Auditory and olfactory abilities of pre-settlement larvae and post-settlement juveniles of a coral reef damselfish (Pisces: Pomacentridae)

The propagules of most species of reef fish are advected from the reef, necessitating a return to reef habitats at the end of the pelagic stage. There is increasing evidence of active attraction to the reef but the sensory abilities of reef fish larvae have not been characterized well enough to fully identify cues. The electrophysiological methods of auditory brainstem response (ABR) and electroolfactogram (EOG) were used to investigate auditory and olfactory abilities of pre- and post-settlement stages of a damselfish, Pomacentrus nagasakiensis (Pisces, Pomacentridae). Audiograms of the two ontogenetic stages were similar. Pre-settlement larvae heard as well as their post-settlement counterparts at all but two of the tested frequencies between 100 Hz and 2,000 Hz. At 100 and 600 Hz, pre-settlement larvae had ABR thresholds 8 dB higher than those of post-settlement juveniles. Both stages were able to detect locally recorded reef sounds. Similarly, no difference in olfactory ability was found between the two ontogenetic stages. Both stages showed olfactory responses to conspecifics as well as L-alanine. Therefore, the auditory and olfactory senses have similar capabilities in both ontogenetic stages. Settlement stage larvae of P. nagasakiensis can hear and smell reef cues but it is unclear as to what extent larvae use these sounds or smells, or both, as cues for locating settlement sites. An erratum to this article can be found at     

Pleistocene diversification of the Pomacentrus coelestis species complex (Pisces: Pomacentridae): historical biogeography and species boundaries

Pleistocene eustatic changes in sea level are often invoked to explain genetic divergence among marine organisms. However, molecular phylogenies have revealed relatively few examples of speciation events dating to the Pleistocene. We present a species-level hypothesis of the timing of evolution for the Pomacentrus coelestis species complex (Pomacentridae), based on the nuclear S7 intron and the mitochondrial Cytb gene, and reconstruct ancestral range distributions across the timetree. Ancestral range reconstruction suggests the complex originated in the Coral Triangle and East Indian Ocean, with subsequent range expansion outward from this region. We suggest that land barriers to dispersal (e.g., Indo-Pacific barrier) may be responsible for the divergence between Indian (P. alleni, P. similis, P. caeruleopunctatus, and P. caeruleus) and Pacific (P. micronesicus, P. auriventris, and P. coelestis) species groups, and subsequent isolation by Pleistocene sea-level fluctuations in certain areas of the Coral Triangle (glacial refugia) may play an important role in the diversification of this species complex. Additionally, our analyses show cryptic lineages within P. micronesicus and highlight the need for comprehensive sampling within and among species to reveal recent speciation events.     

Genetic relationships of the colour morphs of Acanthochromis polyacanthus (Pomacentridae) on the northern Great Barrier Reef

Acanthochromis polyacanthus is an unusual tropical damselfish because it lacks pelagic larvae and has lost the capacity for broad-scale dispersal among coral reefs. On the modern Great Barrier Reef, this fish has multiple colour morphs and we have previously described steep genetic clines in the central region between three colour morphs. A similar transition of colour morphs has been found in the far northern region (Latitude 11°S), where at least five colour morphs and intergrading hybrids were found during a survey of 14 locations on nine adjacent reefs. On two of these reefs, monochromatic white and bicoloured morphs coexist, partitioning the reef slope vertically when sympatric, but occupying the full depth range elsewhere. In areas of overlap, they interbreed, producing hybrids of intermediate colour. Although the discriminatory power of our genetic analysis was weakened by the low level of variability in these populations (only four polymorphic loci), allozyme electrophoresis revealed significant genetic differentiation within and among reefs. There was neither simple linkage between colour and protein variation, nor correlation between genetic similarity and proximity of sampled populations. We suggest that the observed chaotic structure is the result of differences among the founding populations, although the two colour-based stocks have been homogenized through a long history of introgression during multiple episodes of isolation and secondary contact. Despite genetic mixing, phenotypic diversity (i.e. colour morphs) seems to have been preserved in these populations by sexual selection, perhaps reinforced by natural selection. The long-term stability of this outcome, however, is suggested by the presence of unique morphs on two reefs that are detached from the continental shelf and surrounded by very deep water. The strong identity of these populations can be attributed to long occupation of these isolated sites compared with those on the continental shelf which have only been established in the last 15 000 yr and still retain differences arising from the initial founder events. Received: 15 January 1997 / Accepted: 15 August 1997     

Planktonic larval duration of one hundred species of Pacific and Atlantic damselfishes (Pomacentridae)

The plankton larval duration for 100 species of Pacific and Atlantic damselfishes was estimated from daily growth increments on the otolith of juvenile fish collected at various localities between July 1987 and September 1988. For newly-settled fishes, larval duration was determined by counting the entire number of increments present on the otolith, while for older juveniles estimates were made by counting the number of increments between the center of the otolith and a mark corresponding to settlement. We document the development of otolith formation during the period when eggs are incubated on the reef and show that daily increments are only accreted after larvae hatch and enter the planktonic phase. The planktonic larval duration for damselfish is shorter and less variable, both between and within species, compared to other groups of reef fishes such as wrasses and surgeonfishes. Larval duration ranged from 12 to 39 d. Average duration between species ranged from 13.1 to 35.2 d. The time spent in the plankton was not significantly correlated with geographic distribution when evaluated among species, however, genera with confined regional distribution have a shorter mean larval life than do widely distributed genera. Size at settlement was positively correlated with time spent in the plankton among species, but a significant correlation between these variables was only evident within one of ten species. The low variance in planktonic larval duration within species indicates that most damselfish are unable to delay metamorphosis following competency. This inability to postpone settlement limits the potential for dispersal, especially when dispersal time between suitable habitats is greater than about 30 d.     

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

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

Genetic consequences of long larval life in the starfish Linckia laevigata (Echinodermata: Asteroidea) on the Great Barrier Reef

Gene flow between populations of the asteroid Linckia laevigata (Linnaeus) was investigated by examining over 1000 individuals collected from ten reefs throughout the Great Barrier Reef (GBR), Australia, for genetic variation at seven polymorphic enzyme loci. Despite geographic separations in excess of 1000 km, Nei's unbiased genetic distance (0 to 0.003) and standardised genetic variation between populations (F _ST) values (mean 0.0011) were small and not significant. Genetic homogeneity among L. laevigata populations is consistent with the long-distance dispersal capability of its 28 d planktonic larval phase, and is greater than that observed for other asteroid species, including another high-dispersal species, Acanthaster planci, which has a 14 d larval phase. Variation within populations was also higher than previously recorded for asteroids (mean heterozygosity=0.384; number of alleles per locus ranged from 5.1 to 6.0 in each population). Among asteroids, dispersal ability is positively correlated with gene flow and levels of variation, and negatively correlated with levels of differentiation.     

Zooplankton community structure and water flow in the lee of Helix Reef (Great Barrier Reef, Australia)

To investigate which physical processes contribute most in moulding zooplankton community structure in the waters close to coral reefs, light traps moored in a grid pattern were used to collect zooplankton from the sea surface at 16 stations on the downstream side of Helix Reef during three time periods (2100-2200, 2400-0100, and 0300-0400 hours) over three consecutive nights covering the new moon period in January 1992. Two distinct zooplankton communities were present: a community composed primarily of reef-resident, demersal plankton immediately to the south of the reef in an area of reduced flushing, and a community containing coastal and shelf-seas taxa at the more exposed sites in the open flow field. The fauna composition at a number of exposed stations was as rich as that at sheltered stations both in terms of number of taxa and diversity indices but was almost an order of magnitude less abundant. The reef-resident, demersal plankton community was dominated by gammarid amphipods, mysids, and polychaetes, whereas only transient, meroplanktonic forms such as echinoderm and echinopluteus larvae and shelf-seas, holoplanktonic forms such as doliolids and larvaceans were significantly more abundant in the exposed community. Zooplankton associations were apparently formed by a combination of hydrodynamic processes, spatial and temporal distribution patterns of individual taxa, specific behaviours of certain taxa, and the interactions among taxa at different trophic levels.     

Occurrence and characteristics of unusual protistan symbionts from surgeonfishes (Acanthuridae) of the Great Barrier Reef,Australia

The occurrence of unusual symbiotic microorganisms was examined in the intestines of a range of fish from the Great Barrier Reef, Australia. The fish taxa examined included 26 species of the family Acanthuridae, as well as representatives of phylogenetically related and herbivorous taxa. The microorganisms, referred to as protists, were only found in herbivorous and detritivorous members of the Acanthuridae. Protists were not found in planktivorous acanthurids, nor in any members of the families Kyphosidae, Pomacentridae, Scaridae, Zanclidae, Siganidae and Bleniidae we examined. In addition, protists were absent from the herbivorous acanthurids A. xanthopterus and A. nigricans. A range of protist forms, characterized by differences in size (8 to 417 m), shape and mode of cell division (daughter-cell production and binary fission), was observed. The occurrence of these forms appeared to be correlated with host feedingecology. Large forms (>100 m) of the protists were only found in acanthurids which fed over hard-reef substrata. Smaller forms were found in sand-grazing and detritivorous species. One of the protist forms appears identical to protists previously reported from Red Sea acanthurids.     

Locomotion performance of green turtle hatchlings from the Heron Island Rookery,Great Barrier Reef

Locomotion performance plays a vital role in determining hatchling green turtle Chelonia mydas survival in the first few hours after emerging from their nests as hatchlings crawl and swim the gauntlet of predators before reaching the relative safety of the open ocean. Previous laboratory based constant incubation experiments found incubation temperature to influence the size and swimming performance of hatchling green turtles. Here we examine the morphology and crawling and swimming performance of hatchling green turtles as they emerge from nests on Heron Island rookery in the southern Great Barrier Reef to test the hypothesis that nest temperature in the field can influence these attributes. We found inter-nest differences in hatchling mass and dimensions, and that hatchling mass was not correlated with nest temperature. However, hatchlings from warmer nests had smaller carapace dimensions than hatchlings from cool nests suggesting that more yolk was converted to hatchling tissue during embryonic development in cool nests. There was considerable intra- and inter-nest variation in both crawling and swimming performance of hatchlings. Hatchlings from cool nests tended to be faster crawlers than hatchlings from warm nests, but the thrust produced during swimming was not correlated with nest temperature. During the 4 h swimming trial, hatchlings swimming effort decreased significantly during the first 3 h but swimming effort remained relatively constant for the last 1 h. Individual hatchling crawling and swimming performances were not correlated with each other. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Vertical distribution of fish larvae in the Great Barrier Reef Lagoon,Australia

To investigate the natural defenses of Antarctic marine organisms against exposure to ultraviolet (UV) radiation (280 to 320 nm), 57 species (1 fish, 48 invertebrates, and 8 algae) were collected during austral spring 1988 in the vicinity of Palmer Station (Anvers Island, Antarctic Peninsula) and were analyzed for the presence of mycosporine-like amino acids (MAAs), compounds that absorb UV radiation and may provide shielding from these biologically hazardous wavelengths. Nearly 90% of the 57 species examined contained MAAs, and eight specific MAA compounds were identified. Seven of these (palythine, porphyra-334, shinorine, mycosporineglycine, palythene, asterina-330, and palythinol) have been observed previously in marine organisms from temperate and tropical latitudes. A new MAA, mycosporineglycine: valine, was found in the Antarctic fish and in 38 of the invertebrate species examined. This study confirms widespread occurrence of MAAs in Antarctic marine organisms and suggests that these species have some degree of natural biochemical protection from UV exposure.Contribution No. 502 from the Australian Institute of Marine Science     

Spatial,temporal, and behavioral patterns in emergence of zooplankton in the lagoon of Heron Reef,Great Barrier Reef,Australia

A total of 34 zooplanktonic taxa were common in emergence trap, reentry trap, and net-tow samples taken in the lagoon of Heron Reef, Great Barrier Reef, between 27 February and 22 March and between 11 June and 4 July 1985. Twenty-nine of these taxa were classified as demersal (17 taxa), meroplanktonic and larval (10 taxa), or incidental (2 taxa). The remaining five multispecific groups yielded variable results. Differences were observed between two locations separated by 200 m, with emergence 2 to 62 times greater for 22 taxa at a deeper site with larger coral formations. Most zooplankters were more common (2 to 122 times) in samples from 1 m² areas around 0.25 to 0.5 m² patches of branching coral. However, two species of copepods, Pseudodiaptomus colefaxi and Metis holothuriae, were taken in greater numbers (3 to 12 times) from open sand. Seasonal increases (2 to 322 times) were observed for 13 adult taxa and 8 classes of larvae or juveniles in the summer and for 10 adult taxa and one larval group in the winter. Fourteen of the 17 demersal taxa and 4 groups of large larvae or juveniles emerged in numbers 2 to 323 times greater during lunar quarters or new moons. In contrast, 6 larval taxa and 4 groups of small or transparent adults displayed significant emergence during full moons. Five diurnal emergence patterns were shown by 27 taxa, with patterns varying primarily among lunar periods. The variety of taxon-specific patterns observed in this study highlight a need for caution when generalizing about demersal zooplankton.     

Sperm ultrastructure of the giant clam Tridacna maxima (Tridacnidae: Bivalvia: Mollusca) from the Great Barrier Reef

Using transmission electron microscopy, spermatozoa from a member of the Tridacnidae, or giant clams, are described for the first time and compared with spermatozoa of other bivalves, especially other heterodonts. The acrosomal vesicle of Tridacna maxima (Röding, 1798) is short (0.37?μm), blunt-conical, and exhibits a prominent basal ring. A narrow apical elaboration of the nucleus, the nuclear peg, projects deep into the basal invagination of the acrosomal vesicle. Aside from this specialization, the nucleus is a solid elongate-cylindrical structure (7.66?μm) that exhibits several small irregular lacunae. Four or occasionally three round-ovate mitochondria surround a pair of orthogonally-arranged, triplet-substructure centrioles. The proximal centriole is connected to a small indentation of the nuclear base by a thin layer of granular pericentriolar material, whereas the distal centriole is anchored to the plasma membrane by nine terminally-forked satellite fibres. The 9?+?2 pattern axoneme of the tail is continuous with the distal centriole. Comparison with other bivalves indicates a very close relationship between tridacnids and cardiids based on sperm ultrastructure. Specifically, the presence of a nuclear peg links Tridacna spp. with the cardiid genus Cerastoderma, but further information on the many unstudied genera is required to test the exact nature of this relationship. The sperm ultrastructure provides additional support for the recently proposed hypothesis that the Tridacnidae may be no more than a specialized subfamily of the Cardiidae.     

Vertical profiles of bacterial abundance,productivity and growth rates in coastal sediments of the central Great Barrier Reef lagoon

Vertical patterns of bacterial densities, productivity and specific growth rates in coastal muds, quartz sands and muddy sands of the central Great Barrier Reef lagoon were examined in summer (February) and autumn (May) 1988. Variations in these parameters with station location, sediment depth and season were complex, exhibiting significant main and interaction effects in most instances. Some trends were apparent despite the large and complex variations. Bacterial densities did not vary seasonally, ranging from 2.9 to 38.1×10⁹ cells g^-1 dry wt, averaged over sediment depth (0 to 20 cm) and seasons. Trend analysis revealed that densities decreased with increasing sediment depth. Bacterial production (tritiated thymidine incorporation into DNA) was high, ranging from 0.4 to 5.7 gCm^-2 d^-1 (integrated over 10 cm depth), as were specific growth rates (grand mean, =0.25 d^-1; range=0.004 to 1.3 d^-1). Both were generally higher in summer than in autumn. Vertical profiles of productivity and specific growth rates revealed actively growing bacterial assemblages down to 20 cm depth. Factors which may account for these very abundant and productive communities are: (1) subsurface accumulations of detritus exported from adjacent mangrove forests, and (2) physical disturbance from tidal scouring and severe climate (e.g. cyclones, wet-season floods). Disturbance events occur frequently enough to inhibit the development of highly sulphidic conditions, but stimulate production of bacterial types (aerobes, fermenters) capable of incorporating labelled thymidine into their DNA.     

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.     

Karyotypes in two species of the genusHippocampus (Pisces: Syngnatiformes)

Diploid modal numbers of chromosomes 2n=48 forHippocampus ramulosus and 2n=44 forH. hippocampus, collected in the Gulf of Palermo in May 1985, are established. No sex-linked heteromorphic pairs are observed in the two species analyzed. The karyotype ofH. hippocampus seems to be a derived condition.     

Patterns of genetic subdivision in populations of a clonal cnidarian,Zoanthus coppingeri,from the Great Barrier Reef

Samples of an intertidal zoanthid, Zoanthus coppingeri, Haddon and Shackelton, 1891, were collected from three localities in the Great Barrier Reef region during 1992–1993, and subjected to allozyme electrophoretic analysis at seven polymorphic loci. The reduced ratio of observed to expected genotypic diversity indicated that populations were partly clonal, but they were not dominated by a few clones as occurs in some other cnidarians. Regular disturbance by wave action is postulated to prevent the formation of large stands of particular clones by clearing space and mixing genotypes over small scales. The sexual origin of clonal genotypes was confirmed by conformance to Hardy-Weinberg predictions of genotype frequencies at all but one locus. Values of the standardised genetic variance among populations, F _ST , were highly significant between localities and between replicate sites within localities separated by only 50 m. Strong genetic structure has not previously been described in a Great Barrier Reef invertebrate species, and is considered to be the consequence of stochastic changes in gene frequencies as a result of low levels of gene flow. High clonal longevity and low recruitment rates may maintain genetic differences over long periods. Similar effects may be seen in other Great Barrier Reef invertebrate species with comparable reproductive patterns.     

Zoning,a fundamental cornerstone of effective Marine Spatial Planning: lessons learnt from the Great Barrier Reef,Australia

The Great Barrier Reef Marine Park was established to provide for conservation and ecologically sustainable multiple use of 344,400 km² of a large marine ecosystem. Management is based on multiple use, with zoning as a fundamental component of marine spatial planning. The legislative framework, including a specific Act and Regulations, address the objectives of ecosystem-based, integrated management of human uses and impacts consistent with best contemporary understanding of biological diversity. Zoning is one of a suite of management tools that include other spatial and temporal management tools and non-spatial measures including public education, community engagement, codes of environmental best practice, industry partnerships and economic instruments. The first section of the Great Barrier Reef Marine Park came into operation in 1981 and the most recent zoning came into operation in mid 2004. The paper discusses some common misunderstandings about zoning and identifies lessons that appear relevant for others addressing management and use of marine ecosystems and natural resources.     

Effects of autotomy on long-term survival and growth of painted spiny lobster (Panulirus versicolor) on the Great Barrier Reef, Australia

The effects of autotomy (shedding of appendages) on survival and growth rates of painted spiny lobster were investigated at Northwest Island (23^° 18?? S, 152^° 43?? E) during the period 2003?C2006. Adult lobsters were captured, tagged, and classified as either uninjured (n?=?68), minimally injured (n?=?39) or moderately injured (n?=?19) depending on the number and type of appendages that were autotomized during capture and handling. Six to thirty-six months after release, 86 lobsters were recaptured (mean time at large?=?305?days). Recapture rates of uninjured (64.7%), minimally injured (71.8%), and moderately injured lobsters (73.7%) were not significantly different. Similarly, mean annualized growth rates of uninjured, minimally injured, and moderately injured lobsters were not significantly different. This suggests that the energetic cost of a single episode of autotomy is either negligible or exists as a trade-off with some other life history trait, such as reduced reproductive performance. These results support the use of certain management tools (e.g., size limits) that prescribe release of non-legal lobsters, regardless of their injury status.