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Caribbean Journal of Science, Vol. 42, No. 1, 39-52, 2006Copyright 2006 College of Arts and SciencesUniversity of Puerto Rico, Mayagu¨ez A Checklist and Seasonal Account of the Deepwater Rhodophyta
around Cozumel Island on the Caribbean Coast of Mexico
UZ ELENA MATEO-CID *, A. CATALINA MENDOZA-GONZÁLEZ , AND 1Laboratorio de Ficología, Departamento de Botánica, Escuela Nacional de Ciencias Biológicas, I. P. N., Prol. Carpio y Plan de Ayala, 11340, México, D. F. 2Department of Biology, Duke University, Durham, NC 27708-0338 USA. *Corresponding author: lmateoc@ipn.mx ABSTRACT.—Red seaweeds were investigated, using SCUBA, from waters around the island of Cozumel
off the east coast of the Yucatan Peninsula in Mexico. One hundred and eighteen species of Rhodophyta were
collected and identified. Forty-five of these are reported for the first time from Cozumel, and of these 11 are
new records for Mexico. There was a distinct change in the deep water flora between wet and dry seasons;
the dry season had more species with northern ranges in comparison with the flora in general. The wet
season had few north-ranging species. Fifty-four epiphytic or hemiparasitic species were identified, most
frequently on Halimeda tuna, Dictyota cervicornis
and Sargassum spp. This account brings the total mac-
roalgal taxa reported from Cozumel to 385, the richest flora of any locality in the Caribbean region of Mexico.

KEYWORDS.—marine algae, benthic algae, red algae
greater depths. A list of the species col-lected, depths over which they were col- lected, months of collection, a sources for et al. 2000) we reported on the Phaeophyta illustrations and descriptions of each spe- and Chlorophyta collected in the deep wa- cies, and location of voucher specimens are have been seven previous references to the island of Cozumel and for Mexico are indi- cated in the table. Species names are ac- Taylor 1972; Garza 1975; Huerta and Garza- Barrientos 1980; Huerta et al. 1987; and Ma- Ortega et al. (2001) unless otherwise noted.
teo-Cid and Mendoza-González 1991),none of these reports were based on deep- water collections. Mateo-Cid and Men-doza-González reported a total of 168 taxa of Rhodophyta in the shallow waters of Co- zumel. Thirty seven and a half percent of (Table 1). Forty-five of these taxa were new these were present only during the wet sea- son and 6.5% only in the dry season.
new records for Mexico. There were signifi-cant changes in the flora with the change in the seasons (Appendix). Almost a third ofthe 168 taxa of red algae occurred as epi- phytes (Table 2). Notes on the distribution and classification of some taxa are given González et al. (2000). Collections were made from depths of 8-30 m, although ad-ditional plants could be seen in even Titanoderma pustulatum (J. V.
Lamour.) Nägeli ms. received June 8, 2005; accepted August 31, 2005 the genus Lithophyllum but in his notes on TABLE 1. List of deepwater rhodophyta from Cozumel Island, localities, and depths RhodophytaPorphyridiaceaeStylonema alsidii (Zanardini) K. M.
ErythrotrichiaceaeErythrotrichia carnea (Dillwyn) J.
CorallinaceaeAmphiroa fragilissima (L.) J. V.
Haliptilon cubense (Mont. ex Kütz.) Hydrolithon boergesenii (Foslie) Foslie Hydrolithon farinosum (J. V. Lamouroux) Lithoporella bermudensis (Foslie) W. H.
Titanoderma pustulatum (J. V.
Neogoniolithon accretum (Foslie & M.
Neogoniolithon mamillare (Harv.) Setchell Neogoniolithon fosliei (Heydrich) Setchell Neogoniolithon strictum (Foslie) Setchell *Notes: NC: New record from Cozumel Island; NM: New record from MexicoNumbers between [ ]: The ENCB Herbarium numberReferences to illustrations and descriptions of each taxon are listed using the following letter: A: Abbott (1990), B: Abbott (1999), C: Aponte et al. (1997), D: Børgesen (1920), E: Bula-Meyer (1985), F: Coppejans (1983), G:Cormaci and Furnari (1988), H: Cribb (1983), I: Foslie and Howe (1906), J: Fredericq and Norris (1986), K: Harvey(1849), L: Joly (1957), M: Joly (1965), N: Joly (1966), O: Kützing (1862), P: Littler and Littler (2000), Q: Norris andBucher (1982), R: Schmidt (1924), S: Schneider and Searles (1991), T: Taylor (1960), U: Wynne (1997). Reefs:PN-Paraiso North, Ch-Chankanab, To-Tormentos, Y-Yucab, Tu-Tunich, SF-San Francisco, SR-Santa Rosa, P.
Palancar, Co-Colombia.
Footnotes: Reported previously under the following names: 1. Audouinella hallandica (Kylin) Woelkerling, 2.
Audouinella hypneae (Børgesen) Schneider, 3. Amphiroa rigida var. antillana Børgesen, 4. Goniolithon accretum Foslieand M. Howe, 5. Porolithon mamillare var. occidentalis Lemoine, 6. Goniolithon solubile Foslie and Howe, 7. Liagorafarinosa J. V. Lamouroux, 8. Liagora decussata Montagne, 9. Galaxaura lapidescens (Ellis and Solander) J. V. Lamour-oux, 10. Galaxaura oblongata (Ellis and Solander) J. V. Lamouroux, 11. Peyssonnelia atlantica Schneider andReading, 12. Gracilaria cylindrica Børgesen, 13. Gloioderma atlanticum Searles, 14. Ceramium fastigiatum (Roth)Harvey, 15. Dasya arbuscula (Dillwyn) C. Agardh, 16. Martensia pavonia (J. Agardh) J. Agardh, 17. Herposiphoniatenella (C. Agardh) Ambronn.
GelidiaceaeGelidium pusillum (Stackhouse) Le Jolis GelidiellaceaeGelidiella acerosa (Forssk.) Feldmann & LiagoraceaeGanonema farinosum (J. V. Lamouroux) Liagora ceranoides J. V. Lamouroux Trichogloeopsis pedicellata (M. Howe) GalaxauraceaeGalaxaura marginata (J. Ellis & Solander) Galaxaura rugosa (J. Ellis & Sol.) J. V.
Scinaia complanata (Collins) Cotton Tricleocarpa fragilis (L.) Huisman & BonnemaisoniaceaeAsparagopsis taxiformis (Delile) Trevisan DumontiaceaeAcrosymphyton caribaeum (J. Agardh) G.
HypneaceaeHypnea spinella (C. Agardh) Kütz.
KallymeniaceaeKallymenia westii Ganesan NemastomataceaePredaea feldmannii Børgesen PeyssonneliaceaePeyssonnelia boudouresquei Yoneshigue11 Peyssonnelia conchicola Picc. & Grunow *Notes: NC: New record from Cozumel Island; NM: New record from MexicoNumbers between [ ]: The ENCB Herbarium numberReferences to illustrations and descriptions of each taxon are listed using the following letter: A: Abbott (1990), B: Abbott (1999), C: Aponte et al. (1997), D: Børgesen (1920), E: Bula-Meyer (1985), F: Coppejans (1983), G:Cormaci and Furnari (1988), H: Cribb (1983), I: Foslie and Howe (1906), J: Fredericq and Norris (1986), K: Harvey(1849), L: Joly (1957), M: Joly (1965), N: Joly (1966), O: Kützing (1862), P: Littler and Littler (2000), Q: Norris andBucher (1982), R: Schmidt (1924), S: Schneider and Searles (1991), T: Taylor (1960), U: Wynne (1997). Reefs:PN-Paraiso North, Ch-Chankanab, To-Tormentos, Y-Yucab, Tu-Tunich, SF-San Francisco, SR-Santa Rosa, P.
Palancar, Co-Colombia.
Footnotes: Reported previously under the following names: 1. Audouinella hallandica (Kylin) Woelkerling, 2.
Audouinella hypneae (Børgesen) Schneider, 3. Amphiroa rigida var. antillana Børgesen, 4. Goniolithon accretum Foslieand M. Howe, 5. Porolithon mamillare var. occidentalis Lemoine, 6. Goniolithon solubile Foslie and Howe, 7. Liagorafarinosa J. V. Lamouroux, 8. Liagora decussata Montagne, 9. Galaxaura lapidescens (Ellis and Solander) J. V. Lamour-oux, 10. Galaxaura oblongata (Ellis and Solander) J. V. Lamouroux, 11. Peyssonnelia atlantica Schneider andReading, 12. Gracilaria cylindrica Børgesen, 13. Gloioderma atlanticum Searles, 14. Ceramium fastigiatum (Roth)Harvey, 15. Dasya arbuscula (Dillwyn) C. Agardh, 16. Martensia pavonia (J. Agardh) J. Agardh, 17. Herposiphoniatenella (C. Agardh) Ambronn.
Peyssonnelia nordstedtii Weber-van Peyssonnelia stoechas Boudour. & RhizophyllidaceaeOchtodes secundiramea (Montagne) SolieriaceaeFlahaultia tegetiformans HalymeniaceaeCryptonemia crenulata (J. Agardh) GracilariaceaeGracilaria blodgettii Harvey12 ChampiaceaeChampia parvula (C. Agardh) Champia parvula var. prostrata LomentariaceaeLomentaria rawitscheri A. B. Joly RhodymeniaceaeBotryocladia pyriformis (Børgesen) Botryocladia spinulifera W. R. Taylor Coelarthrum cliftonii (Harvey) Kylin *Notes: NC: New record from Cozumel Island; NM: New record from MexicoNumbers between [ ]: The ENCB Herbarium numberReferences to illustrations and descriptions of each taxon are listed using the following letter: A: Abbott (1990), B: Abbott (1999), C: Aponte et al. (1997), D: Børgesen (1920), E: Bula-Meyer (1985), F: Coppejans (1983), G:Cormaci and Furnari (1988), H: Cribb (1983), I: Foslie and Howe (1906), J: Fredericq and Norris (1986), K: Harvey(1849), L: Joly (1957), M: Joly (1965), N: Joly (1966), O: Kützing (1862), P: Littler and Littler (2000), Q: Norris andBucher (1982), R: Schmidt (1924), S: Schneider and Searles (1991), T: Taylor (1960), U: Wynne (1997). Reefs:PN-Paraiso North, Ch-Chankanab, To-Tormentos, Y-Yucab, Tu-Tunich, SF-San Francisco, SR-Santa Rosa, P.
Palancar, Co-Colombia.
Footnotes: Reported previously under the following names: 1. Audouinella hallandica (Kylin) Woelkerling, 2.
Audouinella hypneae (Børgesen) Schneider, 3. Amphiroa rigida var. antillana Børgesen, 4. Goniolithon accretum Foslieand M. Howe, 5. Porolithon mamillare var. occidentalis Lemoine, 6. Goniolithon solubile Foslie and Howe, 7. Liagorafarinosa J. V. Lamouroux, 8. Liagora decussata Montagne, 9. Galaxaura lapidescens (Ellis and Solander) J. V. Lamour-oux, 10. Galaxaura oblongata (Ellis and Solander) J. V. Lamouroux, 11. Peyssonnelia atlantica Schneider andReading, 12. Gracilaria cylindrica Børgesen, 13. Gloioderma atlanticum Searles, 14. Ceramium fastigiatum (Roth)Harvey, 15. Dasya arbuscula (Dillwyn) C. Agardh, 16. Martensia pavonia (J. Agardh) J. Agardh, 17. Herposiphoniatenella (C. Agardh) Ambronn.
CeramiaceaeAglaothamnion cordatum (Børgesen) Aglaothamnion boergesenii (Aponte et Antithamnion decipiens (J. Agardh) Antithamnionella boergesenii (Cormaci Callithamniella tingitana (Schusb. ex Centroceras clavulatum (C. Agardh in Centrocerocolax ubatubensis A. B. Joly caraibicum H. E. Petersen &Børgesen Ceramium luetzelburgii O. C. Schmidt Griffithsia globulifera Harvey. ex *Notes: NC: New record from Cozumel Island; NM: New record from MexicoNumbers between [ ]: The ENCB Herbarium numberReferences to illustrations and descriptions of each taxon are listed using the following letter: A: Abbott (1990), B: Abbott (1999), C: Aponte et al. (1997), D: Børgesen (1920), E: Bula-Meyer (1985), F: Coppejans (1983), G:Cormaci and Furnari (1988), H: Cribb (1983), I: Foslie and Howe (1906), J: Fredericq and Norris (1986), K: Harvey(1849), L: Joly (1957), M: Joly (1965), N: Joly (1966), O: Kützing (1862), P: Littler and Littler (2000), Q: Norris andBucher (1982), R: Schmidt (1924), S: Schneider and Searles (1991), T: Taylor (1960), U: Wynne (1997). Reefs:PN-Paraiso North, Ch-Chankanab, To-Tormentos, Y-Yucab, Tu-Tunich, SF-San Francisco, SR-Santa Rosa, P.
Palancar, Co-Colombia.
Footnotes: Reported previously under the following names: 1. Audouinella hallandica (Kylin) Woelkerling, 2.
Audouinella hypneae (Børgesen) Schneider, 3. Amphiroa rigida var. antillana Børgesen, 4. Goniolithon accretum Foslieand M. Howe, 5. Porolithon mamillare var. occidentalis Lemoine, 6. Goniolithon solubile Foslie and Howe, 7. Liagorafarinosa J. V. Lamouroux, 8. Liagora decussata Montagne, 9. Galaxaura lapidescens (Ellis and Solander) J. V. Lamour-oux, 10. Galaxaura oblongata (Ellis and Solander) J. V. Lamouroux, 11. Peyssonnelia atlantica Schneider andReading, 12. Gracilaria cylindrica Børgesen, 13. Gloioderma atlanticum Searles, 14. Ceramium fastigiatum (Roth)Harvey, 15. Dasya arbuscula (Dillwyn) C. Agardh, 16. Martensia pavonia (J. Agardh) J. Agardh, 17. Herposiphoniatenella (C. Agardh) Ambronn.
DasyaceaeDasya baillouviana (S. G.
Dasya haitiana Fredericq & J. N.
Heterosiphonia crispella var. laxa DelesseriaceaeHypoglossum hypoglossoides *Notes: NC: New record from Cozumel Island; NM: New record from MexicoNumbers between [ ]: The ENCB Herbarium numberReferences to illustrations and descriptions of each taxon are listed using the following letter: A: Abbott (1990), B: Abbott (1999), C: Aponte et al. (1997), D: Børgesen (1920), E: Bula-Meyer (1985), F: Coppejans (1983), G:Cormaci and Furnari (1988), H: Cribb (1983), I: Foslie and Howe (1906), J: Fredericq and Norris (1986), K: Harvey(1849), L: Joly (1957), M: Joly (1965), N: Joly (1966), O: Kützing (1862), P: Littler and Littler (2000), Q: Norris andBucher (1982), R: Schmidt (1924), S: Schneider and Searles (1991), T: Taylor (1960), U: Wynne (1997). Reefs:PN-Paraiso North, Ch-Chankanab, To-Tormentos, Y-Yucab, Tu-Tunich, SF-San Francisco, SR-Santa Rosa, P.
Palancar, Co-Colombia.
Footnotes: Reported previously under the following names: 1. Audouinella hallandica (Kylin) Woelkerling, 2.
Audouinella hypneae (Børgesen) Schneider, 3. Amphiroa rigida var. antillana Børgesen, 4. Goniolithon accretum Foslieand M. Howe, 5. Porolithon mamillare var. occidentalis Lemoine, 6. Goniolithon solubile Foslie and Howe, 7. Liagorafarinosa J. V. Lamouroux, 8. Liagora decussata Montagne, 9. Galaxaura lapidescens (Ellis and Solander) J. V. Lamour-oux, 10. Galaxaura oblongata (Ellis and Solander) J. V. Lamouroux, 11. Peyssonnelia atlantica Schneider andReading, 12. Gracilaria cylindrica Børgesen, 13. Gloioderma atlanticum Searles, 14. Ceramium fastigiatum (Roth)Harvey, 15. Dasya arbuscula (Dillwyn) C. Agardh, 16. Martensia pavonia (J. Agardh) J. Agardh, 17. Herposiphoniatenella (C. Agardh) Ambronn.
RhodomelaceaeAcanthophora spicifera (Vahl) Chondria curvilineata Collins & Chondria dasyphylla (Woodward) C.
Chondria platyramea A. B. Joly & Chondria polyrhiza Collins & Hervey Chondrophycus corallopsis (Montagne) Chondrophycus poiteaui (J. V.
Digenea simplex (Wulfen) C. Agardh Herposiphonia secunda f. tenella (C.
Laurencia intricata J. V. Lamouroux Laurencia obtusa (Hudson) J. V.
Lophocladia trichoclados (C. Agardh) Micropeuce mucronata (Harvey) Kylin Neosiphonia flaccidissima (Hollenberg) Neosiphonia gorgoniae (Harvey) S. M.
Neosiphonia sphaerocarpa (Børgesen) Polysiphonia atlantica Kapraun & J. E.
Polysiphonia ferulacea Suhr ex J.
*Notes: NC: New record from Cozumel Island; NM: New record from MexicoNumbers between [ ]: The ENCB Herbarium numberReferences to illustrations and descriptions of each taxon are listed using the following letter: A: Abbott (1990), B: Abbott (1999), C: Aponte et al. (1997), D: Børgesen (1920), E: Bula-Meyer (1985), F: Coppejans (1983), G:Cormaci and Furnari (1988), H: Cribb (1983), I: Foslie and Howe (1906), J: Fredericq and Norris (1986), K: Harvey(1849), L: Joly (1957), M: Joly (1965), N: Joly (1966), O: Kützing (1862), P: Littler and Littler (2000), Q: Norris andBucher (1982), R: Schmidt (1924), S: Schneider and Searles (1991), T: Taylor (1960), U: Wynne (1997). Reefs:PN-Paraiso North, Ch-Chankanab, To-Tormentos, Y-Yucab, Tu-Tunich, SF-San Francisco, SR-Santa Rosa, P.
Palancar, Co-Colombia.
Footnotes: Reported previously under the following names: 1. Audouinella hallandica (Kylin) Woelkerling, 2.
Audouinella hypneae (Børgesen) Schneider, 3. Amphiroa rigida var. antillana Børgesen, 4. Goniolithon accretum Foslieand M. Howe, 5. Porolithon mamillare var. occidentalis Lemoine, 6. Goniolithon solubile Foslie and Howe, 7. Liagorafarinosa J. V. Lamouroux, 8. Liagora decussata Montagne, 9. Galaxaura lapidescens (Ellis and Solander) J. V. Lamour-oux, 10. Galaxaura oblongata (Ellis and Solander) J. V. Lamouroux, 11. Peyssonnelia atlantica Schneider andReading, 12. Gracilaria cylindrica Børgesen, 13. Gloioderma atlanticum Searles, 14. Ceramium fastigiatum (Roth)Harvey, 15. Dasya arbuscula (Dillwyn) C. Agardh, 16. Martensia pavonia (J. Agardh) J. Agardh, 17. Herposiphoniatenella (C. Agardh) Ambronn.
TABLE 2. Epiphytic species and their host from Isla Cozumel Dictyota cervicornis Kützing, Wrangelia penicillata Chrysymenia enteromorpha, Udotea dixonii Littler & Littler Halimeda goreauii W. R. Taylor, Avrainvillea longicaulis (Kützing) Murray & Boodle Laurencia intricata, Halimeda discoidea Decaisne Udotea flabellum (Ellis et Solander) J. V. Lamouroux Dictyota cervicornis, Halimeda tuna (J. Ellis & Solander) J. V. Lamouroux Champia parvula var. prostrata Champia parvula, Halimeda discoidea, Dictyota cervicornis Coelarthrum albertesii, Amphiroa tribulus Amphiroa tribulus, Chondrophycus poiteaui, Halimeda tuna Polysiphonia ferulacea, Udotea flabellum Dictyota cervicornis, Gelidiopsis intricate Halimeda tuna (Ellis et Sol.) J. V. Lamouroux Champia parvula, Halimeda tuna, Caulerpa cupressoides (H. West in Vahl) C. Agardh Galaxaura rugosa, Sargassum polyceratium Montagne Halimeda incrassata (J. Ellis) J. V. Lamouroux Rhipocephalus phoenix f. longifolius A. Gepp et E. S. Gepp Heterosiphonia crispella var. laxa Amphiroa tribulus, Halimeda discoidea Herposiphonia secunda f. tenella Udotea loensis D. Littler & M. Littler, Halimeda tuna Caulerpa cupressoides, Sargassum vulgare Trichogloeopsis pedicellata, Chondrophycus corallopsis Caulerpa mexicana (Sonder) J. Agardh the genus gave recent references in which Titanoderma is discussed as a genus distinct ported localities are Florida (Bucher et al, from Lithophyllum. Based on a cla- 1990), Puerto Rico (Ballantine 1985), and distic analysis of 18S rRNA gene sequences, Bailey (1999) indicated that Titanoderma isphylogenetically distant from Lithophyllum, Centrocerocolax ubatubensis A. B. Joly This parasite was originally described by Acrosymphyton caribaeum (J. Agardh) Joly (1966) from Brazil and was not known from another locality until its report from Venezuela by Aponte and Ganesan (1990).
from the western Caribbean. Previous rec- This is the third and northernmost record ords were from Puerto Rico (Ballantine and Norris 1989) and Saba Bank (Taylor 1960),Bermuda (Schneider 2003) and Florida described from St. Helena in the South At- lantic by Børgesen (1950). Since then it has been reported from West Africa (John et al.
1977), the Azores (Fredericq et al. 1992), and Mateo-Cid et al. (2002, 2003), but 45 are new records for Cozumel; this represents a (1997) from Puerto Rico. This species ap- recorded less than a 35% increase for the throughout its range. The type collection from St. Helena came from a depth of 50 m, phyta now reported from Cozumel is 216.
and the shallowest depth reported was 10- 27 m in Ghana (Lawson and John 1982).
121 Chlorophyta (Mendoza-González et al.
Gloiocladia atlantica (Searles) R. E. Norris 2000, 2002), total 385 taxa. Thus, CozumelIsland is the richest seaweed area reported species in the Caribbean. It was also re- next richest region reported is the adjacent United States as Gloioderma atlantica (Sear- where Dreckman et al. (1996) reported 219 les 1972) and from Puerto Rico (Ballantine taxa. Littler and Littler (1997) reported 181 1984), and Bermuda (Schneider and Searles in their account of the marine flora of the that this species may be conspecific with Caribbean island for which there is a recent the Pacific G. iyoensis (Okamura) R. E. Nor- floristic report is Puerto Rico (Ballantine ris and thus may have a wider distribution.
and Aponte 1997), and for this larger island471 Phaeophyta, Chlorophyta and Rhodo- Botryocladia spinulifera W. R. Taylor and I.
land are clearly richer in species for all lands (Taylor and Abbott 1973), this alga is three taxonomic divisions than the deeper waters. A significant fraction of the marine flora seems restricted to deeper waters ac- April, and the rainy season is from May to cessible only by diving, or potentially, by January (García 1973). In our study, the dry the taxa reported in this study and by Men- restricted to the wet season and 26 (22%) to only in deep water (23% of the 385 species the dry season (Table 2). In the shallow wa- ter collections from Cozumel of Mateo-Cidand Mendoza-González (1991), sixty-five (39%) of the 168 taxa of Rhodophyta re-ported were found only in the rainy season At a latitude of 20ºN, Cozumel lies well within the tropics, with surface water tem- Seven of the 26 species restricted to deep from Cozumel only in the dry season. Con- versely, of the 32 species collected in deep species, 3% of all deep water red algal taxa, water only in the wet season (spring, sum- were collected in the four seasons: Amphi- mer, or fall), six were recorded by Mateo- roa fragilissima, Neogoniolithon mammillare and Champia parvula. Only 20 (23%) of the their dry season, shallow water collections, 88 taxa present in winter (March) were still leaving 24 species in deep water restricted, present in spring (May). Of the 40 taxapresent in spring, 17 (40%), were still found in summer (June). Thirteen of the 39 sum- mer taxa (33%) were still present in the fall (September), and 27 of the 48 fall taxa (56%) were still present at the end of winter. Fifty- eight taxa were found to occur only during deep water brown algal taxa) and 21 Chlo- spring, 5 in summer and 9 in fall (Table 2).
rophyta (28% of all deep water green algal Certainly some taxa were present, but over- Mendoza-González, 1991), collections were should be noted that the summer collecting most diverse in the wet season with sixty- effort was not as thorough as in the other three (37.5%) of the 168 taxa of Rhodophyta seasons. The presence of Ampiroa rigida in reported found only in the wet season and vening seasons, and Peyssonnelia inamoena season. By comparison, in this study of dry study of phenology of macroalgae in a shal- low water site, just to the north, at Isla Mu- 32 (27%) to the wet season and 26 (22%) to jeres, Quan-Young et al. (2004) also noted multiple peaks in size and reproductive ac- Although seasons in the tropics are char- tivity of taxa studied in detail and some acterized by more subtle differences in tem- species, including Amphiroa rigida, were also only present sporadically. Species rich- compared to temperate and boreal regions, ness of red algae at that site was highest in the data in our collections suggest a strong January and again in June with low points seasonality in the red algal flora of deep waters. This interesting ecological feature of this Caribbean algal flora requires fur- nial species persisting through the winter, and noted that the deepwater flora wascomposed almost entirely of tropical spe- cies with centers of distribution in the Car- groupings of Caribbean seaweeds in theWestern Atlantic: 1. Strictly Caribbean (in- cluding Bermuda), 2. Caribbean and north- In the study area, 54 taxa of epiphytic or ern, 3. Caribbean and southern, and 4.
hemiparasite combinations (Table 3). They lished ranges he reported 31% strictly Car- were most frequently observed on calcified tions), especially on Halimeda tuna. Among spread north and south. The 118 species re- the Phaeophyta, Dictyota and Sargassum were common hosts, particularly D. cervi- (26%) restricted in the Western Atlantic to cornis and S. vulgare. No particular Rhodo- although articulated, coralline red algae Carolina, 37 (31%) from the Caribbean and such as Amphiroa tribulus were hosts to sev- percentages are similar to those of Taylor.
ready observed in the Cozumel Island shal- low water flora (Huerta et al. 1987; Mateo- C i d a n d M e n d o z a - G o n z á l e z 1 9 9 1 , only in the dry or wet season, there is a Mendoza-González and Mateo-Cid 1992).
clear biogeographic signal from those spe- Most epiphytes were taxa with small, often cies with a northward distribution. Of the finely filamentous plants of the Ceramiac- 19 species restricted to the dry season, 31% are strictly Caribbean, 32% Caribbean and be an important ecological strategy where hosts serve as a secondary substrate when the 24 species restricted to the wet season, strated by Hay (1986), the unpalatable na- epiphyte a degree of protection from graz- and south. These data show a strong north- ing, or (Wahl and Hay 1995) an unpalatable ern component in the dry season flora with tion from grazing for a palatable host.
ibbean and northern species present and amarked drop in that component of the flora Acknowledgments.—Collection permits Earle’s (1969), report on the Phaeophyta Instituto Nacional de Ecología of the Secre- of the Eastern Gulf of Mexico, noted sea- sonality in that flora with a distinctive win- rales y Pesca of Mexico. Logistical support was provided by the Armada de México.
with northern distributions. Further north in North Carolina, Hoyt (1920) was the first without the diving assistance of Erin Krell- Schneider (1976) reported that the offshore, thanks the Comisión y Fomento de Activi- deepwater flora has a peak in species rich- dades Académicas del I.P.N. for a fellow- ness in the summer with only a few peren- cil provided funds for travel and diving.
Barrier Reef. I: Rhodophyta. Australian Coral Reef Society, Brisbane. 1-173 + [2] p. Pls. 1-71.
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latum, Champia parvula, Chondria curvilin- eata, Chondrophycus poiteaui, Coelothrix ir- mudensis, Lophocladia trichoclados, Neogonio- regularis, Cryptonemia crenulata, Dasya lithon mamillare, Neosiphonia flaccidissima, rigidula, Dudresnaya crassa, Galaxaura rugosa, Peyssonnelia inamoena, Peyssonnelia nordst- Ganonema farinosum, Gelidiella acerosa, Hy- edtii, Peyssonnelia simulans, Polysiphonia drolithon boergesenii, Hypoglossum hypoglos- ferulacea, Scinaia complanata, Seirospora occi- soides, Jania adhaerens, Jania capillacea, dentalis, Spyridia filamentosa, Trichogloeopsis Laurencia intricata, Laurencia obtusa, Liagora pedicellata, Wrangelia bicuspidata, Wrangelia albicans, Liagora ceranoides, Lophocladia trichoclados, Martensia fragilis, Neogoniolithon Winter: Acanthophora spicifera, Acrochae-
fosliei, Neogoniolithon mammillare, Nithophyl- tium hallandicum, Acrochaetium hypneae, lum adhaerens, Ochtodes secundiramea, Aglaothamnion boergesenii, Aglaothamnion Peyssonelia conchicola, Peyssonelia inamoena, cordatum, Amphiroa fragilissima, Amphiroa Peyssonelia nordstedtii, Peyssonelia stoechas, tribulus, Anotrichium tenue, Antithamnion de- Polysiphonia ferulacea, Scinaia complanata, cipiens, Antithamnionella boergesenii, Aspara- Tricleocarpa fragilis, Trichogloeopsis pedicel- gopsis taxiformis, Botryocladia pyriformis, Bot- ryocladia spinulifera, Centroceras clavulatum, Summer: Aglaothamnion boergesenii, Am-
Centrocerocolax ubatubensis, Ceramium cim- phiroa fragilissima, Amphiroa rigida, An- bricum, Ceramium flaccidum, Ceramium lepto- otrichium tenue, Antithamnionella boergesenii, zonum, Ceramium leutzelburgii, Champia par- Botryocladia spinulifera, Ceramium brevizona- vula, Champia parvula var. prostrata, Champia tum var. caraibicum, Ceramium leutzelburgii, salicornioides, Chondria curvilineata, Chondro- Champia parvula, Champia salicornioides, phycus corallopsis, Chondrophycus poiteaui Chondria polyrhiza, Chondrophycus corallop- Chrysymenia halymenioides, Coelarthron clifto- sis, Coelarthrum cliftonii, Crouania attenuata, nii, Coelothrix irregularis, Crouania attenuata, Dasya mollis, Dasya rigidula, Gelidiopsis in- Dasya baillouviana, Dasya caraibica, Dasya trincata, Griffithsia globulifera, Heterosiphonia haitiana, Dasya hutchinsiae, Dasya mollis, crispella var. laxa, Hydrolithon farinosum, Hy- Dasya ocellata, Dasya rigidula, Dudresnaya poglossum hypoglossoides, Hypoglossum tenui- crassa, Erytrotrichia carnea, Flahaultia tegeti- folium, Jania capillacea, Titanoderma pustula- formans, Galaxaura marginata, Galaxaura ru- tum, Laurencia obtusa, Liagora ceranoides, gosa, Gelidium pusillum, Gloiocladia atlantica, Liagora pinnata, Lomentaria rawitscheri, Mar- Gracilaria blodgettii, Griffithsia globulifera, tensia fragilis, Neogoniolithon accretum, Neo- Griffithsia radicans, Haliptilon cubense, goniolithon mammillare, Neosiphonia sphaero- Haloplegma duperreyi, Herposiphonia bipin- carpa, Nithophyllum adhaerens, Polysiphonia nata, Herposiphonia secunda f. tenella, Hetero- atlantica, Seirospora occidentalis, Spermotham- siphonia crispella var. laxa, Hydrolithon farino- nion investiens, Trichogloeopsis pedicellata, sum, Hypoglossum hypoglossoides, Jania Wrangelia argus, Wrangelia penicillata. adhaerens, Jania capillacea, Kallymenia westii, Fall: Acanthophora spicifera, Acrosymphy-
Laurencia obtusa, Liagora ceranoides, Liagora ton caribaeum, Amphiroa fragilissima, Amphi- pinnata, Liagora valida, Lithoporella bermuden- roa tribulus, Botryocladia pyriformis, Cal- sis, Lomentaria rawitscheri, Martensia fragilis, lithamniella tingitana, Centroceras clavulatum, Mesophyllum mesomorphum, Micropeuce mu- Champia parvula, Champia parvula var. cronata, Neogoniolithon accretum, Neogonio- prostrata, Champia salicornioides, Chondria lithon fosliei, Neogoniolithon mammillare, Neo- curvilineata, Chondria dasyphylla, Chondria goniolithon strictum, Neosiphonia flacci- platyramea, Chondrophycus corallopsis, Chon- dissima, Neosiphonia gorgoniae, Neosiphonia drophycus poiteaui, Crouania attenuata, Dasya sphaerocarpa, Nithophyllum adhaerens, baillouviana, Dasya caraibica, Dasya haitiana, Ochtodes secundiramea, Peyssonnelia boudour- Dasya ocellata, Dasya rigidula, Digenea sim- esquei, Pneophyllum fragile, Predaea feldma- plex, Gelidiopsis intrincata, Gracilaria blodget- nnii, Scinaia complanata, Seirospora occiden- tii, Halodictyon mirabile, Hydrolithon farino- talis, Spyridia filamentosa, Stylonema alsidii, sum, Hypnea spinella, Jania adhaerens, Taenioma nanum, Tricleocarpa fragilis, Wran- Laurencia intricata, Laurencia obtusa, Liagora gelia argus, Wrangelia bicuspidata, Wrangelia albicans, Liagora pinnata, Lithoporella ber-

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