Large-scale spatial variation in epilithic algal matrix cryptofaunal assemblages on the Great Barrier Reef

Spatial variation in the epilithic algal matrix (EAM) cryptofauna was investigated at three locations on the Great Barrier Reef: two inner shelf—Orpheus Island and the Turtle Island group—and a mid-shelf location, Lizard Island. Although the EAM appears to be a relatively simple and consistent habitat, significant differences in cryptofaunal assemblages were found between locations. EAM assemblages from Orpheus Island were markedly different. This appears to be a function of the sediment profile characteristics (grain size >60 µm) at Orpheus Island, as many cryptofaunal taxa displayed positive relationships with sediment volume. However, sediment volumes did not differ significantly between the three locations, highlighting the possibility of cyclonic activity affecting the sediment profile at Orpheus Island in the months preceding the study, in addition to the nutrient input from major terrigenous sources. The results of this study show that EAM cryptofaunal assemblages are not uniform across the Great Barrier Reef and suggest that dissolved nutrients, sediment loads and distance from river systems may be significant drivers of cryptobenthic faunal compositions.     

The distribution of herbivorous fishes on the Great Barrier Reef

The composition and functionality of ecologically important herbivorous fish assemblages were examined throughout much of Australia’s Great Barrier Reef (GBR). Diversity and abundance of surgeonfishes (Acanthuridae), parrotfishes (Labridae) and rabbitfishes (Siganidae) were strongly associated with position on the continental shelf, whilst effects of latitude were weaker and inconsistent. Species distributions varied considerably amongst taxonomic groups; parrotfishes were mostly widespread whilst distributions of surgeonfishes were often restricted. Most inshore environments supported depauperate herbivore assemblages dominated by different taxa and functional groups compared with assemblages in offshore environments. There were also strong cross-shelf transitions in the main taxa performing each functional role. Overall, this study show that the functional contributions of herbivorous fish assemblages to important ecosystem processes and the contributing taxa vary considerably amongst different GBR environments. Additionally, the two most numerically dominant herbivores actively select detritus, not algae, supporting increasing evidence for the importance of detritus in coral reef ecology.     

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.     

Estimates of adult and juvenile mortality for labrid fishes at One Tree Reef,Great Barrier Reef

Few estimates of natural mortality have been reported for coral reef fishes, yet this information is essential for predicting the effects of recruitment fluctuations on adult populations. In this study, mortality of adult (10 species) and juvenile (11 species) labrid fishes resident on ten isolated patch reefs in One Tree Lagoon, southern Great Barrier Reef, was estimated by visual censuses covering 30 mo, from November 1981 to April 1984. Numbers of adults of all species declined linearly over this period. Mortality rates differed significantly among species, ranging from 4.9 to 69.5% per annum. Average life-span for fishes that had survived the first year of life varied from 1.6 to 11.5 yr, depending on species. Average juvenile mortality varied among species from 25 to 91.8% per annum and was highest for abundant schooling species. Different cohorts of the same species also experienced significantly different rates of mortality. No clear relationship emerged between the number of new recruits and the number of fishes surviving to one year of age. The influence of recruitment fluctuations on adult populations of labrid fishes may be difficult to predict.     

Diversity,abundance, and distribution of reef sharks on outer-shelf reefs of the Great Barrier Reef,Australia

Quantifying the distribution and habitat use of sharks is critical for understanding their ecological role and for establishing appropriate conservation and management regimes. On coral reefs, particularly the Great Barrier Reef (GBR), little is known regarding the distribution of sharks across major reef habitat types. In this study, we surveyed shark populations across outer-shelf reefs of the GBR in order to determine the diversity, abundance, and distribution of reef sharks across three major coral reef habitats: (1) the reef slope, (2) the back reef and (3) the reef flat. Model selection revealed that habitat was the principal factor influencing shark distribution and abundance. Specifically, overall shark abundance and diversity were significantly higher on the reef slope (and to a lesser degree, the back reef) than the reef flat. This confirms that shark populations are not homogeneously distributed across coral reefs. Thus, the results presented herein have important implications for shark population assessments. In addition, our results highlight the potential importance of the reef slope, with high levels of live coral cover and structural complexity, for sustaining reef shark populations. As this habitat is highly susceptible to disturbance events, this study provides a useful context for predicting and understanding how environmental degradation may influence reef shark populations in the future.     

Vertical and horizontal distribution of fish larvae near coral reefs at Lizard Island,Great Barrier Reef

Consistent patterns of horizontal distribution of fish larvae from plankton tows were found in shallow waters around Lizard Island, Great Barrier Reef during 1979 and 1980. Few types of larvae were most abundant in Lizard Lagoon, and none of these were old larvae. Forty percent of the 57 types of larvae studied differed in abundance between windward and downwind sides of the island. More types of old larvae were found in greatest abundance off the windward side of the island than the downwind side. Most types of larvae preferred deeper water (>3 m) during the day and moved upward at night, although a few types preferred upper (<3 m) or middle portions of the water column. These latter were more likely to descend at night or to maintain their day-time distribution than to move upward. Windward larvae [those more abundant off the windward (SE) side of the island] were more shallow-living than were downwind larvae, and were more likely to maintain their day-time distribution at night. The current patterns around Lizard Island were favourable for retention of larvae in both Lizard Lagoon and off the windward side of the island, if combined with certain vertical distributions of the larvae. However, while there was evidence for retention on the windward side of the island, there was no evidence for retention in Lizard Lagoon. Currents on the downwind side of the island were not favourable for retention of larvae and there was little evidence that larvae were retained there. Retention may be an accidental result of interaction between currents and larval behaviour, or the result of a strategy of retention by the larvae. These could not be distinguished in the present study.     

The use of clear-water non-estuarine mangroves by reef fishes on the Great Barrier Reef

Within the tropics, mangroves and coral reefs represent highly productive biomes. Although these habitats are often within close proximity, the role and importance of mangrove habitats for reef fish species remains unclear. Throughout the Indo-Pacific, reef fish species appear to have few links with estuarine mangrove habitats. In contrast, clear-water non-estuarine mangrove habitats throughout the Caribbean support many reef fish species and may be fundamental for sustaining reef fish populations. But how important are clear-water non-estuarine mangroves for reef fishes within the Indo-Pacific? Using visual surveys during diurnal high tide, the fish assemblages inhabiting clear-water mangrove and adjacent reef habitats of Orpheus Island, Great Barrier Reef, were recorded. Of the 188 species of fishes that were recorded, only 38 were observed to inhabit both habitats. Of these, only eight were observed more than five times within each habitat. These observations provide little indication that the clear-water mangroves are an important habitat for reef fish species. In addition, although based on just a 3-month survey period, we found little evidence to suggest that these areas are important nurseries for reef fish species. The clear-water mangroves of Orpheus Island may, however, provide an additional foraging area for the few reef fish species that were observed to utilize these habitats during high tide. The difference in the importance of clear-water mangroves for reef fishes within this study compared with clear-water mangrove counterparts within the Caribbean is surprising. Although only preliminary, our observations would support suggestions that the patterns reflect the different hydrological characteristics and evolutionary histories of these two biogeographic regions.     

Composition of the near-reef zooplankton at Heron Reef,Great Barrier Reef

Using a light trap, zooplankton was sampled at three stations at Heron Reef, Great Barrier Reef: (a) a typical patch reef in the Heron lagoon, (b) a site in 8 m water on the southern slope of Heron reef, and (c) a station approximately 300 m south of (b), in the open water of the channel between Heron and Wistari reefs. Samples were taken at the surface and on the substratum at the lagoon and reef-slope stations, and at 3 to 6 m depth at the open-water station. A total of 114 taxa, many recognized as species, were distinguished in the samples. Pronounced differences existed in abundance, diversity, and taxonomic composition of the samples obtained at different stations. Less pronounced differences existed between surface and substratum samples from the same station. Near-reef samples were more similar to one another than to open-water samples. Decapod larvae, amphipods, and cumaceans were all abundant in near-reef samples and very rare in open-water samples. Forams, isopods, mysids and polychaetes were common in near-reef samples, and rare or absent in open-water samples. Copepods were abundant in all samples but the near-reef samples contained predominantly different species than did samples from the open water. The near-reef fauna included 66 taxa which did not occur in open-water samples. Many of these were epibenthic rather than strictly planktonic in behaviour.     

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 herbivory on zonation of Sargassum spp. within fringing reefs of the central Great Barrier Reef

A combination of small-scale transplants and herbivore exclusion was used to test the importance of herbivory, physiological tolerance limits, and recruitment and dispersal in regulating the distribution and abundance of the genus Sargassum on two nearshore fringing reefs of the central Great Barrier Reef, during 1992/1993. Sargassum (predominantly S. oligocystum and S. tenerrimum) were transplanted from reef-flat zones where they normally grow, to a seaward coral zone where they are not normally found. At Great Palm Island, coral-zone transplants only survived if protected from herbivores. At Brook Island, survival of uncaged coral-zone transplants was more variable but not significantly lower than plants returned to the Sargassum zone. Thus herbivory may be a major cause of the zonation patterns of adult Sargassum on these fringing reefs, but the importance of this factor varies between and within reefs. Since protected Sargassum survived and grew for up to 6 mo in the coral zone, the adult algae are not physiologically limited by any physical or chemical differences between zones. However, Sargassum recruitment to the coral zone was very low (mean 2.7 recruits m⁻² over 13 mo), and was not significantly affected by herbivores. Since rates of herbivory were relatively slow, effective exclusion of Sargassum from the coral zone by herbivores may depend on low recruitment of the algae. In a broader context, the distribution of Sargassum may depend on the combined spatial patterns of herbivory and recruitment. Received: 24 January 1997 / Accepted: 12 May 1997     

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     

Variation in emergence of parasitic and predatory isopods among habitats at Lizard Island, Great Barrier Reef

Gnathiid isopods are one of the most abundant groups of ectoparasites on coral reef fishes. They, and other isopods, have been shown to significantly affect the health and behaviour of many reef fish. Whether isopod emergence differs among habitats on coral reefs is not known. In this study, we measured emergence rates of parasitic isopods (Gnathiidea and Flabellifera) in six habitats at two sites at Lizard Island during new moon periods in March and December 2004. Isopods were collected from the periphery and centres of micro-reefs, patch reefs, continuous reefs, and from inter-reefal habitats (sand or rubble) with 1 m² emergence traps. Sites (Casuarina and Coconut Beach) were located on opposite sides of Lizard Island. Live gnathiids were collected with light traps in November 2005 to investigate species differences between sites. At both sites, the most abundant gnathiid species was exclusive to that site. More gnathiid larvae emerged at night, and emergence of fed gnathiids (pranizae) and flabelliferan isopods was almost exclusively nocturnal. Diurnal emergence was greater at Coconut Beach than Casuarina Beach. Although emergence counts were not consistently affected by parameters such as habitat, site, or sampling period, gnathiid size and feeding state were. Where significant differences existed, gnathiids were larger and more often fed over reef borders than centrally. We suggest first stage larvae (Z1) have the largest influence on total abundance and are patchily distributed in accordance with adults from which they have recently hatched. As later stage larvae depend on fish, more successful (fed) and older larvae are found on the edges of reefs where appropriate hosts may be more abundant, or predation is lower. Gnathiids were over-dispersed in all habitats investigated, including apparently homogeneous beds of coral rubble and sand. This indicates that their distributions may be better predicted by very fine scale differences in substrate or that aggregations are the result of gregariousness and may be difficult to predict on the basis of substrate. Emergence traps collected comparatively few parasitic flabelliferan isopods. This community differed greatly from the previously described community of scavenging isopods at Lizard Island. These differences are probably the result of differences in trapping methodology.     

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.     

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     

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.     

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.     

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     

Among-habitat variation in herbivory on Sargassum spp. on a mid-shelf reef in the northern Great Barrier Reef

Herbivory is widely acknowledged as a key process determining the benthic community structure and resilience of coral reefs. Despite numerous studies that have examined herbivory across reef gradients in the Caribbean, few studies have directly quantified this process on Pacific reefs. Bioassays of two species of erect macroalgae (Sargassum swartzii and S. cristaefolium) were used to quantify variation in grazing intensity across seven habitats of varying depth and wave exposure on a mid-shelf reef in the northern Great Barrier Reef. Removal rates of Sargassum varied significantly among habitats, with both species displaying broadly similar patterns. The shallow habitats on the exposed aspect of the reef (i.e. reef crest, flat and back reef) experienced the highest reductions in mass (81.4–91.6% day⁻¹) for both S. swartzii and S. cristaefolium, while the deeper exposed habitats (reef slope and base) displayed the lowest reductions (3.8–13.4% day⁻¹) over a 24 h period. In contrast, the grazing intensity varied between the two species in the three habitats on the leeward aspect of the reef. Reductions in mass remained relatively high for S. swartzii on the patch reef and sheltered reef base and flat (62.7–76.5% day⁻¹) but were considerably lower for S. cristaefolium (37.9–63.5% day⁻¹) across the same habitats. Surprisingly, the rates of removal of Sargassum displayed no relationship with the density or biomass of roving herbivorous fishes or those species known to consume erect macroalgae, either collectively or independently. These results suggest that the relationship between browsing rates and herbivorous fish biomass is complex and may be driven by species that are underestimated in visual surveys. Direct quantification of browsing intensity using assays revealed a different pattern to inferences based on herbivore densities and highlights the potential difficulties of evaluating ecosystem processes based on visual census data alone.     

Reproduction in five species of crinoids at Lizard Island,Great Barrier Reef

Reproduction in five species of crinoids [Himerometra bartschi (A. H. Clark), H. robustipinna (P. H. Carpenter), Cenometra bella (Hartlaub), Colobometra perspinosa (P. H. Carpenter), and Oligometra serripinna (P. H. Carpenter)] was studied at Lizard Island, Queensland, over 24 mo, from April 1981 to March 1983 (except for O. serripinna —June 1981 to March 1983). All five species of crinoids were dioecious, although a few instances of synchronous hermaphroditism were noted in H. bartschi, C. perspinosa, and O. serripinna. Gonad morphology and the sequence of developmental stages were similar in each species. Within individuals, gametogenesis was synchronous in gonads both along and between arms, except in some proximal and distal genital pinnules in which gonads never matured. Within a species, there was a degree of synchrony of gametogenesis in a sample, but the level of gametogenic activity varied considerably between years. A high level of continuous reproductive activity was observed in H. bartschi, H. robustipinna, Cenometra bella, and Colobometra perspinosa from about mid-summer to early/mid-winter, while O. serripinna probably had a bimodal reproductive cycle with peaks around February and June. There was a significant 1:1 male:female sex ratio in the populations sampled, and unsexable individuals were significantly smaller (as determined by maximum arm length) than either sexable males or females.