Settlement, recruitment and potential predators and competitors of juvenile echinoderms in the rocky subtidal zone

Recruitment patterns of marine invertebrates are affected both by settlement and early post-settlement events. This study examined the settlement and recruitment patterns of echinoderms at three sites in the rocky subtidal zone of Bocabec Cove, Bay of Fundy, Canada using artificial turf collectors and quadrats on the natural substrate. Potential predators were quantified at two of the sites along transects and in 1-m² quadrats. Both potential predators and competitors were quantified in 0.0625-m² quadrats. Settlement varied across sites (1.5–3 km apart) and two years of sampling (2004, 2005). The site of most potential settlement differed for the three groups of echinoderms: sea urchins (Strongylocentrotus droebachiensis), sea stars (Asterias spp.) and sea cucumber (Psolus fabricii). Settlement densities on the artificial turf collectors tended to be greater than the densities of settlers on the natural substrate. On the natural substrate, the only significant difference between densities of juveniles over time was that newly settled sea stars were found in July and were not found the following October. Large lobsters and carnivorous worms were potential predators with densities that varied between sites. Potential competitors that differed in abundance between sites were herbivorous gastropods and conspecifics for sea urchins; and carnivorous worms for sea stars. This study suggests that patterns of recruitment are either set up by patterns of settlement or by events during the first few weeks/months on the benthic substrate for these echinoderms.     

Intertidal community structure and oceanographic patterns around Santa Cruz Island, CA, USA

Recent studies suggest that nearshore oceanographic conditions can have important effects on the structure of benthic communities. On Santa Cruz Island (SCI), CA, USA there is a persistent difference in mean annual sea surface temperature (SST) around the island due to its location at the confluence of opposing cold and warm ocean current systems. Over the course of a 4-year study (1997–2001) seawater nutrient and chl-a concentrations, algal tissue C:N ratios, recruitment and growth of filter-feeders (barnacles and mussels), and intertidal community structure were measured at six intertidal sites around the island. There were strong associations between remotely sensed SST and patterns of community structure. Macrophyte abundance was highest at sites with persistently low SST, while recruitment, abundance, and growth of filter-feeding invertebrates were strongly, positively correlated with SST. The cold-water sites were associated with higher nutrient concentrations and lower algal C:N ratios, particularly in the winter months. Values of chl-a were generally low and variable among sites, and were not correlated with the predominant SST gradient. Recruitment of barnacles and mussels was positively correlated with adult abundance across all sites. While detailed experimental studies are needed to further evaluate the mechanisms underlying community dynamics, these results indicate that the confluence of cold- and warm-water masses around SCI may determine the contrasting patterns of intertidal community structure.     

Wave exposure effects on population structure and recruitment in the mussel <Emphasis Type="Italic">Perna perna</Emphasis> suggest regulation primarily through availability of recruits and food,not space

Recruitment and population structure of Perna perna in low shore mussel beds were investigated over 15 months at six sites along the south coast of South Africa. Initial, subjective classification of sites as wave exposed or wave sheltered (three of each) was confirmed using the dissolution of cement blocks to measure average water flux and dynamometers for maximum wave force. Recruitment occurred throughout the year, but recruit (1–5 mm) densities were significantly higher from January to April 1996 on both shore types. Recruit densities were positively correlated with adult (>15 mm) densities for both shore types (P < 0.05) but the correlations were extremely weak (r ² < 0.06 in each case). In areas with 100% cover, adult size (mean and maximum lengths) was greater on exposed sites, but density showed the reverse and was negatively correlated with maximum wave strength (r = −0.84). Despite differences in adult densities and sizes, biomass, which is a product of the two, showed no significant difference between the two shore types (ANOVA P > 0.05). Thus wave exposure dramatically affects density, recruitment and mussel size, but not recruitment timing or biomass where there is 100% cover, and mediates a three-way interaction among food supply, larval supply and intraspecific competition for space. In contrast to shores with saturation recruitment, mussel biomass here appears to be limited by recruit supply and constraints of food, especially on sheltered shores, while density is regulated through intraspecific competition for space primarily on exposed shores and at small spatial scales.     

Spatial and temporal patterns of recuruitment of juvenile coral reef fishes to coral habitats within “One Tree Lagoon”, great barrier reef

Recruitment of juvenile fishes to coral grids, each comprising 4 colonies of 3 species of coral (Acropora formosa, Seriatopora hystrix and Pocillopora damicornis), was examined at 4 widely separated sites within the lagoon of One Tree Reef over two successive summers and intervening months (November 1976–January 1978). Recruitment was highly seasonal, with most recruitment occurring during summer. For many species, numbers settling differed greatly from year to year with total numbers (over all sites) differing as much as an order of magnitude between summers. Many fish species demonstrated marked preference among the three coral species as settling sites. The distribution of each of the 20 commonest species across the 4 lagoon sites differed significantly from a random pattern. Differences of an order of magnitude were common in the numbers of a given species recruited to different sites. Each site was preferred by at least one species. In each of 5 cases examined, the pattern of settlement of the species across the 4 sites changed significantly from one summer to the next. The distribution of recruits of a number of species corresponded to the distribution of adults, but for other species there was no correspondence. It is concluded that, at the spatial scale examined, patterns of recruitment of some taxa are consistently more variable than those of other taxa. The implications of spatial variability of recruitment for the distribution of adult fish is discussed and the importance of being able to discriminate such natural variability from other kinds of change is stressed.     

Coral recruitment: a spatio-temporal analysis along the coastline of Eilat,northern Red Sea

Recruitment rates of stony corals to artificial substrates were monitored for 2 years at 20 sites along the coast of Eilat, northern Red Sea, to compare with those recorded at other coral reef locations and to assess variation in recruitment at several spatial scales. Coral recruitment was low compared to that observed on the Great Barrier Reef in Australia, but was similar to levels reported from other high-latitude reef locations. Pocilloporids were the most abundant coral recruits in all seasons. Recruitment was twofold higher during the first year than during the second year of study. There was considerable spatial variability, with the largest proportion of variance, apart from the error term, attributable to differences between sites, at a scale of 10² m. Spearmans ranked correlation showed consistency in spatial patterns of recruitment of pocilloporid corals between years, but not of acroporid corals. During spring, when only the brooding pocilloporid coral Stylophora pistillata reproduces at this locality, most coral recruitment occurred at central and southern sites adjacent to well-developed coral reefs. During summer, recruitment patterns varied significantly between years, with wide variation in the recruitment of broadcasting acroporid corals at northern sites located distant from coral reefs. Settlement was low at all sites during autumn and winter. This work is the first detailed analysis of coral recruitment patterns in the Red Sea, and contributes to the understanding of the spatial and temporal scales of variation in this important reef process.Communicated by O. Kinne, Oldendorf/Luhe     

Temporal and spatial components of variability in benthic recruitment,a 5-year temperate example

Deployment of artificial substrata is a common method of investigating early community development and recruitment, but rarely are such experiments of long enough duration to include even year time scales. We placed replicate, machined-slate panels (15×15 cm) in the intertidal and at depths of 6 and 12 m at two sites of differing flow rate at Lough Hyne, SW Ireland. These were serially replaced every 30–60 days for a period of 5 years (1997–2002), except in the intertidal (2000–2002). The number and identity of all recruits were recorded. Recruitment varied over several orders of magnitude both on temporal and spatial scales. The greatest source of variability was between the intertidal (with few species or recruit numbers) and the subtidal zones (many species, some with thousands of recruits per panel per 30 days). Highest levels of recruitment occurred at the low-flow site (Labhra Cliff). Here, recruitment was dominated by the serpulid polychaete, Pomatoceros sp., reaching ~4000 individuals per panel per 30 days. Highest species richness occurred, however, at the high flow site (Whirlpool Cliff). At this site more colonial forms (e.g. bryozoans) settled. Season was found to be the dominant pattern explaining subtidal recruit and species number variability. Year, however, was the dominant temporal pattern explaining change in diversity (Shannon–Wiener H). In space, depth explained most variability of recruit numbers, whereas site explained more variation in species richness. Both these spatial factors contributed similarly to variability of diversity (H). Recruitment has long been known to vary considerably over large spatial scales, such as with latitude and isolation, but we that show changes of a similar magnitude in recruitment can occur across small spatial scales. Individual taxa showed varied temporal patterns of recruitment including continuous, regular seasonal fluctuations and irregular pulses in particular years.     

Efficiency and regulation of recruitment during colony emigration by the ant Temnothorax curvispinosus

Recruitment helps insect societies by bringing individuals to places where work needs to be done, but it also imposes energetic and opportunity costs. The net effect depends both on recruitment efficiency and on the ease with which insects can find work sites on their own. This study examined both of these factors for colony emigration by the ant Temnothorax curvispinosus. Emigrations were organized by a corps of active ants who transported the rest of the colony. These active ants either found new sites independently or followed tandem runs led by successful scouts. Although most tandem runs broke apart before reaching their target, even lost followers found the new site faster than did unguided searchers. When the new site was near the old nest, tandem runs were rare and summoned only a small proportion of the transporter corps. When the new site was instead distant and inconspicuous, tandem runs were common and brought roughly one third of the transporters. This pattern likely results from the quorum rule used by individual scouts to decide when to switch from tandem runs to transports. By monitoring how many nestmates have already found the nest, the ants ensure that the costs of recruitment are born only when necessary.     

Larval supply and juvenile recruitment of coral reef fishes to marine reserves and non-reserves of the upper Florida Keys,USA

For marine organisms, decoupling between the planktonic larval stage and the benthic-associated juvenile stage can lead to variable patterns of population replenishment, which have the potential to influence the effectiveness of marine reserves. We measured spatial and temporal variability in larval supply and recruitment of fishes to coral reefs of different protection levels and tested whether protection level influenced the relationship between supply and recruitment. We sampled pre-settlement larvae and newly settled recruits from four reefs (two reserves and two non-reserves) in the Florida Keys National Marine Sanctuary, USA. Replicate point measures of larval supply over 14 months and 17 monthly measurements of recruitment varied significantly among months and sites. Sites with the same protection level had significantly different patterns of larval supply as well as larval and recruit diversity, but recruitment magnitude differed only by protection level, where densities were greater at reserves. Differences in larval supply among sites included two particularly large peaks in larval abundance at one site, possibly associated with the observed passage of small-scale oceanographic features. To examine whether relationships between larval supply and recruitment varied by protection level, we selected one species that was present in both the light trap samples and the monthly recruitment surveys. Recruitment of the bicolor damselfish Stegastes partitus was significantly and positively related to larval supply at three of the four sites thus, protection level did not influence this linkage. Since local variability among sites can lead to spatial differences in population replenishment, characterization of larval supply and recruitment to potential marine reserve sites may help to identify optimal locations in a region and contribute to more effective reserve design.     

Growth and longevity in the crustose red algaPetrocelis middendorffii

Petrocelis middendorffii is a non-calcareous encrusting red alga growing on rocks in the middle and upper intertidal zone. We have photographically followed the fate of 43 individual plants for periods up to 88 months at Waadah Island, near Neah Bay, Washington (USA). Growth is slow, with surface area increasing about 4% year^-1 for those individuals growing. Other plants were observed to decrease in total surface coverage, fragment, and eventually disappear. Plants on neighboring but not contiguous sample sites are not in synchrony, with some subpopulations increasing their total space coverage while others diminished theirs. Age estimates, based on a variety of growth interpretations, suggest that an average sized individual (123 cm²) will be from 25 to 87 years old. Recruitment and mortality rates appear low. The ecological traits ofP. middendorffii may be similar to those of other crustose heteromorphic benthic algae.     

Recruitment and juvenile survival in the sea hare Aplysia juliana (Gastropoda: Opisthobranchia)

The specificity in settling response of larval Aplysia juliana (Quoy and Gaimard, 1832) on species of its food algae, Ulva spp., provided a rare opportunity for the quantitative study of recruitment and early survival rates in a non-sessile marine invertebrate. Post-metamorphic juveniles spend about 3 weeks feeding and growing on the algae before moving to the rocky habitat of the adults. Recruitment and survival were studied by collecting weekly samples of algae and holding them in the laboratory until the recruits were large enough to be seen and reliably counted. Recruitment occurred throughout the year, although the rates were relatively low in late winter and spring. Recruitment was monitored relative to algal weight and bottom area. Variation in larval abundance and algal standing crop influenced recruitment to the study area. Mortality during the juvenile stage was very high in all cases, although the rates and shapes of the survivorship curves varied between algal species and location.     

Temporal and spatial variability in coral recruitment on two Indonesian coral reefs: consistently lower recruitment to a degraded reef

Corals are the primary reef-building organisms, therefore it is key to understand their recruitment patterns for effective reef management. Coral recruitment rates and juvenile coral abundance were recorded in the Wakatobi National Marine Park, Indonesia, on two reefs (Sampela and Hoga) with different levels of environmental degradation (12.5 vs. 44 % coral cover with high and low sedimentation rates, respectively) to examine consistencies in recruitment patterns between years and seasons. Recruitment was measured on multiple panels at two sites on each reef (6–7 m depth) and cleared areas of natural reef. Although coral recruitment was twofold higher in 2008–2009 than in 2007–2008, and seasonal differences were identified, consistent significant differences in recruitment rates were found between the two reefs even though they are separated by only ~1.5 km. Recruitment rates and juvenile abundance were lower on the more degraded reef. These patterns are likely a consequence of differential pre- and post-settlement mortality as a result of the high sedimentation rates and degraded conditions and possibly reduced larval supply.     

Seed and establishment limitation contribute to long-term native forb declines in California grasslands

The effects of exotic species invasions on biodiversity vary with spatial scale, and documentation of local-scale changes in biodiversity following invasion is generally lacking. Coupling long-term observations of local community dynamics with experiments to determine the role played by exotic species in recruitment limitation of native species would inform both our understanding of exotic impacts on natives at local scales and regional-scale management efforts to promote native persistence. We used field experimentation to quantify propagule and establishment limitation in a suite of native annual forbs in a California reserve, and compared these findings to species abundance trends within the same sites over the past 48 years. Observations at 11 paired sites (inside and outside the reserve) indicated that exotic annual plants have continued to increase in abundance over the past 48 years. This trend suggests the system has not reached equilibrium > 250 years after exotic species began to spread, and 70 years after livestock grazing ceased within the reserve. Long-term monitoring observations also indicated that six native annual forb species went extinct from more local populations than were colonized. To determine the potential role of exotic species in these native plant declines, we added seed of these species into plots adjacent to monitoring sites where plant litter and live grass competition were removed. Experimental results suggest both propagule and establishment limitation have contributed to local declines observed for these native forbs. Recruitment was highest at sites that had current or historical occurrences of the seeded species, and in plots where litter was removed. Grazing history (i.e., location within or outside the reserve) interacted with exotic competition removal, such that removal of live grass competition increased recruitment in more recently grazed sites. Abundance of forbs was positively related to recruitment, while abundance of exotic forbs was negatively related. Thus, exotic competition is likely only one factor contributing to local declines of native species in invaded ecosystems, with a combination of propagule limitation, site quality, and land use history also playing important and interactive roles in native plant recruitment.     

Life history of a dominant polychaete, <Emphasis Type="Italic">Polygordius jouinae</Emphasis>, in inner continental shelf sands of the Mid-Atlantic Bight,USA

The life history of the polychaete Polygordius jouinae Ramey, Fiege, and Leander 2006 was studied in the sandy sediments of the LEO-15 research area off the coast of Tuckerton, New Jersey (39°27.69′N, 74°15.81′W). Divers collected sediment cores biweekly, from February 2004 to November 2005 to examine the temporal patterns of abundance and size–frequency distributions of P. jouinae. These parameters were also examined (for the first time) for P. jouinae in samples taken in ambient sediments and in three reciprocal 3 to 5-day sediment transplant experiments that were part of a previous, unrelated study at coarse sand and muddy sand sites (∼3 km apart) at LEO-15 in the summers of 1994 and 1995. Analysis of these previous data along with the current data provided a larger set of values for P. jouinae, as well as further insight into the life history, habitat selection, and dispersal of P. jouinae (not considered in previous studies). Polygordius jouinae is gonochoristic and the population was mostly composed of sexually mature individuals in late May 2004 and 2005. The reproductive period usually occurs from May to August and individuals that spawn generally have a 1-year life span. Non-spawning individuals of P. jouinae could potentially live longer than 1 year and therefore the population may sometimes consist of individuals of two generations. Recruitment begins no later than July as recently settled individuals (≤9 mm body length), which were not present during the first week of June 1995, 2004, and 2005, first appeared in July in all 3 years. The smallest individual was 2.01 mm long and 0.07 mm wide and this provides the first estimate of size at initial recruitment. Although some worms in sediment trays likely settled as larvae, most worms were recently settled juveniles (length categories 6 and 9 mm) transported in from the surrounding sediments, presumably as a consequence of re-suspension or bed load transport. As expected, larger individuals were present later in September. Although it is unknown whether P. jouinae were transported actively or passively these data suggest that recently settled juveniles are capable of habitat selection (prefer sand over mud) and show that post-settlement transport and selection influence community structure.     

Effect of growth factors and light quality on the growth,pigmentation and photosynthesis of two diatoms,Thalassiosira gravida and Phaeodactylum tricornutum

Recruitment, life span and growth rate were investigated in field and experimental populations of Abra alba (Wood) in Kiel Bay, FRG (55°N) from 1975 to 1978 to determine production to biomass (P:B) ratios and to assess the importance of A. alba to production by commercial fish. Life span and growth rates were determined from changes in length frequency modes at each site and from winter rings on the shell. A peak of recruitment usually occurred in August, sometimes followed by a second peak between December and February. Life span was between a little more than one year and two and a half years. Growth rates were highest at the two sites in offshore fishing grounds, where bivalves reached a mean length of 13 to 16 mm at the end of two years. At the inshore control site and in the nearby experimental containers, individuals reached a mean length of 7 mm at the end of two years. Production estimates ranged from 110 to 3.000 mg C m^-2 year^-1, differing markedly among sites and among years. Production was highest during the first year after recruitment, occurring mainly between July and December. Mortality occurred mainly between January and June, and was in approximate balance with production over a three-year period. Annual P:B ratios were from 1.3 to 3.4; a long-term mean P:B ratio of 2.2 is suggested for Kiel Bay populations of A. alba. Annual production by A. alba appears to exceed considerably consumption by fish of commercial size. The significance of A. alba in the food web of Kiel Bay may thus be as food for juvenile fish or for intermediate-level predators that are themselves prey for larger fish.Publication No. 431 of the Sonderforschungsbereich 95 Meer-MeeresbodenPublication No. 1031 of the CSIRO Marine Laboratories     

Patterns of lunar settlement and early recruitment in Caribbean reef fishes at Panamá

Lunar patterns of settlement of 15 Caribbean reef fishes were assessed from daily collections of newly arrived fishes from each of two small (A=63 and B=28 m²) patch reefs at Punta de San Blas (Panamá) during two consecutive 2 yr periods (1984, 1985 and 1986, 1987). In 12 species settlement was lunar-cyclic, with a broad peak of activity around the new moon in 11 cases, and at first quarter in 1 species. However, 3 of these lunar-cyclic species displayed intermittent semilunar periodicity in settlement, with peaks around the quarter moons. One other species had a semilunar cycle of settlement with peaks at both quarter moons. Two species apparently lacked lunar settlement cycles. In some, but not all, paris of congeners: (a) monthly variation in settlement intensity was positively correlated, (b) the duration of settlement pulses in the same month consistently differed, and (c) the timing of settlement pulses in the same month often differed (although one species did not consistently arrive before the other). Recruitment of juvenile fish that survived to the end of the lunar cycle in which they settled was monitored monthly, concurrently with settlement, at a number of large sites scattered up to 2 km apart. The levels of variation in the amount of settlement and recruitment each month differed in only 2 of 17 cases, with settlement variation exceeding recruitment variation in only one of these. The intensity of recruitment was positively correlated with the intensity of settlement in 16 of 17 cases. In 15 of those 16 cases, R ² values for linear regressions of recruitment on settlement exceeded R ² values for curvilinear regressions that would indicate either increasing or decreasing recruitment success with increasing intensity of settlement. Thus it appears that variation in recruitment can be used to estimate variation in settlement, and that recruitment success may be density-independent.     

Mechanisms of habitat segregation between an invasive and an indigenous mussel: settlement,post-settlement mortality and recruitment

The mussel Mytilus galloprovincialis is highly invasive worldwide, but displays varying degrees of local and regional coexistence with indigenous mussels through spatial habitat segregation. We investigated the roles of settlement, post-settlement mortality, juvenile growth and recruitment in partial habitat segregation between the invasive M. galloprovincialis and the indigenous mussel Perna perna on the south coast of South Africa. We used two study locations, Plettenberg Bay and Tsitsikamma, 70 km apart, with two sites (separated by 300–400 m) per location, each divided into three vertical zones. There were no significant effects in Tsitsikamma, where daily settlement and monthly recruitment were significantly lower than in Plettenberg Bay. In Plettenberg Bay, settlement (primary and secondary) and recruitment of both species decreased upshore. Post-settlement mortality was measured over two consecutive 6-day periods during a spring tide and a neap tide. For both species mortality was low on the low-shore. High-shore mortality was consistently low for M. galloprovincialis, but increased dramatically for P. perna during spring tide. No data were obtained for growth of P. perna, but juvenile M. galloprovincialis grew more slowly farther upshore. P. perna recruited mainly in spring and summer, with a peak in summer far greater than for M. galloprovincialis. Recruitment of M. galloprovincialis was more protracted, continuing through autumn and winter. Thus local coexistence is due to a combination of pre- and post-recruitment factors differing in importance for each species. P. perna is excluded from the high-shore by recruitment failure (low settlement, high mortality). High survival and slow growth in juveniles may allow large densities of M. galloprovincialis to accumulate there, despite low settlement rates. With no differences between species in settlement or mortality on the low-shore, exclusion of M. galloprovincialis from that zone is likely to be by post-recruitment processes, possibly strengthened by periodic heavy recruitments of P. perna. At larger scales, larval retention and protracted recruitment contribute to the success of M. galloprovincialis at Plettenberg Bay, while recruitment limitation may explain why M. galloprovincialis is less successful at other sites.     

Local and meso-scale patterns of recruitment and abundance of two intertidal crab species that compete for refuges

Interference competition for limited habitat or refuges is known to produce density-dependent mortality and generate patterns of micro-habitat distribution. While in mobile species the outcome of interference at a local scale can usually be determined from differences in body size and behavior, the population-level consequences of such interactions vary depending on rates of settlement and recruitment at a site, which are not directly correlated to local reproductive success. Previous experimental studies in central Chile demonstrated that interference competition for refuges is the primary factor driving microhabitat segregation between the predatory crabs Acanthocyclus gayi and Acanthocyclus hassleri, with the latter species monopolizing galleries inside mussel beds and excluding A. gayi to rock crevices. Between April 2001 and March 2006 we quantified monthly recruitment rates in artificial collectors at 17 sites over 900 km of the central coast of Chile. Results show that recruitment rates of A. hassleri are almost two orders of magnitude lower than those of A. gayi, and that they are tightly and positively correlated among sites across the region, suggesting that at scales of kilometers larval stages of these species are affected by similar oceanographic processes. Total crab densities per site were also positively correlated between species and strongly associated to mussel cover, with overall low crab densities at all sites where mussel cover was lower than about 60%. At all sites with mussel cover >60%, the ratio of A. gayi to A. hassleri density progressively decreased from recruits (2.6) to juveniles (0.5) to adults (0.04), overcoming initial differences in recruitment rates. The relative success of the inferior competitor at sites with low mussel cover does not appear to provide a potential mechanism favoring regional coexistence through dispersal to other sites (“mass effects”), because their densities were lower than at sites of high mussel cover. Yet, at many sites of low mussel cover the dominant competitor is virtually absent, allowing A. gayi to attain larger population sizes at the scale of the region. Thus, the factors limiting the dominant competitor from successfully utilizing other microhabitats seem to be the most critical factor in promoting both local and regional coexistence between these species.     

Mechanisms of habitat segregation between an invasive (Mytilus galloprovincialis) and an indigenous (Perna perna) mussel: adult growth and mortality

The invasive mussel Mytilus galloprovincialis and the indigenous mussel Perna perna coexist intertidally on the south coast of South Africa through partial vertical habitat segregation: M. galloprovincialis dominates the upper shore and P. perna the lower shore. Recruitment patterns can explain the zonation of P. perna, but not the invasive species. We examined the role of post-recruitment interactions by measuring spatial and temporal differences in adult growth and mortality rates of the two species. Specifically, we tested the hypothesis that interspecific differences in growth and mortality reflect adult distribution patterns. The two study locations, Plettenberg Bay and Tsitsikamma, are 70 km apart with two sites (separated by 300–400 m) per location, each divided into three vertical zones. Growth was measured seasonally using different marking methods in 2001 and 2003. Cumulative adult mortality was measured through summer in 2003/2004. Both species generally grew more slowly upshore, but they showed different effects of season. For P. perna, growth was significantly reduced in winter in the low zone, but unaffected by season in the high zone. For M. galloprovincialis, growth was either unaffected by season or increased in winter, even in the high zone. Thus, growth of P. perna and M. galloprovincialis was reduced under cool winter and warm summer temperatures, respectively; and while growth was more similar between species in summer, M. galloprovincialis grew much faster than P. perna in winter. Mortality of P. perna increased upshore. For M. galloprovincialis, mortality was not zone-dependent and was significantly greater than for P. perna on the low-shore and (generally) across the shore in Tsitsikamma. Both species had higher growth and mortality rates in Plettenberg Bay than in Tsitsikamma. Thus, P. perna seems able to maintain spatial dominance on the low-shore and at certain sites because of higher mortality of M. galloprovincialis. We conclude that seasonality in growth of the two species reflects their biogeographic affinities and that coexistence is possible through pre-recruitment effects that limit the vertical distribution of P. perna and post-recruitment effects that limit M. galloprovincialis.     

Population ecology of the low-shore chitons Onithochiton quercinus and Plaxiphora albida

Populations of two chitons, Onithochiton quercinus (Gould, 1846) and Plaxiphora albida (Blainville, 1825) were sampled at five sites in the low-shore algal community at Cape Banks, New South Wales, Australia, from January 1985 to February 1988. Both species were abundant, reaching densities of 96 and 45 m^-2, respectively and grew to 80 mm in length. O. quercinus and P. albida were found on the solitary ascidian Pyura stolonifera and on the substratum in amongst the foliose macroalgae which dominate this community. Size-frequency data were used to estimate the rates of growth, mortality and longevity, and time of recruitment. Growth of both species was rapid for the first 3 yr and slowed thereafter. O. quercinus and Plaxiphora albida had similar longevities and were estimated to live for at least 6 yr. Reproductive samples were also collected to identify when sexual maturity occurred, to clarify the reproductive periodicity, and to examine the sex ratios of the two species. Both species spawned over a discrete period in early autumn and there was no evidence of the bi-annual spawning reported in a previous study. Recruitment of O. quercinus and P. albida occurred during autumn, but recruits could not be quantified owing to their cryptic nature. The microhabitats occupied by the recruits of both species included the empty shells of the barnacle Austromegabalanus nigrescens, the spaces between the calcareous tubes of the polychaete Galeolaria caespitosa, and the space amongst the algae on Pyura stolonifera. Both species reached reproductive maturity at 2 yr of age. The sex ratios of both species were consistent with the results of a previous study: P. quercinus had a sex ratio of almost 2:1 biased in favour of males, and Plaxiphora albida exhibited a sex ratio of 1:1. The study suggested that populations of O. quercinus and P. albida could persist in the low-shore algal community in the absence of recruitment. This, together with their large size and consumption of macroalgae suggests that they have the potential to affect the structure and dynamics of this intertidal community.     

Coexistence of nine anemonefish species: differential host and habitat utilization, size and recruitment

The region of Madang, Papua New Guinea, has the highest reported species diversity of both anemonefishes (nine species) and their host anemones (ten species). To determine which factors may allow so many anemonefish species to coexist at this location, we studied their patterns of distribution, abundance, and recruitment. Population surveys at three replicate reef sites within four zones situated at varying distances from the mainland (nearshore, mid-lagoon, outer barrier, and offshore) indicated that each species of host anemone and anemonefish lived within a particular range of zones. Each species of anemonefish lived primarily with one species of host. Anemonefish species that lived with the same host species usually had different distribution patterns among zones (e.g., Amphiprion percula occupied Heteractis magnifica in nearshore zones, while A. perideraion occupied H. magnifica in offshore zones). Monitoring of natural populations showed that there were few changes (losses or recruitment) in the number or species of fishes associated with each individual anemone over periods ranging from 3 to 9 months. Recruitment was monitored on anemones with and without residents (resident fishes were removed) within each of three zones (nearshore, mid-lagoon, outer barrier). Significantly more anemonefishes recruited to anemones without resident fishes than to anemones with resident fishes. Each anemonefish species recruited to particular host species and zones. The distribution and abundance of the recruits of each fish species among zones were positively correlated with the distribution and abundance of resident fishes in the benthic habitat. This suggests that the spatial patterns of recruitment among zones strongly determined the distribution and abundance patterns of the benthic populations, and they were not the result of post-recruitment mortality or movement. Coexistence of the nine anemonefish species on the limited anemone resource was considered possible because of niche differentiation (i.e., differences in host and habitat utilization among zones), and the ability of two small species (i.e., Amphiprion sandaracinos and A. leucokranos) to cohabit individual anemones with other anemonefish species. Received: 29 July 1999 / Accepted: 1 September 2000