Symbiotic state influences life-history strategy of a clonal cnidarian.

Along the North American Pacific coast, the common intertidal sea anemone Anthopleura elegantissima engages in facultative, flexible symbioses with Symbiodinium muscatinei (a dinoflagellate) and Elliptochloris marina (a chlorophyte). Determining how symbiotic state affects host fitness is essential to understanding the ecological significance of engaging in such flexible relationships with diverse symbionts. Fitness consequences of hosting S. muscatinei, E. marina or negligible numbers of either symbiont (aposymbiosis) were investigated by measuring growth, cloning by fission and gonad development after 8.5-11 months of sustained exposure to high, moderate or low irradiance under seasonal environmental conditions. Both symbiotic state and irradiance affected host fitness, leading to divergent life-history strategies. Moderate and high irradiances led to a greater level of gonad development in individuals hosting E. marina, while high irradiance and high summer temperature promoted cloning in individuals hosting S. muscatinei and reduced fitness of aposymbiotic anemones. Associating with S. muscatinei may contribute to the success of A. elegantissima as a spatial competitor on the high shore: (i) by offsetting the costs of living under high temperature and irradiance conditions, and (ii) by promoting a high fission rate and clonal expansion. Our results suggest that basic life-history characteristics of a clonal cnidarian can be affected by the identity of the endosymbionts it hosts.

ELLIPTOCHLORIS MARINA SP. NOV. (TREBOUXIOPHYCEAE, CHLOROPHYTA), SYMBIOTIC GREEN ALGA OF THE TEMPERATE PACIFIC SEA ANEMONES ANTHOPLEURA XANTHOGRAMMICA AND A. ELEGANTISSIMA (ANTHOZOA, CNIDARIA)(1).

Symbiotic green algae from two species of intertidal Pacific sea anemones, Anthopleura elegantissima and Anthopleura xanthogrammica, were collected from the northeastern Pacific coast of North America across the known range of the symbiont. Freshly isolated Anthopleura symbionts were used for both morphological and molecular analyses because Anthopleura symbiont cultures were not available. Light and transmission electron microscopy supported previous morphological studies, showing the symbionts consist of spherical unicells from 5 to 10 µm in diameter, with numerous vesicles, and a single bilobed chloroplast. Pyrenoids were not seen in LM, but a thylakoid-free area was observed in TEM, consistent with previous findings. Many algal cells extracted from fresh anemone tissue were observed in the process of division, producing two autospores within a maternal cell wall. The morphology of the green symbionts matches that of Elliptochloris Tscherm.-Woess. Molecular phylogenetic analyses of the nuclear SSU rDNA and the plastid encoded gene for the large subunit of RUBISCO (rbcL) support the monophyly of these green algal symbionts, regardless of host species and geographic origin. Phylogenetically, sequences of the Anthopleura symbionts are nested within the genus Elliptochloris and are distinct from sequences of all other Elliptochloris spp. examined. Given the ecological and phylogenetic distinctions among the green algal symbionts in Anthopleura spp. and the named species of Elliptochloris, we designate the green algal symbionts as a new species, Elliptochloris marina (Trebouxiophyceae, Chlorophyta).

LOW THERMAL LIMIT OF GROWTH RATE OF SYMBIODINIUM CALIFORNIUM (DINOPHYTA) IN CULTURE MAY RESTRICT THE SYMBIONT TO SOUTHERN POPULATIONS OF ITS HOST ANEMONES (ANTHOPLEURA SPP.; ANTHOZOA, CNIDARIA)(1).

Symbiodinium californium (#383, Banaszak et al. 1993) is one of two known dinoflagellate symbionts of the intertidal sea anemones Anthopleura elegantissima, A. xanthogrammica, and A. sola and occurs only in hosts at southern latitudes of the North Pacific. To investigate if temperature restricts the latitudinal distribution of S. californium, growth and photosynthesis at a range of temperatures (5°C-30°C) were determined for cultured symbionts. Mean specific growth rates were the highest between 15°C and 28°C (µ 0.21-0.26 · d(-1) ) and extremely low at 5, 10, and 30°C (0.02-0.03 · d(-1) ). Average doubling times ranged from 2.7 d (20°C) to 33 d (5, 10, and 30°C). Cells cultured at 10°C had the greatest cell volume (821 µm(3) ) and the highest percentage of motile cells (64.5%). Growth and photosynthesis were uncoupled; light-saturated maximum photosynthesis (Pmax ) increased from 2.9 pg C · cell(-1 ) · h(-1) at 20°C to 13.2 pg C · cell(-1 ) · h(-1) at 30°C, a 4.5-fold increase. Less than 11% of daily photosynthetically fixed carbon was utilized for growth at 5, 10, and 30°C, indicating the potential for high carbon translocation at these temperatures. Low temperature effects on growth rate, and not on photosynthesis and cell morphology, may restrict the distribution of S. californium to southern populations of its host anemones.

Nutritional role of two algal symbionts in the temperate sea anemone Anthopleura elegantissima brandt.

The intertidal sea anemone Anthopleura elegantissima in the Pacific Northwest may host a single type of algal symbiont or two different algal symbionts simultaneously: zooxanthellae (Symbiodinium muscatinei) and zoochlorellae (green algae; Trebouxiophyceae, Chlorophyta). A seasonal comparison of zooxanthellate and zoochlorellate anemones showed stable symbiont population densities in summer and winter, with densities of zoochlorellae about 4 times those of zooxanthellae. Photosynthesis-irradiance curves of freshly isolated symbionts show that the productivity (P(max) cell) of freshly isolated zooxanthellae was about 2.5 times that of zoochlorellae during July; comparable rates were obtained in other months. Models of algal carbon flux show that zoochlorellae may supply the host with more photosynthetic carbon per unit anemone biomass than zooxanthellae supply. Zooxanthellate anemone tissue was 2 per thousand ((13)C) and 5 per thousand ((15)N) enriched and zoochlorellate anemone tissue was 6 per thousand ((13)C) and 8 per thousand ((15)N) enriched over their respective symbionts, suggesting that zoochlorellate anemones receive less nutrition from their symbionts than do zooxanthellate individuals. The disparity between predicted contributions from the algal carbon budgets and the stable isotopic composition suggests that short-term measures of algal contributions may not reflect actual nutritional inputs to the host. Isotopic data support the hypothesis of substantial reliance on external food sources. This additional nutrition may allow both algae to persist in this temperate intertidal anemone in spite of differences in seasonal photosynthetic carbon contributions.

Phylogenetic placement of "zoochlorellae" (Chlorophyta), algal symbiont of the temperate sea anemone Anthopleura elegantissima.

At northern latitudes the sea anemones Anthopleura elegantissima and its congener A. xanthogrammica contain unidentified green chlorophytes (zoochlorellae) in addition to dinophytes belonging to the genus Symbiodinium. This dual algal symbiosis, involving members of distinct algal phyla in one host, has been extensively studied from the perspective of the ecological and energetic consequences of hosting one symbiotic type over the other. However, the identity of the green algal symbiont has remained elusive. We determined the phylogenetic position of the marine zoochlorellae inhabiting A. elegantissima by comparing sequence data from two cellular compartments, the nuclear 18S ribosomal RNA gene region and the plastid-encoded rbcL gene. The results support the inclusion of these zoochlorellae in a clade of green algae that form symbioses with animal (Anthopleura elegantissima), fungal (the lichen genus Nephroma), and seed plant (Ginkgo) partners. This clade is distinct from the Chlorella symbionts of Hydra. The phylogenetic diversity of algal hosts observed in this clade indicates a predisposition for this group of algae to participate in symbioses. An integrative approach to the study of these algae, both within the host and in culture, should yield important clues about how algae become symbionts in other organisms.

Spatial distribution of light and nutrients in some coral reefs of Costa Rica during January 1997

The proximity of coral reefs to areas of present and future coastal development in Costa Rica highlights the need for assessing environmental conditions important to maintaining healthy corals. In January 1997 a survey of light penetration, inorganic nutrient concentrations, temperature, and salinity was conducted in the patch reefs of Bahía Culebra (Pacific Ocean) and on the Caribbean coast in the fringing reef at Parque Nacional Cahuita and near Limón. Temperature was 28 degrees C at all sites, and salinity ranged from 33 to 36 psu. Light attenuation coefficients ranged from 0.12 to 0.29 m(-1) in reef areas. Seawater nutrient concentrations were within the range of concentrations reported for tropical reef waters; combined NO3- and NO2- and PO4(3-) were each below 1 microM. NH4+ ranged from 0.2 to 7 microM, representing a significant source of nitrogen. The ratio of total dissolved inorganic nitrogen to phosphate averaged 27 for all reef waters. The high nitrate (3.6 microM) and light attenuation (0.95 m(-1)) values from the surface waters of the La Estrella plume (Caribbean coast) show that this river represents a significant source of nitrogen and light attenuation for the neighboring reefs at Cahuita. This survey provides a useful baseline for future studies, which should monitor these important coastal coral reef areas during both wet and dry seasons.