth 4-6 mm. Petal-limb 8-14 mm, pinkish-purple with darker stripes. DISTRIBUTION. From eastern edge of Psiloritis massif to Mount Giouhtas south of Iraklio in central Crete, 600-800 m. SPECIMENS EXAMINED. Crete: Malevizi: in rupestribus supra Prinias, 4.6.1899, Baldacci 57 bis (BM). Temenos: in rupes- tribus m. Joukta, 6.6.1899, Baldacci 57 (BM); ad cacumen montis Juktas, 700 m, 14.9.1966, Greuter 7517 (K); summit of Mt Giouhtas, 800 m, 25.4.1989, Turland 86 (BM). CRETAN FLORA 2. THE DIANTHUS JUNIPERINUS COMPLEX 169 SUPPLEMENTARY NOTES The correct provenance of Guiol's specimen of Dianthus juniperinus subsp. juniperinus at BM is obscure. The 'Acrotiri' mentioned on the label could refer to the Akrotiri peninsula north-east of the city of Hania, indeed the moun- tains there rise to 528 m on the peak called Skloha, which is almost as high as the lower altitudinal limit of some of the other taxa. However, this region is considerably disjunct from the area of the type locality, and none of the other taxa shows such a disjunction. The Greek word akrotiri means cape or headland, of which there are many around the coast of Crete. One possible explanation is that a particular cape was meant, but the distinguishing epithet omitted. It is also possible that the specimen was mis-labelled. In autumn 1989 a Dianthus obviously belonging to the D. juniperinus complex was observed by the author in the middle part of the Prassiano gorge, south-east of Rethimno in central Crete. Unfortunately, no fertile parts were remaining and no material was collected owing to the inaccessibility of the plants. Subsp. pulviniformis and subsp. idaeus are the nearest taxa geographically. ACKNOWLEDGEMENTS. I should like to thank Chris Hogg for drawing the illustrations, Bob Press for drawing the map and for helpful advice, and Marian West for translating the Latin diagnoses. REFERENCES Boissier, P.E. 1867. Flora Orientals 1. Basileae et Genevae. Greuter, W. 1965. Beitrage zur Flora der Siidagais 1-7. Candollea 20: 167-218. Hahksy, E. von, 1900-1901. Conspectus Florae Graecae 1. Lipsiae. Rum-mark, H. 1980. Studies in the Aegean flora XXIII. The Dianthus fruticosus complex (Caryophyllaceae). Bot. Not. 133: 475-490. Snogerup, S. 1967. Studies in the Aegean flora VIII, IX. Erysimum sect. Cheiranthus A. Taxonomy. B. Variation and evolution in the small- population system. Opera Bot. 13, 14. Tutin, T.G. 1964. Dianthus L. In: T.G.Tutin et al. (Eds.). Flora Europaea 1: 188-204. Cambridge. Bulletin of the British Museum (Natural History) Botany Series Earlier Botany Bulletins are still in print. The following can be ordered backlist is not shown, this may also be obtained from the same address. Volume 5 No. 1 The dryopteroid ferns of Ceylon. W. A. Sledge. 1973. Pp. 1-43, 4 figs. £3.00 No. 2 New Himalayan and Tibetan species of Corydalis (Papaveraceae). F.Ludlow & W.T. Stearn. 1975. Pp. 45-69, 15 plates, 14 figs. £3.85 No. 3 The marine algae of Trinidad, West Indies. W.D.Richardson. 1975. Pp. 71-143, 12 plates, 2 figs. £6.80 No. 4 A revision of the Macaronesian genus Argyranthemum Webb ex Schultz Bip. (Compositae-Anthemideae).C.J. Humphries. 1976. 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Where the complete No. 4 Vascular plant collections from the Tristan da Cunha group of islands. E.W. Groves. Pp. 333-420, 33 figs. £12.50 Volume 9 No. 1 The lichenicolous Coelomycetes. D.L. Hawksworth. 1981. Pp. 1-98, 36 figs. £13.25 No. 2 The genus Callithamnion (Rhodophyta: Ceramiaceae) in the British Isles. P.S. Dixon & J.H. Price. 1981. Pp. 99-141, 5 figs. £6.00 No. 3 . Parmelia subgenus Amphigymnia (lichens) in East Africa. H. Krog & T.F.V. Swinscow. 1981. Pp. 143-231, 31 figs. £12.25 No. 4 The genus Selaginella in tropical South America. A.H.G. Alston, A.C. Jermy & J.M. Rankin. 1981. Pp. 233-330, 18 figs. £13.50 Volume 10 No. 1 Taxonomic studies in the Labiatae tribe Pogostemoneae. J.R. Press. 1982. Pp. 1-89, 33 figs. £12.75 No. 2 The typification of Hudson's algae: a taxonomic and nomenclatural reappraisal. L.M. Irvine & P.S. Dixon. 1982. Pp. 91-105. £2.25 No. 3 Seaweeds of the Faroes. Various authors. 1982. Pp. 107-225, 13 figs. £16.75 No. 4 The lichen genus Steinera. A.M. Henssen & P.W. James. 1982. Pp. 227-256, 24 figs. £4.50 Volume 11 No. 1 The algae of Lightfoot's Flora scotica. P.S. Dixon. 1983. Pp. 1-15, 2 figs. £2.30 No. 2 A taxonomic study of the lichen genus Micarea in Europe. B.J. Coppins. 1983. Pp. 17-214, 57 figs, 28 maps. £27.75 No. 3 The hepatics of Sierra Leone and Ghana. E.W. Jones & A.J. Harrington. 1983. Pp. 215-289, 8 figs. £10.50 No. 4 Studies in the Corallinaceae with special reference to Fosliella and Pneophyllum in the British Isles. Y.M. Chamberlain. 1983. Pp. 291-463, 89 figs. £24.50 Volume 12 No. 1 A revision of the Morinaceae (Magnoliophyta- Dipsacales). M.J. Cannon & J.F.M. Cannon. 1984. Pp. 1-35, 9 figs. £5.40 No. 2 An introduction to fern genera of the Indian subcontinent. C.R. Fraser-Jenkins. 1984. Pp. 37-76, 1 fig. £6.00 No. 3 A revision of African Sphagnales. A. Eddy. 1985. Pp. 77-162, 47 figs. £13.50 No. 4 Studies in the genus Hypericum L. (Guttiferae) 3. Sections 1. Campylosporus to 6a. Umbraculoides. N.K.B. Robson. 1985. Pp. 163-325, 24 figs, 34 maps. £28.80 Volume 13 No. 1 The lichen genus Usnea subgenus Neuropogon. F.J. Walker. 1985. Pp. 1-130, 39 figs. £19.80 No. 2 Cytotaxonomic studies of the ferns of Trinidad. A.C. Jermy & T.G. Walker. 1985. Pp. 131-276, 69 figs. £22.00 No. 3 Some genera of the Biddulphiaceae (diatoms) with interlocking linking spines. R. Ross & P. A. Sims. 1985. Pp. 277-381, 33 plates. £20.00 Volume 14 No. 1 Cytological observations on Indian subcontinent and Chinese Dryopteris and Polystichum (Pteridophyta: Dryopteridaceae). M. Gibby. 1985. Pp. 1-42, 78 figs. £6.00 No. 2 A redisposition of the species referred to the ascomycete genus Microthdia. D.L. Hawksworth. 1985. Pp. 43-181, 73 figs. £24.00 No. 3 A classification of the genus Cryopteris (Pteridophyta: Dryopteridacaee). C.R. Fraser-Jenkins. 1986. Pp. 183-218, 4 figs. £5.60 No. 4 Evolutionary cladistics of marattialean ferns. C.R. Hill & J.M. Camus. 1986. Pp. 219-300, 27 figs. £14.50 Volume 15 No. 1 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 1. Genera A-F. J.H. Price, D.M. John & G.W. Lawson. 1986. Pp. 1-122, 1 fig. £23.50 No. 2 Cytology of the fern flora of Madeira. I. Manton, J.D. Lovis, G. Vida & M. Gibby. 1986. Pp. 123-161, 12 plates. £7.50 No. 3 A revision of the lichen genus Xanthoparmelia in Australasia. J.A. Elix, J. Johnston & P.M. Armstrong. 1986. Pp. 163-362, 42 figs, 117 maps. £38.00 Volume 16 No. 1 Studies in the genus Hypericum L. (Guttiferae) 7. Section 29. Brathys (part 1). N.K.B. Robson. 1987. Pp. 1-106, 14 figs, 25 maps. £20.00 No. 2 The lichen genus Ramallna in Australia. G.N. Stevens. 1987. Pp. 107-233, 15 plates, 31 figs. £22.50 No. 3 An annotated list of vascular plants collected in the valleys south of Mt Everest. G. Miehe. 1987. Pp. 225-268, 4 figs. £8.50 No. 4 Further genera of the Biddulphiaceae (diatoms) with interlocking linking spines. R. Ross & A. Sims. 1987. Pp. 269-311, 13 plates. £8.00 Volume 17 No. 1 Studies in Pseudocyphellaria (lichens) 1. The New Zealand species. D.J. Galloway. 1988. Pp. 1-267, 124 figs. £51.00 Volume 18 No. 1 An illustrated catalogue of the type specimens in the Greville diatom herbarium. D.M. Williams. 1988. Pp. 1-148, 74 plates. £28.00 No. 2 Erik Acharius and his influence on English lichenology. D.J. Galloway. 1988. Pp. 149-194, 18 figs. £8.80 No. 3 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Phodophyta (Florideae) 2. Genera G.J.H. Price, D.M. John & G.W. Lawson. 1988. Pp. 195-273, 1 fig. £14.80 No. 4 Some Cretaceous and Palaeogene Trinacria (diatom) species. P. A. Sims & R. Ross. 1988. Pp. 275-322, 13 plates. £9.10 No. 5 A monograph of Dryopteris (Pteridophyta: Dryopteridaceae) in the Indian subcontinent. C.R. Fraser-Jenkins. 1989. Pp. 323-477, 79 figs. £30.00 No. 6 Corydalis (Papaveraceae: Fumarioideae) in Nepal. M. Liden. 1989. Pp. 479-538, 26 figs. £11.20 Volume 19 A new species of Maytenus (Celastraceae) in Ethiopia. Sebsebe Demissew. 1989. Pp. 1-3, 1 fig. Central American Araliaceae - a precursory study for the Flora Mesoamericana. M.J. Cannon & J. P.M. Cannon. 1989. Pp. 5-61, 36 figs. A revision of the Solarium nitidum group (section Holo- phylla pro parte): Solanaceae. S. Knapp. 1989. Pp. 63-102, 21 figs. Six new species of Solanum sect. Geminata from South America. S. Knapp. 1989. Pp. 103-112, 8 figs. The application of names of some Indian species of Ocimum and Geniosporum (Labiatae). J.R. Press & V.V. Sivarajan. 1989. Pp. 113-116, 4 figs. Revision of Piper (Piperaceae) in the New World 1. Review of characters and taxonomy of Piper section Macrstachys. M.C. Tebbs. 1989. Pp. 117-158, 41 figs. £48.00 Volume 20 No. 1 Studies in the genus Hypericum L. (Guttiferae) 8. Sections 29. Brathys (part 2) and 30. Trigynobrathys. N.K.B. Robson. 1990. Pp. 1-151, 22 figs, 46 maps. £45.00 No. 2 The marine algal flora of Namibia: its distributions and affinities. G.W. Lawson, R.H. Simons and W.E. Isaac. 1990. Pp. 153-168, 1 fig, 7 plates. The infragenic classification of Gentiana (Gentianaceae). T.-N. Ho and S.-W. Liu. 1990. Pp. 169-192, 13 figs. Revision of Piper (Piperaceae) in the New World. 2. The taxonomy of Piper section Churumayu. M.C. Tebbs. 1990. Pp. 193-236, 49 figs. £25.00 Volume 21 No. 1 Historical and taxonomic studies in the genus Titanoderma (Rhodophyta, Corallinales) in the British Isles. Y.M. Chamberlain. 1991. Pp. 1-80, 247 figs. No. 2 Early collections of the Holy Thorn (Crataegus monogyna cv. Biflora). A.R. Vicky. 1991. Pp. 81-83. 1 fig- A taxonomic study of the species referred to the ascomycete genus Leptorhaphis . B. Aguirre-Hudson. 1991. Pp. 85-192. 76 figs. The typification and identification of Calymperes crassilimbatum Renauld & Cardot (Musci: Calymperaceae). L.T. Ellis. 1991. Pp. 193-194. 1 fig. Volume 22 No. 1 An account of southern Australian species of Lithophyllum (Corallinaceae, Rhodophyta). Wm. J. Woelkerling and S.J. Campbell. 1992. Pp. 1-107, 63 figs. £37.00 British Museum (Natural History) Publications THE CORALLINE RED ALGAE An analysis of the genera and subfamilies of the nongeniculate Corrallinaceae W.J. Woelkerling Nongeniculate coralline red algae (Corallinales, Rhodophyta) are both abundant in and vital to coral reefs, cementing the coral animal skeletons into a durable reef framework. In colder waters they form pavements of many square kilometres on the sea bed. It is one of the aims of this study to help resolve outstanding difficulties relating to taxonomy of the group. Extremely well illustrated with over 60 pages of scanning electron micrographs. 1988, 245 x 176 mm, 268 pp, 82 scanning electron micrographs, 259 figs, 19 tables. Co-published with Oxford University Press. 0 198 54249 6 Hardback £40.00 TAXONOMY AND BIOGEOGRAPHY OF MACQUARIE ISLANDS SEAWEEDS R.W. Ricker 1987, 175 x 250 mm, 344 pp, 126 figs. 0 565 00998 2 Hardback £40.00 SEAWEEDS OF THE BRITISH ISLES Vol. 1. Part 1. Rhodophyta. Introduction, Nemaliales, Gigartinales. P.S. Dixon & L.M. Irvine. 1977, xi + 252 pp, 90 figs. 0 565 00781 5 Paperback £15.00 Vol. 1. Part 2A. Cryptonemiales (sensu stricto), Palmariales, Rhodymeniales. L.M. Irvine. 1983, xii + 115pp, 28 figs. 0 565 00871 4 Paperback £15.00 Vol. 2. Chlorophyta. E.M. Burrows. 1991,256pp. 0 565 00981 8 Paperback £27.00 Vol. 3. Part 1. Fucophyceae (Phaeophyceae). R.L. Fletcher. 1987, 145 x 210 mm, 359 pp, 90 figs, 15 plates. 0 565 00992 3 Paperback £30.00 Vol. 4. Tribophyceae (Xanthophyceae). T. Christensen. 1987, 145 x 210 mm, 36 pp, 8 figs. 0 565 00980 X Paperback £7.50 CONTENTS 109 Palynological evidence for the generic delimitation of Sechium (Cucurbitaceae) and its allies J.L Alvarado, R. Lira-Saade and J. Caballero 123 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 3. Genera H-K J.H. Price, D.M. John and G.W. Lawson 147 Two new species of Solanum section Geminata (Solanaceae) from Cerro del Torra in western Colombia S. Knapp 153 Fissidens ceylonensis Dozy & Molkenb. (Musci = Fissidentaceae) and some allied taxa from southern India L T. Ellis 157 New species of Piper (Piperaceae) from Central America M. Tebbs 159 Studies on the Cretan flora 1. Floristic notes N.J. Turland 165 Studies on the Cretan flora 2. The Dianthus juniperinus complex (Caryophyllaceae) N.J. Turland illetin British Museum (Natural BOTANY SERIES Vol. 22, No. 2, November 1992