Howard R. Lasker

DNA fingerprints of a gorgonian coral : a method for detecting clonal structure in a vegetative species

Clonal reproduction, a common life history strategy among sessile marine invertebrates, can lead to high local abundances of one to a few genotypes in a population. Analysis of the clonal structure of such populations can provide insight into the ecological and evolutionary history of the population, but requires markers that can identify individual genets. Forensic and demographic studies have demonstrated that DNA fingerprinting can provide markers that are unique for an individual genotype. We have generated DNA fingerprints for over 70 colonies of the clonal gorgonian, Plexaura A (Plexaura sp. A) collected from June 1990 through July 1991 in the San Blas Islands, Panama. DNA fingerprints within a singic individual were identical and fingerprinting resolved multiple genotypes within and among reefs. On one reef in the San Blas Islands, Panama, 59% of the colonies sampled were of one genotype and this genotype was not found on any other sampled reefs. A previous study using tissue grafts identified 13 putative clones on these reefs, while DNA fingerprints of the same colonies differentiated 17 genotypes. The present study demonstrates the utility of DNA fingerprinting for distinguishing clones and for identifying clonal structure of marine invertebrate populations.

Sediment rejection by reef corals: The roles of behavior and morphology in Montastrea cavernosa (Linnaeus)

Abstract The sediment rejection abilities of colonies of Montastrea cavernosa (Linnaeus) were monitored in a series of field and laboratory experiments. Under conditions of natural sediment deposition, colonies were capable of removing virtually all sediment from their surfaces. Equivalent sections of dead coral skeleton accumulated as much as 345 mg sediment · 25 cm−2 · day−1. Laboratory experiments show the sediment rejection process to consist of passive and active phases. Passive movement of sediment off colonies occurs at the time of initial sediment impact. Fine grain size, tall polyps, and convex colonies all enhance passive removal. Active removal involves the behavioral responses of the colony and is independent of sediment type or colony morphology. The presence of appropriate morphologies can increase the proportion of sediment passively removed and reduce the amount of work a colony must perform to keep its surface clear of sediment.

Bleaching of reef coelenterates in the San Blas Islands, Panama

Starting in June 1983, 25 species of hermatypic corals, gorgonians, hydrocorals, anemones and zoanthids in the San Blas Islands, Panama, began showing signs of a loss of colour leading in some cases to a white “bleached” appearance. Histologic examination of six coral species indicated that bleaching was associated with drastic reductions in the density of zooxanthellae and with the atrophy and necrosis of the animal tissue. The severity of the bleaching varied among species and many species were unaffected. The species most extensively affected were: Agaricia spp., which became completely bleached and frequently died; Montastraea annularis which bleached and continued to survive; and Millepora spp. which bleached white but quickly regained their colouration. Shallow reefs dominated by Agaricia spp. suffered the most extensive bleaching. At one site, Pico Feo, 99% of the Agaricia (32% of the living cover) was bleached. On fore reers, which were dominated by Agaricia spp. and M. annularis, the proportion of M. annularis bleached ranged from 18 to 100% and that of Agaricia spp. from 30 to 53%. Transects at Sail Rock and House Reef were surveyed in August 1983 and January 1984. At those sites, 53% of the Agaricia cover died between August and January. The remaining living cover of Agaricia and of all other species exhibited normal colouration in January. Salinity and temperature were monitored every second day at 4 m depth between May 10 and August 28, 1983 at one of the localities. Bleaching was first observed within two weeks of a 2 °C rise in temperature which occurred in late May 1983. Temperatures remained at or above 31.5 °C for the following 3 weeks and were at or above 30 °C for an additional 4 weeks. The bleaching of corals in the San Blas was most likely due to those elevanted temperatures.

Sexual reproduction and external brooding by the Caribbean gorgonianBriareum asbestinum

The reproductive biology of the gonochoricBriareum asbestinum (Pallas), a common Caribbean gorgonian, was studied in the San Blas Islands (Panama), La Paguera (Puerto Rico) and the Bahamas. Of 721 colonies examined at the height of the reproductive season, during the summers of 1985 to 1988, all were either male or female. No hermaphroditic colonies or polyps were observed. The sexratio of Panama populations was significantly biased in favor of males (2.1:1,n=454) over all four years of the study. A significant male bias was also found in populations sampled in Puerto Rico (2.3:1,n=143) and in the Bahamas (2.5:1,n=124). The consistency of this geographically widespread skewed sex-ratio is unique among octocorals and scleractinians. Ooyctes appear annually in September and reach maximum diameter (600 to 900µm) in June and July of the following year. The 3 to 5 mo developmental cycle of spermaries is shorter than that of females and begins in March. Spawning in males is synchronous and occurs following the full moons of June and July. Fertilization is internal. Embryos are released from polyps very early in development, often whilst still in the process of cleaving. Embryos remain attached to the outside of the colony for 3 to 5 d, many becoming entrapped in mucous sheets secreted by the colonies. Larvae are negatively buoyant and settle rapidly once displaced from the colony. Males become reproductive at smaller branch sizes than females. Reproductive effort (gonads polyp−1 and total gonad volume polyp−1) for both males and females increased with increasing branch size and in polyps away from the tip and base of the branch.

POPULATION STRUCTURE OF A CLONAL GORGONIAN CORAL: THE INTERPLAY BETWEEN CLONAL REPRODUCTION AND DISTURBANCE

Clonality is a common feature of plants and benthic marine organisms. In some cases clonal propagation results in a modest increase in population density, while in other cases dense populations may be generated by the propagation of only a few clones. We analyzed the population structure of the clonal gorgonian Plexaura kuna across several reef habitats with a range of disturbance regimes in the San Blas Islands, Panama, and the Florida Keys, U.S.A. Using multilocus DNA fingerprinting to distinguish clones, we estimated that clones ranged in size from single individuals to 500 colonies. The number of genotypes identified on nine reefs ranged from three to 25. Population density and clonal structure varied markedly among reefs with GO:GE ranging from 0.03 to 1.00. On some reefs vegetative reproduction transformed P. kuna from a rare species to the numerically most abundant gorgonian. The effect of clonal propagation on P. kuna population structure was dependent on interactions between fragmentation and the reef environment (disturbance regime, substratum). We present a generalized model relating population structure of clonal species to disturbance and the mode of vegetative propagation. Disturbance promotes colony propagation and skews the size‐frequency distribution of clones among P. kuna and many species that propagate via fragmentation. Propagation of these species is promoted by disturbance (disturbance sensitive), and they tend to have clones that are dispersed across local sites. Species that fragment and have dispersed clones, have high genotypic diversity in habitats with low levels of disturbance. Genotypic diversity then decreases at intermediate disturbance and increases again at the highest disturbance levels. Clonal species that do not rely on disturbance for vegetative propagation (disturbance insensitive) generally do not disperse and form aggregated clones. Among these taxa disturbance has a greater affect on individual survival than on propagation. Genotypic diversity is directly related to the level of disturbance until very high levels of disturbance, at which time genotypic diversity declines.

The Reproductive Cycle and Spawning in a Caribbean Gorgonian

The reproductive biology of Plexuara A, a common but undescribed Caribbean gorgonian, was studied in the San Blas Islands, Panama. Oocytes were present in the polyps throughout the year, though a seasonal developmental cycle is evident. Early stage oocytes appeared at the base of the polyps in November. In January total egg volume per polyp began to increase and reached a maximum in early May. Total egg volume per polyp then decreased through the summer as mature eggs were released. Spawning occurred over a period of 4-7 days following each full moon in May, June, and July. Egg release was synchronous, starting at approximately 18:30 and lasting 90 minutes. Larval development is initiated just prior to or at the time of release. Initiation of larval development at the time of release without brooding is unknown for any gorgonian or scleractinian. Of 265 colonies examined from 6 reefs near San Blas Point, all but 3 contained gonads and were female. No male or hermaphroditic colonies have been found in the ...

In situ Rates of Fertilization Among Broadcast Spawning Gorgonian Corals

Fertilization rates among marine benthic taxa have implicitly been assumed to be uniformly high in most analyses of life history evolution, but in situ fertilization rates during natural spawning events are rarely measured. Fertilization rates of the Caribbean gorgonians Plexaura kuna and Pseudoplexaura porosa were measured at a site in the San Blas Islands, Panama, by collecting eggs downstream of colonies during synchronous spawning events during the summer months in the years 1988-1994. Eggs collected by divers were incubated, and the proportion of eggs that developed was determined. Proportions of eggs developing suggest fertilization rates that vary from 0% to 100%. Monthly means ranged from 0% to 60.4%. Failure of gametes to develop can be attributed to sperm limitation, as eggs collected during spawning had higher fertilization rates if incubated with an excess of sperm. Plexaura kuna fertilization rates were highest during the July spawning events. Fertilization of Plexaura kuna eggs was usually lower during the first two nights of the 4-6 night spawning event. The proportion of eggs being fertilized when collected from a given place and time was highly variable, with one peak in the frequency distribution at or below 20% fertilization, and a second group of samples with greater fertilization rates. High variance in fertilization rates is evident at all levels of analysis: between replicate samples, times within nights, and among nights and months. This variance can be attributed to a combination of the effects of heterogeneity in the water column as gametes are diluted, spawning behavior of the gorgonians, and the current regime. Fertilization rates are often low and may represent a limiting step in recruitment during some years. Low fertilization rates may also be an important component of the life history evolution of these species.

Repeated loss of coloniality and symbiosis in scleractinian corals

The combination of coloniality and symbiosis in Scleractinia is thought to confer competitive advantage over other benthic invertebrates, and it is likely the key factor for the dominance of corals in tropical reefs. However, the extant Scleractinia are evenly split between zooxanthellate and azooxanthellate species. Most azooxanthellate species are solitary and nearly absent from reefs, but have much wider geographic and bathymetric distributions than reef corals. Molecular phylogenetic analyses have repeatedly recovered clades formed by colonial/zooxanthellate and solitary/azooxanthellate taxa, suggesting that coloniality and symbiosis were repeatedly acquired and/or lost throughout the history of the Scleractinia. Using Bayesian ancestral state reconstruction, we found that symbiosis was lost at least three times and coloniality lost at least six times, and at least two instances in which both characters were lost. All of the azooxanthellate lineages originated from ancestors that were reconstructed as symbiotic, corroborating the onshore–offshore diversification trend recorded in marine taxa. Symbiotic sister taxa of two of these descendant lineages are extant in Caribbean reefs but disappeared from the Mediterranean before the end of the Miocene, whereas extant azooxanthellate lineages have trans-Atlantic distributions. Thus, the phyletic link between reef and nonreef communities may have played an important role in the dynamics of extinction and recovery that marks the evolutionary history of scleractinians, and some reef lineages may have escaped local extinction by diversifying into offshore environments. However, this macroevolutionary mechanism offers no hope of mitigating the effects of climate change on coral reefs in the next century.

Microsatellite variation reveals high levels of genetic variability and population structure in the gorgonian coral Pseudopterogorgia elisabethae across the Bahamas

The primary mechanism of gene flow in marine sessile invertebrates is larval dispersal. In Pseudopterogorgia elisabethae, a commercially important Caribbean gorgonian coral, a proportion of the larvae drop to the substratum within close proximity to the maternal colony, and most matings occur between individuals in close proximity to each other. Such limited dispersal of reproductive propagules suggests that gene flow is limited in this gorgonian. In this study, we characterized the population genetic structure of P. elisabethae across the Bahamas using six microsatellite loci. P. elisabethae was collected from 18 sites across the Bahamas. Significant deviations from Hardy–Weinberg equilibrium due to deficits of heterozygotes within populations were detected for all 18 populations in at least one of the six screened loci. Levels of genetic structure among populations of P. elisabethae were high and significant. A distance analysis placed populations within three groups, one formed by populations located within Exuma Sound, a semi‐isolated basin, another consisting of populations located outside the basin and a third group comprising two populations from San Salvador Island. The patterns of genetic variation found in this study are concordant with the life‐history traits of the species and in part with the geography of the Bahamas. Conservation and management plans developed for P. elisabethae should considered the high degree of genetic structure observed among populations of the species, as well as the high genetic diversity found in the San Salvador and the Exuma Sound populations.

Reproductive success in the Caribbean octocoral Briareum asbestinum

For most sessile marine invertebrates the number of offspring produced by an individual is often determined indirectly from counts of eggs present, with the assumption that all or most become viable offspring. Few field data on the actual number of offspring produced per individual (reproductive success) have been reported. We examined reproductive success in the octocoral, Briareum asbestinum (Pallas), by measuring embryo densities on 40 female colonies on two reefs (Pinnacles and House reefs) in the San Blas Islands, Panama from 1986 to 1988. The percentage of female colonies with mature eggs which subsequently released embryos was low, ranging from 46.6% in 1986 to 5.0% in 1988. Reproductive success was significantly different between reefs. Pinnacles reef had significantly more successful colonies (those releasing embryos) than House reef in all three years and significantly higher embryo densities in 1986 and 1987. There was a significant positive correlation between female reproductive success and the density and proximity of nearby males in both 1986 and 1987. In addition, fertile female branches placed 25 cm away from male branches produced significantly more embryos than female branches placed 50 cm away. In 1988 two groups of female branches were placed at distances of <50 cm and 5 m from four large male colonies at Pinnacles reef. Of the 20 female branches placed 5 m from the males none released embryos while 35% (7 of 20 branches) of the branches <50cm from males released embryos. These data indicate that simple counts of mature eggs present within female colonics prior to the reproductive season provide a poor estimate of reproductive success in B. ashestinum and that female reproductive success is positively influenced by the close proximity of males.