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.     

Acanthochromis polyacanthus,a fish lacking larval dispersal,has genetically differentiated populations at local and regional scales on the Great Barrier Reef

Acanthochromis Gill is a monotypic genus within the damselfish family Pomacentridae, erected for an unusual species [A. polyacanthus (Bleeker)] that uniquely lacks larval dispersal. Instead, offspring are reared in the parental territory, in the manner of cichlids, and fledged into the surrounding habitat. Phenotypic and genotypic variation was surveyed on the basis of body colouration and 7 polymorphic loci in 19 populations from 5 regions of the central and southern Great Barrier Reef (GBR). Variation in both characters was found at regional and local scales. Two colour morphs were recognised: a bicoloured morph from the three northern regions and a uniform dark morph from the two southern regions. Isozyme analysis showed a similar pattern with greatest variation between the different morphs, but also with significant variation at both regional and local scales within morphotypes. Heterozygosity was maximal in the central populations, which, together with other measures of variability, suggests a mixing of separate gene pools in this region and denies species status to the two morphotypes despite numerous fixed differences in allele frequencies between the most distant populations. The presence of fixed differences in multiple alleles between populations separated by 1000 km indicates negligible gene flow over such distances and long isolation of these gene pools. These patterns may reflect recolonisation of the GBR after the last sea-level rise by fish from two stocks. Founder effects and random drift in small populations after colonisation are probably the major sources of the local and regional variations observed at smaller spatial scales. This diversity has been maintained among populations at all scales by the very low levels of gene flow possible without an effective strategy for larval dispersal between coral reefs.     

Genetic structure of<Emphasis Type="Italic"> Linckia laevigata</Emphasis> and<Emphasis Type="Italic"> Tridacna crocea</Emphasis> populations in the Palawan shelf and shoal reefs

Allozyme variation of 10 populations of Linckia laevigata at 8 polymorphic loci and 13 populations of Tridacna crocea at 6 polymorphic loci were analyzed to compare genetic variability and genetic affinities among reefs in Palawan, Philippines. Two to five populations were sampled from each of four regions: the shelf reefs in (1) northern Palawan and (2) southern Palawan and the offshore reefs in (3) the Kalayaan island group (KIG) in the South China Sea and (4) the Tubbataha shoals in the Sulu Sea. Heterozygosity was highest in populations of L. laevigata from the south shelf of Palawan and populations of T. crocea from the Tubbataha shoals of the Sulu Sea. The lowest heterozygosity estimates were from the reefs of the KIG in the South China Sea, for both species. Overall F _ST values for both species were significant, with an estimated average number of effective migrants per generation (N _EM) of 4.85 (~5 individuals) for L. laevigata and 3.54 (~4 individuals) for T. crocea. Within-region comparisons showed N_EM ranging from 6.29 to 92.34 for L. laevigata and from 3.40 to 6.30 for T. crocea. The higher gene flow among L. laevigata populations relative to T. crocea is consistent with the greater dispersal potential of the former species. Finer scale genetic structuring was evident in T. crocea populations. For both species, the Tubbataha reefs in the Sulu Sea have higher genetic affinity with the populations from the southern shelf of Palawan, while the reefs in the KIG had higher affinity with the northern Palawan shelf reefs. The north and south shelf populations have the least genetic affinity. Genetic patchiness among reefs within regions suggests the importance of small-scale physical factors that affect recruitment success in structuring populations in small island and shoal reef systems in Palawan.     

Ecological and physiological differences between two colour morphs of the coral Pocillopora damicornis

Pocillopora damicornis (Linnaeus) is a ubiquitous branching coral found throughout the Indo-Pacific region. Like many other species of coral, P. damicornis displays a large range of morphologies. At One Tree Island, it occurs as two distinct morphs that are easily distinguished by the presence or absence of pink pigmentation. The two colour morphs of P. damicornis were found to differ in their distribution and abundance in the One Tree Island Lagoon. The brown morph was more abudant than the pink morph in the shallows (<1 m),whereas the pink morph was more abundant at deeper sites (>3 m). The two morphs also differed physiologically. The brown morph tended to have a greater calcification rate than the pink morph, regardless of environmental conditions. However, the difference in the calcification rate between the two morphs became non-significant under shaded conditions (5% full sunlight), indicating some degree of physiological plasticity of the morphs. The pink colour in P. damicornis was due to a hydrophilic pigment with a major peak absorbance at 560 nm. The expression of pink pigment had both genetic and phenotypic components. The brown morph has a reduced genetic capacity to express the pigment relative to the pink morph. On the other hand, pigment expression could be induced by light in the pink morph. Although genetic differences ultimately determine the differences between the two morphs of P. damicornis, the extent of pigment expression is under some degree of environmental influence.     

Genetic differentiation between morphotypes of the marine sponge Suberites ficus (Demospongiae: Hadromerida)

Sponges of three morphotypes of Suberites ficus (Johnston, 1842) were collected during February and March 1985 off the south-west of the Isle of Man, and were compared by using spicule size distributions and genetic allele frequencies of isozyme loci. The populations did not show any significant differences of spicule size or type, but could be easily differentiated into three separate species based on isozyme patterns. Samples of pale orange S. ficus growing on gastropod shells inhabited by hermit crabs (Pagurus spp.) were reproductively isolated from the redorange and the pale yellow colour morphs encrusting the bivalve Chlamys opercularis. These latter two colour morphs were genetically similar, but significant differences were observed at two of the 19 gene loci assayed. All the sponges studied were sympatric, and therefore the genetic differences, indicating reproductive isolation, are strong evidence for separate gene pools and, hence, that they are different species. The genetic identity between the two colour morphs of S. ficus on C. opercularis shells was 0.977, whilst between each of these and S. ficus on hermit crabs it was about 0.65. In all three species genetic variability was high, with mean expected and observed heterozygosity values per locus ranging from 0.17 to 0.36.     

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.     

No apparent genetic basis to demographic differences in scarid fishes across continental shelf of the Great Barrier Reef

 Two species of parrot fish, Scarus frenatus and Chlorurus sordidus, are known to exhibit demographic and life-history differences across the continental shelf of the northern Great Barrier Reef (GBR). DNA sequences from the mitochondrial (mt) control region were analysed to determine whether there were genetic differences between the populations from the mid- and outer-shelf reefs. Analysis of molecular variance (AMOVA) indicated high levels of gene exchange for both species at a local scale between reefs on mid- and outer continental shelf positions (20 km apart) and at a broader scale along the length of the GBR province (>1000 km apart). There was no evidence to suggest that local differences in scarid life-history characteristics on the northern GBR have a genetic basis. Rather it appears more likely that phenotypically plastic responses to prevailing social and environmental conditions explain differences in the life-history characteristics of both taxa. However, analysis of genetic variation and historical demography revealed striking differences between the two species. S. frenatus haplotypes differed from one another at relatively few nucleotide sites (mean = 3.30), and the pairwise mismatch distribution suggested this species has undergone a population expansion within the limit of the resolution of the marker. C. sordidus haplotypes, however, differed from one another at a number of sites (mean = 7.67). Mismatch distribution analysis suggested that the population size of this species has remained at equilibrium over time. These patterns could also reflect differences in the metapopulation sizes or generation times between taxa. Some of the implications for fisheries management are discussed. Received: 28 January 2000 / Accepted: 9 July 2000     

Genetic structure of crown-of-thorns starfish (Acanthaster planci ) on the Great Barrier Reef, Australia: comparison of two sets of outbreak populations occurring ten years apart

Six populations of the crown-of-thorns starfish, Acanthaster planci, showing large increases in population size in 1994 to 1995, were examined by allozyme electrophoresis of 100 to 300 individuals collected from each population in November 1995 to February 1996. Analysis of nine loci, used to determine the structure of outbreak populations in the early-mid-1980s, demonstrated no significant differentiation among the 1996 outbreak populations in any age class (2 to 6 yr old). No significant variation was detected between age classes in any of the six populations. These data are consistent with the recent outbreak populations having been derived from one genetic source, and suggest no change in the source of recruits to these reefs between 1989/1990 to 1994/1995. In a multi-dimensional scaling analysis, the 1996 samples clustered with the eight 1986 outbreak populations in a small part of the genetic space spanned by the 1986 non-outbreak populations, suggesting that the outbreak populations were derived from the same genetic source in both 1986 and 1996. Statistically significant, but small, differences in gene frequencies detected between the 1986 and 1996 outbreak populations are thought to result from genetic drift, but are unlikely to provide a means of distinguishing sets of outbreak populations because of the large sample sizes required to detect the small shifts in gene frequency. Received: 21 March 1997 / Accepted: 2 April 1997     

Genetic structure of local populations and divergence between growth forms in a clonal invertebrate,the Caribbean octocoral Briareum asbestinum

Although the genetic structure of many populations of marine organisms show little deviation from panmixia, in those marine species with limited larval dispersal, patterns of microgeographic genetic differentiation may be common. The octocoral Briareum asbestinum should show local population differentiation because colonies reproduce asexually by fragmentation, most matings occur between colonies in very close proximity, and the sexually produced larvae and sperm appear to disperse only short distances. Variability in secondary chemistry of individual B. asbestinum colonies from different populations in close proximity also suggests local population differentiation. We determined the genetic composition of local populations by surveying allozyme variation of three shallow and two deep populations within a 300 m² area at San Salvador Island, Bahamas and at a site 161 km away on Little San Salvador, Bahamas in July 1990. As B. asbestinum occurs as either an erect branching form or an encrusting mat often at the same sites, we sampled both morphs to examine the extent of genetic exchange between them. Five of 21 loci were polymorphic and most populations showed a deficit of heterozygotes. Allele frequencies differed significantly between morphs at each site where they occurred together. The mean genetic distance (D=0.065) between morphs is consistent with the interpretation that the two morphs are genetically isolated. Despite the close spatial proximity of the San Salvador populations, both the branching and encrusting morphs showed significant genetic heterogeneity among neighboring populations. Similarly, pooled allelic frequencies for samples collected from the islands of San Salvador and Little San Salvador differed significantly at 1 locus for the branching morph and at 3 out of 5 loci for the encrusting morph.     

Microsatellite analysis of plaice (Pleuronectes platessa L.) in the NE Atlantic: weak genetic structuring in a milieu of high gene flow

Many species of marine fish are typified by large population sizes, strong migratory behavior, high fecundity, and pelagic eggs and larvae that are passively transported by ocean currents, all features that tend to increase gene flow, and hence reduce genetic partitioning, among localized populations. The plaice, Pleuronectes platessa, is a commercially important demersal species that exhibits all of these characteristics. We analyzed genetic variation at eight microsatellite loci in samples of spawning adults (N = 348) from the coasts of Ireland, Iceland, and, for the first time, from the Baltic Sea. Significant differentiation was observed between Iceland and Irish and Baltic Sea samples. However, there were no genetic differences between Irish and Baltic Sea samples, which contrast with the significant differentiation reported between Baltic Sea and North Sea/Atlantic populations of other flatfish species. To increase the data set, we carried out a cross-calibration exercise, allowing us to perform a joint analysis of data with an earlier study on adult and juvenile plaice (N = 480) collected over a broad geographic range, using six microsatellite loci in common to the two studies. Significant differentiation was observed between fish collected at the northern (Iceland, Faeroes, Norway) and southern (Bay of Biscay) parts of the species range. In contrast, the results showed little evidence of genetic structuring over much of the continental shelf of Europe. We believe that bathymetric and hydrographic barriers are the major factors shaping genetic structure, while lack of structure over much of the European continental shelf may be explained by a combination of past historical events, population structure, and dynamics of the species.     

A molecular approach to the ecology of a mussel (Mytilus edulis – M. trossulus) hybrid zone on the east coast of Newfoundland, Canada

Mytilus trossulus Gould and M. edulis L. coexist in mixed populations in Atlantic Canadian waters. Although there is evidence that the two species hybridize in natural populations and that hybrids produce progeny through backcrosses, no study of the microgeographic distribution of the two forms and their hybrids has been made. Here we examine subtidal samples of mussels taken in July 1997 from two locations in eastern Newfoundland (Canada) and from wave-exposed and protected environments within each location. Shell lengths ranged from 15 to about 90 mm. Mussels were classified as pure forms or hybrids (F₁, F₂ and from backcrosses) based on four diagnostic markers, two allozyme loci (Mpi and Est-D) and two nuclear PCR-based DNA markers (ITS and Glu-5). In addition, a PCR-based mtDNA marker (COIII) was used to characterize the distribution of mtDNA mitotypes among pure and hybrid individuals. There were differences in the proportions of pure M. edulis and M. trossulus between sites and between environments at one location. M. trossulus was the predominant species at one of the two exposed sites. In all four samples, M. trossulus was also the predominant form among small individuals. The frequency of hybrids was 26% overall and did not differ among samples. Hybrids consisted mostly of backcrosses that were M. trossulus-biased among small mussels and M. edulis-biased among large ones. We conclude that both intrinsic genetic factors and extrinsic environmental factors influence the relative frequency of M. edulis, M. trossulus and their hybrids. Received: 29 June 1998 / Accepted: 4 November 1998     

Genetic structure of dictyoceratid sponge populations on the western Coral Sea reefs

Allozyme variation at six polymorphic loci was examined in foliose dictyoceratid sponges from isolated reefs in the western Coral Sea. Four major genetic groups corresponding to the species Phyllospongia lamellosa, P. alcicornis, Carterospongia flabellifera and Collospongia auris were examined. A further two rare morphotypes from individual reefs formed genetic outliers to the P. lamellosa group, and may represent further taxa related to P. lamellosa. Gene frequencies in individual reef populations were largely in Hardy-Weinberg equilibrium, suggesting that random mating occurred in local populations of all four common species. Genetic variability was high and observed heterozygosities within populations ranged from 0.13 to 0.40. All four taxa showed significant genetic differentiation among populations (F _ST=0.05 to 0.36). Genetic distances (Nei's D) among populations within species ranged from 0 to 0.723 and increased with increasing geographical separation. There was evidence that genetic differentiation between populations to the north and to the south of the southern limit of the South Equatorial Current (SEC) divergence was greater than expected on the basis of their geographical separation. The SEC divergence may form a partial barrier to gene flow among populations of these ecologically important sponges on the submerged Queensland Plateau. Levels of migration among populations of three of the species was less than those required to prevent divergence of the populations through genetic drift (Nm<1). Restricted migration among populations may provide a mechanism to explain the occurrence of highly divergent populations of dictyoceratid sponges whose specific identity is not clear, and may allow them additionally to develop partial reproduction isolation from other populations.     

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     

Genetic differentiation of bicolor damselfish (Eupomacentrus partitus) populations in the Florida Keys

Electrophoretic variation in proteins encoded by 23 loci revealed substantial genetic differentiation among populations of bicolor damselfish (Eupomacentrus partitus) collected from four coral reefs in the Florida Keys, USA, during 1986–1988. Genetic differentiation was concentrated between a sample collected from Little Grecian Rocks Reef (LGR) and the remaining samples, including fish from a reef only 600 m distant (Grecian Rocks Reef). Genetic distinction of the LGR sample derived from significantly heterogenous allelic frequencies at six of eight polymorphic loci. Aco-1 (aconitase); Ada (adenosine deaminase); Gpi-2 (glucosephosphate isomerase); Ldh-2 and Ldh-3 (lactate dehydrogenase); and Me-1 (malic enzyme); nevertheless, differentiation at cytosolic aconitase (Aco-1) far exceeded that observed for other loci (fixation index, F _ST=0.482), and differences in Aco-1 allele frequencies were largely responsible for large genetic distances (0.20) between LGR and the other reefs. Paradoxically, estimates of numbers of migrants exchanged between reefs per generation (mN _e=17.47) indicated the potential for extensive gene flow. The extent of genetic differentiation among these populations is evaluated relative to models of population genetic structure based on equilibrium between gene flow and natural selection or genetic drift.     

Western Mediterranean immature loggerhead turtles: habitat use in spring and summer assessed through satellite tracking and aerial surveys

An aerial survey was conducted in early spring 2002 over the continental shelf of the Balearic Archipelago to study the distribution of neritic loggerhead turtles. Furthermore, five juvenile loggerhead turtles [straight carapace length (SCL) range 37.1–48.7 cm], were instrumented with transmitters and monitored during 2003 by satellite tracking to study habitat use over a broader geographical range. The distribution of the turtles over the continental shelf matched habitat availability, as defined by depth. However, those tracked by satellite spent most of the time in the oceanic waters of the Algerian basin and generally avoided continental shelf areas. In these turtles, average speed of travel and mean cosine of turning angle did not significantly differ between habitats, indicating that avoidance of shelf areas is not due to active habitat selection. On average, tracked turtles spent 35.1±19.7% of the time at the surface, although surface time was much greater in the turtle with the shortest carapace length, suggesting that this individual had limited swimming capacity. We conclude that the transition between passive drifting and active habitat selection occurs at an SCL of about 40 cm. The turtles followed tracks that matched prevailing currents, but on some occasions they also swam upstream. Hence, the distribution of late juvenile loggerhead turtles in the southern and central western Mediterranean may reflect a combination of passive drifting and active habitat selection.Communicated by O. Kinne, Oldendorf/Luhe     

Spatio-temporal patterns in the genetic structure of recently settled blue mussels (<Emphasis Type="Italic">Mytilus</Emphasis> spp.) across a hybrid zone

Previous studies of a hybrid zone between the mussels Mytilus edulis Linnaeus and M. galloprovincialis Lamarck have not resolved the relative importance of the genetic composition of settling larval cohorts versus post-settlement selection in determining the distribution of the parental species and their hybrids. In the present study, recently settled mussels (spat) were collected from 20 sites in southwest England throughout the summer and fall (May–October) in 1998 and 1999. This study investigated the spatio-temporal patterns of settlement and genetics of mussel spat by genetically identifying M. edulis, M. galloprovincialis and their hybrids using the diagnostic PCR marker Glu-5. Settlement was observed earlier in populations of M. edulis than in populations of M. galloprovincialis. Settlement occurred in hybrid populations at times intermediate to and overlapping with both of the parental populations. Temporal genetic variation within years was rare at most sites, while there was some variation between the two years. Spatial genetic variation, however, was common among spat settling within the hybrid populations and matched that observed in small, sub-adults at the same sites. No consistent directional changes in allele frequency were observed over the course of several weeks after settlement. These data suggest that the observed spatial variation in the adult populations is the result of spatial variation in settling larval cohorts and not of either temporal genetic variation or of selection soon after settlement.Communicated by J.P. Grassle, New Brunswick     

Limited gene flow in the brooding coral Favia fragum (Esper, 1797)

Understanding population connectivity in corals is particularly important as these organisms are increasingly threatened by abiotic and biotic factors. This study examined the population genetic structure of the brooding coral Favia fragum across four locations in the Caribbean and Western Atlantic using mitochondrial and nuclear markers. Morphological features were also compared to test whether genetic diversity corresponds with skeletal morphology. When comparing across distantly related Caribbean and Bermudian locations, F _ST values were high and significant, indicating strong genetic structure. At a local scale, significant genetic structure was found among reefs in Panama, while no genetic structure was found among reefs within Barbados, Bermuda or Jamaica. Surprisingly, a single haplotype for each of the three markers examined was found in Bermuda, where samples varied significantly from all other locations in three out of four morphological features analyzed. These data indicate that gene flow of F. fragum may occur locally among reefs but is highly restricted at distant locations. Furthermore, isolated populations, such as that of Bermuda, must be self-seeding to maintain the observed genetic uniformity.     

Rates of nitrogen fixation on coral reefs across the continental shelf of the central Great Barrier Reef

This study was undertaken in 1981 to determine whether there were major variations in potential rates of nitrogen fixation on apparently bare coralline substrate from reefs across the continental shelf of the central Great Barrier Reef. Nitrogen fixation, measured as rates of ethylene production (nmol cm^-2h^-1), was significantly lower on substrata from two inner-shelf reefs, (0.46 and 1.07) than on two middle-shelf reefs (2.10 and 2.97) and on two outer-shelf reefs (3.20 and 3.81). By contrast, algal biomass (mg cm^-2) on experimental substrate was significantly higher on inner-shelf reefs (80.8 and 59.4) than on middleshelf (27.1 and 23.8) and outer-shelf reefs (26.4 and 22.4). The rate of nitrogen fixation was positively correlated with the proportion of bare substratum and significantly higher concentrations of dissolved inorganic nitrogen were found in waters over the reefs than in water flowing onto those reefs. The abundance of algal-grazing fishes was reported previously to be significantly lower on inner-shelf reefs. It is suggested that this cross-shelf variation in the activity of algal-grazing fishes may be a determinant of the observed cross-shelf variations in potential nitrogen fixation.Contribution No. 233 from the Australian Institute of Marine Science     

Effects of extreme variation in female morph frequencies on the mating behaviour of male damselflies

Female-limited polymorphism is often attributed to selection to avoid excessive male mating attempts. It is encountered in various taxonomic groups, but is particularly common in damselflies, where one female morph (andromorph) typically resembles the conspecific male in colour pattern, while the other(s) (gynomorph(s)) do not. Two sets of theories have been proposed to explain the phenomenon in damselflies, which can be classified as the learned mate recognition (LMR) and male mimicry (MM) hypotheses. To test predictions of these hypotheses, we evaluated the rate of male sexual response towards female morphs and conspecific males in the damselfly Nehalennia irene. The LMR hypothesis predicts that males should respond sexually to andromorphs at greater rates in populations containing a higher relative frequency of andromorphs. The MM hypothesis predicts that males respond more often sexually to both andromorphs and males as the ratio of andromorphs to males increases. While LMR predicts that the rate of mating attempts towards gynomorphs should vary, the MM predicts that it should be relatively fixed. On experimentally presenting live specimens to focal males in five different populations with extreme variation in female morph frequencies, we observed that as the andromorph frequency and ratio of andromorphs to males increased, the proportion of male mating attempts increased on both andromorphs and males, whereas it decreased on gynomorphs. While the simplest form of the MM hypothesis is rejected, the results support specific predictions of both hypotheses and suggest that future studies should not treat these hypotheses as mutually exclusive.     

Population structure of the widely dispersing marine bryozoan Membranipora membranacea (Cheilostomata): implications for population history, biogeography, and taxonomy

Morphologically plastic, cryptic, or geographically widespread species pose similar challenges to the evolutionary biologist: their taxonomic status is often unclear yet must be known to study almost any aspect of their biology, ecology, evolution, or biogeography. The marine bryozoan Membranipora membranacea (L.) is morphologically plastic and geographically widespread in temperate oceans of the Northern and Southern Hemispheres, and its taxonomy is unclear. This study examined genetic relationships among allopatric populations and sympatric morphs of this species, or species complex. Colonies were collected from 1992 to 1995. Allozymes were used to elucidate the relationships among four widely separated populations, two in the North Atlantic and two in the North Pacific Ocean. Allozymes and mtDNA sequencing were used to clarify the genetic relationships among three sympatric morphs that might correspond to the species M. villosa Hincks and M. membranacea in the northeastern Pacific (Washington State). Populations in the North Atlantic and North Pacific had no fixed allelic differences at the loci tested but were separated by an average Nei's genetic distance of 0.581, suggesting their near-sibling species status. Populations from Friday Harbor (Washington) and Catalina Island (California) were not significantly differentiated, which was attributed to high gene flow. Populations on either side of the North Atlantic were genetically indistinguishable, which is most likely due to the recent establishment of the West Atlantic populations from European founders. At Friday Harbor, sympatric morphs varying in their spination and spine inducibility were genetically indistinguishable, supporting the hypothesis that M. villosa is an induced phenotype of M. membranacea and not a distinct species in the northeastern Pacific. Since such phenotypic plasticity is common in cheilostome bryozoans, the morphospecies concept must be used with caution. Received: 31 August 1998 / Accepted: 10 August 1999