Pit relocation by antlion larvae: A simple model and laboratory test

Summary We generated a computer model to analyse the effects of shadow competition for sit-and-wait predators, particularly antlion larvae. The model used a simple foraging assessment rule to determine the quality of an antlion's location, and antlions relocated randomly in their habitat when a location proved to be of low quality. Shadow competition, or competition for food caused when one sit-and-wait predator intercepts moving prey before a second sit-and-wait predator is encountered, was incorporated into the model by restricting antlions to a bounded arena, and having prey for the antlions enter from the arena periphery. Antlions responded to shadow competition by relocating their pits to peripheral areas of their habitat. This peripheral accumulation of pits was most pronounced when antlion densities were high, and when prey availabilities were intermediate. An experimental test with the antlionMyrmeleon immaculatus supported the importance of shadow competition as a cause of observed pit distributions. Only the treatment which incorporated shadowing had pit distributions near the periphery, while the pit distributions in the control treatments did not differ from randomly generated distributions. We conclude that shadowing can influence sit-and-wait predator distributions when the prey distributions and movement patterns generate the conditions necessary for shadowing. But when prey availability is unpredictable, making assessment of patches difficult, or when prey do not originate in the periphery of the habitat, other factors, such as temperature or moisture, could be more important.     

Composition and distribution of meiofauna,including nematode genera,in two contrasting Arctic beaches

The meiofauna of two tidal beaches, one exposed and one more sheltered, on Bjornoya (Bear Island) was investigated in summer 2000. Both meiofaunal densities and composition seem to be controlled by physical properties of the sediment, which in turn are controlled by exposure. The moderately and poorly sorted sediments in the sheltered beach were more abundant in terms of meiofaunal densities than the well sorted sediments in the exposed beach (254–481 individuals in 10 cm² vs 7–269 individuals in 10 cm², respectively). In total, seven higher meiofaunal taxa were found. Turbellaria were the numerically dominant taxon in the exposed beach. In the sheltered beach, Turbellaria also dominated, followed by Nematoda and Harpacticoida. The vertical distribution of the meiofauna was in accordance with what has been reported from other intertidal beaches. Nematoda were studied in detail and their densities ranged over 0.7–7.7 individuals in 10 cm² in the exposed beach and 2.7–186.0 individuals in 10 cm² in the sheltered beach. Nematodes were identified to genus level and a total of eight nematode genera were found. Sediment community respiration, measured as oxygen consumption, ranged between 2.3 cm³ O₂ m^–2 h^–1 in the exposed beach and 7.3 cm³ O₂ m^–2 h^–1 in the sheltered beach (respectively, the equivalent of 24 mg and 75 mg of organic carbon metabolised per day). Values from the sheltered site are within the range of results registered in much warmer localities.     

An experimental study on the foraging behavior of a pit-building antlion larva,Myrmeleon bore

Foraging behavior of a pit-building antlion larva, Myrmeleon boreTjeder was investigated experimentally to elucidate the relation between the feeding level and pit relocation.

1. In artificial sands constructed in the field the 3rd instar larvae of M. bore rarely changed the positions of their pits, though several antlions had moved actively until they constructed pits. The average feeding rate was 0.3 prey/day/pit, and about 60% of prey captured were ants.
2. To examine whether or not M. bore larvae concentrate into the area where they can capture more prey, 8 antlions were released into each of 6 boxes filled with sand. I divided the sand surface of each box into two half areas, then gave prey to the pits built in a half area and gave no prey to the pits built in the other half. During the 50-day observation period, nonfed antlions never moved into the area where prey were given.
3. The 3rd instar larvae were reared separately without food. Even under starved conditions they rarely relocated their pits until dealth. The average duration of survival period was 83.9 days.
4. The experimental results indicate that M. bore larvae adopt a tactic of sedentary ambushing. These larvae exhibit low movement rates which are independent of prey capture rates.

     

The effect of sand depth,feeding regime,density, and body mass on the foraging behaviour of a pit‐building antlion

Abstract 1. Pit‐building antlions are small sit‐and‐wait arthropod predators, which dig conical pits in sandy soils. We studied how biotic (conspecific density and feeding regime) and abiotic (sand depth) factors affect pit diameter and depth, while taking into account the larval body mass. 2. Pit diameter increased with larval body mass at a decelerating rate. In addition, larger larvae tended to relocate less frequently than smaller ones. 3. Sand depth positively affected overall pit size, while increasing conspecific density had a weaker but negative effect on pit size. 4. Feeding the antlions resulted in an increase in pit diameter compared with an unfed control group. However, as prey size increased this positive effect diminished. This result suggests that the existence of prey provides information about the quality of the microhabitat, triggering pit extension. However, similarly to the reduction in the foraging effort of saturated predators, antlions provided with large prey invested only little effort in pit enlargement. 5. Antlions were previously shown to be sensitive to prey and conspecific vibrations in the sand. We thus expected the feeding regime of the neighbour to affect antlion behaviour – surrogate of discriminating between local and global shortage of prey. Nevertheless, antlions with fed neighbours (a local prey shortage) did not show different behaviour compared with a control group in which both antlions were unfed (a global prey shortage).     

Intraspecific competition of Glyptotendipes paripes (Diptera: Chironomidae) larvae under laboratory conditions

Glyptotendipes paripes larvae were reared in wells of tissue culture plates, in groups of 2, 4, 8, 16, and 32 (representing densities of about 1,300, 2,600, 5,200, 10,400, and 20,800 larvae per m², respectively). Larval groups were supplied with one of two concentrations (low or high) of food and larvae were individually observed to evaluate the effects of density on mortality, growth, development, behavior, and adult body size. Increased larval densities resulted in higher mortality, as well as slower larval growth and development. The distribution of developmental time became flatter at higher density, with a wider range of values, or even became bimodal. This was a consequence of the most rapidly developing individuals at higher densities emerging as adults sooner than the fastest developing individuals at lower densities, although overall mean developmental time was longer at higher densities. At higher densities, growth and development of smaller larvae were slowed, based on the relative difference in body length between competitors. When larger competitors emerged as adults or died, the growth of smaller larvae may have accelerated, resulting in increased variability of developmental times. The effect of larval density on adult body size was complex, with the largest body size found at the lowest density and a second peak of adult size at high-middle densities, with smaller adult body sizes found at low-middle, and high densities. Similarly, as with developmental time, the range of body size increased with increasing density. Examined food concentrations had no effect on larval mortality, but significantly affected developmental time, growth rate, and adult body size. At higher densities, larvae spent more time gathering food and were engaged in aggressive or antagonistic behaviors.     

Some effects of insecticide deposit patterns on the parasitism of Trialeurodes vaporariorum by Encarsia formosa

The functional response of Encarsia formosa parasitising Trialeurodes vaporariorum on Phaseolus vulgaris was a typical type II functional response (Holling, 1959) where, with increasing host density, the numbers of hosts parasitised by a single parasitoid show a negatively accelerating rise to an upper plateau. A mean maximum parasitism of 6.6 per female per 24 h reached at host densities near 80 larvae per leaf was observed. When presented with insecticide (‘Pynosect 30 ULV’) deposits, the parasite's search pattern changed, conforming to a typical Sigmoid functional response curve. With a uniform cover spray deposit of 27.6 × 10^-7 ml/cm² searching began only at host densities of c. 80 larvae per leaf and the mean maximum parasitism was only 1.5 per female per 24 h. When the insecticide deposit of 27.6 × 10^-7 ml/cm² was in the form of discrete groups of 100 μm droplets, searching began at host densities of c. 40 larvae per leaf but the mean maximum parasitism was low at 2.2. However by using the same discrete deposit pattern but reducing the deposit level to 2.65 × 10^-7 ml/cm² it was possible to obtain the same high mean maximum parasitism as with the untreated control.     

Seasonal, altitudinal and host plant-related variation in the abundance of aphids (Insecta, Hemiptera) on sub-Antarctic Marion Island

Although aphids are the most species-rich group of invasive alien insects across the Southern Ocean Islands, their biology on the islands is poorly known. In this study, host plant-related, seasonal and altitudinal variation in the abundance of the three aphid species (Macrosiphum euphorbiae, Myzus ascalonicus, Rhopalosiphum padi) found on sub-Antarctic Marion Island is examined. Myzus ascalonicus is the most abundant of the species, reaching densities of c. 2,400 individuals per 1,200 cm² on the indigenous Acaena magellanica (Rosaceae) and 479 individuals per 1,200 cm² on the indigenous Cotula plumosa (Asteraceae), with lower numbers at elevations up to 300 m and a pronounced seasonal peak in December-February. Macrosiphum euphorbiae occurs on the same plant species in densities of between c. 11 individuals per 1,200 cm² on C. plumosa and c. 200 individuals per 1,200 cm² on A. magellanica, and with a pronounced summer peak in December. In both species, small numbers of winged forms are found (c. 1–5%), although they are permanently parthenogenetic on the island. Rhopalosiphum padi is also permanently parthenogenetic on the island and likewise shows a December peak in abundance. The current data suggest that low ambient temperatures, which are close to the lower development threshold of R. padi, and other temperate aphid species, limit development in winter, thus leading to a decline in abundance.     

Laboratory study of conspecific avoidance by host-colonizingDendroctonus frontalis Zimm. (Coleoptera: Scolytidae)

Bark/phloem disk sandwiches (100 cm²) were colonized with one to six mating pairs of the southern pine beetle, Dendroctonus frontalis Zimm., to observe gallery construction and evaluate if avoidance behavior occurs between beetles. Frequencies of turn aways, touches, and crosses (three characteristics used to describe conspecific interaction) all increased significantly with beetle density. A threshold of three mating pairs within the 100- cm² phloem area was the density at which avoidance behavior became evident. At a threshold of five mating pairs the number of gallery crossings was significantly greater than at lower densities. Single and paired female beetles introduced into artificial galleries crossed empty neighboring galleries significantly more often than when neighboring galleries were occupied with other D. frontalis. The data suggest that D. frontalis females can regulate gallery spacing within the host through their ability to detect nearby beetles and their galleries. Possible spacing mechanisms are discussed.     

Variation in pit size of antlion (Myrmeleon carolinus) larvae: the importance of pit construction

Abstract.Larvae of several antlions build pits that vary in size across and within species. The influence of food limitation and pit building experience on variation in pit size of the larvae of Myrmeleon carolinus was investigated in the laboratory. Unfed larvae that were allowed to build pits had smaller pit diameters than fed larvae. However, fed antlions that had been previously prevented from pit building, initially did not build larger pits than unfed antlion larvae that, too, had been prevented from pit building. Therefore, physiological constraints associated with food limitation alone are not sufficient to explain the reduction in pit size of food limited antlions of this species.     

Modeling the succession of barnacles and mussels on a sandstone reef in Pernambuco State, Brazil

Settlement and recruitment rates, biological interactions, environmental parameters, and larval supply are factors that influence community structure in areas where barnacles and mussels are the dominant fauna. This study modeled the succession populations of two barnacle species, Tetraclita stalactifera (Lamarck, 1818) and Chthamalus bisinuatus (Pilsbry, 1916), and a bivalve, Brachidontes solisianus (d’Orbigny, 1846), on a sandstone reef at Piedade Beach, Pernambuco State, Brazil. The different stages of the life cycles of these invertebrates are represented by adult and immature forms in this model and are simulated as numbers of individuals per 100 cm². Recruitment rates are represented as forcing functions based on field recruitment experiments with time-steps of 1 day. In addition to intraspecific competition, the model takes into account interspecific competition based on reciprocal effects and carrying capacities. Field data from field succession experiments were used to calibrate the model. Recruitment and succession data were collected from random 10 × 10-cm plots within a 1-m transect on a Brachidontes bed from April to December 2007. Correlation coefficients between field data and simulation results were statistically significant for both populations; sensitivity analysis showed that increases and decreases in invertebrate population densities were always smaller than 10% compared to baseline model values. Patch closure by mussels population growth was one of the main succession processes observed during the rainy season. However, barnacles were the dominant species at the end of both field and simulation succession experiments, with an average density of close to 100 individuals per 100 cm². Cirripedia individuals may inhibit bivalve re-colonization by settling over shells and more efficiently occupying free space in addition to having higher recruitment rates than bivalves during the dry season.     

Effect of nematode Panagrellus redivivus density on growth, survival, feed consumption and carcass composition of bighead carp Aristichthys nobilis (Richardson) larvae

The study aimed to determine the optimum density of free‐living nematodes in feeding bighead carp, Aristichthys nobilis, larvae. In the first experiment, carp stocked at 25 larvae L^?1 were fed varying levels of nematodes (50, 75, 100, 125 and 150 per ml) twice a day for 21 days from the start of exogenous feeding. Final body weight was significantly higher (P < 0.05) in larvae fed 125 and 150 nematodes per ml than in those fed 50 and 75 per ml, but survival was low (61.8 and 63.6%, respectively). Survival rate was highest in larvae fed 100 nematodes ml^?1 (81.3%). Carcass analysis showed that larvae fed 125 and 150 nematodes ml^?1 had significantly lower body protein and higher body lipid than those fed other nematode densities. Carcass ash was similar for larvae fed 50–100 nematodes ml^?1 but it decreased significantly at the higher nematode densities. Carp larvae in a subsequent experiment were given 50, 75 and 100 nematodes ml^?1 per feeding. Newly hatched Artemia was the control feed. Nematode consumption and growth of the larvae were determined. Larvae were sampled at intervals of 2–4 days and the nematodes in the gut were counted and measured. At each nematode density, the number of nematodes present in the gut of the larvae increased significantly with time. At each sampling day, the number of nematodes in the gut did not differ significantly among treatments (P > 0.05) although it tended to increase with nematode density at day 2 and day 4 but decrease at day 7 onward. The carp larvae consumed significantly shorter nematodes on day 2 and day 4 than on the succeeding sampling days regardless of nematode density. However, the length of nematodes in the gut of the larvae did not differ significantly among the nematode densities. The final body weight of larvae increased with increasing nematode density. The body weight of larvae fed 100 nematodes ml^?1 did not differ significantly from that of larvae given Artemia nauplii. Results show that bighead carp larvae should be fed 100 free‐living nematodes per ml at each feeding time.     

Effects of Infection Density on the Size of Barley and Wheat Leaf Rust Colonies before and on the Size of Uredia after the Start of Sporulation

Three barley genotypes were exposed to four different inoculum densities of barley leaf rust, Puccinia hordei. Colony area well before first sporulation and uredia size well after the start of sporulation were measured. In a second series two barley and two wheat cultivars were exposed to four different inoculum densities of barley and wheat leaf rust (P. recondita f. sp. tritici), respectively. Before the sporulation is initiated the colony size was independent of the uredia density. This was valid for densities ranging from approximately 7 to over 200 uredia per cm² leaf area. After the sporulation had started the uredia size was strongly dependent on the uredia density. The size of the uredia was approximately halved when the uredia density increased from about 10 to about 150 per cm². The urediospore production per uredium decreased much stronger with increased uredia density.     

Horizontal distribution and abundance of cysts of several large branchiopods in temporary pool and ditch sediments

Cysts of three species of anostracans (Branchipusschaefferi, Chirocephalus diaphanus, and Tanymastigites perrieri), two species of Notostracans (Triops numidicus and Lepidurus apus) and one species of Spinicaudata (Leptestheriamayeti) were collected from the upper 2 cm of dry sediments at intervals along transects from a temporary pool in an arid zone of Morocco and from two ditches in France. The horizontal distribution of viable cysts was patchy and is discussed in relation to several ecological parameters prevailing during the aquatic phase. These include vegetation distribution, depth and morphometry of the pond, dominant wind pattern, and granularity of the sediments. The densities reached more than 1700 cysts of C. diaphanus per 100 cm², and 1150 cysts of L. apus per 100 cm² in the two ditches. The mean densities of cysts per 100 cm² calculated for each transect ranged from 250 to 450 for C. diaphanous and 240 to 260 for L. apus (in the ditch populations) and about 1–6 for T. perrieri and L. mayeti, 3–15 for B. schaefferi, and16–58 for T. numidicus in the pool in Morocco. This revised version was published online in August 2006 with corrections to the Cover Date.     

Evolutionarily Stable Relocation Strategy in an Antlion Larva

Antlion pits are often spatially aggregated even though there are potential negative effects associated with the aggregation (e.g., heightened competition and predation risk). This study investigated the possibility that a strategy leading to aggregation can be an evolutionarily stable strategy (ESS). In particular, the strategy considered was ‘decreasing relocation tendency when there are neighbors’. An individual based model showed that the strategy can be the unique ESS when the spatial distribution of prey is not completely random and antlions can learn it from their past foraging experiences. A laboratory experiment was conducted to examine the effects of the presence of neighbors and foraging success on the relocation behavior of antlion larvae. Antlions reduced their relocation tendency with respect to these factors, consistent with the predicted ESS. The results suggest that pit aggregations are formed because antlions reduce their relocation tendency when neighbors exist, and this strategy is an ESS.     

Ovipositional response ofVenturia canescens (Grav.) (Hymenoptera: Ichneumonidae) to various host and parasite densities

Summary Ovipositional response inVenturia canescens (Grav.) parasitizingAnagasta kühniella (Zeller) is analyzed for all combinations of five parasite densities (2, 5, 10, 25, and 50), four host densities (25, 50, 100, and 200), and three floor area universe sizes (76 cm², 160 cm², and 345 cm²). Over this range, no mutual interference (as measured by decreasing parasite fecundity) was observed, althoughNicholson’s area of discovery decreased significantly with parasite density increase as a result of a non-Poisson distribution of parasite eggs; use of this parameter is therefore not a suitable means of determining if mutual interference is present. Other parameters studied, and for which significant correlations were obtained, include mean parasite eggs per host, percentage parasitization, percentage of parasite eggs wasted, and negative binomialk approximations. This study was conducted as part of a broad investigation into the processes operating in the dynamics of arthropod populations under a grant toC. B. Huffaker by the U.S. Public Health Service, National Institutes of Health.     

Intertidal meiofauna of a high-latitude glacial Arctic fjord (Kongsfjorden, Svalbard) with emphasis on the structure of free-living nematode communities

Meiofauna communities of four intertidal sites, two sheltered and two more exposed, in Kongsfjorden (Svalbard) were investigated in summer 2001 at two different tidal levels (i.e. the low-water line and close below the driftline, referred to as mid-water (MW) level). A total of seven meiofaunal higher taxa were recorded with nematodes, oligochaetes and turbellarians being numerically dominant. Mean total meiofaunal densities ranged between 50 ind. 10 cm⁻² and 903 ind. 10 cm⁻². Our data showed a clear decrease in total meiofaunal densities with increasing coarseness of the sediment. Total meiofaunal biomass varied from 0.2 g dwt m⁻² to 2 g dwt m⁻² and, in general, was high even at low meiofaunal densities, i.e. larger interstitial spaces in coarser sediments supported larger meiofauna, especially turbellarians. The results on the vertical distribution of meiofauna contrasted sharply with typical meiobenthic depth profiles on other beaches, probably in response to ice-scouring and concomitant salinity fluctuations. Oligochaetes were the most abundant taxon, with a peak density of 641 ind. 10 cm⁻² at Breoyane Island. They were mainly comprised of juvenile Enchytraeidae, which prohibited identification to species/genus level. Nematode densities ranged between 4 ind. 10 cm⁻² and 327 ind. 10 cm⁻². Nematodes were identified up to genus level and assigned to trophic guilds. In total, 28 nematode genera were identified. Oncholaimus and Theristus were the most abundant genera. The composition of the nematode community and a dominance of predators and deposit feeders were in agreement with results from other arctic and temperate beaches. Nematode genus diversity was higher at the more sheltered beaches than at the more exposed ones. Low-water level stations also tended to harbour a more diverse nematode communities than stations at the MW level. Differences in nematode community structure between low- and MW stations of single beaches were more pronounced than community differences between different beaches and were mainly related to resources quality and availability.     

Spatial and temporal distribution of Antarctic fur seals (Arctocephalus gazella) on the breeding grounds at Bird Island,South Georgia

Summary The number of Antarctic fur seals (Arctocephalus gazella) in the area of the Scotia Sea has been increasing. Observation of their distribution on the breeding grounds are important to help design and interpret censuses. These seals are highly polygynous. In the study area at Bird Island, South Georgia, females give birth and are mated in densely packed colonies located along the shore close to the tideline. Males establish territories in this area, but are also found further inland. This study examined the density-dependent processes regulating the instantaneous size of the male breeding population; the distribution of males in relation to space and the number of females available to be mated and the effect of gregarious behaviour of females on male dispersion. Males were only territorial on the beaches in the areas where most females gave birth and subsequently had their post-partum oestrus. There was an apparent lower (19–20 m²) and upper (40 m²) limit to territory size. Males were excluded from the beach areas when the average density on the beaches was greater than 5 males per 200 m². An asymptotic density of 10–11 males per 200 m² was reached on the beaches and 4–5 males per 200 m² elsewhere. These two asymptotic densities may represent the upper and lower limits of density for a territorial system of dispersion. A model of the temporal changes in female numbers suggested that the total number of females occupying an area of beach during the mating period was approximately twice the number at the peak of the season. There was no indication that males compensated for low female density by increasing territory size. Females and pups became more dispersed after the end of the mating period. It is suggested that one function of gregariousness in females is as a mechanism for mate selection. This study also has implications for methods used to measure population size.     

The influence of larval density, food availability and habitat longevity on the life history and population growth rate of the midge Chironomus riparius

Despite long‐standing interest in the forms and mechanisms of density dependence, these are still imperfectly understood. However, in a constant environment an increase in density must reduce per capita resource availability, which in turn leads to reduced survival, fecundity and somatic growth rate. Here we report two population experiments examining the density dependent responses under controlled conditions of an important indicator species, Chironomus riparius. The first experiment was run for 35 weeks and was started at low density with replicate populations being fed three different rations. Increased ration reduced generation time and increased population growth rate (pgr) but had no effect on survival, fecundity and female body weight in the first generation. In the second generation there was a six‐fold increase in generation time, presumably due to the greatly reduced per capita resource availability as the estimated initial densities of the second generation were 300 times greater than the first. Juvenile survival to emergence, fecundity, adult body weight and pgr declined by 90%, 75%, 35% and 99%, respectively. These large between‐generation effects may have obscured the effects of the threefold variation in ration, as only survival to emergence significantly increased with ration in the second generation. These results suggest that some chironomid larvae survive a reduction in resource availability by growing more slowly. In the ephemeral habitats sometimes occupied by C. riparius, the effects of population density may depend crucially on the longevity of the environment. A second experiment was therefore performed to measure pgr from six different starting densities over an eight‐week period. The relationship between pgr and density was concave, viewed from above. At densities above 16 larvae per cm², less than 1% of the population emerged and no offspring were produced. Under the conditions of experiment 2 – an 8‐week habitat lifespan – carrying capacity was estimated as 8 larvae per cm².     

Supercolonies of the invasive yellow crazy ant, Anoplolepis gracilipes, on an oceanic island: Forager activity patterns, density and biomass

Summary. Ants have the capacity to reach unusually high densities, mostly in their introduced ranges. Numerical dominance is often cited as key to the ability of exotic ants to displace native ant species, reduce the abundance of invertebrates and negatively impact upon bird, land crab and other vertebrate populations. On Christmas Island, Indian Ocean, the yellow crazy ant, Anoplolepis gracilipes (Jerdon), forms supercolonies, where extremely high densities of foraging ants have contributed to ‘invasional meltdown’ in rainforest areas. Densities of up to 2254 foraging ants per m² and a biomass of 1.85 g per m² were recorded, and nest densities reached 10.5 nest entrances per m². Populations of A. gracilipes can overcome and kill red endemic land crabs (Gecarcoidea natalis) over 100 times their own biomass. This is the highest recorded density of foraging ants, and adds another element to the definition of ‘supercolony’ of unicolonial ants. This paper documents one extreme in a continuum of densities of unicolonial, invasive ant species and highlights the need to incorporate forager densities into invasive ant research.Received 17 November 2004; revised 14 February 2005, accepted 21 February 2005.     

Recovery of the coral Montastrea annularis in the Florida Keys after the 1987 Caribbean “bleaching event”

Many reef-building corals and other cnidarians lost photosynthetic pigments and symbiotic algae (zooxanthellae) during the coral bleaching event in the Caribbean in 1987. The Florida Reef Tract included some of the first documented cases, with widespread bleaching of the massive coral Montastrea annularis beginning in late August. Phototransects at Carysfort Reef showed discoloration of >90% of colonies of this species in March 1988 compared to 0% in July 1986; however no mortality was observed between 1986 and 1988. Samples of corals collected in February and June 1988 had zooxanthellae densities ranging from 0.1 in the most lightly colored corals, to 1.6x10⁶ cells/cm² in the darker corals. Minimum densities increased to 0.5x10⁶ cells/cm² by August 1989. Chlorophyll-a content of zooxanthellae and zooxanthellar mitotic indices were significantly higher in corals with lower densities of zooxanthellae, suggesting that zooxanthellar at low densities may be more nutrientsufficient than those in unbleached corals. Ash-free dry weight of coral tissue was positively correlated with zooxanthellae density at all sample times and was significantly lower in June 1988 compared to August 1989. Proteins and lipids per cm² were significantly higher in August 1989 than in February or June, 1988. Although recovery of zooxanthellae density and coral pigmentation to normal levels may occur in less than one year, regrowth of tissue biomass and energy stores lost during the period of low symbiont densities may take significantly longer.