381 West et af.: Caloglossa apomeiolica sp. nov. Botanica Marina Vol. 37, pp. 381-390,1994 , i Offprint Caloglossa apomeiotica sp. nov. (Ceramiales, Rhodophyta) from Pacific Mexico J. A. West l , G. C. Zuccarello*, F. F. Pedroche**' and U. Karsten*** School of Botany, University of Me/bollme, Parkville, Victoria 3052, Allstralia * Department of Biology, University of Califa mia, Santa Crllz, California 95064, U. S. A. ** Departamenta de Hie/robialagia, Universidad Au/6nama Metropolitana-I, Aparwdo Poswl 5j-535, Mexico DF 09340, Mexico *** J\1ax-Planck-Instiwtjiir Marine l\1ikrobiologie, Fahrenhei/s/l: 1, 28359 Bremen, FR. o/Germany (Accepted 25 March 1994) Abstract A new species, Caloglossa apomeio/iea West el Zuccarello, is described based on material from San Carlos, Bahia Magdalena, Bajo Califomia Sur. The species is morphologically similar to C. lepriellrii but characterised by the production of bisporangia and, less commonly, tetrasporangia, and the lack of sexually reproducing plants. In eight culture isolates from throughout the geographic range of the species only the bisporangia were viable. The present known distribution of Caloglossa apomeialiea is from Bahia Magdalena, Baja California Sur to BalTa de Navidad, Jalisco in Pacific Mexico. It is suggested that C. apo111eioliea is derived from C. leprieurii populations flllther south in the Americas through a loss of sexual reproduction. Based on our field collections in March, 1993 the nearest populations of sexually reproducing C. leprieurii are 1200 kl1l away in Estero las Garzas, Chiapas, Mexico and extend south into Guatemala. Introduction Caloglossa is one of the most widely distributed red algal genera on tropical and temperate coasts, often associated with mangroves, Sparlil/a and other substrata in marine estuarine habitats as well as in freshwater streams. Post (1936) summarized the systematics and geographic distribution of Caloglossa species and in later papers (Post 1943, 1963, 1967) considered the morphology and anatomy as well as providing further notes on the ecology and distribution. In an extensive investigation on the morphology and anatomy of Caloglossa, Papenfuss (1961) provided a complete analysis of vegetative and reproductive development in C. leprieurh (Montagne) 1. Agardh. Most recently King and Puttock Address for correspondence Present address: University Herbarium, University of California, Berkeley, CA 94720, U.S.A. I 2 Botanica Marina / Vol. 37 / 1994/ Fase. 4 Copyright © 1994 by Walter d.G Gruyter . Berlin· New York (1994) have compiled a monograph of the genus and redefined several species. Although the genus is widely distributed and considered to be characteristic of mangrove habitats it has been recorded only few times in Pacific Mexico (Ortega el al. 1987, Mateo-Cid and Mendoza-Gonzalez 1991, West and Zuccarello 1990, West el al. 1992 b), where mangroves extend south from Bahia San Ignacio (Jat. 27°00' N, long. 113° I0' W) on the Pacific Ocean side and from Bahia de Los Angeles (lat. 28°55' N, long. 113°30' W) of the Baja California peninsula (Robel1s 1989) as well as from Puel10 Lobos, Sonora (Iat. 30°15 1 N, long. 112°4Y W) south along the mainland of Pacific Mexico (West 1977, Flores-Verdugo el al. 1992). In these areas BosllJ'chia radicalls (Montagne) Montagne is the dominant red alga associated with the pneumatophores of Avieennia germiJlans (L.) L. and Lagul1ciliaria racemosa (L.) Gael1h and the prop roots of Rhi- 382 zophora mangle L. (Dawson 1962, Cordeiro-Marino el al. 1992). In the first report of Caloglossa for the coast of Baja California, Pacific Mexico, West and Zuccarello (1990) noted the presence of sporophytes and the absence of gametophytes at Bahia Balandra and Bahia Magdalena. Earlier investigators (Dawson 1962, Huerta and Mendoza-Gonzalez 1985) working in these areas had not recorded Caloglossa. When living plants from Baja were returned to laboralory, microscopic examination revealed the presence of bisporangia and tetrasporangia. Culture investigations showed that the bispores germinated readily and the successive generations also formed bisporangia. Tetrasporangia were also seen in the same plants and no gameto- phytes developed in culture. Subsequent collections and culture work in 1991 and 1992 (West el al. 1992 b) from other localities (Table I) confirmed these initial observations. Previous records for Caloglossa along the Pacific mainland coast of Mexico (Ortega el al. 1987, Mateo-Cid and Mendoza-Gonzalez 1991) did not consider reproductive phases. Material and Methods Collection and culture methods are as described in West el al. (1992 b). Collections were made at the localities and dates indicated in Table I. Temperature, salinity and irradiance levels were taken respectively, with a standard stainless-steel-jacketed centigrade thermometer, Reichert temperature-compensated refractometer calibrated in parts per thousand (%0) (Warner-Lambert West et at.: Caloglossa apomeiOfica sp. nov. Techn., Inc., Buffalo, NY) and a QSI-I40B integrating quantum scalar irradiance meter (Biospherical Instruments, Inc., San Diego, CA, USA). In the field, specimens were first examined with a lOX hand lens and then with a Nikon H field microscope at 100X to verify their reproductive state. Herbarium mounts and liquid- preserved (5% formaldehyde in sea water) specimens were prepared at most sampling sites. Voucher speci- mens are deposited in the UC Herbarium, Berkeley, the UAMIZ Herbarium (Universidad Autanoma Metropolitana-I) and the UABCS Herbarium (Universidad Autanoma Baja California Sur). A culture of the holotype (isolate 3025) collected at San Carlos, Bahia Magdalena, Baja California Sur, is deposited at the Culture Collection of Algae, Department of Botany, University of Texas, Austin, TX 78713-7640. Cytological and anatomical preparations were made as described by Goff and Coleman (1987,1990) and West and Calumpong (1988). To obtain chromosome counts cultured plants were fixed at two-hour intervals in the dark and light cycles (12 : 12, L: D) but the mitotic division rates were so low that no satisfactory preparations were obtained. For D-mannitol analyses algal samples were air-dried at 25°C in the laboratory or oven-dried at 70 DC, both overnight. Each 10-15 mg dry weight (dw) sample was extracted in I mL 70% ethanol (v/v) for 3 -4 h in a waterbath at 70°C. Following extraction, samples were centrifuged at 4500 g for 10 min. The supernatants (700 JlL) were evaporated to dryness under vacuum (Speed Vac Concentrator SVC 100H, Bachofer Ltd, Germany). Dried samples were re-dissolved in 700 JlL distilled Table I. Collection locality and dare, culture number and reproduction of Ca/og/ossa apomeiolica sp. nov. Locality and date Culture No. Reproduction San Carlos, Bahia Magdalena, Baja California Sur (8. C. S.), Mexico 6 January 1990 3025 bisporangialtetrasporangia Bahia Balandra. B. C. S., Mexico 6 January 1990 3033 bisporangialtetrasporangia Bahia Magdalena, B. C. S. Mexico 25 March 1991 313t bisporangialtetrasporangia Barra de Navidad, Jalisco, Mexico 19 March 1992 3243 bisporangialtetrasporangia Barra de Navidad, Jalisco, Mexico 19 March 1992 3244 bisporangialtelrasporangia San Bias, Nayarit, Mexico 22 March 1992 3245 bisporangialtetrasporangia Teacapan, Sinaloa, Mexico 20 March 1992 3246 bisporangia/tetrasporangia Isla Espiritu Santo, B. C. S., Mexico 19 May 1992 3276 bisporangia/tetrasporangia Botanica Marina I Vol. 37 I 1994 I Fasc. 4 West ttl 383 al.: Caloglossa apomeiotica sp. nov. water and directly utilized for analysis. The D-mannitol was quantified by HPLC with a refractive index detector. All analytical conditions are described by Karsten el al. (1991). Observations and Discussion Caloglossa apomeiolica J. West el G. Zuccarello sp. nov. Diagnosis Plantae tanfum in statu spomphytieo existentes, 10- 30 mm 10llgae, 0.5 - 1.5 mm latae, planae, bi/ureatae, ad nodos leviter eonstrictae. Nodi rhizoidea non nisi basaliter instructi. Laminae cum 10-20 alis-cellulis in quoque la/ere cellulae axialis. Bisporangia tetrasporangiave 6-8 in quoque segmento axiali, S01"OS distinctos formantes, bisporangiis maluris ellipsoidibus 30-44 X 40-62 f.tln el telrasporangiis 40-49 X 45-68 pm. Formatio evolutioque sporangialis Caloglossae leprieurii similis. Bisporae uninucleatae 50-64 pm diam., facientes post germinationem pla11las bisporangiales/ tetra5porangiales. Tetrasporae 36-42 jJm, non germ inantes. Plantae in Mexico Pacifico ex Baja California Sur ad Jaliseo inventae. Plants know only as sporophytes (gametophytes lacking), 10-30 mm long and 0.5-1.5 mm wide, flat, bifurcate, slightly constricted at nodes; nodes usually lacking rhizoids except near base where attached; blades with 10-20 wing cells on each side of the axial cell; 6-8 bisporangia/tetrasporangia in each axial segment on each side of the axial cell; forming distinct sori; mature bisporangia ellipsoidal 30-44 X 40-62 fim and tetrasporangia 40-49 X 45-68 fim; sporangial formation and development as described for Caloglossa lepriellrii except that bisporangia and tetrasporangia are formed; bispores uninucleate 50-64 fim diam. and germinate to form further generations of bisporangiaVtetrasporangial plants; tetraspores 36-42 fim diam., fail to germinate; geographic distribution ranging from Baja Calfomia Sur Morphology Mature plants in culture (Fig. I) are characteristically pseudodichotomous with rhizoids generally not arising from the nodes except near the base. At some nodes one or more secondary endogenous blades develop that remain repressed in growth. Branching of the main axis is exogenous. Secondary branches are out of the plane of the thallus and arise endogenously from the first axial cell above the node. Adventitious blades occasionally arise from the wing cells at the blade margin as a result of physical damage. At the nodes the first lateral adaxial pericentral is absent. These characters are all similar to those of Caloglossa leipriellrii as presently defined by King and Puttock (1994). Although blade formation in C. leprieurii was well described by Papenfuss (1961) secondary pit connection formation and nuclear fre- quency and secondary pit formation in the pericentral and wing cells have not been considered. In C. apomeiolica the axial cells remain uninucleate throughout the entire blade. However, the four pericentral cells of each axial cell quickly become multinucleate through nuclear transfer by conjunctor cells formed when the three secondary pit connections occur vertically between adjacent pericentral cells (Fig. 13). In addition it appears that one or more nuclei in each pericentral cell divide without conjunctor cell fonnation, and this results in a total of 8-10 nuclei per cell. The lateral pericentral cells each produce two wing cells, each of which contains up to 6 nuclei that are also derived through secondary pit connection development witb wing cells of adjacent rows. The additional wing cells that extend to the blade margin also become multinucleate in the same manner. The number of nuclei per cell decreases corresponding with the decrease in secondary pit connection formation towards the margin. Consequently, the marginal cells that lack secondary pit connections are uninucleate. Figure 16 is a diagrammatic view of these events in a ma- ture blade segment below the 40th axial cell from the apex. to Jalisco in Pacific Mexico. Holotype: on Rhizophora mangle prop roots, San Carlos, Bahia Magdalena, Baja California Sur., Mexico (Wesl 3025, 6. i. 1990, UC 1199678). ReproduClion Additional colleclions: See Table I. any other Caloglossa species evident. In all localities sporophytes were seen with only bisporangial/tetrasporangial sori. The sporangiaI sori of field-collected plants were generally larger and more numerous than those in laboratory-cultured plants, presumably because the blades of field specimens were longer and wider than those in laboratory-cultured plants. In field-collected plants incompletely divided tetrasporangia were seen, but discharged tetrads of spores were never observed. Dislribwion: States of Baja California Sur, Sinoloa, lalisco and Nayarit in Pacific Mexico. See Table I and Figure 17. Comments: In Karsten el al. (1992 a), Karsten and West (1993) West el af. (1993) this species was referred to as Caloglossa apomeiotica nom. provo In West et at. (1992 b) it was designated as Caloglossa sp. I. Botanica Marina I Vol. 37 / 1994 I Fasc. 4 Gametophytes were never present in any of the localities where Caloglossa apomeiotica was collected nor were 384 West et of.: Ca/og/ossa apomeiOlica sp. nov. 5mm 11 Figs I ~ 15. LaboralOI)'-culturt'd plants of' Ca/oglossa apomeiolica sp. nov. Fig. I. Habit of 2-mollth-old plant (3033) bearing sporangia! sari. Fig. 2. Bisporangial sorus of 3131. Note discharge of older sporangia nearest main axis. Fig. 3. Bisporangia wilh uninucleate bispores (3033) stained with Wittman's iron-haematoxylin. Fig. 4. Abortive tctrasporangiulll (3245) with two spores lacking a nucleus. Wittman's iron-haematoxylin. Same scale as shown in Figure 3. Fig. 5. Plnnl (3245) wilh bisporangia and tctrasporangia. Two tetrads (*) and one diad of bisporcs (arrowheads) released. Same scale as shown in Figure 2. Fig. 6. Twcmy-four-h-old four-celled bispore gennling (3131) with rhizoidal pole formed. Fig. 7. Twenty-four-h-old eight-celled bispore germ ling (3131) with two-celled rhizoid. Same scale as shown in Figure 6. Fig. 8. Seventy-two-h-old unattached bispore germling with elongating rhizoid and pericentral ce,lls formed by middle three axial cells. Same scale as shown in Figure 2. Fig. 9. Ninety-six-h-old bispore germling (3131) with basal disc developing. Cf Figure 8, Fig. 10. One-month-old bispore germling (3033) with first node (an·ow). Secondary rhizoids and blades developing at base of plam (arrowhead). Fig. II. Secondary blade with sporangial sorus, developing from elongate basal rhiozoid (3025). Same scale as shown in Figure 9. Fig. 12. Spermatangial sorus (arrowhead) developing below bisporangial sorllS (3025). Same scale as shown in Figure 9. Fig. 13. Pericentral cells of axial cell no. 40 from apex with 3 pit connections at each pole and 6-8 nuclei per cell (3033). Witlman's ironhaematoxylin. Same scale as shown in Figure 3. Fig. 14. Uninucleate bisporangia (3033). DAPI-slaining. Same scale as shown in Figure 3. Fig. 15. Uninucleate and binucleate bisporangia (3033). DAPI-slaining. Same scale as shown in Figure 3. In the laboratory all isolates formed bisporangia and tetrasporangia and these were often intermixed in the same sari. Uninucleate sporangial parent cells (Fig. 14) were clearly evident in developing sori. Mature sori lISlIally contained bisporangia rather than tetrasporangia (Fig. 2). These were binucleate (Figs 3 and 15). ComBotanica Marina / Vol. 37 / 1994/ Fasc. 4 West el 01.: Calog/ossa apomeiolica sp. nov. • • ••• • ••• ~/0000 /-06M -o6&G •• Fig. 16. Diagrammatic scheme of nuclear and pil connection numbers in axial cells, peri central cells and wing cells from center to margin. pletely divided retrasporangia with 4 nuclei were seen in most stained preparations. There was also a high incidence of incompletely divided tetrasporangia with only two nuclei (Fig. 4). Many tetrasporangia released tetraspores (Fig. 5), but these were never observed to germinate. By contrast, bispores released from the bisporangia of the same plants (Fig. 5) showed nearly 100% gennination. Even though the sporangia differentiated in an acropetal sequence along the blade axis and from axis center to margin in each axial segment the sporangia did not necessar- 385 ily divide in this same sequence. Completely divided sporangia frequently occurred among undivided sporangia. Senescent, yellowing, undivided and clearly abonive sporangia were also common in most sori. Sporangia derived from pericentral cells were often larger than those derived from wing cells. Occasionally bisporangia were also formed by the pericentral cells on the dorsal or ventral side of the blade in most isolates. in situ germination was not seen in field collections but occurred sometimes on plants grown in stationary culture. Spore germination characteristic of the Ceramiales is bipolar with the initial formation of the primary rhizoid followed by the secondary fonnation of the erect blade axis (Figs 6 and 7). Within 3 days the blade sector initiates pericentral and wing cell fomlation (Fig. 8) and if the sporeling is attached to the substratum a digitate disc fOl111s witbin 4 days (Fig. 9). Witbin one month, by which time the plant is about 0.4 mm long, the first node develops (Fig. 10). As rhizoids on older parts elongate intercalary cells develop short lateral cells that f0l111 new blades which often become reproductive before the first node arises (Fig. II). Mixed-phase reproduction occurred only once (September 1992) in a single blade of isolate 3025 bearing bisporangial sori in the upper sector and spermatangial sari in the lower sector (Fig. 12). No procal])s or cystocarps were seen on any field-collected or laboralOry-culrured sporophytes. The only chromosome count (n = 30, 2n = 60) for C. leprieurii was reported by Rao (1967). Microspectro- Barra de Navidad Iw 114' Iw 110' Fig. 17. Map of Pacific Mexico indicating locations where C. apomeiofica was collected. Botanica Marina / Vol. 37/1994/ Fasc. 4 386 fluorometry has been used to elucidate life histories within the red algae (Goff and Coleman 1987, 1990, Gonzalez and Goff 1989) and is especially useful in determining the sites of meiosis and karyogamy. The fluorochrome DAPI (4',6-diamidino-2-phenylindole) binds quantitatively to DNA. In an attempt to determine the ploidy levels of several Caloglossa apomeiolica isolates for comparison with C. leprieurii isolates. haematoxylin staining for chromosome counts and DAPI staining for relative DNA levels were used in vegetative and reproductive cells; but neither method provided reliable results. Because of the great variation in DNA levels of nuclei throughout the plant it was not possible to obtain reliable comparative values for Calog/ossa apomeiotica and C. lepriel/rii using DAPI methodology. Consequently, further investigations with these methods are necessary before we can resolve the relative ploidy levels in these two species. The distinct difference in reproductive biology of Caloglossa populations in Pacific Mexico from typical Caloglossa lepriel/rii with its sexual Polysiphonia-type life history (see Yarish and Edwards 1982) suggested that they are different genetically isolated taxa. Thus far, bisporangia are not recorded for Caloglossa or for any other genus of the Delesseriaceae although both apomeiotic and meiotic bisporangia are well documented in most orders of the Rhodophyta (Guiry 1990). In the Ceramiales there are three recently described taxa based primarily on their bisporic reproduction: Aglaothamnion diaphanl/m L'Hardy-Halos el Maggs (1991), Bosllychia bispora West ef Zuccarello (West el al. 1992 a) and Ceramil/m bisporl/m Ballantine (1990). Preliminary observations by West el al. (1993) also indicate the occurrence of monosporangia in one Ca/og/ossa isolate from South Carolina and bisporangial and sexual plants in one isolate from Connecticut. The only other Caloglossa species for which a Polysiphonia-type life history is known based on laboratory culture investigations is C. ogasawaraensis Okamura (Tanaka and Kamiya 1933, West 1991, Pedroche el al. unpublished observations). Habital and Ecophysiology In all localities where Ca/og/ossa apomeiotica was recorded the saliniry measured at the time of collection was 20~ 38%0. Caloglossa apomeiotiea was not present in the upper estuaries of mainland Pacific Mexico which are subjected to a much greater freshwater influx and lower salinities (0-150/00). This contrasts with C. leprieurii and other species which often occur in freshwater streams. Nonetheless C. /eprieurii and C. apomeiolica investigated in laboratory culture appear well adapted to a wide salinity, showing similar increases in D-mannitol levels as the salinity increases to 70%0 and showing opti- West et al.: Ca/og/ossa apomeiotica sp. nov. mum growth rates at 15%0 (Karsten and West 1993). The water temperature range in the collection sites of mainland Mexico was 27-30°C and on the Pacific Ocean side of Baja California Sur temperatures were 17-22°C. Caloglossa species are osmotically adapted to the rapid salinity changes in estuarine habitats, partially through the production of D-mannitol (Kasten el al. 1992 a). West ef al. (1992 b, Pedroche ef al. unpublished observations) also verified this pattern in field-collected populations of Caloglossa from mainland Pacific Mexico. The other common mangrove-dwelling red algae Bosoychia and Sliclosiphonia are similarly adapled through D-sorbitol and D-dulcitol accumulation (Karsten el al. 1992 b). It is important to point out that polyol synthesis is a slow process relative to rapid salinity changes. In laboratory culture, stable polyol levels are reached 48-72 h after the plants are placed in a controlled salinity series. It appears that Caloglossa adapts to rapid salinity changes first with ion water influxes and expansion or contraction of the cell wall (Mostaert and King 1994). Analyses of D-mannitol were made for Caloglossa apomeiotica specimens from field and culture (Table II). In field-collected samples D-mannitol ranged from 245 to 468 nmol kg-I dw (mean 360 mmol kg-I kw) in salinities of 35 - 360/00. Field specimens were usually obtained while the algae were exposed to air, but it was not possible to determine the duration of exposure. The laboratory cultured specimens were maintained continually immersed in 30%0 medium, and D-mannitol levels were somewhat lower (327-331 mmol kg-I dw). These data are similar to those obtained for C. apomeiotiea in salinity shock experiments in 5, 15, 31, 50 and 700/00 with D-mannitol values extending from a low of 180 mmol kg-I dw at 5%0 to a high of 750 mmol kg-I dw (Karsten el al. 1992 b). Even though C. apomeiolica tolerates shOtt-term salinity extremes in the laboratory it appears that the natural salinity range tolerance is aboul 25350/00. Surprisingly, in short-term laboratory growth experiments the optimum salinity range was 5-30%0 with a specific growth rate of 8- 12% d- I (Karsten and West 1993). Natural Caloglossa lepriel/rii populations in tropical estuarine rivers appear to have a much greater salinity tolerance than C. apomeiolica (e. g. Mosich 1993). In Baja California Sur (I. Espiritu, Bahia Balandra, Bahia Magdalena) Caloglossa apomeiofica was present only on the shaded backside of Rhizophora mangle prop roots about 5-10 em lower than the maximum biomass level of Bos/lychia radicans. None occurred on LagufIell/aria or Avicennia pneumatophores. By contrast in two mainland Mexico sites near the estuary entrances Botanica Marina / Vol. 37 11994/ Fasc. 4 West et 01.: Caloglossa apomeiotica sp. nov. 387 Table II. D-Mannitol content of field and culture specimens of C. opomeiotica. • Collection locality and date Culture No. D-Mannitol (mmol kg- l dw) *San Carlos, Bahia Magdalena, Baja California Sur (B. C. S.), Mexico 6 January 1990, 35%0 3025 327.2 (n ~ I) *Bahia Balandra, B. C. S., Mexico 6 January 1990, 34%0 3033 331.0 (n ~ I) tTeacapim, Laguna Teacapan, Sinaloa, Mexico 20 March 1992,350/00 field 395.8 :t 18.7 (n ~ 4) tBarra de Navidad, Jalisco, Mexico 19 March 1992, sun, 350/00 field 360.1 :t 137.4 (n ~ 4) tBarra de Navidad, Jalisco, Mexico 19 March 1992, shade, 35%0 field 373 (n ~ 4) Bahia Balandra, B. C. S., Mexico 17 May 1992,35%0 field 468.3 :t 26.5 (n ~ 4) Estero Bahia Fals, B. C. S., Mexico 18 May 1992, 35%0 field 244.8 :::t 30.7 (n ~ 4) Isla Espiritu Santo, 8. C. S., Mexico 19 May 1992, 36%0 field 350.4 :t 55.2 (n ~ 4) San Carlos, Bahia Magdalena, B. C. S., Mexico 21 May 1992, 35%0 field 321.6 :t 18.1 :t 105.7 (n~4) * data from Karsten et oJ. 1992 a t data from West ef oJ. 1992 b (Barra de Navidad and San Bias) C. apamelOtlCa was abundant on lightly shaded Avicennia trunks and pneumatophores but was absent in heavily shaded Rhizophora thickets. At Teacapan only one population of C. apomeiolica was observed, and that occurred on a lightly shaded Rhizophora prop root. Caloglossa apomeiotica displayed a light-saturated net photosynthesis (Pm,,) of 64 ~mol Oz mg- I Chi a h- I at 200 ~mol photons m- z S-I and a specific growth rate of 15% d- I at 40 ~mol m- z S-I whereas C. leprieurii exhibited a light-saturated net photosynthesis of 128 ~mol Oz mg- I Chi a h- I at 200 ~ml photons m- z S-I and a specific growth rate of 10% d- I at 40 ~mol m- z S-I (Karsten and West 1993). These irradiance levels for Pmax are somewhat greater than usually measured in the natural habitats of shaded mangrove systems. In 1993 we made extensive field collections south of Jalisco in the mangroves of Colima, Guerrero, Oaxaca and Chiapas, Mexico and the Pacific coast of Guatemala. No Caloglossa specimens were observed in Colima, Guerrero or Oaxaca. The northernmost reproductive gametophytes and tetrasporophytes of Cologlossa leprieurii were observed in Estero las Garzas, Ghiapas, nearly 1200 km south of Barra de Navidad, where the southernmost population of C. apomeiotica was observed. Gametophytic and sporophytic plants of C. leprieurii were also seen in all coastal sites throughout Chiapas and Botanica Marina / Vol. 37 /1994 / Fasc. 4 Guatemala. Culture isolates have been established from most of these collections (Table III). General considerations Apomixis is widespread in higher plants, and as Asker and lerting (1992) point out it is difficult to detect traces of sexuality in highly apomictic taxa. However, even those species with predominantly asexual reproduction such as the cactus Lophocereus schoUii Britton et Rose (Parker and Hamrick 1992) maintain the same or greater level of genetic diversity as sexually reproducing species. Among several fern families there are taxa in which the gametophyte is uncoupled from the sporophyte, reproducing by asexual gemmae and having a greater tolerance of lower temperatures than the sporophytes, thus extending the geographic range. Farrar (1990) suggested that the gametophytes of the fern Vittaria spp. merit recognition as distinct species. Farrar and Mickel (1991) described a new species, Vittaria applachiana Farrar el Mickel, based on an asexually reproducing gametophyte lacking a sporophyte as an alternate generation. Farrar (1992) also described a new species, 7i'ichomanes il1tricatum Farrar, that is known only in the gametophyte phase. This pattern of genetic uncoupling and loss of sexual reproduction in ferns may have parallels with asexually reproducing red algal species that are present as either asexual gametophytes or sporophytes. Mogie (1992) suggested that in plants apomixis may result 388 West el al.: Caloglossa apomeiOlicll sp. nov. Table III. Collection locality. date, reproduction in field and culture numbers of Caloglossa spp. collected in Guatemala/Mexico. March, 1993. Species. locality and date Culture C ogasawarael1sis Colonia IXlapa, Guatemala 21 March 1993 3374 No. Reproduction in field tetraspophyte tetrasporophyte C. lepriellrii Likin, Guatemala 22 March 1993 C. leprie/lrii Boca Rio Naranjo. Guatemala 23 March 1993 3376 tctrasporophyte, male, female C. lepriellrii Rio San Juan Estero, Chiapas. Mexico 25 March 1993 3377 3378 tctTasporophyte. male. female C. Jeprieurii Rio Huehueatlan, Chiapas Mexico 24 March 1993 3379 female vegetative C. leprieurii Boen Rio San Juan, Estero las Garzas, Chiapas, Mexico 25 March 1993 C. sripilara Post Likin, Guatemala 22 March 1993 3375 vegetative, letrasporophyte from either polyploidy or hybridization, but presently we have no evidence to indicate that these were factors leading to asexual reproduction in Caloglossa apomeiotica. Many of the same arguments made for recognition of Bosllychia bispora (West el at. 1992 a) can be made for the taxonomic recognition of Caloglossa apomeiot- Iished observations), Costa Rica (Post 1967), Panama (Earle 1972), Colombia (Schneiter and Bula-Meyer 1982) and Peril (West 1991). For EI Salvador Dawson (1961) showed one figure of a possible sporophyte but did not mention this in the text of the paper. The algal ica. are poorly explored and no records of Caloglosso are It is very difficult to detern,ine the geological age of the known to us. mangrove populations and their associated algal flora in Pacific Mexico. In their treatment of the island biogeography and geology of the Sea of Conez (Gulf of California), Case and Cody (1983) provide no paleontological evidence concerning the mangroves. \Voodrofe and Grindrod (1991) generally conclude that .... records suggest significant changes in mangrove distribution in the Pacilic in the last few thousand years', a conclusion based on palynological evidence of the mangrove distribution in insular and mainland coastal areas throughout the Pacific. It may be assumed that mangrove-associared algae were introduced to the Pacific American coastline with RhizopllOra mangle and other genera after the Pliocene (for a discussion of this matter see Ellison 1991). A period of I -2 million years would be sufficient to isolate the genetically different asexual species, Caloglossa apomeiolica, from the sexually reproducing species C. lepriellrii to the south. Along the Pacific coast of Latin America the nearest populations of sexually reproducing C. leprieurii are those in Chiapas, Mexico, Guatemala (Bird and Mchltosh 1979, Pedroche ef al. unpub- floras of the Pacific coasts of Honduras and Nicaragua Dispersal of asexual species of algae and other plants is achieved through spore recruitment and vegetative propagules. Even without nonnal sexual reproduction Caloglossa apomeiOlica may have been able to spread quickly by vegetative fragmentation and bisporic reproduction linked to the simultaneous dispersal of the mangrove host species. TIle brown alga Sargassul1l I1llllicum (Yendo) Fensholt provides an excellent example of rapid dispersal. In just 50 years it has spread principally through vegetative propagules from British Columbia, Canada to La Paz, Mexico - a distance of more than 4000 km (Critchley el al. 1990, Riosmena, personal communication). Caloglossa apomeiofica occupies a similar habitat and has similar morphological and physiological characters to C. leprieurii: yet there is clearly a need to assess the genetic diversity of C. apomeiofica and to compare this with the genetic diversity of sexually reproducing populations of C. leprieurii that are nearest neighbors. Botanica Marina / Vol. 37/ 1994/ Fasc. 4 • 389 West et 01.: Caloglossa apomeiotica sp. nov. Acknowledgements , Fernando Garza Sanchez assisted in collections made at Isla Espiritu Santo. Rafael Riosmena Rodriguez provided infonnation about the distribution of Sargasswn muaCUI1l in Baja California Sur. Prof. Yoshiaki Hara and the Mombusho Foundation of Japan provided financial support for the work at Isla Espiritu Santo and The Hardmann Foundation for Conservation provided funds for the field research in mainland Mexico during 1992 and 1993. In 1993 the Universidad Autonoma Metropolitana provided a vehicle for work in Guatemala and Mexico. 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A field and cultural investigation of the horizontal and seasonal distribution of estuarine red algae of New Jersey. Phyc%gia 1/: 112-124. Botanica Marina / Vol. 37/ 1994/ Fasc. 4 View publication stats