A comparative analysis of the photobiology of zooxanthellae and zoochlorellae symbiotic with the temperate clonal anemone Anthopleura elegantissima (Brandt)

The temperate anemone Anthopleura elegantissima hosts two phylogenetically different symbiotic microalgae, a dinoflagellate Symbiodinium (zooxanthellae, ZX) and a chlorophyte (zoochlorellae, ZC), throughout certain regions of its latitudinal range. Because of the broad intertidal and geographic range of this anemone, we examined the role of irradiance to ascertain which specific symbiotic parameters are affected and whether light intensity governs the observed distributions of natural populations of ZX and ZC. Irradiance appears to be a key factor in regulating both the photophysiology and metabolism of this alga-cnidarian association. Regardless of light intensity, algal densities remained stable for anemones harboring ZX or ZC, whereas the mitotic indices of ZX and ZC both varied directly with light intensity. The chlorophyll content of ZX remained fairly constant regardless of irradiance; in contrast, ZC chlorophyll content was inversely proportional to light intensity. Regardless of irradiance, the carotenoid content of both symbionts was constant; however, ZX carotenoid levels were higher than those of ZC. Net photosynthesis was directly related to light intensity for both algal symbionts and ZX photosynthetic rates were consistently higher than those of ZC. Similarly, the potential carbon contribution of ZX and ZC to animal respiration (CZAR) displayed a direct relationship with light intensity, peaking at 800 µmol·m^-2·s^-1, then subsequently declined. Lower ZX growth rates, coupled with higher photosynthetic rates and higher CZAR estimates, compared to ZC, suggest that the ZX should be the dominant symbiont as light intensity increases; this may explain the high densities of anemones in the field containing ZX where the levels of irradiance are naturally high. These results support the interpretation that irradiance is a significant environmental parameter that dictates the microhabitat and latitudinal distribution of the two symbiotic algal taxa. This is the second in a series of papers examining the physical parameters that influence the distribution of ZX- and ZC-bearing A. elegantissima.     

A comparative analysis of the photobiology of zooxanthellae and zoochlorellae symbiotic with the temperate clonal anemone <Emphasis Type="Italic">Anthopleura elegantissima</Emphasis> (Brandt). III. Seasonal effects of natural light and temperature on photosynthesis and respiration

The sea anemone Anthopleura elegantissima hosts two phylogenetically different symbiotic microalgae, a dinoflagellate Symbiodinium (zooxanthellae, ZX) and a chlorophyte (zoochlorellae, ZC). The photosynthetic productivity (P), respiration (R), and contribution of algal carbon translocated to the host (CZAR) in response to a year’s seasonal ambient changes of natural light and temperature are documented for both ZX- and ZC-bearing anemones. Light and temperature both affect photosynthesis, respiration, and CZAR, as well as various algal parameters; while there are evident seasonal differences, for the most part the relative effects on P, R, and CZAR by the two environmental variables cannot be determined. Net photosynthesis (P_n) of both ZX and ZC was significantly higher during spring and summer. During these seasons, the P_n of ZX was always greater than that of ZC. Regardless of algal symbiont, anemone respiration (R) was significantly higher during the spring and summer. The annual net carbon fixation rate of anemones with ZX and ZC was 325 and 276 mg C anemone⁻¹ year⁻¹, respectively, which translates to annual net community productivity rates of 92 and 60 g C m⁻¹ year⁻¹ for anemones with ZX or ZC, respectively. CZAR did not show a clear relationship with season; however the CZAR for ZX was always significantly greater than for ZC. Lower ZX growth rates, coupled with higher photosynthetic rates and higher CZAR estimates, compared to ZC, suggest that if A. elegantissima is simply carbon limited, ZX-bearing anemones should be the dominant symbiont in the field. However ZC-bearing anemones persist in low light and reduced temperature microhabitats, therefore more than the translocation of carbon from ZC must be involved. Given that global climate change will increase water temperatures, the potential for latitudinal range shifts of both ZC and ZX (S. californium and muscatinei) might be used as biological indicators of thermal shifts in the littoral zone of the Pacific Northwest.     

Acetate incorporation into the lipids of the anemone Anthopleura elegantissima and its associated zooxanthellae

Anthopleura elegantissima containing zooxanthellae, as well as isolated zooxanthellae, incubated with acetate-1-¹⁴C under both light and dark conditions readily incorporate radioactivity into their total lipid pools. In both cases, the specific activity was greatly increased in the light. Dark-incubated anemones and isolated zooxanthellae incorporate activity predominantly into polar lipid; the remainder being present principally in the triglyceride moiety. Light-incubated organisms, however, show a dramatic redistribution of isotope towards greatly increased triglyceride and was ester incorporation, with a concomitant drop in polar lipid. onder light conditions, 70 to 75% of the radioactivity found in the fatty acids of the total zooxanthellae lipid was present in hexadecanoic (16:0) and octadecenoic (18:1) fatty acids. These are also the two major fatty acids by mass. Octadecanoic acid (18:0) is less than 5% by mass. Isotope incorporation patterns suggest that octadecenoic acids arise by elongation of hexadecenoic acids and that this conversion is blocked in the dark. Isotope incorporation patterns for anemones suggest that fatty acids, primarily in the form of saturated or monoenoic fatty acids, are translocated from algal to animal cells. No activity was found in either octadecadienoic (18:2) or octadecatrienoic (18:3) acids. The significance of these data is discussed.     

Feeding behavior and acquisition of zooxanthellae by planula larvae of the sea anemone Anthopleura elegantissima

Symbiotic associations between cnidarians and photosynthetic dinoflagellates (i.e., zooxanthellae) are common in the marine environment. Many symbiotic cnidarians produce offspring that are initially nonsymbiotic. These new hosts must acquire symbiotic algae from environmental sources. We examined zooxanthella acquisition by laboratory-reared planula larvae of the temperate sea anemone Anthopleura elegantissima. Larvae ingested zooxanthellae while they were feeding. However, the signal that prompted larval feeding behavior did not originate from the symbiotic algae; the addition of algal cells to larval cultures never elicited a feeding response. In contrast, the addition of macerated animal tissue from several sources invariably generated a strong feeding response, which resulted in the larvae indiscriminately ingesting any particulate matter that was present, including zooxanthellae or other unicellular algae. Ingested zooxanthellae were incorporated into endodermal cells, where they remained undigested, while all other ingested material was digested or expelled within 24 h. Our results provide evidence that one source of zooxanthellae likely to serve as a route of infection in the natural environment is zooxanthella-laden mucus egested by anemones. This egested material fulfilled both of the criteria necessary for successful infection: it prompted larvae to begin feeding and provided an abundant supply of zooxanthellae that were ingested and taken up into endodermal cells of the new host.     

Genetic relationships within and between clonal and solitary forms of the sea anemone Anthopleura elegantissima revisited: evidence for the existence of two species

Along the temperate Pacific coast of North America, the actiniarian sea anemone Anthopleura elegantissima exhibits two discrete life-history phenotypes. Although both forms sexually produce planula larvae, the clonal morph can also asexually propagate by fission, whereas the solitary morph does not. Whether the two forms constitute one or two species has long been contested. Hand originally designated the two forms as conspecifics, whereas Francis – on the basis of differences in microhabitat, biogeographic range and phenotypic frequencies – argued that the two forms constituted a sibling-species pair. From the results of an electrophoretic survey in which they pooled allelic frequencies across several geographic locations, Smith and Potts subsequently argued that the two forms were not genetically differentiated, and therefore represented a single species. We re-examined the relationship between the forms electrophoretically, substantially extending the geographic range and doubling the sample sizes beyond those used by Smith and Potts, and not pooling allelic frequencies in our analyses. Our analysis of patterns of genetic variation at ten highly polymorphic allozyme loci shows that although no fixed genetic differences distinguish the two forms, there are significant differences in allele frequencies between clonal and solitary A.␣elegantissima at every site we sampled throughout their range of sympatry (over 1000 km); within each form, however, there is little detectable genetic differentiation among populations. We therefore conclude that the two forms represent recently reproductively isolated taxa, and propose that the clonal form retain the binomial A. elegantissima (Brandt, 1835), whereas the solitary form be described and named a new species, Anthopleura sp. Received: 28 August 1996 / Accepted: 25 September 1996     

Oxygen as a limiting factor in phototaxis and in intraclonal spacing of the sea anemone Anthopleura elegantissima

Phototaxis in Anthopleura elegantissima, a sea anemone symbiotic with zooxanthellae, was investigated with special reference to oxygen as a possible controlling factor. Under high oxygen concentrations in seawater, movement towards light was not observed for symbiotic anamones as it was under normal oxygen concentrations. Both aposymbiotic and symbiotic anemones demonstrated movement towards high oxygen concentrations in seawater. Oxygen is, therefore, implicated as a controlling factor in phototaxis. Under laboratory conditions, increased intraclonal spacing occurred with low oxygen concentrations in seawater. In the field, individuals in symbiotic clones were spaced significantly closer than in aposymbiotic clones. Since intraclonal spacing is controlled by oxygen in the laboratory, spacing may also be affected in the field by oxygen; symbiotic clones may be spaced closer because they have better oxygen availability than do aposymbiotic clones.     

The effects of symbiotic state on heterotrophic feeding in the temperate sea anemone <Emphasis Type="Italic">Anthopleura elegantissima</Emphasis>

The temperate sea anemone Anthopleura elegantissima is facultatively symbiotic with unicellular algae. Symbiotic A. elegantissima can supplement heterotrophic feeding with excess photosynthate from their algal partners, while asymbiotic individuals must rely solely on heterotrophy. A. elegantissima individuals were collected from Swirl Rocks, Washington (48°25′6″ N, 122°50′58″ W) in July 2010, and prey capture and feeding characteristics were measured to determine whether asymbiotic individuals are more efficient predators. Feeding abilities were then measured again after a 3-week exposure to full sunlight or shaded conditions. Freshly collected asymbiotic anemones had larger nematocysts, but symbiotic individuals showed greater nematocyte sensitivity. Sunlight enhanced digestion and reduced cnida density in all anemones regardless of symbiotic state. Results suggest that the phototropic potential of A. elegantissima, as influenced by symbiotic condition, has little effect on heterotrophic capacity. The anemones appear to maximize heterotrophic energy input independent of the presence or identity of their algal symbionts.     

Effect of feeding regime and irradiance on the photophysiology of the symbiotic sea anemone Aiptasia pulchella

To determine how the animal and algal components of the symbiotic sea anemone Aiptasia pulchella respond to changes in food availability and culture irradiance, sea anemones from a single clone were maintained at four irradiance levels (320, 185, 115, and 45 E m^-2 s^-1) and either starved or fed for 5 wk. Changes in protein biomass of sea anemones maintained under these conditions were not related to the productivity of zooxanthellae, since the protein biomass of fed A. pulchella decreased with increase in irradiance and there was no difference in protein biomass among starved sea anemones at the four irradiance levels. Except for the starved high-light sea anemones, the density of symbiotic zooxanthellae was independent of culture irradiance within both starved and fed. A. pulchella. Starved sea anemones contained over twice the density of zooxanthellae as fed sea anemones. Within both starved and fed individuals, chlorophyll per zooxanthella increased with decreasing culture irradiance while algal size remained constant (in fed sea anemones) at about 8.80 m diameter. Chlorophyll a: c ₂ ratios of zooxanthellae increased with decreasing culture irradiance in zooxanthellae from starved sea anemones but remained constant in zooxanthellae from fed sea anemones. As estimated from mitotic index data, the in situ growth rates of zooxanthellae averaged 0.007 d^-1 and did not vary with irradiance or feeding regime. Photosynthesis-irradiance (P-I) responses of fed A. pulchella indicated an increase in photosynthetic efficiency with decreasing culture irradiance. But there was no consistent pattern in photosynthetic capacity with culture irradiance. Respiration rates of fed sea anemones also did not vary in relation to culture irradiance. The parameter I _k , defined as the irradiance at which light-saturated rates of photosynthesis are first attained, was the only parameter from the P-I curves which increased linearly with increasing culture irradiance. The daily ratio of net photosynthesis to respiration for A. pulchella ranged from 1.6 to 2.8 for sea anemones maintained at the three higher irradiances, but was negative for those maintained at 45 E m^-2 s^-1. Since the final protein biomass was greatest for sea anemones maintained at the lowest irradiance, these results indicate that sea anemone growth cannot be directly related to productivity of zooxanthellae in this symbiotic association.     

Cellular and ultrastructural changes in the endoderm of the temperate sea anemone Anemonia viridis as a result of increased temperature

Exposure of the temperate sea anemone Anemonia viridis Forskål to increased seawater temperature (from 16 to 26°C) reduced the lysosomal latency of coelenterate tissues. Lysosomes in the mesenterial filaments of anemones were destabilised by increased temperature, with greater destabilisation in heat-shocked symbiotic anemones than in heat-shocked aposymbiotic anemones in the early stages of the experiment. Lysosomal enzyme activity in zooxanthellae from heat-shocked symbiotic anemones was associated with the algal membranes and the cytoplasm of degenerate algal cells. While the relationship between host coelenterate and symbiotic alga may confer many benefits under normal conditions, comparison of the responses of symbiotic and aposymbiotic anemones to heat shock suggests that there may be disadvantages for symbiotic anemones under stress.     

Autecology and clonal variability of the marine centric diatom Thalassiosira rotula (Bacillariophyceae) in response to light,temperature and salinity

The influence of 49 combinations of salinity (10–40 S, at 5 S intervals) and temperature (0°–30°C, at 5C° intervals) on the maximum daily division rate (K) and 18 combinations of light intensity (six levels) and temperature (5°, 15°, and 25°C) on photosynthesis, cell division, and chlorophyll a was examined using two clones of Thalassiosira rotula Meunier isolated from the upwelling area of Baja California (clone C8) and from Narragansett Bay, Rhode Islands (clone A8). Physiological differences appear to characterize these to clones with regard to their temperature tolerance (C8 5°–30°C, A8 0°–25°C), maximum growth rate (C8 K=2.9, A8 K=2.4), chlorophyll a content, and in the rates of growth and photosynthesis in response to light intensity and temperature. Optimum salinity for both clones (25–30 S) was generally independent of temperature, while chlorophyll a content decreased with temperature. T. rotula is a cosmopolitan paractic species; experimental studies indicate that it is eurythermal and moderately euryhaline. Comparison of five additional Narragansett Bay isolates of T. rotula reveal minimal spacial or temporal variability in genetically determined physiological characteristics within this local population.     

Agriotes proximus and A. lineatus (Coleoptera: Elateridae): a comparative study on the pheromone composition and cytochrome c oxidase subunit I gene sequence

The presence of geranyl octanoate, previously found in pheromone gland extracts of Agriotes lineatus females, was also demonstrated in gland extracts of A. proximus females. Similar to A. lineatus, geranyl butanoate was present only in trace amounts in A. proximus female gland extracts. In air entrainment samples of female A. lineatus and A. proximus beetles, the relative ratio of geranyl butanoate and geranyl octanoate was higher than that in gland extracts. In addition, comparison of a segment of the mitochondrial cytochrome c oxidase subunit I gene of feral specimens of A. lineatus and A. proximus showed >99% similarity. Both pheromone profile and nucleotide sequence analysis delineate close relationship between the investigated taxa and postulate taxonomic revision. Further studies on sympatric populations of A. lineatus and A. proximus are underway to investigate and better understand possible processes of species diversification.     

Effect of temperature on life cycles of nematodes associated with the mangrove (Rhizophora mangle) detrital system

The effect of temperature on the life history of various representatives of the meiofauna associated with decaying mangrove (Rhizophora mangle) leaves was investigated. Life cycles, recorded in days, for 6 species of marine nematodes cultured at a temperature of 24°C are as follows: Rhabditis marina 2 1/4; Diplolaimelloides sp., 7; Diplolaimella ocellata, 11 1/2; Enoplus paralittoralis, 22; Oncholaimus sp., 29; Haliplectus dorsalis, 34. In general, life cycles become shorter with increased temperatures; however, as temperatures approach the upper limits which support reproduction, life cycles become slightly lengthened. For most species, the ability to complete a life cycle was inconsistent within the temperature range of 33° to 35°C. Studies with two harpacticoid copepod and two foraminifera species tend to support the 33° to 35°C range as being the thermal stress zone.Contribution No. 1697 from the University of Miami, Rosentiel School of Marine and Atmospheric Science, 10 Rickenbacker Causeway, Miami, Florida 33149, USA.     

A comparative study of the blood of four Tilapia species (Pisces)

The blood of 4 species of Tilapia has been examined. Mean erythrocyte count and size, hemoglobin content, and hematocrit value varied markedly among the species examined. A tendency towards a reciprocal relationship between the number of erythrocytes and their size was observed. In general, their is a correlation between the erythrocyte counts, hemoglobin content, and hematocrit value among the species examined.     

Effect of temperature on growth and reproduction of the epigeic earthworm,Eudrilus Eugeniae (Kinberg)

Influence of temperature on growth and reproduction of Eudrilus eugeniae has been investigated by laboratory culturing at regulated 25.0 degrees C, 30.0 degrees C, 37.5 degrees C and 40.0 degrees C and in fluctuating (22.7-27.3 degrees C) room temperature of prevailing (winter) season over 16 weeks. All worms died during first and tenth week at 40.0 degrees C and 37.5 degrees C respectively. Weight (biomass) and growth of worms cultured at different temperature varied significantly (P<0.01). The mean growth (mg/g live weight/day) at 25.0 degrees C., 30.0 degrees C, 35.0 degrees C, 37.5 degrees C and in fluctuating temperatures was 1,074.04 +/- 6.07, 1,554.01 degrees 192.37, 148.1 +/- 15.28, 192.83 +/- 25.8 and 1450.4 +/- 162.1 respectively. Growth declined after maturity drastically with coccon production. At 25.0 degrees C though worms are sexually mature, they failed to produce cocoons within 16 weeks whereas, at 35.0 degrees C and 37.5 degrees C they did not sexually mature. Worms attained sexual maturity at a mean weight of about 1000 mg/worm. The mean per cent maturity was higher and earlier in fluctuating temperatures and at 30.0 degrees C than at 25.0 degrees C. Cocoon production was observed only at 30.0 degrees C and in fluctuating temperatures with a mean of 0.9 and 1.5 cocoons/ wom/week and the cumulative cocoon number of 10.8 and 14.7/worm over 16th week respectively. The fluctuating temperature of uncontrolled room environment and 30.0 degrees C were favorable for various life activities of the worms. Eudrilus eugeniae appears to have range of temperature optima more than 25.0 degrees C and less than 28 degrees C. The climatic conditions prevailing in whole of the peninsular India during winter season are favourable for employing this worm in intensive field scale vermiculturing practices.     

Effect of temperature and salinity on larval development of southern king crab (Lithodes antarcticus)

Larvae of Lithodes antarcticus Jacquinot were reared in October, 1981 from hatching to the glaucothoe stage at 16 temperature/salinity combinations (5.5°; 7.5°; 9.5° and 13.5°C; 26, 29, 32 and 35 S) to determine optimal environmental conditions for larval development. The highest survival percentage was obtained in the culture at 7.5°C and diminished according to temperature increase or decrease. High temperature cultures significantly shorten the larval life duration, but produce large mortalities. At 5.5°C mortality occurred almost exclusively during the moult to glaucothoe stage. Higher survival percentages were obtained as salinity was increased. In the lowest salinity culture (26 S) no zoea reached the post-larvae stage at culture temperatures. The best T/S combination was obtained at 7.5°C and 35 S, with a survival percentage of 29%. The shortest zoeal developments were obtained at 32 S in all culture temperatures. Salinity also affects larvae coloration: there is a pigment concentration on erythrophores, which causes a color decrease.     

Effect of temperature fluctuations and food supply on the growth and metabolism of juvenile sea scallops (Placopecten magellanicus)

On the eastern shore of Nova Scotia late summer atmospheric systems cause upwelling of shelf water; the associated temperature variations of 10 °C with a 6 to 8 d period are comparable in magnitude to the seasonal variation. A laboratory study was undertaken to assess the effects of these temperature fluctuations on sea scallop (Placopecten magellanicus) growth and metabolism. In a factorial design, scallops were subjected to constant (10 °C) or a variable (6 to 15 °C) 8 d temperature cycle, and either a low (seston in filtered seawater) or high (seston supplemented with cultured phytoplankton) food diet. During the 48 d experiment scallop mortality was low and growth positive in all treatments. Shell and total tissue growth rate did not differ between temperature treatments, but growth in the high food treatments was 40 to 50% higher than in the low food treatments. However, soft tissue (excluding adductor) growth did show a temperature treatment effect; growth rates were significantly higher in the fluctuating temperature treatment, due in part to greater gonad development. Weight-standardized rates of scallop oxygen consumption (V _sO₂ , μmol O₂ g⁻¹ h⁻¹) were 20 to 25% higher in high food than in low food treatments, consistent with the expected increase in respiration due to the higher growth rates. Scallop metabolism did not acclimate to the fluctuating temperature cycle; V _sO₂ and ammonium excretion (V _sNH⁺ ₄, μmol O₂ g⁻¹ h⁻¹) remained dependent on ambient temperature throughout the experiment. V _sNH⁺ ₄ Q₁₀ (2.77) was higher than V _sO₂ Q₁₀ (2.01) which was reflected in a decrease in the O:N ratio at 15 °C, indicating a shift toward increased protein catabolism and a stressed state. At 10 °C, V _sO₂ and V _sNH⁺ ₄ in the variable temperature treatments were 15 to 18% lower than in the constant temperature treatments, a difference that was not detected in growth measurements. Results demonstrate that the metabolism of Placopecten magellanicus, unlike some bivalve species, is tightly coupled to fluctuations in ambient temperature. Although an absence of compensatory acclimation had a minimal effect on growth in this study, if high temperatures were combined with low food conditions a reduction in scallop production could result. Received: 23 June 1998 / Accepted: 8 February 1999     

Experimental ecology of the temperate scleractinian coral Astrangia danae I. Partition of respiration,photosynthesis and calcification between host and symbionts

Calcification, photosynthesis and respiration of the scleractinian coral Astrangia danae were calculated from the changes in total alkalinity, pH, calculated total CO₂, and oxygen concentration produced by colonies incubated in glass jars. A correction for changes in ammonia, nitrate and nitrite was taken into account and the method evaluated. The fluxes of oxygen and CO₂ were highly correlated (r=0.99). The statistical error of alkalinity determinations was less than 10% of the changes observed in the slowest calcifying samples. Metabolism of polyparium alone was estimated by difference after removal of tissue and reincubation of bare corallum. Zooxanthellae concentration in the polyps was obtained from cell counts made on homogenates of polyp tissue. The calculated photosynthetic rate of the zooxanthellae in vivo was 25 mol O₂ (10⁸ cell)^-1 h^-1 at a light intensity of 120 Ein m^-2 s^-1. In corals having 0.5x10⁹ zooxanthellae/dm² of colony area up to 8% of the total photosynthesis was attributed to the corallum microcosm. Polyp respiration, photosynthesis, and CaCO₃ uptake rates were all much higher than rates previously reported from A. danae, apparently because in these experiments the organisms were better fed. This increased photosynthesis in turn enhanced calcification still further. The symbiosis therefore appears to provide a growth advantage even to fed corals, under the conditions of these experiments.     

A qualitative analysis of the stability of the root-microorganism soil system I. carbon-nitrogen status and nitrogen mineralization

This paper demonstrates that if soil microorganisms are devoted suppliers of nitrogen, plant roots can coexist with them in the soil, i.e. if nitrogen net-mineralization is positive, the root-decomposer system is asymptotically stable. On the other hand, if net mineralization is negative, the system evolves in time towards the annihilation of the roots.Available data from an old pine stand system show that net mineralization is sometimes positive and sometimes negative during the course of a year. It is shown here that this situation is compatible with a state of dynamic stability in the root-decomposer system. The restrictions imposed on the validity of this analysis are also deduced.     

Effect of temperature and food availability on reproductive responses of Streblospio benedicti (Polychaeta: Spionidae) with planktotrophic or lecithotrophic development

Streblospio benedicti (Webster) from Tar Landing North Carolina (NC), USA with either planktotrophic or lecithotrophic development were reared under two food levels and three temperature regimes (two mimicking seasonal cycles in NC and one at constant 20°C). During the eight-month experiment no females switched reproductive mode and no significant differences in survivorship or reproductive activity were observed between reproductive types. However, reproductive activity and fecundity-related parameters were subject to influence by food and temperature. Survivorship, body size, and larval production was greater in winter-spring than summer-fall regimes. Higher food levels produced increased survivorship, reproductive activity and egg production in adults with lecithotrophic development but no change in those with planktotrophic development. Body size, egg size, egg number, numbers of larvae per brood pouch, and brood size were strongly correlated in female S. benedicti and most correlation coefficients were similar (or identical) in individuals having planktotrophic and lecithotrophic development. A comparison of egg size and brood size in females from Tar Landing suggests that individuals with the two forms of development package offspring differently but expend approximately equivalent reproductive effort. Larval trophic mode is best viewed as a genetic polymorphism in S. benedicti. Individuals with planktotrophic and lecithotrophic development exhibit similar reproductive responses to environmental variation and there is no evidence for speciation.