Phenotypic assortative mating and genetic population structure in the crab Trapezia digitalis

Colonies of Pocillopora spp. containing the crab Trapezia digitalis were collected from Hawaii in summer of 1979 and of 1980. Individuals of T. digitalis in Hawaii vary extensively in coloration of the carapace: crabs were separated into phenotypic classes by their degree of carapace reticulation. When presented with two females of different phenotype, males preferentially paired with the female which was similar to them in reticulation. The genetic population structure of T. digitalis was unusual by comparison with other species of Trapezia. Previously reported homozygote excesses at esterase-4, hexokinase-1, phosphoglucoseisomerase, and general protein-2 cannot be explained by the single-locus Wahlund effect, since there was not significant genetic differentiation among phenotypic classes, as measured by Wright's F _ST statistic. There was significant variation among classes in allele frequencies at a fifth locus, hexokinase-2. Two pairs of loci were in significant gametic disequilibrium. Gametic disequilibrium was not detected in seven other species of Trapezia. It is suggested that multi-locus genetic analysis and an increased geographic sampling range are needed to determine the genetic population structure of T. digitalis in relation to observed variation in coloration.     

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.     

Biochemical genetic variation and evidence of speciation in Chthamalus barnacles of the tropical eastern Pacific Ocean

Gene-enzyme variation was studied electrophoretically within and between barnacle populations of the genus Chthamalus from 8 intertidal stations from central California to the Pacific coast of Panamá. Horizontal starch-gel electrophoresis separated and resolved 18 enzymes from 512 individual barnacles. A maximum of 25 gene-enzyme systems was interpretable from the resulting zymograms. Electrophoretic phenotypes and patterns of phenotypic variation generally conformed with those observed in other organisms. The amount of genetic variation within barnacle populations varied; average heterozygosity, for example, ranged from a low of 3.4% in a Mexican population of the C. fissus group to a high of 10.4% in C. anisopoma from the Gulf of California. Observed and expected average heterozygosities agreed in all population samples, indicating that these species are outbreeding. In contrast to the prediction stemming from a hypothesis that trophic stability regulates the amount of genetic variation in marine species, average heterozygosity tends to be positively correlated with latitude. Data from these and other barnacle species may support a hybrid environmental heterogeneity-trophic diversity model recently proposed to explain genetic variation in decapod crustaceans. Juxtaposition of individuals from different localities revealed numerous genetic differences among populations of the C. fissus group. At least three partially sympatric sibling species are separated by genetic distances as large as those observed between the C. fissus group and the distinct species C. dalli and C. anisopoma. A cladistic analysis places C. anisopoma close to the Mexican and Panamanian sibling species, with C. fissus from San Diego and C. dalli successively farther removed.     

Energetics,competency, and long-distance dispersal of planula larvae of the coral Pocillopora damicornis

Pocillopora damicornis (Linnaeus) were collected from Enewetak Atoll, Marshall Islands, in 1980–1981) and Kaneohe Bay, Hawaii, in 1982. Their planula larvae contained 17% protein, 70% lipid, and 13% carbohydrate by dry weight. Calculations based on stored energy reserves and daily metabolic expenditure indicate that planulae could survive approximately 100 d and still settle successfully. Competency experiments demonstrated that larvae settled and metamorphosed after 103 d. This period of time is sufficient to allow immigration of larvae from the Central Pacific to the eastern Pacific, and supports the hypothesis of long-distance dispersal of larvae for the origin of present eastern Pacific populations of P. damicornis.     

Genetic differentiation between populations of Talitrus saltator and Talorchestia deshayesii (Crustacea: Amphipoda) from coastal areas of the north-western European continent

Electrophoretic studies of gene-enzyme variation in the littoral talitrids Talitrus saltator (Montagu) and Talorchestia deshayesii (Audouin) were undertaken to estimate the amount of divergence among geographically separated populations. Samples of both species were taken from sandy beaches over a transect of approximately 3 500 km along the coast of the European continent including Baltic, North Sea and Atlantic locations. A total of 22 T. saltator and 15 T. deshayesii populations were analysed for genetic variation at various enzyme loci. Both amphipods revealed relatively low levels of polymorphism and heterozygosity. Among the loci studied, phosphoglucose isomerase (Pgi) and phosphoglucomutase (Pgm) were highly polymorphic. Patterns of micro- and macrogeographic variation in terms of distributions of allele frequencies at these particular loci are compared. Interpopulation allozymic variation was shown to be lower in T. deshayesii than in T. saltator. As demonstrated by T. saltator populations sampled in coastal sites ranging from Denmark to western France, clinal variation in frequencies of two alleles became evident at the PGI locus, exhibiting a steady increase in the level of polymorphism with decreasing latitudes. It is argued that limited gene flow and, to some extent, random genetic drift may account for the gene pool structure of the talitrid species investigated.     

Electrophoretic and biometric variability in the abyssal grenadier Coryphaenoides armatus of the western North Atlantic,eastern South Pacific and eastern North Pacific Oceans

Specimens of the abyssal grenadier Coryphaenoides armatus (Hector, 1875), from the western North Atlantic and eastern North Pacific Oceans were compared electrophoretically at 27 presumptive gene loci. At 6 of the 7 polymorphic loci there were only minor differences in allelic frequencies but a nearly fixed difference was found at one locus, phosphogluconate dehydrogenase. Eastern North Pacific grenadiers typically have a narrower interorbital space, a shorter dorsal interspace, more soft rays in the 1st dorsal fin (9–10 versus 8–9) and more pelvic fin rays (21–23 versus 18–21) than grenadiers from the western North Atlantic (as well as grenadiers from the eastern South Pacific, which were included in the biometric analysis). There is an apparent disjunction in the distribution of C. armatus in the eastern Pacific at the Gulf of Panamá which coincides with the change of morphology. It is suggested that North Pacific grenadiers comprise a subspecies, C. armatus variabilis Günther, 1878, which is morphologically distinct from the subspecies C. armatus armatus (Hector, 1875) of the other areas.     

Life-history variation in three coexisting cheilostome bryozoan species of the genus Stylopoma in Panamá

Life-history variation was studied in three sympatric species of Stylopoma Levinsen [S. spongites (Pallas), S. projecta Canu and Bassler and Stylopoma n. sp. 15] in Panamá. Bryozoan colonies were collected from 27 reefs along 300 km of the Caribbean coast of Panamá. The distribution and abundance of each species were very patchy, but with broad overlap in occurrence among localities and depths. Nevertheless, species differed considerably in colony size, size at first reproduction and numbers of brooding larvae; implying that interspecific differences in life-history traits may contribute to their coexistence. To examine closely this variation in life-history patterns, we grew, in a common garden experiment, the offspring of the two most common species that were obtained from parent colonies collected from several reefs. There were highly significant differences in growth rates and the timing and extent of sexual reproduction which corresponded well to patterns observed in individuals from the field. Other factors, including size of larvae, extent of secondary zooidal calcification and numbers of avicularia were also correlated with differences in life histories. Despite this additional complexity, however, ecological consequences of trade-offs in life history among modular species such as Stylopoma spp. appear very similar to those among unitary species.     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panamá, and Galápagos Islands (Ecuador). III. Agariciidae (Pavona gigantea and Gardineroseris planulata)

The reproductive ecology of Pavona gigantea Verrill and Gardineroseris planulata (Dana) was investigated in the equatorial eastern Pacific region from 1985 to 1994. These zooxanthellate scleractinian corals were adversely affected in this region during the 1982–1983 El Niño warming event. Both species were hermaphroditic, with individual colonies showing sequential cosexual development, thus resulting in dominantly outbreeding reproduction. Sexuality was mixed, with high percentages of gonochoric and hermaphroditic colonies in both species. Approximately 1:1 male-to-female gonad ratios were found in gonochoric and hermaphroditic colonies combined. Broadcast spawning was observed in P. gigantea in the Galápagos Islands, and the sudden disappearance of mature gametes and the presence of spent gonads suggest that G. planulata is also a broadcast spawner. Colonies of both species with 200 cm² (10 cm diam) live tissue were nonreproductive. Estimated ages of the youngest reproductive colonies were 11 yr for P. gigantea and 20 yr for G. planulata. The percentage of all colonies of P. gigantea with gonads at nonupwelling sites (Caño Istand, Costa Rica and Uva Island, Panamá) ranged from 37 to 47%, respectively; colonies with gonads from upwelling environments (Saboga and Taboga Islands, Panamá) ranged from 31 to 39%, respectively, and reproductively active colonies from the thermally variable Galápagos islands comprised 40% of the collections. Compared with P. gigantea, the numbers of sexually active G. planulata colonies were roughly onehalf at nonupwelling Caño Island (20%) and Uva Island (25%) sites, or less (10%) at the upwelling Saboga Island site. Peak reproductive activity in P. gigantea occurred during the rainy season at all study sites. In the nonupwelling Costa Rican (Caño Island) and Panamainan (Uva Island) sites, mean monthly sea-surface temperatures (SSTs) were high (28 to 29°C), but slightly lower than in the dry season (29°C). In the upwelling Gulf of Panamá (Saboga and Taboga Islands), reproduction occurred after mean monthly SSTs increased from 24 to 28–29°C. In the Galápagos Islands, reproductive activity peaked during sea warming, when mean monthly SSTs reached 25°C. Sexually active colonies of G. planulata, present only at the main collection sites of Caño and Uva Islands, were also observed during the wet season. The presence of mature or spawned gonalds in both species mostly around new and full lunar phases suggests that spawning is at least weakly synchronized with moon phase. Fecundity estimates disclosed the following nonsignificant differences between sites for P. gigantea, expressed as egg production cm^-2 colony surface surface yr^-1: Galápagos (10 300 to 30 800), Uva Island (4900 to 9800), Caño Island (1800 to 7400), Saboga Island (600 to 1300) Taboga Island (1200 to 2400). Fecundity estimates for G. planulata were considerably lower: Uva Island (700 to 1400), Caño Island (500 to 1000). The sexual recruitment of P. gigantea into El Niño-Southern Oscillation (ENSO) 1982–1983-disturbed, equatorial eastern Pacific coral communities has been low, with only moderate recovery evident since 1983. G. planulata has revealed no sexual recruitment where seed populations are absent or rare (Caño Island, Galápagos Islands), and only low recruitment (Panamá) in areas with colonies that survived the ENSO disturbance.     

Allozyme variation in global populations of striped mullet, Mugil cephalus (Pisces: Mugilidae)

The striped mullet, Mugil cephalus Linnaeus, 1758, is one of the few species of marine shore fish with a worldwide circumtropical distribution. Because of this distribution and the dependency of M. cephalus on coastal waters during various phases of its life cycle, as well as nearshore living habits, questions have been raised regarding levels of genetic divergence and gene flow among transoceanic populations. To cast more light on this, allozyme variation at 27 presumptive gene loci was investigated in ten globally diverse populations. The observed heterozygosity ranged from 0.018 (Hawaii) to 0.081 (Florida), averaging 0.050. The proportion of polymorphic loci showed a similar trend. Several populations were characterised by fixed allelic differences. Estimated gene diversities were very high, the allele frequency variation among populations was found to be 68%; genetic distances reached 0.242, with an average of 0.117. Estimated rates of gene flow were high among Mediterranean populations (Nm = 7.26), and between Mediterranean and East Atlantic populations (Nm= 2.86), but extremely low between non-contiguous populations within the Indian, Pacific and Atlantic Oceans, where Nm ranged from 0.03 to 0.05. Received: 7 April 1997 / Accepted: 4 February 1998     

Effects of temperature on photosynthesis and respiration in hermatypic corals

Photosynthesis and respiration rates of the reef corals Pocillopora damicornis (Linn.), Montipora verrucosa (Lamarck), Porites compressa Dana and Fungia scutaria Lamarck were measured under controlled temperatures. Results indicate that coral metabolism is closely adapted to ambient temperature conditions. Tropical corals measured at Enewetak, Marshall Islands, showed greater primary production compared to maintenance requirements at elevated temperatures than did subtropical varieties of the same species in Hawaii. Photosynthesis: respiration (P:R) ratios were significantly and negatively related with temperature between 18° and 31°C for all Hawaiian corals, whereas at Enewetak this ratio generally showed a curvilinear relationship for this temperature range. Extrapolations of P:R regressions on temperatures to a value of 2.0 (estimated as a minimum required for long-term functional autotrophy) coincide for Hawaiian specimens with published upper lethal temperatures. Extrapolation of P:R regressions for Enewetak specimens at temperatures above 25°C suggests lethal temperatures for these corals to be 2 to 5 C° higher than for Hawaiian corals, in good agreement with recent experimental findings. Interspecific differences in P:R temperature regressions for Hawaiian corals correlating with upper lethal temperature tolerances are described.Contribution No. 505 of the Hawaii Institute of Marine Biology.     

Close genetic similarity of Atlantic and Pacific skipjack tuna (Katsuwonus pelamis) demonstrated with restriction endonuclease analysis of mitochondrial DNA

Restriction endonuclease analysis of mitochondrial DNA indicated a surprisingly high degree of genetic similarity between skipjack tuna (Katsuwonus pelamis) from the Atlantic and Pacific Oceans. The present results (1983) support the findings of previous morphological and electrophoretic studies. Evidently, since the uplift of the Panamá land bridge about 3.1 million years ago, there has been continued genetic contact between Atlantic and Pacific skipjack tuna, presumably via the Southern Ocean.     

Comparative species richness,composition, and abundance of invertebrates in Caribbean seagrass (Thalassia testudinum) meadows (Panamá)

The results of a year-long study in which epibenthic invertebrates were collected monthly from seagrass (Thalassia testudinum) meadows along the Caribbean coast of Panamá and the Panamá Canal Zone are described in this paper. Differences in species composition and abundance among sites were primarily due to the proximity of surrounding habitats, especially coral reefs, which contain a number of species that utilize the seagrass meadows. In contrast to many previous characterizations of tropical marine habitats, important seasonal fluctuations in both species number and abundance took place at each of the sites. Data on breeding activity among several species of decapod crustaceans indicate year-round reproduction, although considerable seasonal differences occur in the percentage of ovigerous females. These interspecific differences in observed reproductive output may be explained by differences in life-cycle length, a factor not often considered in discussions of seasonal breeding patterns in tropical marine invertebrates. Overall species composition was qualitatively similar to that reported in comparable studies of tropical and subtropical seagrass meadows elsewhere, although caridean shrimp and xanthid orab species were reduced in number and total abundance were much lower than in previous studies.     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panamá, and Galápagos Islands (Ecuador). II. Poritidae

A comparative study of the reproductive ecology of the zooxanthellate, scleractinian corals Porites lobata Dana and P. panamensis Verrill was conducted from 1985 to 1991 in eastern Pacific reef environments that were severly impacted by the 1982–1983 El Niño warming events. P. lobata, a presumed broadcast spawner of large colony size, is widely distributed in the equatorial eastern Pacific, whereas P. panamensis, a brooder of small colony size, is abundant only on some reefs in Panamá. Both species were gonochoric with nearly 1:1 sex ratios in large study populations except for P. lobata at Caño Island that had 14% hermaphroditic colonies. Mature, unfertilized oocytes contained numerous zooxanthellae in both Porites species, and all planula developmental stages contained zooxanthellae in P. panamensis. Year-round sampling revealed high proportions of colonies with gonads, ranging from 30 to 68% in P. lobata and from 60 to 68% in P. panamensis. No clear relationship between numbers of reproductive colonies and the thermal stability of the habitat was evident in P. lobata: percent colonies with gonads at non-upwelling sites was 48 to 68% at Caño Island (Costa Rica) and Uva Island (Panamá), and at upwelling sites 30 to 50% at Saboga Island and Taboga Island (Panamá), and the Galápagos Islands (Ecuador). Similarly, 90% of all P. panamensis colonies were reproductive at Uva Island (a non-upwelling site), and 86% were reproductive at Taboga Island (an upwelling site). Upwelling at Taboga Island is seasonal, nevertheless P. panamensis produced mature gonads or planulae over most of the year (11 mo), whereas P. lobata exhibited reproductive activity during only 2 mo (May and June). No clear lunar periodicity was observed in P. panamensis (Taboga Island), but a high proportion of P. lobata showed increased gonadal development around full and new moon, especially at Caño and Uva Islands. Estimated fecundities were relatively high for P. lobata at Caño (4000 eggs cm^-2 yr^-1) and Uva (5200 eggs cm^-2 yr^-1) Islands, and notably low (70 to 110 eggs cm^-2 yr^-1) in the Galápagos Islands. P. panamensis mean fecundity at Taboga Island was 720 planulae cm^-2 yr^-1 or 4.0 mm³ cm^-2 yr^-1, which was lower than the egg volume production of P. lobata at Caño and Uva Islands (7.0 to 10.0 mm³ cm^-2 yr^-1). The capacity of P. lobata and P. panamensis to reproduce sexually supports the notion that eastern Pacific coral reef recovery may not be dependent on long-distance dispersal from central Pacific areas. However, sexual recruits of P. lobata are absent or uncommon at all eastern Pacific study sites while recruits of P. panamensis were common to abundant only at the Uva Island study site. Asexual fragmentation in P. lobata augments recruitment locally, but plays no role in P. panamensis recruitment.     

Population dynamics of Diadema antillarum (Echinodermata: Echinoidea) following mass mortality in Panamá

In 1983, Diadema antillarum suffered mass mortality throughout the Caribbean Sea and the western Atlantic Ocean. I followed the dynamics of populations at the San Blas Islands, Panamá from April 1983 to November 1987. Density measurements indicate that populations of D. antillarum have not recovered from the die-offs that killed nearly 97% of the individuals. There was recruitment to the 1 to 1.5 cm class immediately after the mass mortality, but there has been little additional influx of juveniles since then. The low number of observable juveniles could not be attributed to elevated rates of predation on very small individuals. Rates of recruitment did not differ between reefs with artificially increased densities of D. antillarum and reefs kept free of sea urchins; thus, the lack of recruitment did not arise from absence of adults that could provide settlement cues to the larvae or protection to newly settled juveniles. Other species of sea urchins did not show a clear pattern of increase after the demise of D. antillarum. Therefore, interspecific competition directed towards D. antillarum juveniles did not increase after the mass mortality. Two reefs where Echinometra viridis, Eucidaris tribuloides and Lytechinus williamsi, were removed showed no significant differences in recruitment of D. antillarum relative to two reefs where these species were allowed to remain at their natural densities. Resident D. antillarum after the mass mortality produced gametes with the same per capita intensity and lunar synchrony as before the mass mortality. However, it is possible that the probability of fertilization of their gametes decreased because of low population density. The most likely explanation for lack of recruitment is that the reduced numbers of reproducing adults at Panamá and upstream locations resulted in levels of larval supply that were inadequate to sustain recruitment on Panamanian reefs.     

Daily otolith increments and recruitment in two coral-reef wrasses,Thalassoma bifasciatum and Halichoeres bivittatus

Increments on the otoliths of two common coral reef fishes, the bluehead wrasse Thalassoma bifasciatum and the slippery dick Halichoeres bivittatus, were demonstrated by mark-recapture experiments to be daily. Otoliths were marked in two ways; by depriving fish of light, food, and temperature cycles and also by supplemental feeding in the field. Both experiments were performed in late 1980 in the San Blas Islands of Panamá. A mark corresponding to settlement of the planktonic larva onto the reef was found on the otoliths of the bluehead wrasse. This settlement mark was used to calculate the dates of settlement of a collection of juveniles of this species taken from a patch reef in the San Blas Islands of Panamá in 1981. Settlement occurred in short and irregular bursts. The number of daily increments before the settlement mark indicates a planktonic larval life of 40 to 72 d.     

Systematic status of two oyster populations of the genus Tiostrea from New Zealand and Chile

A systematic examination of Tiostrea chilensis endemic to the Pacific coast of South America and of T. lutaria endemic to new Zealand was conducted using protein-gel electrophoresis. Also, ecological and life-history traits were compared between these two documented species. Our study revealed that although three different ecomorphs could be distinguished from various environmental and ecological conditions, the life-history traits between the two species were remarkably similar. Estimates were made of levels of genetic variation and similary based on 29 structural loci among individuals from each documented species. The percentage of loci polymorphic was estimated at 17.2 and 17.6, while the percentage of loci heterozygous was 2.1 and 6.7 for T. chilensis and T. lutaria, respectively. The genetic similarity between the two species was computed as 0.991, where 1.0 indicates no genetic difference among structural loci. The overwhelming evidence of this study indicates that T. chilensis and T. lutaria are different geographical populations of one species and by priority this species should be T. chilensis.     

Sand disposal of the painted ghost crab Ocypode gaudichaudii (Decapoda: Ocypodidae): A possible role in courtship

In 1991/1992, we studied the sand disposal behavior of the painted ghost crabs Ocypode gaudichaudii on the Pacific coast of Panamá. O. gaudichaudii either kick, dump or tamp sand they excavate from their burrows. Here we relate these three kinds of sand disposal to burrow structure and distribution, as well as to crab size and sex. Our objective was to determine whether tamping may be a male courtship signal. Burrows whose owners tamped sand were on average longer, deeper, and higher on the beach than were burrows whose owners kicked or dumped sand. Five burrow shapes were distinguished, with half-spiral and spiral shapes being most common among tamped burrows. All crabs excavated from tamped burrows were males. Tamped burrows peaked in abundance around full and new moons. These observations, together with what is known about mating and breeding behavior of other ghost crabs, suggest that tamping may be involved in O. gaudichaudii courtship.     

Extensive gene flow among mytilid (Bathymodiolus thermophilus) populations from hydrothermal vents of the eastern Pacific

Prior studies of the hydrothermal vent mussel Bathymodiolus thermophilus (Bivalvia: Mytilidae), provided conflicting predictions about the dispersal ability and population structure of this highly specialized species. Analyses of morphological features associated with its larval shells revealed a feeding larval stage that might facilitate dispersal between ephemeral vent habitats. In contrast, an allozyme study revealed substantial genetic differentiation between samples taken from populations 2370 km apart on Galápagos Rift (Latitude 0°N) and the East Pacific Rise (13°N). To resolve the discrepancy between these studies, we examined allozyme and mitochondrial (mt) DNA variation in new samples from the same localities plus more recently discovered sites (9° and 11°N) along the East Pacific Rise. Although analysis of 26 enzyme-determining loci revealed relatively low levels of genetic variation within the five populations, no evidence existed for significant barriers to dispersal among populations. We estimated an average effective rate of gege flow (Nm) of 8 migrants per population per generation. Two common mtDNA variants predominated at relatively even frequencies in each population, and similarly provided no evidence for barriers to gene flow or isolation-by-distance across this species' known range. Larvae of this species appear to be capable of dispersing hundreds of kilometers along a continuous ridge system and across gaps separating non-contiguous spreading centers.     

Spatial autocorrelation analysis of small-scale genetic structure in a clonal soft coral with limited larval dispersal

The philopatric larval dispesal and small effective population sizes characteristic of many clonal species should promote the development of significant small-scale genetic structure within populations as a result of isolation-by-distance. We used spatial autocorrelation statistics to detect genetic structure, arising from both clonal reproduction and philopatric dispersal of sexual propagules, for five allozyme loci within populations of the soft coral Alcyonium sp. In a population on Tatoosh Island, Washington, USA, sampled in 1991/1992, we found significant positive spatial autocorrelation at all loci among individuals separated by <40 cm, reflecting the presence of significant smallscale genetic structure due to associations among clonemates. For 4 of 5 loci, however, we detected no significant spatial autocorrelation among the different clones within this population over distances of 1 to 40 m. Analysis of soft-coral populations from six additional, topographically diverse sites in the north-east Pacific also did not reveal significant spatial autocorrelation among clones at any loci. This general lack of spatial autocorrelation of genotypes among clones suggests that significant small-scale genetic structure has not arisen in populations of Alcyonium sp. as a consequence of isolation-by-distance.     

Hybridization and introgression between Indo-Pacific species of Diadema

We present the first case of hybridization between echinoid species evaluated through genetic markers and morphology. We examined the three tropical Indo-Pacific species of the genus Diadema: D. paucispinum A. Agassiz, D. savignyi (Audouin) Michelin and D. setosum (Leske). Specimens morphologically intermediate between two of these species, D. savignyi and D. setosumhave previously been noted. Fertile hybrids have also been produced in the laboratory. To determine extent of hybridization, we first assayed the allozyme products of 22 loci in individuals which, on the basis of morphology and collection locality, could be unambiguously assigned to one of the three species. We found four loci that were either diagnostic or semi-diagnostic between D. setosum and the other two species, and one locus semi-diagnostic between D. savignyi and D. paucispinum.We then assayed individuals of intermediate morphology to find out whether they had hybrid genotypes. In the Ryukyu Islands, where D. setosum and D. savignyi coexist, we found one specimen which on the basis of all four diagnostic loci was an F₁ hybrid, and several individuals that could be either F₂ (or later-generation) hybrids or progeny of backcrosses.We also found one individual that on both genetic and morphological grounds appeared to belong to D. paucispinum (even though this spcies has only been reported from Hawaii and Kiribati) and three other individuals that carried alleles characteristic of D. paucispinum. Thus, previous reports of hybridization between D. setosum and D. savignyi were correct; it is also possible that larvae of D. paucispinum occasionally arrive at localities outside Hawaii, reach sexual maturity, and hybridize with the other two species. Counts of pure and hybrid morphotypes in other populations across the western tropical Pacific revealed a low but widespread incidence of apparent F₁ hybrids and backcrosses of D.savignyi and D. setosum. However, the existence of diagnostic or semi-diagnostic loci, low interspecific gene-flow estimates based on F _st statistics, and the lack of Hardy-Weinberg or linkage disequilibria among individuals of pure morphology all suggest that gene introgression between the three species is limited.