Ecology of a corallivorous gastropod,<Emphasis Type="Italic"> Coralliophila abbreviata</Emphasis>, on two scleractinian hosts. II. Feeding,respiration and growth

Coralliophila abbreviata is a corallivorous gastropod that has been observed to cause large feeding scars on reef-building corals on Floridian and Caribbean reefs. We detected differences in the population structure (length-frequency distribution and sex ratios) of C. abbreviata populations living on two coral host taxa, Acropora palmata and Montastraea spp., in the Florida Keys in a previous study. We hypothesize that diet (host) has a major influence on snail population structure and, thus, we characterize metabolism, feeding and growth for snails residing on these coral taxa. Here, we present results of a reciprocal transplant experiment demonstrating that the taxon of the host influences snail growth rates, as indicated by changes in shell and body tissue weight. Regardless of the host from which they were drawn, snails resident on A. palmata grew faster than those resident on Montastraea spp. Thus, diet influences snail population structure. However, the tissue of Montastraea spp. provides more N and C per area of tissue than that of A. palmata. Respiration rates and tissue composition of snails collected from the two host taxa did not differ. Therefore, snails feeding on Montastraea spp. should have to consume less tissue per day to satisfy their metabolic requirements compared to snails feeding on A. palmata. Feeding rates for snails on A. palmata were measured in the laboratory over 48 h (1–9 cm² coral tissue snail⁻¹ day⁻¹) and estimated from feeding scars observed in the field (weekly mean rate of 1.07 cm² coral tissue snail⁻¹ day⁻¹). The lack of definition of snail feeding scars on Montastraea spp. required the calculation of coral tissue consumption rates based on estimated minimum carbon requirements. Calculated feeding rates for C. abbreviata were 0.13–0.88 cm² coral tissue day⁻¹ snail⁻¹, when feeding on Montastraea spp., and 0.44–3.28 cm² coral tissue day⁻¹ snail⁻¹, when feeding on A. palmata. The calculated range for the latter is consistent with measured rates. Thus, C. abbreviata exhibits high variation in growth parameters in response to environmental variability and/or food source. At mean levels of snail density on reefs off Key Largo, Fla., 20% of A. palmata colonies lose between 1.32 and 9.84 cm² tissue day⁻¹, while 50% of Montastraea spp. colonies lose between 1.04 and 7.04 cm² tissue day⁻¹. Together with published coral tissue regeneration rates, these results suggest that if sustained, such rates of predation could have a serious effect on the viability of these coral populations on Florida's reefs.     

Ecology of a corallivorous gastropod,<Emphasis Type="Italic"> Coralliophila abbreviata</Emphasis>, on two scleractinian hosts. I: Population structure of snails and corals

Despite their potential importance in structuring reef communities, invertebrate corallivores and their population structures are poorly understood. We found distinct differences in the population structures (length-frequency distribution and sex ratio) of the corallivorous gastropod Coralliophila abbreviata residing on two coral-host taxa, Montastraea spp. and Acropora palmata, in the Florida Keys. In each of two survey years, around 50% of the Montastraea spp. colonies were infested, with a mean snail density of eight snails per infested colony (range 1–45), while around 20% of A. palmata colonies harbored three snails per infested colony (range 1–23). Variation in patterns of snail occurrence was also observed within a host taxon. A. palmata occurred in low- and high-density stands (0.4 and 1 colony m^–2, respectively, at the initial survey) at different sites. Hurricane Georges struck the area in September 1998. When resurveyed in 1999, density of colonies in low-density stands had decreased by 75% to 0.1 colonies m^–2. This decrease was accompanied by a doubling in the proportion of colonies infested with snails (from 19% to 46%) and an increase in snail density per infested colony (from 3.7±3.3 SD to 5.4±4.6 SD) as snails apparently concentrated on surviving A. palmata. In contrast, sites with high density A. palmata stands (thickets) retained colony densities of about ~1 colony m^–2 among years, while snail infestation increased only from 9% to 14% of colonies surveyed and snail density essentially remained unchanged (from 2.7±1.8 to 2.9±1.9 snails per infested colony). Snails collected from Montastraea spp. were shorter than those from A. palmata in low-density stands and were longest on A. palmata in thickets. On both host taxa, female snails were longer than males. The sex ratio of snails on Montastraea spp. hosts was even (1:1), while that of snails on A. palmata was skewed (70% males). Factors that could explain observed differences in size structure and sex ratio between Coralliophila populations on the two coral host taxa include: differential susceptibility to predators, influence of host tissue nutritional quality and/or secondary metabolite content, and genetic differences (cryptic species). The host-specific characteristics of C. abbreviata populations imply that the impact of gastropods on reef communities will vary with the coral species composition.     

Three party symbiosis: acoelomorph worms,corals and unicellular algal symbionts in Eilat (Red Sea)

Epizoic worms were found to occur on certain coral colonies from reefs off the coast of Eilat (Red Sea). We identified 14 coral species infested by acoelomorph worms at a depth range of 2–50 m. The host corals were all zooxanthellate and included both massive and branching stony corals and a soft coral. Worms from all hosts were identified as belonging to the genus Waminoa and contained two distinct algal symbionts differing in size. The smaller one was identified as Symbiodinium sp. and the larger one is presumed to belong to the genus Amphidinium. Worm-infested colonies of the soft coral, Stereonephthya cundabiluensis, lacked a mucus layer and exhibited distinct cell microvilli, a phenotype not present in colonies lacking Waminoa sp. In most cases, both cnidarian and Acoelomorph hosts displayed high specificity for genetically distinctive Symbiodinium spp. These observations show that the epizoic worms do not acquire their symbionts from the “host” coral.     

Fine-scale patchiness and genetic heterogeneity of recruits of the corallivorous gastropod Drupella cornus

The size-frequency distributions and genetic composition of recruits of the corallivorous snail Drupella cornus (Röding, 1798) were examined in outbreak populations collected from Ningaloo Reef, Western Australia in August 1990. The recruits are found in groups on digitate Acropora spp. corals. Among coral colonies, mean lengths of recruits in our samples ranged from <9 to 22 mm, but the snails within a group were generally similar in size. Despite the fact that D. cornus has planktonic larvae, there were marked genetic differences between groups of recruits on different coral colonies. The relatively large genetic subdivision among groups of recruits within sites over distances <80 m was measured as a value of F _ST (standardized variance in allelic frequencies) of 0.044. This was three times the value from comparisons of pooled samples of recruits from areas up to 119 km apart, and six times as great at the genetic subdivision among populations of adults over a distance of 180 km. Much of the genetic heterogeneity among groups of recruits is associated with mean size of the snails. Taken together, the size-frequency distributions and the genetic differences indicate that recruits within the same coral colony shared a common history of settlement, suggesting a cohesiveness of groups of larvae. Although the mechanisms for this patchiness are not understood, one implication is that studies of size-frequency distributions and genetic composition of cohorts of D. cornus must treat the group, not the individual snail, as the unit of replication.     

Neoplasia,regeneration and growth in the reef-building coral Acropora palmata

Abnormal processes of calcification, such as regenerating lesions and neoplasia, situated near the tips(<25 cm) of colonies of Acropora palmata (Lamarck) suppressed normal linear growth. Branches having neoplasia at a larger distance from the tip do not grow significantly differently from controls. This indicates a functional minimal area in terms of energy supply. Neoplasia are pure aragonite and have the same coenosteal structure as regenerative skeletal material. Regeneration of tissue as well as tissue+skeleton lesions involves the simultaneous formation of tissue and regenerative skeleton, trapping foreign material under the regenerated surface. Recovery of a damaged surface slows down with time and this may, in other coral species, result in permanent lesions. A. palmata recovered from all lesions (n=32) within 80 d and appears to be a superior regenerator among Caribbean corals. This is consistent with other life-history characteristics of this highly specialized coral species.     

Effects on growth and reproduction of the coral Stylophora pistillata by the mutualistic damselfish Dascyllus marginatus

Although coral dwelling fishes are common on coral reefs, the nature of their effect on the host corals is poorly understood. The present study, conducted in the Gulf of Eilat (Red Sea) between July 1989 and August 1990, demonstrated that the branching coral Stylophora pistillata (Esper) benefits, in two components of coral fitness, from the presence of the damselfish Dascyllus marginatus (Rüppell), an obligate coral dweller. The growth rate of damselfish-inhabited corals was significantly higher than that of corals without damselfish. This was observed, using two growth assessment methods, in long-term (>7 mo) comparisons between: (1) corals where the damselfish were experimentally removed versus corals with unaltered fish groups; and (2) naturally inhabited versus non-inhabited corals. The presence of damselfish did not affect the coral's specific (per surface area) reproductive output, whether it was assessed by the number of female gonads per polyp or by the number of planulae released cm^-2 surface area d^-1. However, the more rapid increase in branch size in damselfish-inhabited corals resulted in an apparent increase in the total reproductive output, with age, in growing corals. These findings demonstrate that the association between the damselfish D. marginatus and its host coral, S. pistillata, is mutualistic.     

Depth limits of Bermudan scleractinian corals: a submersible survey

Depth distribution, zonation pattern and growth morphology of 17 hermatypic and 4 ahermatypic coral species were investigated at eight different locations along the Bermuda platform with the research submersible GEO and by SCUBA diving in August–September 1983. Hermatypic coral growth occurs to a depth of 50 to 70 m, with a single Montastrea cavernosa growing at 78 m. Dominant forms in shallow-water coral communities are Diploria sp. and Porites astreoides, while M. cavernosa, Agaricia fragilis and Scolymia cubensis occur in deep-water associations below 60 m. Vertical visibilities (up to 178 m) and distribution of the photosynthetically active radiation revealed good light penetration values (1% level at about 100 m depth), which should favour hermatypic coral growth to a much greater depth than it actually occurs. Nor should the prevailing temperatures limit the depth of coral growth. Most deep-water hermatypes observed grow on remnants of Pleistocene reefs down to about 60 m. The vast areas of large massed rhodolith nodules below 50 to 60 m are unsuitable bottom for coral colonisation. Macroalgae growth seems to be the strongest factor controlling coral growth in deep water. Bermuda stony corals have a low growth form diversity. Various intraspecific morphs may occur at the same as well as at different depths, with a general trend towards flatter shapes with depth. Comparison with a similar study on Red Sea corals suggests that annual distribution of radiant energy on the most northern Atlantic reefs of Bermuda may be responsible for the occurrence of flat and cuplike growth forms in relatively shallow water, and for the shallower depth limits of hermatypic growth.     

Monthly skeletal extension rates of the hermatypic corals<Emphasis Type="Italic"> Montastraea annularis</Emphasis> and<Emphasis Type="Italic"> Montastraea faveolata</Emphasis>: biological and environmental controls

Monthly skeletal extension rates were measured in colonies of Montastraea annularis and M. faveolata growing at Mahahual and Chinchorro Bank, in the Mexican Caribbean. Temperature, light extinction coefficient (k_d), sedimentation rate, dissolved nutrients and wave energy were used as indicators of environmental conditions for coral growth. Zooxanthella density and mitotic index, nitrogen, phosphorous and protein in coral tissue, and living tissue thickness were measured during periods of high-density-band (HDB) and low-density-band (LDB) formation. To test their value as indirect measures of competition between zooxanthellae and host, as well as coral health and performance in both species, these biological parameters were also measured, during the HDB-formation period, in corals collected at La Blanquilla. This reef is located in the Gulf of Mexico, in an area of suboptimal environmental conditions for coral growth. M. faveolata had a significantly higher skeletal extension rate than M. annularis. Corals growing in Mahahual had significantly higher skeletal extension rate than those living in Chinchorro Bank. This is consistent with inshore–offshore gradients in growth rates observed by other authors in the same and other coral species. This is probably due to less favorable environmental conditions for coral growth in near shore Mahahual, where there is high hydraulic energy and high sedimentation rate. Contrary to observations of other authors, skeletal extension rate did not differ significantly between HDB- and LDB-formation periods for both species of Montastraea. Both species produced their HDB between July and September, when the seawater temperatures are seasonally higher in the Mexican Caribbean. Tissue thickness indicated that environmental conditions are more favorable for coral health and performance during the HDB-formation period. Mitotic index data support the idea that zooxanthellae have competitive advantages for carbon over the host during the LDB-formation period. So, corals, during the LDB-formation period, with less favorable environmental conditions for coral performance and at a disadvantage for carbon with zooxanthellae, add new skeleton with little or no opportunity for thickening the existing one. This results in an equally extended skeleton with lower density, and the stretching response of skeletal growth, proposed for M. annularis growing under harsher environmental conditions, also occurs during the LDB-formation period.Communicated by P.W. Sammarco, Chauvin     

Annual density banding in massive coral skeletons: result of growth strategies to inhabit reefs with high microborers’ activity?

Porites and Montastraea are the major reef-building massive coral genera in the Indo-Pacific and Atlantic oceans, respectively. They are also the most commonly used genera in sclerochronological studies. Despite the marked differences in the way these genera use calcareous material to construct their skeletons (growth strategies) and in their skeletal architectural structure, they form annual high and low density bands in their skeletons, that result from the positive relationship of coral calcification rate with sea surface temperature and seasonal changes of the latter. Evidence in the literature suggests that the different growth strategies allow these organisms to construct denser skeletons far from terrigenous inputs, on reefs where microborers’ activity is high. It seems quite probable that this has consequences for the evolution, diversity, distribution and abundance of reef corals.     

Distribution,abundance and survival of juvenile hermatypic corals (Scleractinia) and the importance of life history strategies in the parent coral community

The distribution and abundance of juvenile corals were examined at depths from 3 to 37 m on the reefs of Curaçao and Bonaire (Netherlands Antilles). Juveniles of Agaricia agaricites were most abundant (60.6%), followed by Helioseris cucullata (8.3%). The large massive corals such as Montastrea annularis, M. cavernosa and branched species such as Madracis mirabilis and Acropora palmata had few juveniles. This, combined with species characteristics, shows that these species employ very different life history strategies. In some species the abundance of juveniles over the reef paralleled that of larger colonies, but not for example in Agaricia agaricites. The composition of the coral community was apparently no direct function of juvenile abundance. A change in angle of settlement of A. agaricites juveniles with increasing depth, from vertical to horizontal surfaces, seems to reflect the preferred light intensity. We studied the survival of juvenile corals during a half-year period. One-third remained unharmed, one-third died or disappeared, and one-third was limited in growth by factors such as spatial competition. This was the same for all depths, but factors influencing survival varied with depth.     

Predators selectively graze reproductive structures in a clonal marine organism

Although the fitness consequences of herbivory on terrestrial plants have been extensively studied, considerably less is known about how partial predation impacts the fitness of clonal marine organisms. The trophic role of Caribbean parrotfish on coral reefs is complex: while these fish are important herbivores, as corallivores (consumers of live coral tissue), they selectively graze specific species and colonies of reef-building corals. Though the benefits of parrotfish herbivory for reef resilience and conservation are well documented, the negative consequences of parrotfish grazing for coral reproductive fitness have not been previously determined. We examined recently grazed colonies of Montastraea annularis corals to determine whether grazing was positively associated with coral reproductive effort. We measured gonad number, egg number and size, and proportional reproductive allocation for grazed and intact coral colonies 2–5 days prior to their annual spawning time. We found that parrotfish selectively grazed coral polyps with high total reproductive effort (number of gonads), providing the first evidence that parrotfish selectively target specific tissue areas within a single coral colony. The removal of polyps with high reproductive effort has direct adverse affects on coral fitness, with additional indirect implications for colony growth and survival. We conclude that chronic grazing by parrotfishes has negative fitness consequences for reef-building corals, and by extension, reef ecosystems.     

Association of <Emphasis Type="Italic">Waminoa</Emphasis> sp. (Acoela) with corals in the Wakatobi Marine Park,South-East Sulawesi,Indonesia

This is the first quantitative study on the prevalence of epizoic Waminoa sp. acoel worms and their association with corals in the Wakatobi Marine National Park (WMNP), South-East Sulawesi, Indonesia. Three replicate transects were laid on the reef crest, flat and slope at six sites in 2006 and eight sites in 2007. Four of the sites were common in both years. In total 69 transects were surveyed in 2006, and 87 transects in 2007. A total of 4.8% of all observed hard corals were associated with acoel worms in 2006 and 2.6% of hard and soft corals in 2007. Acoels were present on 16 and 21 of the coral taxa studied in 2006 and 2007 respectively. The worms were strongly associated with the azooxanthellate coral Tubastrea spp. and were rare or absent on the most abundant coral genera Montipora and Porites. The mean number of corals having acoels was highest on reef slopes, whereas acoels were virtually absent on reef flats. Corals that had a high and a medium cover of worms were more common in 2007 than in 2006. No significant trend in the adaptation of the zooxanthellae of Waminoa sp. to different depths at different sites was revealed. The impact of the worm on the coral is unknown, but high numbers may have a shading effect and a negative impact on the coral’s photophysiology. This acoel merits more study of its life cycle, its photophysiology, and its impact on its host corals. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effect of epibiosis on the fitness of the sandy shore snail<Emphasis Type="Italic"> Batillaria zonalis</Emphasis> in Hong Kong

Batillaria zonalis is a common snail on Hong Kong sandy shores, occupying the low-mid shore. Shells of B. zonalis (basibiont) are often fouled with the rock oyster Saccostrea cucullata and the barnacle Balanus reticulatus (epibiont). This epibiotic interaction has a negative impact on the fitness of B. zonalis in terms of impairing fecundity, mobility and metabolism of fouled individuals. During the reproductive season, fouled snails had lower Gonadosomatic Index and fewer egg-bearing individuals than non-fouled snails. Fouled snails crawled in a more tortuous pathway and with lower speed when compared with non-fouled snails. Reduction in oxygen consumption and metabolic rate in fouled snails further suggested the restriction in mobility of B. zonalis by the epibionts. Glycogen content in the foot muscle tissues and hepatopancreas of fouled and non-fouled snails was, however, similar, revealing epibiosis does not influence the energy storage of B. zonalis. Mortality of fouled and non-fouled snails was similar under laboratory conditions, suggesting epibiosis does not influence the survivorship of B. zonalis in Hong Kong. Impairment in the fitness of fouled B. zonalis indicates that epibiosis can influence the population dynamics of B. zonalis in Hong Kong.Communicated by T. Ikeda, Hakodate     

Host effect on size structure and timing of sex change in the coral-inhabiting snail<Emphasis Type="Italic"> Coralliophila violacea</Emphasis>

The distribution, size and reproductive characteristics of the snail Coralliophila violacea (Lamarck), which inhabits the surface of both the branching coral Porites nigrescens and the massive corals P. lobata and P. lutea, were surveyed to examine the host effect on: (1) population structure and (2) reproductive characteristics, including the size at sex change of symbionts. On branching hosts, most snails were solitary, whereas on massive hosts, most had formed multiple-snail patches. Significantly smaller snails as well as proportionally more females were found on branching than on massive hosts. Furthermore, the fecundity of the females on the branching hosts was significantly lower than that on the massive hosts. The size at sex change (male to female) of the snails was smaller on the branching hosts than on massive hosts. Patch composition differences can partly explain the smaller size at sex change for snails on branching hosts; however, there was also evidence that host morphology had a significant effect on the timing of sex change.Communicated by T. Ikeda, Hakodate     

Dynamics of a coral reef community after mass mortality of branching <Emphasis Type="Italic">Acropora</Emphasis> corals and an outbreak of anemones

Dynamics of a coral reef community at Tiao-Shi Reef, southern Taiwan were studied using permanent transects to examine coral recovery and successive cascades to collapse stage resulting from chronic anthropogenic impacts and typhoons. Three distinct zones were recognized within a relatively small study area (250 m across) formerly dominated by large stands of branching Acropora corals. The first zone still retains the dominance of branching Acropora corals, although they show a significant decreasing tendency. The second zone exhibits recovery with a significant increase in branching Montipora stellata, which is recruited and grows faster than branching Acropora corals. The third zone is occupied by anemone, Condylactis sp., and demonstrates a stable phase of coral deterioration without recovery. Such differences in coral reef community dynamics within a small spatial scale illustrate mosaic dynamics which have resulted from degradation of the water quality, patchy mortality of large branching Acropora thickets caused by typhoons, the rapid asexual fragmentation and growth of M. stellata making it a successful colonizer, and occupation by anemone, Condylactis sp., together with unstable remnants of dead Acropora rubbles have not allowed coral recruits to survive.     

Plasticity in size and age at maturity in a monogamous fish: effect of host coral size and frequency dependence

The number and maximum body size of the gobioid fish, Paragobiodon echinocephalus, increase with the size of its obligate host coral, Stylophora. Only the largest two individuals breed monogamously in each coral head, and the reproductive success of each spawning is positively correlated with body size. In this study, the plasticity in size and age at maturity in P. echinocephalus was examined. We analyzed life history data from gobies 15–20 mm TL (total length) at their initial marking. Gobies found in larger corals were of lower rank in size order and began to breed later at a larger size, usually upon moving to other corals. The size at maturity ranged widely from 17.2 to 36.0 mm TL. After maturation, growth rates decreased. Mortality, however, was not affected by the timing of maturation. The host coral size did not affect the growth and mortality of marked fish, but the mortality rate of juveniles prior to marking appeared to be higher in smaller corals. The estimated lifetime reproductive success did not differ between the gobies inhabiting corals of different size. Thus the plasticity in size and age at maturity in this species may be maintained by frequency-dependent selection in choosing a host coral size that affects an individual’s social status. Received: 5 April 1995/Accepted after revision: 18 February 1996     

Size and structure of echinoderm populations associated with different coexisting coral species at Aldabra Atoll,Seychelles

The associated echinoderm fauna of 125 living heads of massive microatolls of Porites spp., branched heads of Porites nigrescens Dana and small, densely packed heads of the hydrocoral Millepora exacea Forskål of a shallow sublittoral lagoon flat at Aldabra Atoll, Seychelles was examined. The weight and number of echinoderms increased with coral size in all cases. The microatolls were the largest coral heads and sheltered the most diverse, underlying echinoderm cryptofauna comprising the largest bodied species, regardless of class or family. The infauna in heads of P. nigrescens was less diverse and smaller bodied than the echinoderms under microatolls. Only with P. nigrescens did species number increase with increasing coral size. Heads of M. exacea were the smallest and provided a habitat of compact interstitial spaces for the most depauperate, smallest bodied infauna which included juveniles of larger species. Ophiuroids dominated the coral-associated echinoderm faunas whereas larger bodied classes, such as echinoids and holothuroids, were less well represented. Large bodied ophiuroids such as certain ophiodermatids occurred only under microatolls, whereas the usually small bodied ophiotrichids dominated in the heads of M. exacea. The generally ubiquitous ophiocomids were well represented in, or under all corals. Heads of M. exacea could be nursery areas for recently metamorphosed juvenile ophiuroids and echinoids which, as they grow, seek shelter in corals with larger interstitial spaces such as P. nigrescens. Significant interspecific ophiuroid associations were related to space availability for larger species under microatolls and in heads of P. nigrescens. The possibility of microhabitat partitioning in corals among some ophiuroids is suggested.     

Population structure and sex-change in the coral-inhabiting snailCoralliophila violacea at Hsiao-Liuchiu,Taiwan

Surveys of the coral-inhabiting snailCoralliophila violacea (Lamarck) (=C. neritoidea Kiener) were made on shallow fringing reefs (<8 m deep) around Hsiao-Liuchiu, Taiwan, between July and October 1990. The snails were aggregated into patches on the surface of massive poritid coral colonies. Coral colonies >40 cm in diameter were more likely to bear patches of snails than smaller colonies, and also to have more snails. The coralliophilids ranged from 5 to 30 mm in aperture length. The sex ratio of the population was biased toward males (539:279), with only a few small individuals of indistinguishable sex. Snails between 6 and 10 mm were all males, while most snails with aperture lengths 20 mm were females. Judging from the distinct size ranges of males and females within patches and from the observed degeneration of the penis, the snails may have changed sex from male to female with increasing size. Sex-change may occur across a wide size range (10 to 20 mm). The correlation of smallest female size and largest male size among patches indicates that snail size at sex-change is peculiar to each individual patch. Those females in patches with a single female (but many males) were significantly smaller than females in multiple-female patches. It is likely that in the absence of females males change sex at a smaller size, whereas in the presence of large females males delay sexchange until they have reached a larger size. The plasticity of size at sex-change may be adaptive and a result of natural selection at the individual level.     

Stimulation of fat-body production in the polyps of the coralPocillopora damicornis by the presence of mutualistic crabs of the genusTrapezia

A mutualism exists between the xanthid crabs of the genusTrapezia and their host corals,Pocillopora damicornis. It has previously been established that these obligate coral residents benefit the coral hosts by defending them against echinoderm predators and by increasing the survival of polyps located deep between the coral branches. In turn, the corals apparently benefit the crabs by producing lipid-filled structures on which the trapezid crabs feed; these fat bodies may contain some of the lipid which in previous studies of coral metabolism has been termed excess. It was determined by experiments conducted at the Hawaii Institute of Marine Biology that the presence of crabs in colonies ofP. damicornis stimulates the polyps to produce the lipid-filled fat bodies; removal of crabs causes corals to cease producing fat bodies. A structure very similar to the fat bodies ofP. damicornis has been reported inAcropora durvillei. Both of these coral genera ordinarily possess xanthid-crab mutualists. This association between branching corals and crustaceans may have evolved because corals of these genera provide shelter among their branches and because these shallow-water corals are evidently capable of releasing lipid which is excess to the corals' metabolic needs, but which can be utilized by the crabs.