Ultraviolet photosensitivity and feeding in larval and juvenile coral reef fishes

The ability of young coral reef fishes to feed using solely ultraviolet-A (UV-A) radiation during ontogeny was examined using natural prey in experimental tanks. Larvae and juveniles of three coral reef fish species (Pomacentrus amboinensis, Premnas biaculeatus and Apogon compressus) are able to feed successfully using UV-A radiation alone during the later half of the pelagic larval phase. The minimum UV radiation intensities required for larval feeding occur in the field down to depths of 90–130 m in oceanic waters and 15–20 m in turbid inshore waters. There was no abrupt change in UV sensitivity after settlement, indicating that UV photosensitivity may continue to play a significant role in benthic juveniles on coral reefs. Tests of UV sensitivity in the field using light traps indicate that larval and juvenile stages of 16 coral reef fish families are able to detect and respond photopositively to UV wavelengths. These include representatives from families that are unlikely to possess UV sensitivity as adults due to the UV transmission characteristics of the ocular media. Functional UV sensitivity may be more widespread in young coral reef fishes than in the adults, and may play a significant role in detecting zooplanktonic prey.     

Regional differences in duration of the planktonic larval stage of reef fishes in the eastern Pacific Ocean

Regional variation in the duration of the planktonic larval phase of three species of reef fishes, Thalassoma lucasanum (Labridae), Stegastes flavilatus, and Microspathodon dorsalis (Pomacentridae) was investigated between 1982 and 1991 at several sites in the tropical eastern Pacific over a distance of 3500 km, encompassing virtually their entire range of distribution. Durations of the larval phase, determined from counts of daily otolith increments, were significantly different (1.3 to 1.6 x) between sites. Populations of all three species had a consistently shorter larval life at the most northern site, Cabo San Lucas (Mexico) compared to Panamá and the offshore islands of Galápagos and Cocos. Analyses of otolith increment width over the precompetent period revealed that this disparity in larval duration primarily reflected differences in larval growth rates: faster growing fish spent less time in the plankton. In T. lucasanum, some of the variation in larval duration between Panamá and offshore sites (Galápagos Islands and Cocos Island) may be accounted for by a higher frequency of individuals delaying metamorphosis at the offshore sites. These data indicate that conditions in the planktonic environment are not homogeneous throughout the tropical eastern Pacific and may have a profound effect on aspects of the larval ecology of reef fishes in this region.     

Interactions during feeding among certain coral reef fishes in Elat

From July 1983 to March 1984 coral reef fishes in Elat (Red Sea) were fed in novel feeding situations. Thalassoma rüppelli had the shortest latency to first feed, Chaetodon paucifasciatus had the longest latency, while Sufflamen albicaudatus, Coris aygula and Lethrinus sp. had intermediate latencies. The mean number of T. rüppelli feeding at the beginning of experiments was higher than that of the other species. The difference decreased rapidly and disappeared within 90 s of the beginning. Latencies to first feed, and the number of fish feeding, were not correlated with the number of fish of each species in the study area. Aggression was directed predominantly by S. albicaudatus and by C. aygula towards Lethrinus sp. and T. rüppelli. It is suggested that these species which suffer more disturbances during their feeding and which receive more aggression are more likely to approach and feed more quickly in novel feeding situations.     

Reproductive isolation in temperate reef fishes

Opportunity and recognition isolation can lead directly to reproductive isolation, the former via divergence in the location and timing of breeding, and the latter via differential mate preferences. We describe the potential significance of these factors in the maintenance of reproductive isolation in a clade of triplefin fishes that occur sympatrically around coastal New Zealand. Specifically, we investigate the roles of spawning time and nesting habitat in promoting opportunity isolation, and of interspecific variation in male body length and breeding colouration in promoting recognition isolation. The triplefin species investigated are reproductively active over several months and show high overlap in breeding times, thus rejecting temporal isolation as a mechanism. Differences in nesting habitats resulted in a reduced probability of encounter between some species, especially between sister-species pairs. Interspecific colour differences generally decreased during the reproductive period, and males of sister-species pairs showed no interspecific colour differences in the ultraviolet light spectrum, thus mate selection based on male colour patterns is unlikely to lead to premating isolation. Finally, males of closely related triplefin species differed in body length, a secondary sexual trait often involved in assortative mating. Thus, spatial differences in nesting habitats reduce the chances of encountering allospecific mates, which may facilitate opportunity isolation and differences in male length, possibly related to species-specific female selection on male body size, may lead to recognition isolation. The combination of limited spatial overlap in nesting habitat and differences in male body size may facilitate species assortative mating in sympatry or parapatry.     

Demographic plasticity in tropical reef fishes

We use age-based analyses to demonstrate consistent differences in growth, mortality, and longevity of coral reef fishes from similar habitats (exposed reef crests) 20 km apart. On outer-shelf reef crests of the northern Great Barrier Reef (GBR), size in four taxa of reef fishes (Chlorurus sordidus, Scarus frenatus, and S. niger and the acanthurid Acanthurus lineatus) was systematically and significantly smaller when compared with the same taxa on adjacent mid-shelf reef crests. Differences in size could be attributed to differences in growth between habitats (shelf positions). On outer reef crests the species examined had consistently lower size at age profiles and also reduced life spans compared with populations from mid-shelf reefs. To confirm this relationship, two of the most abundant species (C. sordidus and S. frenatus) were selected for more detailed spatial analysis of demographic patterns. Sampling adults of both taxa from reef crests on three mid- and three outer-shelf reefs revealed that most of the variation in growth was explained by shelf position, although C. sordidus also displayed differences in growth among mid-shelf reefs. We conclude that differences in body sizes across the continental shelf of the northern GBR are primarily determined by these trends in growth. Strong spatial patterns also existed in the mean ages of populations and longevity estimates for C. sordidus and S. frenatus between shelf positions. Both species on outer-shelf reefs displayed less variable cohort sizes, significantly reduced mean ages, and foreshortened longevity compared with populations on mid-shelf reefs. Furthermore, differences in these parameters were rare among replicate reefs within mid- and outer-continental-shelf positions. Age-based catch curves suggested that rates of S. frenatus natural mortality on the outer shelf were nearly twice as high as on the mid shelf. Visual surveys indicated that total scarid densities on outer-shelf reef crests are on average fourfold higher than for equivalent mid-shelf habitats. This fact, coupled with significantly reduced growth rates, reduced mean ages, and increased mortality rates, suggests that density-dependent processes may be responsible for observed differences among localities.     

The relationship between population genetic structure and pelagic larval duration in coral reef fishes on the Great Barrier Reef

Pelagic larval duration (PLD) is a commonly used proxy for dispersal potential in coral reef fishes. Here we examine the relationship between PLD, genetic structure and genetic variability in geographically widespread and ecological generalist species from one coral reef fish family (Pomacentridae) that differs in mean larval duration by more than a month. The genetic structure was estimated in eight species using a mitochondrial molecular marker (D-loop) and in a sub-set of five species using nuclear molecular markers (ISSRs). Estimates of genetic differentiation were similar among species with pelagic larvae, but differed between molecular markers. The mtDNA indicated no structure in all species except one, while the ISSR indicated some structure between the sampling locations in all species. We detected a relationship between PLD and genetic structure using both markers. These relationships, however, were caused by a single species, Acanthochromis polyacanthus, which differs from all the other species examined here in lacking a larval phase. With this species excluded, there was no relationship between PLD and genetic structure using either marker despite a range of PLDs of more than 20 days. Genetic diversities were generally high in all species and did not differ significantly among species and locations. Nucleotide diversity and total heterozygosity were negatively related to maximum PLD but again these relationships were caused by A. polyacanthus and disappeared when this species was excluded. These genetic patterns are consistent with moderate gene flow among well-connected locations and indicate that at this phylogenetic level (i.e., within family) the duration of the pelagic larval phase is unrelated to the patterns of genetic differentiation.     

Extremes, plasticity, and invariance in vertebrate life history traits: insights from coral reef fishes

Life history theory predicts a range of directional generic responses in life history traits with increasing organism size. Among these are the relationships between size and longevity, mortality, growth rate, timing of maturity, and lifetime reproductive output. Spanning three orders of magnitude in size, coral reef fishes provide an ecologically diverse and species-rich vertebrate assemblage in which to test these generic responses. Here we examined these relationships by quantifying the life cycles of three miniature species of coral reef fish from the genus Eviota (Gobiidae) and compared their life history characteristics with other reef fish species. We found that all three species of Eviota have life spans of < 100 days, suffer high daily mortality rates of 7-8%, exhibit rapid linear growth, and matured at an earlier than expected size. Although lifetime reproductive output was low, consistent with their small body sizes, short generation times of 47-74 days help overcome low individual fecundity and appear to be a critical feature in maintaining Eviota populations. Comparisons with other coral reef fish species showed that Eviota species live on the evolutionary margins of life history possibilities for vertebrate animals. This addition of demographic information on these smallest size classes of coral reef fishes greatly extends our knowledge to encompass the full size spectrum and highlights the potential for coral reef fishes to contribute to vertebrate life history studies.     

Comparative study of metamorphosis in tropical reef fishes

This study explores the types of changes in pigmentation and morphology that occur immediately after settlement in 13 families of tropical reef fishes encompassing 34 species. The morphology of individual fishes was recorded daily from when they were first caught at night as they came into the vicinity of a reef to settle. Changes in pigmentation and morphology were species specific and often varied greatly among species within a family or genus. Pigmentation changes were typically rapid (<36 h) and dramatic. Morphological changes involved the elongation and regression of fin spines and changes in head shape and body depth. Eighteen percent of species experienced changes in snout shape and dorsal spine length of greater than 5%. Similarly, 15% experienced changes in pectoral fin length and head length of greater than 5%. Changes typically occurred gradually over 6 or more days, although in about 44% of the species the major change in one of the measured body dimensions occurred rapidly (within 36 h). Moderately strong positive relationships were found between both growth and developmental rates and the extent of metamorphosis in the damselfishes (Pomacentridae) (r=0.48 and 0.63, respectively). This suggests there may be a minimum level of development necessary to be a fully functional demersal juvenile. Although many of the changes that occur are subtle compared to the preceding development, these changes occur at an important ecological transition. Published online: 16 August 2002     

Microstructure of settlement-marks in the otoliths of tropical reef fishes

The morphology and ultrastructure of the otolith settlement-mark was examined in 44 tropical reef-fish species spanning nine families. A classification scheme based on similar otolith characteristics is presented. Three major categories are identified based on changes in increment width and optical qualities of the settlement-mark. Of the 44 species examined, 39 possessed “abrupt” settlement-marks (Type I) characterised by a rapid decrease in increment width (up to 50% reduction) over settlement. Type I settlement-marks were found in all nine families examined. The 39 species spanned the whole range of possible larval durations (Pomacentrus moluccensis, 15 d ± 0 SE; Naso hexacanthus, 91.2 d ± 2.97 SE). Four of the 44 species possessed “zonal” settlement-marks (Type II), featuring a band of increments that are wider than pre-settlement increments. Species in this category are the labrids Corisaygula, Thalassoma bifasciatum, T. lunare and an unidentified acanthurid (Acanthurus sp. 2). One species of acanthurid (N. brevirostris) possessed a “gradual” settlement-mark (Type III), manifest as a gradual decrease in increment width during the settlement period. A possible fourth type was identified from the literature. Gnatholepis thompsoni and Coryphopterus glaucofraenum possessed a settlement-mark with increment widths that increased post-settlement. Available data suggest a poor relationship between the structure of the settlement-mark and the magnitude of metamorphosis (previously reported as internal and external morphological change). Evidence suggests that the increment profile over early development and the increment transitions associated with the settlement event are taxon-specific and may enable late-larval stage fishes to be identified to species level. Received: 21 May 1997 / Accepted: 3 February 1999     

Climate-induced reaction norms for life-history traits in pythons

Climate change modelers predict increasingly frequent "extreme events," so it is critical to quantify whether organismal responses (such as reproductive output) measured over the range of usual climatic conditions can predict responses under more extreme conditions. In a 20-year field study on water pythons (Liasis fuscus), we quantified the effects of climatically driven annual variation in food supply on demographic traits of female pythons (feeding rate, body size, body mass, and reproductive output). Reaction norms linking food supply to feeding rates and residual body mass were broadly linear, whereas norms linking food supply to female body size became curvilinear when a dramatic (flooding-induced) famine reduced the mean body size at sexual maturity. Thus, the reaction norms recorded over 16 years of mal" (albeit highly variable) climatic conditions gave little insight into the population's response to a more extreme nutritional crisis.     

Annual fluctuation of life-history traits in the mud crab Macrophthalmus japonicus

Interannual fluctuation of life-history traits in the mud crab Macrophthalmus japonicus (De Haan) was studied for a 7-yr period (1981 to 1987) in an intertidal mudflat in the estuary of the Tatara-Umi River, Fukuoka, Japan. Density of juveniles fluctuated greatly, and maximum annual density was low in 1981 to 1984 (22.0 to 44.0 m^-2) and very high in 1985 to 1987 (98.5 to 188.6 m^-2). In 1980, recruits may have been very few, though there is no data on or before 1980, because no crab of the 1980 cohort was found during the study period. Density of adult crabs was almost stable (11.0 to 32.6 m^-2), perhaps because of density-dependent mortality. Under the high density conditions prevailing from 1985 to 1987 crabs grew slowly and longevity was shorter. Male chela and female abdomen were dwarfish, and females produced few and small broods. The changes of life-history traits under high density conditions were thought to result from strong intraspecific competition for space.     

Video-monitored predation by Caribbean reef fishes on an array of mangrove and reef sponges

Although predation by fishes is thought to structure benthic invertebrate communities on coral reefs, evidence to support this claim has been difficult to obtain. We deployed an array of eight sponge species on Conch Reef (16 m depth) off Key Largo, Florida, USA, and used a remote video-camera to record fish activity near the array continuously during five daylight periods (6 h for 1 d, at least 11.5 h for 4 d) and one night period (11 h). Of the eight sponge species, four were from adjacent reefs (Agelas wiedenmayeri, Geodia neptuni, Aplysina fistularis, and Pseudaxinella lunaecharta), and four were from a nearby mangrove habitat (Chondrosia collectrix, Geodia gibberosa, Halichondria sp., andTedania ignis). Each species of reef sponge was chosen to match the corresponding mangrove species in form and color (black, brown, yellow, and red, respectively). Predation events only occurred during daylight hours. Tallies of the number of times fishes bit sponges revealed intense feeding by the expected species of sponge-eating fishes, such as the angelfishHolacanthus bermudensis, H. tricolor, andPomacanthus arcuatus, the cowfishLactophrys quadricornis, and the filefishCantherhines pullus, but surprisingly also by the parrotfishSparisoma aurofrenatum andS. chrysopterum. Of 35 301 bites recorded, 50.8% were taken by angelfish, 34.8% by parrotfish, and 13.7% by trunkfish and filefish. Mangrove sponges were preferred by all reef fishes; 96% of bites were taken from mangrove species, with angelfish preferringChondrosia collectrix and parrotfish preferringGeodia gibberosa. Fishes often bit the same sponge repetitively, and frequently consumed entire samples within 30 min of their deployment. Sponge color did not influence fish feeding. Two of the four mangrove sponge-species deployed on the array were also found living in cryptic habitats on adjacent reefs and were rapidly consumed by fishes when exposed. Our results demonstrate the importance of fish predation in controlling the distribution of sponges on Caribbean reefs.     

Biotic passage through the Panama Canal,with particular reference to fishes

The effectiveness of the Gatun Lake fresh-water barrier to fish migration of the Panama Canal was examined, based on literature records and recent collections. Six species have migrated to the Pacific and three to the Atlantic Oceans. New records include: Hypleurochilus aequipinnis, Barbulifer ceuthoecus and Oostethus lineatus of Atlantic origin, and Gnathanodon, speciosus of Pacific origin. The majority of Atlantic migrants are known in the Pacific only from the Miraflores Third Lock, a unique ectogenic meromictic lake attached to the Pacific entrance of the canal. The hydrography and biota (including Atlantic algal and crustacean species previously unknown as canal migrants) of the lake are discussed. It is presumed that certain fish migrants transited the canal by associating with fouling material on the underside of ships. The euryhaline species, H. aequipinnis, Lupinoblennius dispar, Lophogobius cyprinoides and Omobranchus punctatus survived in freshwater (0.0‰S) for periods longer than réquired for ship transit of the canal (ca. 8h). The stenohaline migrant Gobiosoma nudum died after 2 h in freshwater, but survived more than 50 h at 2.5‰S. Plans to increase Panama Canal ship transits through the pumping of seawater into Gatún Lake might remove the biological barrier and allow the migration of euryhaline and stenohaline species. We find those plans unwise, and the potential consequences dangerous.     

Differential movement patterns and site fidelity among trophic groups of reef fishes in a Hawaiian marine protected area

We tracked the long-term movements of 70 parrotfishes, surgeonfishes and goatfishes captured inside a small (1.3 km²) marine protected area (MPA: Kealakekua Bay Marine Life Conservation District, Hawaii) by implanting them with small transmitters and deploying underwater monitoring devices inside the bay and along 100 km of the adjacent west Hawaii coastline. Individual fish were detected inside Kealakekua Bay for up to 612 days but many were detected for much shorter periods (median = 52 days). There were species-specific differences in the scale of movements and habitats used, but most fish utilized between 0.2 and 1.6 km of coastline, and individuals of each species showed some degree of diel habitat shift. A wide variety of reef fishes captured inside the MPA swam back and forth across an MPA boundary intersecting continuous reef (i.e., this boundary was porous to reef fish movements), but only 1 of 11 species tagged crossed a wide sandy channel inside Kealakekua Bay suggesting that this feature may function as a natural barrier to movements. Results indicate relatively small MPAs (<2 km of coastline) could provide effective, long-term protection for multi-species assemblages of reef fishes provided that boundaries are situated along major habitat breaks (e.g., large sand channels between reefs) that may serve as natural barriers to reef fish movements. It is crucial that a multi-species approach be used when assessing MPA effectiveness.     

Diel patterns of abundance of presettlement reef fishes and pelagic larvae on a coral reef

Most presettlement reef fish settled at night at One Tree Island, Great Barrier Reef. Fish were sampled day and night using channel nets located on the reef crest, and a plankton-mesh purse-seine net in the lagoon (1992–1994). Catches of fish at night were generally tens to hundreds of times greater than those taken during the day. Preflexion fish, as well as postflexion and pelagic juveniles, were taken in greater numbers at night. Preflexion forms were a combination of those that had hatched from demersal eggs and later stages that had been transported over the reef crest. Highest numbers of postflexion and pelagic juvenile forms of Apogonidae, Blenniidae, Gobiesocidae, Gobiidae, Labridae, Lutjanidae, Mugiloididae, Mullidae, Pomacentridae, Scaridae, Serranidae and Tripterygiidae were found at night. Observations, while SCUBA diving, and purse-seine samples in the lagoon indicated that the only resident larvae were of the genera Spratelloides and Hypoatherina; most of the fishes caught in nets, therefore, were immigrants. Patch reefs, sampled for new settlers early in the morning and late in the day, indicated that the majority of apogonids (Apogon doederleini, >95%) settled at night. Although greater numbers of pomacentrids were found in morning counts (e.g. Pomacentrus wardi), if data were converted to an hourly rate, many pomacentrids showed a similar hourly rate of settlement day and night. Depth-stratified sampling in waters near One Tree Island (to 20 m) indicated that some taxa rise to the surface at night. This behaviour, perhaps combined with avoidance of diurnal predators may explain on-reef movement of potential settlers soon after dark. Studies on settlement cues, therefore, need to focus on night-related phenomena. Received: 3 March 2000 / Accepted: 20 June 2000     

Variability in the chemical defense of the sponge Chondrilla nucula against predatory reef fishes

Chondrilla nucula is a common Caribbean demosponge that grows in a range of habitats, from coral reefs to mangrove swamps. On reefs, C. nucula grows as a thinly encrusting sheet, while in mangrove habitats it surrounds submerged mangrove roots as fleshy, lobate clumps. Previous feeding experiments using predatory reef fish revealed a high degree of variability in the chemical defenses of C. nucula. The present study was undertaken to determine whether a relationship exists between habitat, growth form, and chemical defense of C. nucula. Both laboratory and field feeding-assays of crude extracts confirmed that C. nucula possesses a chemical defense with high intercolony variability, but there was no significant variation in feeding deterrency between reef and mangrove habitats at either geographic location (Bahamas and Florida). Extracts of C. nucula collected during September and October 1994 from the Bahamas were significantly more deterrent than those collected during August 1993, May 1994, and May 1995 from Florida, and extracts of these spring and summer Florida collections were more deterrent than extracts of C. nucula collected in December 1994 and February 1995 in the same locations. There was no evidence that deterrent compounds were concentrated in the surface tissues of the sponge, or that chemical defense could be induced by simulated predation. Laboratory and field assays of the fractionated crude extract revealed that feeding deterrency was confined to the most polar metabolites in the extract. Field transplants were used to determine whether predation influenced the growth form of C. nucula. Uncaged sponges transplanted from the mangrove to the reef were readily consumed by spongivorous reef fishes. Lobate mangrove sponges became thinner after being caged on the reef for 3 mo, but encrusting reef sponges did not become thicker after being caged in the mangroves for the same period of time. Reef sponges that were caged for 3 to 15 mo thickened by only a small amount (<1 mm) compared to uncaged and open-caged (i.e. in cages lacking tops) sponges. Simulated bite marks on both reef and mangrove sponges were repaired at a rapid rate (0.8 to 1.6 mm d⁻¹). Fish predation has an important impact on the distribution and abundance of C. nucula, but the thin growth form common to reef environments may be more the result of hydrodynamics than of grazing by spongivorous fishes. Received: 6 October 1997 / Accepted: 19 March 1998     

Association of color and feeding deterrence by tropical reef fishes

Summary. While many marine molluscs have been suggested to use aposematic coloration to avoid predation, few studies have tested the ability of marine predators to learn to associate colors with distasteful prey. In field experiments, we tested the ability of two populations of reef fishes to discriminate among red, yellow, and black artificial nudibranch models when one color was paired with a feeding deterrent. We offered fishes (1) the models without any feeding deterrents, (2) the models with a feeding deterrent coated onto one color, and (3) the models without deterrents again. If reef fishes learn to associate colors with noxious prey, we expected the color paired with the feeding deterrent to be eaten less frequently in the final assay than the initial assay. In both populations, fishes formed clear associations between color and feeding deterrence. However, when the experiment was repeated in one population, changing the color paired with the feeding deterrent, fishes did not form an association between color and feeding deterrence. In this case, prior learning may have affected subsequent trials. Our study indicates that common colors of nudibranchs are recognizable by fishes and can be associated with noxious prey. Received 24 September 1998; accepted 18 December 1998.     

Marker technique for investigating gut throughput rates in coral reef fishes

Artemia sp. shells were evaluated to determine their accuracy for tracing the passage of algal filaments through the gut of the damselfish Pomacentrus amboinensis Bleeker, 1868 (family Pomacentridae), an omnivorous coral reef fish. An automatic faeces-collection apparatus enabled the quantitative collection of markers and faeces in the laboratory. Defecation rates were similar for light and heavy doses of Artemia sp. shells and controls, indicating no detrimental effects of Artemia sp. shells on the gut throughput rate of P. amboinensis. In addition, similar rates and patterns of the passage of Artemia sp. shells and the algal markers Enteromorpha sp. and Lyngbya sp. indicated that Artemia sp. shells provide a reliable representation of the throughput rate of algal filaments. The mean throughput time of P. amboinensis was 4.6 h ±0.3 SE, with a modal recovery time of 4 h. Laboratory throughput estimates were validated by comparing the distribution patterns of Artemia sp. shells in the dissected gut of specimens administered markers in the laboratory and field. In addition, the retention of markers in the stomach of P. amboinensis suggested a likely site of prolonged processing. Received: 24 October 1996 / Accepted: 18 March 1997     

Patterns of use of space in a guild of territorial reef fishes

There exists on Heron Reef, at the southern end of the Great Barrier Reef, an 8-species guild of ecologically very similar, territorial, herbivorous reef fishes. All individuals of these species maintain territories on rubble substrata throughout juvenile and adult life. Territories are defended from all other guild members. Three rubble patches, each containing residents of 3 guild species: Pomacentrus apicalis, P. wardi, and Abudafduf lachrymatus, have been monitored for 12 to 18 months. This paper examines the patterns of use of space as shown by the antecedent and subsequent histories of sites chosen by 43 new colonists, and sites vacated through the loss of 34 residents on the rubble patches. In addition, variation through time is examined in the total amount of space held in territories on each rubble patch. The 3 species show similar preferences for space as colonists, although adult and juvenile colonists behave differently. Numbers of colonists detected on rubble patches are not proportional to the resident populations of the 3 species. Residents of the 3 species are equal in their abilities to enter spaces vacated through mortality, although they differ slightly in methods used to enter sites. They do not respond preferentially to sites previously occupied by any particular species. P. wardi shows a higher rate of mortality than the other species, and residents of this species are more often dislodged by new colonists. The total amount of space held on any rubble patch did not vary during the year from September, 1972 to October, 1973. The data are interpreted as supporting the hypothesis that the 3 species do not differ in their requirements for space on rubble patches, and that they are in competition for a short supply of such space. Differences exist in their strategies for obtaining and holding such space. These differences are important for explaining the continued presence of P. wardi on rubble patches. P. apicalis and a. lachrymatus are both specialists in holding territories on the upper reef slope. No differences have been detected in their requirements or competitive abilities. P. wardi is a fugitive species on the upper reef slope, coexisting because it maintains a refuge from competition by occupying some un-preferred sites off rubble patches.