Genetic differentiation among populations of a broadcast spawning soft coral, Sinularia flexibilis, on the Great Barrier Reef

The genetic structure of 12 reef populations of the soft coral Sinularia flexibilis (Octocorallia, Alcyoniidae) was studied along the Great Barrier Reef (GBR) at a maximum separation of 1,300 km to investigate the relative importance of sexual and asexual reproduction, genetic differentiation and gene flow among these populations. S. flexibilis is a widely distributed Indo-Pacific species and a gamete broadcaster that can form large aggregations of colonies on near-shore reefs of the GBR. Up to 60 individuals per reef were collected at a minimum sampling scale of 5 m at two sites per reef, from December 1998 to February 2000. Electrophoretic analyses of nine polymorphic allozymes indicated that genotypic frequencies in most populations and loci did not differ significantly from those expected from Hardy–Weinberg predictions. Analysis of multi-locus genotypes indicated a high number of unique genotypes (N _go) relative to the number of individuals sampled (N) in each reef population (range of 0.69–0.95). The maximum number of individuals likely to have been produced sexually (N*) was similar to the number of individuals sampled (i.e. N*:N ˜ 1), suggesting that even repeated genotypes may have been produced sexually. These results demonstrated a dominant role of sexual reproduction in these populations at the scale sampled. Significant genetic differentiation between some populations indicated that gene flow is restricted between some reefs (F _ST=0.026, 95% CI= 0.011 − 0.045) and even between sites within reefs (F _ST=0.041, 95% CI=0.027 − 0.055). Nevertheless, there was no relationship between geographic separation and genetic differentiation. Analyses comparing groups of populations showed no significant differentiation on a north-south gradient in the GBR. The pattern in the number of significant differences in gene frequencies in pairwise population comparisons, however, suggested that gene flow may be more restricted among inner-shelf reef populations near to the coast than among mid/outer-shelf populations further from the coast. Received: 10 July 2000 / Accepted: 5 October 2000     

Allozyme electrophoresis indicates high gene flow between populations of Holothuria (Microthele) nobilis (Holothuroidea: Aspidochirotida) on the Great Barrier Reef

 Genetic variation in 15 Holothuria (Microthele) nobilis (Selenka, 1867) populations on the Great Barrier Reef was studied at seven polymorphic allozyme loci. Although populations were separated by distances up to 1300 km, there were no apparent restrictions to gene flow (F _ST values were not significantly different from 0) and the maximum Nei's unbiased genetic distance was 0.003. Populations were in Hardy–Weinberg equilibrium at all loci. The estimated maximum sexual input [this is the ratio of the number of sexually produced individuals (N*) to the sample size (N _i )] and the minimum sexual input [this is the ratio of the number of genotypes (N _go ) over the sample size (N _i )] were used as estimators for the amount of asexual reproduction. Both parameters suggested that H. nobilis reproduces solely by sexual means (N*:N _i : = 1; N _go :N _i  = 0.74 to 1). The allozyme data indicated high gene flow between populations, but the possibility that allozyme frequencies may not be at equilibrium means that it was not possible to distinguish whether the patterns reflect present-day dispersal or dispersal that occurred in the past. Received: 27 March 2000 / Accepted: 29 June 2000     

Mode of reproduction,recruitment, and genetic subdivision in the brooding sponge<Emphasis Type="Italic"> Haliclona</Emphasis> sp.

Sponges display a variety of reproductive strategies that have the potential to influence population genetic structure. Histological examination of ten reproductive individuals of the Western Australian sponge Haliclona sp. showed that this species broods embryonic larvae that are potentially limited in dispersal capabilities. Because sponges have the potential to propagate in a number of modes, allozyme electrophoresis was used to assess the relative importance of asexual and sexual reproduction to recruitment, and to quantify genetic subdivision over different spatial scales. Tissue samples from 227 sponges were collected from reefs within two areas 400 km apart: Hamelin Bay and Rottnest Island. Contrary to expectations for highly clonal populations, genotypic diversity within sites was high, no linkage disequilibrium was found, and there was no evidence of genotypic clustering within reefs. There was no genetic evidence that asexual reproduction is important for the maintenance of populations. Genetic comparisons were consistent with mixing of sexually produced recruits within reefs, on a scale up to a few hundred metres, but significant genetic subdivision between reefs (F_ST=0.069 at Hamelin Bay, 0.130 at Rottnest Island) indicated that water gaps of several hundred metres are effective at preventing dispersal. Subdivision between the two areas, separated by 400 km, was moderately greater (F_ST=0.142) than within, but the same alleles were predominant in the two areas. These genetic patterns are consistent with limited dispersal capabilities of brooded larvae.Communicated by G.F. Humphrey, Sydney     

Population genetics of the fissiparous holothurians Stichopus chloronotus and Holothuria atra (Aspidochirotida): a comparison between the Torres Strait and La Réunion

Collections of about 50 individuals from each of five populations of the fissiparous holothurian species Stichopus chloronotus and four populations of Holothuria atra were made in 1999. These populations were located in the Torres Strait (western Pacific) and La Réunion (western Indian Ocean). Allozyme electrophoretic surveys of five (S. chloronotus) and six (H. atra) loci were conducted to compare patterns of asexual reproduction and to investigate connectivity between regions separated by large geographic distances. Deviations from genotype frequencies expected under Hardy-Weinberg equilibrium, mostly heterozygote excesses, were observed in all populations of both species. The maximum contribution of sexual reproduction (calculated as the maximum number of sexually produced individuals: sample size=N*/N_i) was similar for all S. chloronotus (58-64%) and H. atra (76-92%) populations, and on the same level as previously reported for midshelf reefs of the Great Barrier Reef. The higher values in the latter species indicated greater contributions of asexual reproduction to S. chloronotus populations. Variability was strongly reduced in S. chloronotus populations at La Réunion, with only one locus being variable in that population. When the dataset was reduced to one representative per multi-locus genotype per population to reduce the effect of asexual reproduction on calculations on gene flow, F_ST values were not significantly different from zero, suggesting high gene flow between these regions. However UPGMA cluster analyses using Rogers' genetic distance, roughly clustered populations by region. In the case of H. atra, pooled populations within each region were significantly different from those of the other region. Thus, although some restrictions in gene flow and greater genetic distances between the regions may exist, those differences are distinctly less than those reported in previous studies on echinoderms over similar geographic scales. Despite the importance of asexual reproduction for the maintenance of local population size, this study also confirmed that the potential for widespread dispersal mediated by sexually produced larvae is large.     

Panmixia in Pocillopora verrucosa from South Africa

The genetic structure of six local collections of Pocillopora verrrucosa from six coral reefs in KwaZulu-Natal, South Africa, was examined using allozyme electrophoresis. The six separate reefs lie within two different reef complexes. Twenty-two enzymes were screened on five buffer systems, but only five polymorphic loci (Gpi-1, Gdh-1, Lgg-2, Lpp-1, Est-1) could be consistently resolved. No significant differences in allelic frequencies were detected among the six sites. All local collections were genotypically diverse, with evidence of only very limited clonal replication at each site. Indeed, the ratio of observed to expected genotypic diversity (mean Go:Ge=0.64ǂ.05 SD), the ratio of observed number of genotypes to the number of individuals (mean Ng:N=0.65ǂ.04 SE), and deviations from the Hardy-Weinberg equilibrium indicate that sexual reproduction plays a major role in the maintenance of the populations. No genetic differentiation was found either within (FSR=0.026ǂ.003 SE) or between (FRT=0.000ǂ.001 SE) reef complexes. The homogeneity of the gene frequencies across the six reefs strongly supports the assumption that the KwaZulu-Natal reef complexes are highly connected by gene flow (Nem=44). The reefs in the southern and central reef complexes along the northern Maputaland coastline can therefore be considered part of a single population.     

Genetical structure within populations of the coral Pocillopora damicornis

Genetic variation of allozymes within populations of Pocillopora damicornis from southwestern Australia was consistent with a primary role of local asexual proliferation of clones in population maintenance. Populations were composed typically of two to four multilocus genotypes accounting for 40 to 80% of individuals, with the remainder assigned to genotypes occasionally in twos or threes but more commonly singly. In the three populations where recruitment was examined genetically, 84% of all first-year recruits was assigned to clones represented in the population's resident adults. The majority of these recruits came from the most highly-replicated of the adult clones. The observed genotypic diversity was, on average, about half that calculated to occur for the same allelic frequencies in a sexually-reproducing population with free recombination. Despite the prevalence of asexual reproduction, both through planulae and fragments, the existence of a sexual mode of reproduction was inferred from the high level of variation produced by pooling populations, the existence of novel genotypes and the concordance of clonal gene frequencies at many sites with the predictions of Hardy-Weinberg equilibria.     

Genetic variation and clonal structure in the scleractinian coral Pocillopora damicornis in the Ryukyu Archipelago, southern Japan

Samples of the scleractinian coral Pocillopora damicornis were collected from six sites located around four islands in the Ryukyu Archipelago, southern Japan, and subjected to allozyme electrophoresis. Seven polymorphic loci were examined for their allelic patterns. The ratio of observed to expected genotypic diversity (0.30 < G _o :G _e  < 0.64), the ratio of the observed number of genotypes to the number of individuals (0.47 < N _g :N _i  < 0.75), and deviations from Hardy–Weinberg equilibrium indicated that asexual reproduction plays a major role in the maintenance of established populations. However, populations were not completely dominated by a single or a few clones, and most clones were represented by only a few individual samples. The high frequency of typhoons in the region suggests that, in P. damicornis, fragmentation caused through occasional exposure to powerful waves is a major mode of asexual reproduction, but asexual production of planulae may also be contributing to the maintenance of populations. A significant genetic differentiation (F ST) was found between the six populations examined (0.027 < F ST < 0.092, average F ST = 0.056). The moderate gene flow is discussed according to characteristics of the larval stage of the species, and to circulation patterns in the region. Received: 7 August 1998 / Accepted: 18 May 1999     

Protection of Genetic Diversity and Maintenance of Connectivity among Reef Corals within Marine Protected Areas

Abstract: High‐latitude coral reefs (HLRs) are potentially vulnerable marine ecosystems facing well‐documented threats to tropical reefs and exposure to suboptimal temperatures and insolation. In addition, because of their geographic isolation, HLRs may have poor or erratic larval connections to tropical reefs and a reduced genetic diversity and capacity to respond to environmental change. On Australia's east coast, a system of marine protected areas (MPAs) has been established with the aim of conserving HLRs in part by providing sources of colonizing larvae. To examine the effectiveness of existing MPAs as networks for dispersal, we compared genetic diversity within and among the HLRs in MPAs and between these HLRs and tropical reefs on the southern Great Barrier Reef (GBR). The 2 coral species best represented on Australian HLRs (the brooding Pocillopora damicornis and the broadcast‐spawning Goniastrea australensis) exhibited sharply contrasting patterns of diversity and connectedness. For P. damicornis, the 8‐locus genetic and genotypic diversity declined dramatically with increasing latitude (N_a= 3.6–1.2, H_e= 0.3–0.03, N_g:N = 0.87–0.06), although population structure was consistent with recruitment derived largely from sexual reproduction (G_o:G_e= 1.28–0.55). Genetic differentiation was high among the HLRs (F_ST[SD]= 0.32 [0.08], p < 0.05) and between the GBR and the HLRs (F_ST= 0.24 [0.06], p < 0.05), which indicates these temperate populations are effectively closed. In contrast for G. australensis, 9‐locus genetic diversity was more consistent across reefs (N_a= 4.2–3.9, H_e= 0.3–0.26, N_g:N = 1–0.61), and there was no differentiation among regions (F_ST= 0.00 [0.004], p > 0.05), which implies the HLRs and the southern GBR are strongly interconnected. Our results demonstrate that although the current MPAs appear to capture most of the genetic diversity present within the HLR systems for these 2 species, their sharply contrasting patterns of connectivity indicate some taxa, such as P. damicornis, will be more vulnerable than others, and this disparity will provide challenges for future management.     

Variation in the genetic composition of coral (Pocillopora damicornis and Acropora palifera) populations from different reef habitats

The genetic structure of populations of the corals Pocillopora damicornis and Acropora palifera was examined in three habitats at One Tree Island during March and April 1993, using electrophoretically detectable variation at six allozyme loci. There were significant genetic differences among populations of P. damicornis within each of the reef crest, lagoon and microatoll habitats. The level of differentiation among populations was similar in each of the habitats. Differences between populations of P. damicornis from lagoon and microatolls were no greater than that within habitats, but genetic differentiation of these from crest populations was much higher. There was no difference in the genetic composition of A. palifera populations within or between the lagoon and microatolls, the only habitats where this species was found. Both coral species had observed:expected (G _O:G_E) genotypic diversity rations >0.80, indicating predominantly sexual reproduction. These data, the high genotype diversity and general conformance of genotype frequencies to those expected under conditions of Hardy-Weinberg, suggested panmixis at each site. The high degree of sexual reproduction in the P. damicornis populations is unusual for a species where asexual reproduction has been the dominant mode of reproduction reported to date. Gene flow in both species was considerable between the lagoon and the closed microatolls. The genetic differences between populations of P. damicornis in these habitats and the reef crest may reflect the relative isolation of all populations within the closed One Tree Lagoon from those outside. However, local currents appear to offer effective means of dispersal between the habitats, suggesting that the genetic differences result from natural selection in the different environments within One Tree Lagoon and the reef crest.     

Genetic structure of the scleractinian coral<Emphasis Type="Italic"> Platygyra sinensis</Emphasis> in Hong Kong

Genetic structure was studied in Platygyra sinensis from six sites in Hong Kong by allozyme electrophoresis using an isoelectric focusing gel (IEF). Seven variable loci were detected using six enzyme systems. The number of alleles per locus ranged from 3.3 to 3.6 in the six samples, and heterozygosity from 0.429 to 0.540. Genotype frequencies were generally close to Hardy–Weinberg expectations. The values of unique multilocus genotype to the number of individuals for each sample (N_G/N) and observed genotypic diversity to expected genotypic diversity (G_O/G_E) were high, indicating a high level of sexual reproduction. Variation in allele frequencies among sites was low (F_ST=0.017), as was Nei's unbiased genetic distance (D), suggesting genetic similarity among sites.Communicated by T. Ikeda, Hakodate     

Seasonality of asexual reproduction in Holothuria (Halodeima) atra, H. (H.) edulis and Stichopus chloronotus (Holothuroidea: Aspidochirotida) on the Great Barrier Reef

Asexual reproduction by fission was monitored for 18 mo in populations of Holothuria (Halodeima) atra, H. (H.) edulis and Stichopus chloronotus on three nearshore fringing reefs and one midshelf reef in the Great Barrier Reef. Fission in S. chloronotus occurred exclusively between March and October, with a peak value of 31% recently divided individuals in one population in July. H. atra showed a similar pattern, with maxima of between 16 and 26% from May to July. In H. edulis, asexual reproduction occurred only between March and July, with a maximum of 17% recently divided individuals in March. Fission rates of H. atra and S. chloronotus in winter were significantly higher than in all other seasons. For H. atra, at least 76% of all individuals at Fantome Island were estimated to undergo fission per year, whereas only 9% undergo fission on the midshelf reef. Highest annual fission rates (43%) for S. chloronotus were found in a dense population on Great Palm Island. The lower-density midshelf reef population exhibited comparatively lower annual fission rates (19%). About 24% of H. edulis undergo fission each year. Annual fission rate and population density were positively correlated in the four populations of S. chloronotus and H. atra studied. Received: 21 March 1997 / Accepted: 26 March 1997     

Genetic study of the extent and consequences of sexual and asexual reproduction in the deep-sea epizoic anemones Amphianthus inornata and Kadosactis commensalis (Cnidaria: Anthozoa)

The anemone Amphianthus inornata is found at bathyal depths living on colonies of the gorgonian Acanella arbuscula. Previous studies of the morphology and reproductive stage of this anemone, during different times of the year, have indicated that it reproduces sexually on a seasonal basis. A small proportion of the study population were also reported to be undergoing asexual reproduction by fission. The anemone Kadosactis commensalis is also bathyal, but lives mainly on the holothurian Paroriza prouhoi. Previous morphological studies have indicated that K. commensalis is a protandrous hermaphrodite that exhibits non-seasonal sexual reproduction only. In the present study, allozyme electrophoresis was used to examine the prevalence and genetic consequences of asexual reproduction in a population of Amphianthus inornata from 2 200 m in the Rockall Trough, North Atlantic Ocean. Genetic evidence, from five randomly selected polymorphic enzyme loci, for asexual reproduction in this species was weak. Exact tests indicated that genotype frequencies did not differ significantly from those expected under Hardy–Weinberg equilibrium. F _IS (correlation of homologous alleles with reference to local population, assuming random mating) values also did not differ significantly from zero, and observed heterozygosity (H _o =0.446) and genotypic diversity (G _o =17.0387) were very similar to Hardy–Weinberg expected frequencies (H _e =0.446; G ^* _e =17.0010). Evidence suggests that the contribution of asexual reproduction to recruitment in the study population of A. inornata is low. For a single population of K. commensalis from 4 850 m on the Porcupine Abyssal Plain, North Atlantic Ocean, the hypothesis that inbreeding due to reduced occurrence of outcrossing between anemones on a single holothurian host was examined by electrophoresis of ten randomly selected enzyme loci. Single-locus genotypic frequencies were significantly different from expected frequencies for one locus P≤ 0.05, hexokinase-1 (Hex-1)]. F _IS values were significantly different from zero for two enzyme loci (Hex-1 and Hex-2, P≤ 0.01 and P≤ 0.05, respectively), and the overall observed heterozygosity was lower than the expected heterozygosity (H _o =0.125, H _e =0.140). The hypothesis of inbreeding could not be rejected by the present study, although sample size was small (N=55), leading to possible bias in tests of significance. Genetic variation in A. inornata was higher than that recorded for most eukaryotes, although interlocus sampling error for only five loci is high. High genetic variability has been found in other sea anemones, and has been related to high longevity and mixed reproductive modes. Genetic variation in K. commensalis was in the higher range of that found in other eukaryotes, and is not unusual for anemones. Received: 5 August 1996 / Accepted: 11 December 1996     

Genetic structure of a scleractinian coral, Mycedium elephantotus, in Taiwan

The genetic structure of Mycedium elephantotus (Pallas, 1766) populations from three regions around Taiwan was examined using allozyme electrophoresis. Eight loci were scored from seven enzyme systems. Seven loci were polymorphic under the 95% criterion. The high ratio of observed to expected genotypic diversities at the collection sites (G _O:G _E=0.8 to 1.0) indicate that M. elephantotus propagates predominantly by sexual reproduction. Allele frequencies of M. elephantotus differed significantly among regions (D=0.024 to 0.256, F _ST=0.032 to 0.218, p < 0.001), while populations among collection sites within each region were homogeneous (D=0.000 to 0.015, F _ST=0.010 to 0.022, p > 0.05). Genetic differentiation between populations from southern Taiwan and the Penghu Islands is greater than that between southern and northern Taiwan populations, although the former two regions are much closer geographically. The UPGMA dendrogram based on Nei's unbiased genetic distance showed a clear subdivision of populations into two groupings, northern Taiwan/Penghu Islands and southern Taiwan. A higher level of gene flow was found between M. elephantotus populations in northern Taiwan and the Penghu Islands (N _m=7.56) than that between populations in southern Taiwan and other regions (N _m=0.90 to 1.72). The pattern of genetic subdivision among regions is consistent with the pattern of ocean currents, indicating that genetic differentiation is likely driven by surface circulation vicariance. Received: 2 December 1997 / Accepted: 15 September 1998     

Allozyme electrophoresis demonstrates that the scleractinian coral Montipora digitata is two species

Allozyme electrophoresis at five loci was used to clarify the species status of two morphs of the scleractinian coral Montipora digitata (Dana, 1846), using specimens collected from three locations off the Queensland coast between October 1991 and April 1993. The two morphs, occurring sympatrically, were distinguished by one fixed gene difference at Locus LT-2 and frequency differences at three other loci, indicating reproductive isolation between them. Populations of both morphs were generally in Hardy-Weinberg equilibrium, suggesting that sexual reproduction does occur. Despite sampling designed to reduce the chance of collecting clonemates, the genetic diversity ratio (G _O:G_E) indicated that asexual reproduction also occurs. This is in agreement with the known modes of reproduction for this species. There was significant genetic differentiation between populations of one of the morphs of M. digitata. This could have been influenced by greater degrees of asexual reproduction in this morph, selection within sites, or variation in larval survival and recruitment patterns. Relatively low F _st (differentiation among populations) values found for a broadcast spawning species during this study are in accordance with the hypothesis that brooded planulae are adapted for rapid settlement, whereas planulae produced by broadcast spawners are adapted for widespread dispersal.     

Population structure in the sexually reproducing sea anemone Oulactis muscosa

The intertidal sea anemone Oulactis muscosa (Drayton) is dioecious and most individuals are sexually mature throughout the year. Biochemical genetic evidence was used to determine the genetic structure of populations and to infer the relative contributions of sexual and asexual reproduction to recruitment. Data were collected for six enzyme-encoding loci from local populations spread along 735 km of the south east coast of Australia. The genetic structure of each of the nine local populations studied was consistent with recruitment by sexually produced individuals. In almost all cases, the observed single-locus genotypic frequencies closely matched those expected for hardy-Weinberg equilibria, however, consistent deficits of heterozygotes were detected for all loci. No apparent subdivision of the population was detected within the sampling area. Low levels of genetic differentiation were found between local populations and standardised variance (F _ST ) values were similar to those for other species with widespread planktonic dispersal of larvae.Contribution No. 60 from the Ecology and Genetics Group of the University of Wollongong     

Asexual production of planulae in the coral Pocillopora damicornis

The reproduction of scleractinian corals through planular larvae has traditionally been viewed as a strictly sexual process. Here, the results of an electrophoretic study of a ubiquitous Indo-Pacific coral, Pocillopora damicornis, show an exact inheritance of parental genotypes by brooded planulae, demonstrating the existence of an asexual mode of production of planular larvae. Comparisons of the genetical structure of a number of populations with structures predicted for sexual reproduction suggest that, although there is probably also a sexual form of reproduction, asexually produced planulae can be of major importance in the maintenance of populations of this species.     

Fissiparity and population genetics of Coscinasterias calamaria

The effect of asexual reproduction on the population genetics of the fissiparous seastar Coscinasterias calamaria was examined at Rottnest Island and the adjacent mainland, Western Australia, in 1985. Field samples and laboratory observations on growth rate and regeneration showed that fission is common at all 14 sites examined. Electrophoretic analysis of six polymorphic enzymes revealed low genotypic diversity and strong genetic disequilibrium at each site, confirming the highly clonal structure of local populations. Striking variation in clonal composition over distances as short as 50 m emphasizes the very localized subdivision of these populations. In the combined sample from all sites, however, the expected range of multilocus genotypes was found in the proportions expected from random mixing of sexually produced larvae, confirming that clonal diversity results from sexual reproduction. Local genotypic diversity was not correlated with fission, emphasizing the difficulty of determining the short-term roles of asexual and larval recruitment in the maintenance of populations. Subtidal populations, however, appear to have both lower incidence of fission and lower larval recruitment than do intertidal populations.     

Spatial and temporal variation in coral recruitment on the Great Barrier Reef: Implications for dispersal hypotheses

Over 15 000 coral recruits were counted on settlement plates from three mid-shelf reefs and six fringing reefs in the northern section of the Great Barrier Reef during two summers (1986 and 1987) and one winter (1987). The density of coral recruits on some settlement plates from a fringing reef was up to 4.88 cm^–2, the highest value ever reported. Mean density of recruits was greater on fringing reefs (81.1 recruits/settlement plate) than on mid-shelf reefs (15.6 recruits/settlement plate), but there was greater spatial variation in abundance of recruits between the fringing reef sites. Other differences between the mid-shelf reefs and the fringing reefs were that different taxa were dominant, and that settlement orientation differed, with mid-shelf recruits settling preferentially on horizontally oriented surfaces and fringingreef recruits preferring vertical surfaces. Of the three midshelf reefs, Green Island reef recorded the highest recruitment rate for each of the two summers, despite having a depauperate adult coral population following predation by the asteroidAcanthaster planci. This suggests that coral larvae frequently travel between reefs. In contrast with an earlier study, there was no consistent difference in abundance of recruits between forereef and backreef locations. Overall, the results indicated great spatial variation in the availability of coral larvae, both on the scale of whole reefs and within-reef habitats.     

Reef recovery 20 years after the 1982–1983 El Niño massive mortality

For over 20 years the El Niño-Southern Oscillation (ENSO) has caused damage to the coral reefs of the eastern Pacific and other regions. In the mid-1980s scientists estimated that coral cover was reduced by 50–100% in several countries across the region. Almost 20 years (2002) after the 1982–1983 event, we assessed the recovery of the virtually destroyed reefs at Cocos Island (Costa Rica), previously evaluated in 1987 and reported to have less than 4% live coral cover. We observed up to fivefold increase in live coral cover which varied among reefs surveyed in 1987 and 2002. Most new recruits and adults belonged to the main reef building species from pre-1982 ENSO, Porites lobata, suggesting that a disturbance as outstanding as El Niño was not sufficient to change the role or composition of the dominant species, contrary to phase shifts reported for the Caribbean. During the 1990s, new species were observed growing on the reefs. Notably, Leptoseris scabra, considered to be rare in the entire Pacific, was commonly found in the area. Recovery may have begun with the sexual and asexual recruits of the few surviving colonies of P. lobata and Pavona spp. and with long distance transport of larvae from remote reefs. We found an overall 23% live coral cover by 2002 and with one reef above 58% indicating that Cocos Island coral reefs are recovering.     

Demography and interannual variability of salp swarms (Thalia democratica)

Swarms of the pelagic tunicate, Thalia democratica, form during spring, but the causes of the large interannual variability in the magnitude of salp swarms are unclear. Changes in asexual reproduction (buds per chain) of T. democratica populations in the coastal waters of south-east Australia (32–35°S) were observed in three austral springs (October 2008–2010). T. democratica abundance was significantly higher in 2008 (1,312 individuals m^?3) than 2009 and 2010 (210 and 92 individuals m^?3, respectively). There was a significant negative relationship (linear regression, r ² = 0.61, F _1,22 = 33.83, P < 0.001) between abundance and asexual reproduction. Similarly, relative growth rates declined with decreasing abundance. Generalised additive mixed modelling showed that T. democratica abundance was significantly positively related to preferred food >2 μm in size (P < 0.05) and negatively related to the proportion of non-salp zooplankton (P < 0.001). Salp swarm magnitude, growth, and asexual reproduction may depend on the abundance of larger phytoplankton (prymnesiophytes and diatoms) and competition with other zooplankton.