Reproduction and growth of Holothuria atra (Echinodermata: Holothuroidea) at two contrasting sites in southern Taiwan

Reproductive periods and growth of two populations of Holothuria atra Jaeger distinctly different in body size at two sites of southern Taiwan were determined. Individuals examined in the present study were collected between march 1990 and February 1992. At Nanwan (21° 57N, 120° 45E), large individuals (351 to 1400 g wet wt) spawned from June to September. At Wanlitung (22°N, 120° 42E), a small proportion of frequently dividing individuals (<190 g wet wt) have mature gonads in May, June and September, but histological examination revealed no sign of spawning. Sexual recruits, defined as small individuals <5 g wet wt without sign of regeneration, were not found at either site during this 2-yr study. After the peak of fission at Wanlitung, 40% of the population showed signs of external regeneration. At Nanwan, small individuals transferred from Wanlitung grew from 6 g (n=6) to 166±8 g within 8 mo, and from 48±4 g (n=50) to 324±16 g within 1 yr, with a 6.8-fold biomass increase in 1 yr. At Wanlitung, the monthly average body weight of H. atra is between 33 and 62 g, apparently due to frequent fission, and the biomass increased only 2.9-fold in 1 yr. In southern Taiwan, sexual reproduction of H. atra occurs in large individuals. Asexual reproduction, occurring in small individuals, is the chief mechanism for population maintenance and increase, but it may decrease sexual reproductive potential.     

Comparison of light- and SST-based geolocation with satellite telemetry in free-ranging albatrosses

Light-based archival tags are increasingly being used on free-ranging marine vertebrates to study their movements using geolocation estimates. These methods use algorithms that incorporate threshold light techniques to determine longitude and latitude. More recently, researchers have begun using sea surface temperature (SST) to determine latitude in temperate regions. The accuracy and application of these algorithms have not been validated on free-ranging birds. Errors in both geolocation methods were quantified by double-tagging Laysan (Phoebastria immutabilis Rothschild) and black-footed (P. nigripes Audubon) albatrosses with both leg-mounted archival tags that measured SST and ambient light, and satellite transmitters. Laysan albatrosses were captured and released from breeding colonies on Tern Island, northwestern Hawaiian Islands (23°52N, 166°17W) and Guadalupe Island, Mexico (28°31N, 118°10W) and black-footed albatrosses from Tern Island. Studies were carried out between December 2002 and March 2003. For all birds combined, the mean ± SD great circle (GC) distance between light-based locations and satellite-derived locations was 400±298 km (n=131). Errors in geolocation positions were reduced to 202±171 km (n=154) when light-based longitude and SST-based latitude (i.e. SST/light) were used to establish locations. The SST/light method produced comparable results for two Laysan albatross populations that traveled within distinctly different oceanic regions (open ocean vs more coastal) whereas light-based methods produced greater errors in the coastal population. Archival tags deployed on black-footed albatrosses returned a significantly higher proportion of lower-quality locations, which was attributed to interference of the light sensor on the tag. Overall, the results demonstrate that combining measures of light-based longitude and SST-based latitude significantly reduces the error in location estimates for albatrosses and can provide valid latitude estimates during the equinoxes, when light-based latitude measurements are indeterminate.Communicated by J.P. Grassle, New Brunswick     

Relationship of daily and circatidal activity rhythms of the fiddler crab,<Emphasis Type="Italic"> Uca princeps</Emphasis>, to the harmonic structure of semidiurnal and mixed tides

Intertidal organisms may employ circatidal rhythms to track the tidal cycle, but tidal patterns may vary within a species range and necessitate adaptation to the local tides. Circatidal rhythms were examined in populations of the eastern Pacific fiddler crab Uca princeps (Smith) from four sites with differing tidal characteristics, La Paz (24°10N; 110°21W), San Blas (21°33N; 105°18W) and Manzanillo (19°6N; 104°24W), Mexico (lower amplitude, mixed semidiurnal tides) and Mata de Limon, Costa Rica (9°55N; 84°43W) (high-amplitude, semidiurnal tides). Local tides were characterized by harmonic constants of M₂, S₂, K₁, and O₁, partial tides that largely determine their semidiurnal and diurnal features. Rhythmic structure in continuously recorded locomotor activity of individual crabs held under laboratory conditions was described by cosinor and periodogram methods of time-series analysis. Both daily and circatidal rhythms were found in crabs studied in light–dark cycles set to local conditions at the time of collection. Crabs at all four sites shared a tendency toward bimodality, with a mid-morning activity peak and varying degrees of nocturnal activity. Circatidal rhythms closely matching the period of the 12.42-h M₂ partial tide were consistently present at all sites except Manzanillo. At Mata de Limon, the circatidal rhythm clearly dominated locomotor activity, but was strongly modulated by a daily rhythm in a repeating pattern at a semilunar interval. In contrast, the amplitude of the daily rhythm was higher than that of the circatidal rhythm in crabs from the three mixed tide sites on the Mexican coast, where the tidal pattern is dominated by a diurnal inequality arising from the diurnal K₁ and O₁ partial tides. These results suggest that populations of U. princeps use both daily and circatidal timing systems to track local forms of the tide generated by their M₂, S₂, K₁, and O₁ geophysical counterparts.Communicated by J.P. Grassle, New Brunswick     

Effect of freezing on photosynthesis of intertidal macroalgae: relative tolerance of Chondrus crispus and Mastocarpus stellatus (Rhodophyta)

The effect of freezing on photosynthetic metabolism was studied in the red algae, Chondrus crispus and Mastocarpus stellatus. Plants of both species were collected from the intertidal at Chamberlain or Kresge Point, Maine, USA (43°56N, 69°54W) between February and March 1987. Photosynthetic rates were measured immediately after freezing at-20°C and following recovery periods in seawater. Photosynthesis in C. crispus declined rapidly following freezing, falling to 70% of control values within 1 h and 30% after 3 h exposure. Minimum photosynthetic rates (7 to 9% of controls) occurred following freezing exposures of 12 h or more. Full photosynthetic recovery in C. crispus after 3 h at-20°C required 48 h. Photosynthesis in C. crispus did not fully recover in plants frozen for 6 h or more. In contrast, photosynthesis in M. stellatus was relatively unaffected by freezing exposures of <12 h. Twelve hours or more at-20°C reduced photosynthesis to 55% of controls. Photosynthesis in M. stellatus fully recovered from 24 h at-20°C within 24 h. In both species the reduction of photosynthesis by freezing was associated with damage to the plasma membrane and reduced efficiency of energy transfer from phycobilisomes to chlorophyll a, but did not appear to involve ribulose-1,5-bisphosphate carboxylase oxygenase activity. The freezing tolerance of C. crispus and M. stellatus positively correlates with their respective intertidal distributions, suggesting that freezing may be involved in controlling the distributions of these species on the shore.     

Reproductive cycle of Golfingia margaritacea,a bipolar sipunculan,in subantarctic water

The reproductive cycle of the bipolar sipunculan Golfingia margaritacea collected from Ushuaia, Argentina (54°49 S; 68°13W) in 1986 and 1987 is reported. The population sampled showed a sex ratio of 1:1. Males, however, frequently outnumbered females. Measurements of coelomic oocytes delineated an annual reproductive cycle. Small oocytes were present in the coelom all year: a subpopulation began to increase size about the middle of spring and reached its maximum size in late summer. Spawns may occur from the end February to April. May is characterized by resorption of large oocytes. Monthy monitoring of the gonads revealed mitotic activity and a continuous release of gametocytes to the coelom. Parasites were detected in the gonads. Large coelomic oocytes were also present in the coelom during the same season in the Northern and Southern Hemispheres, but egg sizes were significantly larger in boreal populations.     

Transformation of soft coral (Coelenterata: Octocorallia) terpenes by Ovula ovum (Mollusca: Prosobranchia)

The faecal pellets from specimens of the prosobranch mollusc Ovula ovum found feeding on the soft coral Sarcophyton sp. at Eclipse Island, Palm Island Group (18°46S; 146°33E) in November 1980 were analysed. The only terpene present in the faeces, 7,8-deoxysarcophytoxide, differed from the major constituent of the soft coral, sarcophytoxide, suggesting that the latter had been transformed into the former within the cowrie. This transformation is not trivial, and could not be produced simply by acid catalysis. Subsequent analysis of tissues dissected from different regions of O. ovum indicates that the transformation is probably effected by enzymes in the digestive diverticula stomach region of the prosobranch. The transformed compound is significantly less toxic to the mosquito fish Gambusia affinis Baird and Girard than the ingested compound.     

Re-appraisal of age composition,growth and survivorship of the deep-sea brittle star Ophiura ljungmani from size structure in a sample time series from the Rockall Trough

Samples were taken from a 2 900 m deep permanent station (centred on 54°40N; 12°16W) in the southern Rockall Trough, from November 1975 to September 1980, inclusive. Revision of juvenile size frequencies was necessary after removal of postlarvae of an annual non-viable settlement of Ophiocten gracilis (G. O. Sars) that were mistakenly included in a previous analysis. The present study of the revised data shows that settlement of Ophiura ljungmani (Lyman) occurs in summer when disk-size frequencies are dominated by the newly metamorphosed postlarvae. The likely age composition in larger sizes was suggested from the progression of modes along the size axis in juvenile size classes in consecutive were developed from Gaussian mixtures fitted by constrained computer search using, as starting values, component parameter values suggested from the frequency histograms. Choice between competing solutions was made on the basis both of goodness-of-fit and biological plausibility. However, identification of year-class structure amongst sizes >3 to 4 mm disk diameter remained conjectural owing to the low frequencies of adults in the samples. Comparison of the fitted component-means in the time series suggested a marked seasonal growth pattern, with maximal growth in spring. Probable age composition amongst adult sizes was extrapolated from von Bertalanffy growth parameters estimated from the means of juvenile year classes.Communicated by J. Mauchline, Oban     

What is the sex ratio of harpacticoid copepods in the deep sea?

The sex ratios of deep-sea harpacticoids have been thought to be greatly skewed toward females. The representation of males is notably more equitable (approximately 1:2) at three deep-sea sites (San Diego Trough: 32°52.4N, 117°45.5W, 1 050 m, January 1987; Porcupine Seabight: 51°36.85N, 12°57.30W, 1 369 m, August 1984; northwest Atlantic, 40°27N, 62°20W, 4 820 m, July 1982, June 1983). This ratio conceals extensive variation at the species level. For some species, males are unknown. For other species, males are more numerous than females. Sampling bias against males can occur and may explain the more extreme reports of male rareness in the literature.     

Response of a soft-bottom harpacticoid community to stingray (Dasyatis sabina) disturbance

The recolonization by harpacticoid copepods of patches disturbed by stingrays [Dasyatis sabina (LeSeur)] feeding in a subtidal (2–3 m depth) site off the Florida gulf coast (Lat. 29°54.55N, Long. 84°31.45W) was followed. Samples from disturbed patches could no longer be distinguished from control samples in terms of numbers of harpacticoids 29 h after disturbance. No species became significantly disproportionately abundant during the recovery suggesting that small-scale natural disturbances are not an important organizing force in this community.     

Response of gametophytes of Ecklonia radiata (Laminariales) to temperature in saturating light

Gametophytes of Ecklonia radiata (C.Ag.)J.Ag. from two New Zealand locations with different field temperature ranges were exposed to temperatures of 5° to 26°C in saturating light. Plants from Goat Island Bay (Lat. 36° 16S, Long. 174°48E) grew in 9.3° to 25°C and reproduced in 9.3° to 24°C. There was no growth at 8°C and plants died at 26°C. Plants from the cooler location, Houghton Bay (Lat. 41°20S, Long. 174°40E), grew from 8° to 24°C and reproduced up to 15°C but not at 21.5°C. The plants did not grow at 6°C and died at 26°C. The timing of the first cell division and subsequent growth rate were retarded close to the upper and lower tolerance limits. Reproduction was a broad optimum of roughly 12° to 20°C. Within this range, fertile female gametophytes grown at lower temperatures had fewer, larger cells and thus fewer potential ova than those grown at higher temperatures.     

Daily energy requirements and trophic positioning of the sand shrimp<Emphasis Type="Italic"> Crangon septemspinosa</Emphasis>

Sand shrimp, Crangon septemspinosa Say, are important to the trophic dynamics of coastal systems in the northwestern Atlantic. To evaluate predatory impacts of sand shrimp, daily energy requirements (J ind.^–1 day^–1) were calculated for this species from laboratory estimates of energy losses due to routine (R_R), active (R_A), and feeding (R_SDA) oxygen consumption rates (J ind.^–1 h^–1), coupled with measurements of diel motile activity. Shrimp used in this study were collected biweekly from the Niantic River, Connecticut (41°33N; 72°19W) during late spring and summer of 2000 and 2001. The rates of shrimp energy loss due to R_R and R_A increased exponentially with increasing temperature, with the magnitude of increase greater between 6°C and 10°C (Q₁₀=3.01) than between 10°C and 14°C (Q₁₀=2.85). Rates of R_R doubled with a twofold increase in shrimp mass, and R_SDA was 0.130 J h^–1+R_R, irrespective of shrimp body size. Shrimp motile activity was significantly greater during dark periods relative to light periods, indicating nocturnal behavior. Nocturnal activity also increased significantly at higher temperatures, and at 20°C shifted from a unimodal to a bimodal pattern. Laboratory estimates of daily metabolic expenditures (1.7–307.4 J ind.^–1 day^–1 for 0.05 and 1.5 g wet weight shrimp, respectively, between 0°C and 20°C) were combined with results from previous investigations to construct a bioenergetic model and make inferences regarding the trophic positioning of C. septemspinosa. Bioenergetic model estimates indicated that juvenile and adult shrimp could meet daily energy demands via opportunistic omnivory, selectively preying upon items of high energy content (e.g. invertebrate and fish tissue) and compensating for limited prey availability by ingesting readily accessible lower energy food (e.g. detritus and plant material).Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by J.P. Grassle, New Brunswick     

Avoiding predation: alarm responses of Caribbean sea urchins to simulated predation on conspecific and heterospecific sea urchins

Alarm responses to the extracts of conspecifics and hetero-specifics were measured for the Caribbean sea urchins Echinometra viridis, E. lucunter, Lytechinus variegatus, L. williamsi, Tripneustes ventricosus, Diadema antillarum, and Eucidaris tribuloides collected along the Caribbean coast (9°3314N; 78°5523W) during October 1984 and July–December 1985. Responses to seawater and extracts of the gnathostomate echinoid Clypeaster sybdepressus were used as controls. The intensity of the response resulting from exposure to sea-urchin extracts was measured by: (1) the percentage of individuals that responded by moving away from the extract and/or towards shelter, and (2) the mean distance moved. Echinometra viridis, E. lucunter, and L. williamsi responded to sea-urchin extracts by moving towards nearby shelter sites. The distance that individuals of each species moved in the first minute following exposure to conspecific extracts was correlated with the distance that species moved from shelter while foraging. L. variegatus and D. antillarum, living in microhabitats not providing protection from predators, responded to extracts of conspecifics and heterospecifics by moving away from the direction of the extract. Eucidaris tribuloides did not exhibit alarm responses to the extracts of con- or hetero-specifics. E. tribuloides secures itself with its stout spines into protected sites within corals. Similarly, L. variegatus living in long, dense seagrass that provided protection from detection by predators, and D. antillarum occupying crevices, showed no alarm responses to extracts of conspecifics. Presumably, in these situations, sea urchins cannot increase their defenses against predation by moving away from an injured neighbor. T. ventricosus showed a weak response to extracts of L. variegatus, but no response to extracts of other species including conspecifics. The reasons for this lack of an alarm response are unclear. For the five species that demonstrated an alarm response to sea-urchin extracts, the intensity of the response varied depending on the type of extract used. L. variegatus, L. williamsi, and D. antillarum responded most strongly to extracts from conspecifics, while Echinometra viridis and E. lucunter responded strongly to extracts from both conspecifics and congeners. The weakest responses were shown to the extracts of T. ventricosus and Eucidaris tribuloides. Habitat overlap, overlap in predators, and phylogenetic relationships did not consistently explain patterns of alarm responses to the extracts of heterospecific sea urchins.     

Fouling organisms in the port of Mar del Plata (Argentina). I. Siphonaria lessoni: ecological and biometric aspects

The present paper deals with ecological aspects of the pulmonate gastropod Siphonaria lessoni (Blainville, 1824), a common member of fouling communities in Mar del Plata's port (38°0817 S, 57°3118 W). Populations on colonized panels of an experimental raft were studied during three annual cycles, to determine feeding habits, growth and various factors which restrict their distribution on floating substrata. Although S. lessoni is harmless as a fouling organism, its grazing on the algal belt clears a part of the substratum's surface, which is then quickly colonized by truly aggressive organisms such as Balanus amphitrite.     

Procedures adopted for the laboratory cultivation of Trichodesmium erythraeum

The present paper describes the procedures adopted for axenic or pure cultivation of the marine blue-green alga Trichodesmium erythraeum (Ehr.)¹ in the laboratory. Strains of T. erythraeum were isolated from the nearshore waters of the Bay of Bengal (11°29 N 79°49 E) during the pre-bloom period in March, 1965, at a depth of 9 to 12m. Modified, enriched, Erdschreiber medium was used for laboratory cultivation of T. erythraeum. A differential concentration of streptomycin 6 mg/100 ml, tetracycline 5 mg/100 ml and sulfadiazine 4 mg/100 ml was used to inhibit the microbial growth in cultures. Difficulties experienced in the initial culture stages of T. erythraeum and the necessary specific instructions employed are described in detail.     

Two new records of Planctosphaera larvae (Hemichordata: Planctosphaeroidea)

Two specimens of the rarely found Planctosphaera larva have been taken in the North Atlantic near Challenger Bank north of the Azores and in the North Equatorial Current, latitude 19°33 N, longitude 40°01 W.This research was supported by Grant 6027-X from the U.S. National Science Fonndation and is Contribution No. 2380 from the Woods Hole Oceanographic Institution.     

Spatial mesoscale patterns of West Pacific picophytoplankton as analyzed by flow cytometry: their contribution to subsurface chlorophyll maxima

Vertical distributions of picophytoplankton (ppp) (<2 m) were studied by ship-board flow cytometry during two cruises in Western Pacific waters to Palau and to Australia in 1990. Weak red-fluorescing small ppp, supposed to be free-living prochlorophytes (Chisholm et al. 1988), were abundant in the area surveyed. These ppp, designated the prochlorophytes, were abundant in the surface waters (>10⁴ cells ml^-1) at the northern region (27°03N; 7°11N) in November, whereas in December at the southern tropical stations (0°23.54S; 9°20.30S; 13°50.6S), they formed subsurface maximum layers (>10⁵ cells ml^-1) on a nitracline at a depth of 3.5 to 5.4% surface irradiation. Their fluorescence intensity increased with depth below 10% surface irradiation. The prochlorophytes at a depth of 1% surface irradiation had ten times higher fluorescence than those at the surface layer. The total fluorescence intensity of the prochlorophytes accounted for 32 to 63% of the sum of the total fluorescence intensity of all fluorescing phytoplankton detected at subsurface chlorophyll maxima in the tropical area. These results suggest that distribution of the prochlorophytes is greatly affected by nitracline and by light intensity and that their chlorophyll is a major contributor to the subsurface chlorophyll maximum in the pelagic West Pacific Ocean.     

Algal overgrowth of alcyonacean soft corals

Colonies of the soft coral Lobophytum pauciflorum (Ehrenberg, 1834) (Coelenterata: Octocorallia: Alcyonacea: Alcyoniidae), some of which were heavily overgrown by the algae Ceranium flaccidum (Kuetzing) Ardissone and Enteromorpha sp., and other minor epizoites, were collected at Taylor Reef (17°50S; 146°35E) in the Great Barrier Reef. Overgrown colonies contained the diterpene 2-epi-sarcophytoxide as the major secondary metabolite, while conspecific colonies with clean polyparies contained two diterpenes in approximately equal amounts: 14-hydroxycembra-1,3,7,11-tetraene and 15-hydroxycembra-1,3,7,11-tetraene. By contrast, twenty conspecific pairs of overgrown and clean colonies of other alcyoniid soft corals collected from Pelorus Channel, Palm Island Group (18°34S; 146°29E), showed no chemical differences within in the pairs. Cultures of a common species of Ceramium [C. codii (Richards) Mazoyer] were incubated with different concentrations of nine soft-coral-derived diterpenes and significant algal growth inhibition was observed in many cases. It appears that terpenoids from soft coral may contribute to the lack of epizoic organisms on soft-coral polyparies.     

Epizoic diatoms on the euphausiidNyctiphanes australis: Consequences for gut-pigment analyses of whole krill

We report the first record of infestation by the epizoic pennate diatoms (Licmophora sp.) of the euphausiidNyctiphanes australis (G. O. Sars, 1883) collected from Otago Harbour and coastal waters, New Zealand (45°50S; 170°37E) between October and April 1986–1989: at times up to 50 to 70% of the krill examined were infested. Infestation by epizoic diatoms introduces a significant source of error into determinations of egestion rates using gut fluorescence of whole individuals, and prevents measurement of background fluorescence in krill. Diatom infestation can lead to overestimation of ingestion rates by a factor of 2 to 5. We suggest that investigators using the gut fluorescence method to measure grazing rates should determine the occurrence of epizoic diatoms in samples prior to analysis. If diatoms are present, it is necessary to hold krill in the laboratory until diatoms are shed at moult before makingin vitro gut-pigment measurements. Alternatively, the stomachs must be dissected for analyses, negating the benefits of rapid sample preparation and quantitative recovery of pigments that are the advantage of the technique. Epizoic diatoms can significantly contribute to variability of apparentin situ ingestion rates inN. australis.     

Delayed recovery of giant kelp near its southern range limit in the North Pacific following El Niño

The northern distribution limit of giant kelp, Macrocystis pyrifera (Agardh), along the Pacific coast of North America is relatively stable near Point Año Nuevo, California, but its southern limit has varied over hundreds of kilometers along the Baja California peninsula during the past 20 years. The factors that drive this variability in the southern limit were examined by: (1) quantifying the abundances of M. pyrifera and the subsurface stipitate kelp, Eisenia arborea (Areschoug), near M. pyriferas then southern limit at Punta San Roque (27°15N; 114°42W) on ten occasions between August 1997 and June 2002 (prior to and following the 1997/1998 El Niño Southern Oscillation); (2) initiating a field manipulation at Punta San Roque in June 2000 to examine competition between these two kelp species; and (3) surveying 20 additional sites near Punta San Hipólito (27°01N; 114°00W) in June 2002 to determine whether M. pyrifera populations had recovered to their pre-1982/1983 El Niño southern range limit after ~20 years of absence. These observations and the experimental results suggest that M. pyrifera is removed from its southern limit by the high temperatures and large waves associated with El Niños, while E. arborea is able to survive and recruit in high densities and thereby delay or even prevent M. pyriferas recovery. An inverse relationship was observed between these two species over the 5-year study. The field experiment showed that at both 4 months and 1 year after areas occupied by E. arborea were cleared, substantial recruitment of M. pyrifera occurred, while recruitment was negligible in uncleared areas. This study suggests that the southern range limit of M. pyrifera is determined by increased mortality and recruitment failure following the warm temperatures and low nutrient conditions associated with El Niños, limited substrate availability, and by competition with the more persistent populations of E. arborea.Communicated by J.P. Grassle, New Brunswick     

Stable isotopic investigation of physiological and environmental changes recorded in shell carbonate from the giant clam Tridacna maxima

The aragonitic shell of the photosymbiont-bearing bivalve Tridacna maxima contains a record of the physiological and environmental changes the organism has experienced during its lifetime. This record is preserved as chemical and microstructural variations throughout the shell. Stable isotopic analyses of oxygen (¹⁸O/¹⁶O) and carbon (¹³C/¹²C) in shell carbonate were combined with growth increment studies to interpret the shell record of specimens collected from the Rose Atoll (Lat. 14°31S; Long. 168°10W) in April 1982. The seasonal water temperature cycle is recorded in the oxygen isotopic signature of the clams, permitting the recognition of annual cycles in the ¹⁸O profile. The total number of these cycles corresponds to the age of a specimen, while the cycle length is a measure of the yearly growth rate. Large-amplitude cycles, reflecting year-round calcification, characterize the early portion of the growth record. With the onset of sexual maturity and slower growth at an age of approximately ten years, the cycles decrease in amplitude and become more erratic. During this later growth phase calcification is limited to the cooler months of the year, perhaps in response to a re-ordering of energy priorities between growth and gametogenesis. A growth curve developed from the ¹⁸O profile indicates rapid juvenile shell growth followed by slower growth thereafter producing a lifespan of several decades. Carbon isotopic analyses of T. maxima were compared to analyses of the symbiont-barren gastropod Terebra areolata collected from the same locality in April 1984. A 2 depletion in the ¹³C composition of T. maxima shell carbonate is attributed to a symbiontenhanced metabolic rate and an increased flow of isotopically light, respired CO₂ into the carbon pool used in calcification. Such a depletion may prove useful in identifying the presence of photosymbionts in extinct species of fossil mollusks.