Discarded queen conch (<Emphasis Type="Italic">Strombus gigas</Emphasis>) shells as shelter sites for fish

Within the Caribbean millions of queen conch (Strombus gigas Linnaeus) are harvested each year and shells discarded randomly or as middens. Fish use of discarded conch shells was investigated in four different habitat types: sand, seagrass beds, mangrove forests, and coral reefs. The study was carried out in the waters off South Caicos, Turks and Caicos Islands (TCI), between October 2003 and January 2004. The density of discarded shells was greatest near coral reefs; however, the percentage of shells occupied by adult fish was higher in isolated shells on sand and in mangrove habitats. Juvenile fish also showed a preference for sheltering in conch shells relative to other microhabitat types on sandy plains and in mangrove and seagrass habitats. Differences in use of single shells by fish in different habitats were attributed to differences in piscivore abundance and habitat complexity. Although not all isolated shells were occupied by fish, all conch middens deposited by fishermen had fish inhabitants. Examination of fish use of conch middens in three habitat types and conch piles of one, three, and five shells constructed on sand found both fish diversity and abundance increased on conch middens of increasing size. This study suggests that disposal of conch shells as large middens in habitats of low complexity will increase the amount of shelter present and may enhance fish populations in these habitats.Communicated by J.P. Grassle, New Brunswick     

Safety in numbers and the spatial scaling of density-dependent mortality in a coral reef fish

In coral reef fishes, density-dependent population regulation is commonly mediated via predation on juveniles that have recently settled from the plankton. All else being equal, strong density-dependent mortality should select against the formation of high-density aggregations, yet the juveniles of many reef fishes aggregate. In light of this apparent contradiction, we hypothesized that the form and intensity of density dependence vary with the spatial scale of measurement. Individual groups might enjoy safety in numbers, but predators could still produce density-dependent mortality at larger spatial scales. We investigated this possibility using recently settled juvenile bluehead wrasse, Thalassoma bifasciatum, a small, aggregating reef fish. An initial caging experiment demonstrated that juvenile bluehead wrasse settlers suffer high predation, and spatial settlement patterns indicated that bluehead wrasse juveniles preferentially settle in groups, although they are also found singly. We then monitored the mortality of recently settled juveniles at two spatial scales: microsites, occupied by individual fish or groups of fish and separated by centimeters, and sites, consisting of approximately 2400-m2 areas of reef and separated by kilometers. At the microsite scale, we measured group size and effective population density independently and found that per capita mortality decreased with group size but was not related to density. At the larger spatial scale, however, per capita mortality increased with settler density. This shift in the form of density dependence with spatial scale could reconcile the existence of small-scale aggregative behavior typical of many reef fishes with the population-scale density dependence that is essential to population stability and persistence.     

Patterns of co-occurrence and interactions between age classes of the common triplefin,Forsterygion lapillum

Many marine organisms have pelagic larval stages that settle into benthic habitats occupied by older individuals; however, a mechanistic understanding of intercohort interactions remains elusive for most species. Patterns of spatial covariation in the densities of juvenile and adult age classes of a small temperate reef fish, the common triplefin (Forsterygion lapillum), were evaluated during the recruitment season (Feb–Mar, 2011) in Wellington, New Zealand (41°17′S, 174°46′E). The relationship between juvenile and adult density among sites was best approximated by a dome-shaped curve, with a negative correlation between densities of juveniles and adults at higher adult densities. The curve shape was temporally variable, but was unaffected by settlement habitat type (algal species). A laboratory experiment using a “multiple-predator effects” design tested the hypothesis that increased settler mortality in the presence of adults (via enhanced predation risk or cannibalism) contributed to the observed negative relationship between juveniles and adults. Settler mortality did not differ between controls and treatments that contained either one (p = 0.08) or two (p = 0.09) adults. However, post hoc analyses revealed a significant positive correlation between the mean length of juveniles used in experimental trials and survival of juveniles in these treatments, suggesting that smaller juveniles may be vulnerable to cannibalism. There was no evidence for risk enhancement or predator interference when adults were present alongside a heterospecific predator (F. varium). These results highlight the complex nature of intercohort relationships in shaping recruitment patterns and add to the growing body of literature recognizing the importance of age class interactions.     

Terrestrial habitat selection and strong density-dependent mortality in recently metamorphosed amphibians

To predict the effects of terrestrial habitat change on amphibian populations, we need to know how amphibians respond to habitat heterogeneity, and whether habitat choice remains consistent throughout the life-history cycle. We conducted four experiments to evaluate how the spatial distribution of juvenile wood frogs, Rana sylvatica (including both overall abundance and localized density), was influenced by habitat choice and habitat structure, and how this relationship changed with spatial scale and behavioral phase. The four experiments included (1) habitat manipulation on replicated 10-ha landscapes surrounding breeding pools; (2) short-term experiments with individual frogs emigrating through a manipulated landscape of 1 m wide hexagonal patches; and habitat manipulations in (3) small (4-m2); and (4) large (100-m2) enclosures with multiple individuals to compare behavior both during and following emigration. The spatial distribution of juvenile wood frogs following emigration resulted from differences in the scale at which juvenile amphibians responded to habitat heterogeneity during active vs. settled behavioral phases. During emigration, juvenile wood frogs responded to coarse-scale variation in habitat (selection between 2.2-ha forest treatments) but not to fine-scale variation. After settling, however, animals showed habitat selection at much smaller scales (2-4 m2). This resulted in high densities of animals in small patches of suitable habitat where they experienced rapid mortality. No evidence of density-dependent habitat selection was seen, with juveniles typically choosing to remain at extremely high densities in high-quality habitat, rather than occupying low-quality habitat. These experiments demonstrate how prediction of the terrestrial distribution of juvenile amphibians requires understanding of the complex behavioral responses to habitat heterogeneity. Understanding these patterns is important, given that human alterations to amphibian habitats may generate extremely high densities of animals, resulting in high density-dependent mortality.     

Age-and sex-dependent patterns of aggregation in the tropical gastropod Strombus luhuanus

The spatial dispersion of the tropical coral sand-dwelling mesogastropod Strombus luhuanus Linné, 1758 was studied at several sites in two localities (Australian Great Barrier Reef and southern Papua New Guinea), from September 1980 to February 1983. S. luhuanus usually occurred in local aggregations, many of which were relatively dense with discrete boundaries: Four types of aggregation are described: mixed age-class, juvenile, mating, and clusters. The former two are termed colonies, because they persisted over time and exhibited some coordinated movements. Mixed age-class colonies contain individuals of all ages, but within them the different ageclasses are frequently spatially segregated from each other. This was sometimes due to the younger age-classes' preference for shallower areas, but also occurred in areas of constant depth. Juvenile colonies are relatively smaller in area but higher in density, and are dominated by a single size-class. Mating aggregations are concentrations of copulating individuals, in which competition between males for access to females occurs. Clusters are concentrations of inactive individuals (juvenile and adult) piled together in close contact. All types of aggregation were concentrated in only a part of what appeared to be the suitable habitat. Most aggregations' boundaries shifted over time within the habitat. We conclude that intraspecific attraction must be an important factor which maintains the spatial structure of all aggregations, although habitat specificity sets the larger area in which they move. Processes which may underlie this pattern, and some of its implications, are discussed.     

Growth costs of territorial overlap: experiments with juvenile lizards (Anolis aeneus)

Summary Several theoretical models have suggested that increases in territory overlap with conspecifics should result in lower rates of resource accumulation by territorial residents. This assumption was tested using juvenile Anolis aeneus lizards, which exhibit variable degrees of territory overlap in both the field and laboratory. In a series of laboratory trials under controlled density and resource conditions, juvenile growth rates were negatively related to the degree of territory overlap. This effect aplied to juveniles of all but the lowest status groups, although higher status juveniles had more exclusive territories than did lower status juveniles. Focal studies indicated that food depletion in the zones of overlap was the most likely reason for the negative relationship between territory overlap and growth. It is suggested that juveniles in the field may tolerate extensive territory overlap in spite of growth penalties because of a general shortage of predator-safe homesites in undisturbed habitats.     

Spatial repartition and ontogenetic shifts in habitat use by coral reef fishes (Moorea,French Polynesia)

This study explores the extent to which ontogenetic habitat shifts modify spatial patterns of fish established at settlement in the Moorea Island lagoon (French Polynesia). The lagoon of Moorea Island was divided into 12 habitat zones (i.e. coral seascapes), which were distinct in terms of depth, wave exposure, and substratum composition. Eighty-two species of recently settled juveniles were recorded from March to June 2001. Visual censuses documented changes in the distribution of juveniles of each species over time among the 12 habitats. Two patterns of juvenile habitat use were found among species. Firstly, some species settled and remained in the same habitat until the adoption of the adult habitats (i.e. recruitment; e.g. Chaetodon citrinellus, Halichoeres hortulanus, Rhinecanthus aculeatus). Secondly, others settled to several habitats and then disappeared from some habitats through differential mortality and/or post-settlement movement (e.g. 65–70 mm size class for Ctenochaetus striatus, 40–45 mm size class for Epinephelus merra, 50–55 mm size class for Scarus sordidus). A comparison of the spatial distribution of juveniles to that of adults (61 species recorded at both stages) illustrated four patterns of subsequent recruitment in habitat use: (1) an increase in the number of habitats used during the adult stage (e.g. H. hortulanus, Mulloidichthys flavolineatus); (2) a decrease in the number of habitats adults used compared to recently settled juveniles (e.g. Chrysiptera leucopoma, Stethojulis bandanensis); (3) the use of different habitat types (e.g. Acanthurus triostegus, Caranx melampygus); and (4) no change in habitat use (e.g. Naso litturatus, Stegastes nigricans). Of the 20 most abundant species recorded in Moorea lagoon, 12 species modified the spatial patterns established at settlement by an ontogenetic habitat shift.Communicated by T. Ikeda, Hakodate     

Otolith analysis of juvenile walleye pollock Theragra chalcogramma from the western Gulf of Alaska

Growth rate and hatch date distributions were estimated for juvenile walleye pollock Theragra chalcogramma (Pallas, 1814) collected in autumn 1987 from the western Gulf of Alaska. Mean juvenile growth rates varied geographically by as much as 45%. A trend for slower growth around Unimak Pass and the Shumagin Islands and faster growth upstream in the Alaska Coastal Current towards Kodiak Island was noted; the fastest growth did not occur in the main habitat region. Juvenile hatch date distributions were compared to identify regional differences that might reflect stock structure. Juvenile hatch dates were compared with hatch dates of the same cohort sampled as larvae to test for selective mortality. Regional differences in hatch dates, along with other information, indicated several minor spawning populations located around Kodiak Island and near Unimak Pass. For the main aggregation of pollock in the Shumagin Island region, hatch date distributions were not significantly different among the early larval cohort sampled in late May, the late larvae sampled in mid-June to early July, and the juveniles sampled in autumn. Neither growth-rate nor size-dependent mortality of pollock between the larval and juvenile stages appears to be a dominant factor in determining survival patterns. An alternative test was attempted, by which lengths-at-age during larval life were back-calculated from juvenile otoliths and compared with lengths-at-age of the population sampled as larvae in May and June. Pollock surviving as juveniles in autumn were not larger as larvae than the general larval population. This result is an example of the observation that back-calculated lengths are almost always smaller than the lengths of fish sampled at age (Lee's phenomenon). Problems in determining survival patterns based on otolith back-calculations and comparison of hatch date distributions are discussed.     

Consequences of patch reef spacing for density-dependent mortality of coral-reef fishes

The spatial configuration of habitat patches can profoundly affect a number of ecological interactions, including those between predators and prey. I examined the effects of reef spacing on predator-prey interactions within coral-reef fish assemblages in the Bahamas. Using manipulative field experiments, I determined that reef spacing influences whether and how density-dependent predation occurs. Mortality rates of juveniles of two ecologically dissimilar species (beaugregory damselfish and yellowhead wrasse) were similarly affected by reef spacing; for both species, mortality was density dependent on reef patches that were spatially isolated (separated by 50 m), and density independent on reef patches that were aggregated (separated by 5 m). A subsequent experiment with the damselfish demonstrated that a common resident predator (coney) caused a substantial proportion of the observed mortality, independent of reef spacing. Compared to isolated reefs, aggregated reefs were much more likely to be visited by transient predators (mostly yellowtail snappers), regardless of prey density, and on these reefs, mortality rates approached 100% for both prey species. Transient predators exhibited neither an aggregative response nor a type 3 functional response, and consequently were not the source of density dependence observed on the isolated reefs. These patterns suggest that resident predators caused density-dependent mortality in their prey through type 3 functional responses on all reefs, but on aggregated reefs, this density dependence was overwhelmed by high, density-independent mortality caused by transient predators. Thus, the spatial configuration of reef habitat affected both the magnitude of total predation and the existence of density-dependent mortality. The combined effects of the increasing fragmentation of coral reef habitats at numerous scales and global declines in predatory fish may have important consequences for the regulation of resident fish populations.     

Feeding ecology of the three juvenile phases of the spiny lobster<Emphasis Type="Italic"> Panulirus argus</Emphasis> in a tropical reef lagoon

The three juvenile phases of the spiny lobster Panulirus argus (algal phase: 5-15 mm carapace length, CL; postalgal phase: 15-45 mm CL, and subadults: 45-80 mm CL) occur in the reef lagoon at Puerto Morelos, Mexico. The algal phase abounds in this lagoon, which is covered by extensive seagrass-algal meadows, but the density of postalgal and subadult juveniles is low, owing to the scarcity of crevice-type shelters suitable for these phases. The feeding ecology of the three juvenile phases was investigated to examine whether spatial or temporal differences in food intake, diet composition, or nutritional condition occurred among phases and could partially account for the low abundance of the larger juveniles. Juveniles were collected by divers at night, from January to November 1995, throughout the mid-lagoon and back-reef zones. Percent stomach fullness, relative weight of the digestive gland (RWDG, an index of nutritional condition), percent frequency of occurrence and percent volume of food categories in the diet were compared between sexes, juvenile phases, molt stages (postmolt, intermolt, premolt), seasons, and sampling zones (mid-lagoon and back-reef zones). Significant differences in stomach fullness occurred only among molt stages, mainly because postmolt individuals had emptier stomachs. The main food categories in all juvenile phases were crustaceans (mostly hermit crabs and brachyurans) and gastropods, but the food spectrum was wide, including many other animal taxa as well as plant matter. In June 1995, the epibenthic macrofauna was sampled in five sites in the lagoon that differed in their amount of vegetation. The most abundant taxa in all sites were decapods and gastropods, but density and diversity measures showed that the distribution of these potential prey taxa for juvenile P. argus was rather patchy. Diet overlap in juvenile lobsters was high between sexes, juvenile phases, sampling zones, seasons, and molting stages, indicating that all juveniles fed on the same general food categories throughout time. The only factor that affected the RWDG was the juvenile phase. RWDG was significantly lower in subadults than in algal and postalgal phases, suggesting a poorer nutritional condition in the largest juveniles. This may be related to the scarcity of suitable shelters for large juveniles throughout the lagoon, which may preclude subadults from exploiting food resources in areas of the lagoon where shelter is limited.     

Home range, activity and distribution patterns of a temperate rocky-reef fish, Cheilodactylus fuscus

Red morwong (Cheilodactylus fuscus; Cheilodactylidae: Perciformes), are large temperate rocky reef fish that are patchily distributed in local aggregations of 3 to >100 fish. The home ranges and aggregating behaviour of red morwong were investigated using external tags (n = 114, over 35 mo) and ultrasonic transmitters (n = 9 over 20 d) at coastal and estuarine sites in south-east Australia. Adult red morwong had a mean home range during the day of 1865 m² (SE = 268). Night movement, determined by ultrasonic telemetry, indicated a significantly greater mean home range of 3639 m² (SE = 416). Home range did not differ significantly with size or sex. Fish movements were greatest and aggregation sizes varied most during the non-reproductive mid-summer period. Tagged fish (N = 20) displaced 200 to 900 m returned in 1 to 3 d to the point of capture, often traversing open sand habitat and other aggregations. During the day, fish were commonly found on bouldered habitat. During the night, these site-specific aggregations fragmented as fish dispersed over a variety of substrata, with crepuscular peaks in activity. Therefore, estimates of habitat-specificity and feeding patterns collected only by day may give misleading results. Received: 28 June 1996 / Accepted: 8 July 1998     

Interactions between sea urchins (Strongylocentrotus droebachiensis) and their predators in field and laboratory experiments

Field observations and manipulative experiments in a nearshore cobble bed (2 to 3 m below mean low water) at Eagle Head, Nova Scotia, Canada, between 1984 and 1986, showed that small juveniles ofStrongylocentrotus droebachiensis (3 to 6 mm diam) sheltering beneath cobbles had a refuge from predators such as rock crabs, small lobsters, and fish. Sea urchins gradually outgrew these refuges and small adults (25 to 30 mm) required larger rocks as shelter from predators, particularly large cancrid crabs. Small juveniles were usually solitary and well dispersed beneath cobbles, whereas small adults tended to aggregate on the undersides and in the interstices of boulders. These aggregations may develop passively as sea urchins accumulate in suitablysized refuges. Chemotaxis experiments indicate that juvenileS. droebachiensis are repelled by waterborne stimuli from conspecifics. In a factorial experiment, effects of the presence of potential predators (rock crabs and lobsters) and/or food (kelp) on the behaviour of large juvenile (10 to 15 mm) and small adult sea urchins were examined in flowing seawater tanks. Both size classes formed exposed feeding aggregations when kelp was provided as food, irrespective of the presence or absence of predators. In the absence of kelp, each size class responded differently to the presence of a predator: juveniles became more cryptic, whereas adults aggregated on the tank sides. Increased movement to the sides of a tank in the presence of a predator may reflect a flight response, since chemotaxis experiments indicated thatS. droebachiensis is repelled by waterborne chemical stimuli from predators. Observational and experimental data suggest that predation at the late juvenile and early adult stages may influence population structure, distribution and abundance ofS. droebachiensis.     

Oviposition,larval abundance,in situ larval growth and recruitment of the herbivorous gastropodLacuna vincta in kelp canopies in Barkley Sound,Vancouver Island (British Columbia)

We investigated recruitment of the herbivorous gastropodLacuna vincta (Montagu, 1803) in the canopies ofMacrocystis integrifolia andNereocystis luetkeana beds in Barkley Sound, Vancouver Island (British Colombia), from 1987 to 1989. Four factors influencing intensity and patterns of recruitment were studied: (1) seasonality of oviposition, (2) larval abundance, (3) growth of larvae in the field and (4) larval settlement. Egg masses were abundant on low intertidal algae but were scarce in kelp canopies. Although egg masses could be found almost year-round, a distinct and intense period of oviposition occurred during winter and spring. Intracapsular development lasted 2.5 to 3.5 wk before planktotrophic veligers emerged. The duration of the planktonic period, 7 to 9 wk, was determined through an in situ study of cohorts ofLacuna spp. larvae present in the plankton between January and June 1988. The general timing of the onset of the spring peak recruitment period was predicted from these cohorts. Primary periods of recruitment ofL. vincta in the canopy occurred in April–May (average density up to 383.9 juveniles m^–2 blades), with a second period of lower intensity in the late summer—fall period. We observed similar trends between abundance of advanced larvae (> 500µm) in the plankton and recruitment rates in kelp canopies. Although adults were occasionally observed in the canopy, newly metamorphosed juveniles consistently dominated the habitat. The persistance of small juveniles (0.7 to 1.5 mm), rapid declines in density shortly after recruitment, and SCUBA observations of drifting individuals suggest that juveniles migrate to the under-canopy or low intertidal area after a brief period of growth on kelp blades.     

Predation, habitat complexity, and variation in density-dependent mortality of temperate reef fishes

Density dependence in demographic rates can strongly affect the dynamics of populations. However, the mechanisms generating density dependence (e.g., predation) are also dynamic processes and may be influenced by local conditions. Understanding the manner in which local habitat features affect the occurrence and/or strength of density dependence will increase our understanding of population dynamics in heterogeneous environments. In this study I conducted two separate field experiments to investigate how local predator density and habitat complexity affect the occurrence and form of density-dependent mortality of juvenile rockfishes (Sebastes spp.). I also used yearly censuses of rockfish populations on nearshore reefs throughout central California to evaluate mortality of juvenile rockfish at large spatial scales. Manipulations of predators (juvenile bocaccio, S. paucispinus) and prey (kelp, gopher, and black-and-yellow [KGB] rockfish, Sebastes spp.) demonstrated that increasing the density of predators altered their functional response and thus altered patterns of density dependence in mortality of their prey. At low densities of predators, the number of prey consumed per predator was a decelerating function, and mortality of prey was inversely density dependent. However, at high densities of predators, the number of prey killed per predator became an accelerating response, and prey mortality was directly density dependent. Results of field experiments and large-scale surveys both indicated that the strength of density-dependent mortality may also be affected by the structural complexity of the habitat. In small-scale field experiments, increased habitat complexity increased the strength of density-dependent mortality. However, at large scales, increasing complexity resulted in a decrease in the strength of density dependence. I suggest that these differences resulted from scale-dependent changes in the predatory response that generated mortality. Whether increased habitat complexity leads to an increase or a decrease in the strength of density-dependent mortality may depend on how specific predatory responses (e.g., functional or aggregative) are altered by habitat complexity. Overall, the findings of this study suggest that rates of demographic density dependence and the resulting dynamics of local populations may largely depend upon attributes of the local habitat.     

Hawksbill sea turtles in seagrass pastures: success in a peripheral habitat

Hawksbill sea turtles, Eretmochelys imbricata, are closely associated with coral reef and other hard-bottom habitats. Seagrass pastures are peripheral habitats for Caribbean hawksbills. With the decline in quality and quantity of coral reefs, seagrass habitats may become more important for hawksbills. We use data from a 30-year mark-recapture study of hawksbills and green turtles, Chelonia mydas, in the southern Bahamas to assess the quality of a seagrass habitat for hawksbills. Size distribution, residence times, and body condition index for the seagrass hawksbill aggregation are similar to those of hawksbill aggregations over Caribbean reefs. Somatic growth rates of seagrass hawksbills are in the upper range of those reported for reef hawksbills. Based on these parameters, peripheral seagrass habitats can support healthy, productive hawksbill aggregations. During the 30-year study, a sixfold variation in green turtle density in the study area did not affect the productivity or body condition of hawksbills.     

Population dynamics and growth of juveniles of the velvet swimming crab Necora puber (Decapoda: Portunidae)

Recruitment variability plays a critical role in determining local population densities of benthic organisms, but extreme vulnerability at the onset of juvenile life is a trait that is largely responsible for population survivorship trends. The aim of the present study was to determine the role of juvenile recruitment in the population structure of Necora puber. Juveniles of N. puber were collected from the lower intertidal of rocky shores of Plymouth Sound (southwest coast of the UK) and monthly size–frequency distribution were used to determine the dynamics and the growth of the population. The parameters of the von Bertalanffy growth function were estimated (K=0.281 year⁻¹; t ₀=0.043; C=0.103; and t _s=0.268) assuming a L _∞=105 mm. Growth was markedly seasonal and present results indicated a slower juvenile growth rate than described previously for N. puber. The recruitment period was extensive and was two times higher in 2001 than in 2000 at the start of the 1+ year, but levelled off at the end of the 1+ year class on three of the four shores studied. Instantaneous mortality as high as 5.1 year⁻¹(99.4% year⁻¹) was observed during the higher recruitment year. Early juvenile mortality appears to be density dependent and a demographic bottleneck appears to limit the number of juveniles on some shores.     

Suitability of estuarine nursery zones for juvenile weakfish (Cynoscion regalis): effects of temperature and salinity on feeding,growth and survival

Juvenile weakfish, Cynoscion regalis (Bloch and Schneider, 1801), exhibit significant spatial diffrences in growth rate and condition factor among estuarine nursery zones in Delaware Bay. The potential influence of temperature and salinity on the suitability of estuarine nursery areas for juvenile weakfish was investigated in laboratory experiments by measuring ad libitum feeding rate, growth rate and gross growth efficiency of juveniles collected in Delaware Bay in 1990 (40 to 50 mm standard length; 1.4 to 2.1 g) in 12 temperature/salinity treatments (temperatures: 20, 24, 28°C; salinities: 5, 12, 19, 26 ppt) representing conditions encountered in different estuarine zones during spring/summer. Feeding rates (FR) increased significantly with temperature at all salinities, ranging from 10 to 15% body wt d^-1 at 20°C to 33–39% body wt d^-1 at 28°C. Specific growth rates (SGR) ranged from 1.4 to 9.4% body wt d^-1 (0.3 to 1.5 mm d^-1) and gross growth efficiencies (K ₁) varied from 13.6 to 26.4% across temperature/salinity combinations. Based on nonlinear multiple regression models, predicted optimal temperatures for SGR and K ₁ were 29 and 27°C, respectively. Salinity effects on SGR and K ₁ were significant at 24 and 28°C where predicted optimal salinity was 20 ppt. At these warmer temperatures, SGR and K ₁ were significantly lower at 5 than at 19 ppt despite higher FR at 5 ppt. Therefore, maximum growth rate and growth efficiency occurred under conditions characteristic of mesohaline nurseries. This finding is consistent with spatial patterns of growth in Delaware Bay, implying that physicochemical gradients influence the value of particular estuarine zones as nurseries for juvenile weakfish by affecting the energetics of feeding and growth. Laboratory results indicate a seasonal shift in the location of physiologically optimal nurseries within estuaries. During late spring/early summer, warmer temperatures in oligohaline areas permit higher feeding rate and faster growth compared to mesohaline areas. By mid-late summer, spatial temperature gradients diminish and mesohaline areas provide more suitable physicochemical conditions for growth rate and growth efficiency whereas oligohaline areas become energetically stressful. Substantial mortality occurred at 5 ppt and 28°C, providing additional evidence that oligohaline conditions are stressful during late summer. Furthermore, juveniles provided a choice among salinities in laboratory trials preferred those salinities which promoted higher growth rates. The extensive use of oligohaline nurseries by juvenile weakfish despite the potential for reduced growth rate and growth efficiency suggests this estuarine zone may provide a substantial refuge from predation.     

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     

Field estimates of growth and mortality of juvenile banana prawns (Penaeus merguiensis)

Postlarval and juvenile Penaeus merguiensis de Man from the Embley River estuary on the north-eastern Gulf of Carpentaria were sampled every 2 wk from September 1986 until August 1989, using a small beam trawl. Settlement of planktonic postlarvae peaked during the pre-wet season (October to December), and declined through the wet season (January to March). Using length-frequency analysis between 12 and 14 cohorts of juvenile prawns were identified each year. Length-frequency analysis and modal progression were used to derive growth rates during the estuarine phase of the life cycle. Growth rates, which could be described by a linear model, ranged from 0.63 to 1.65 mm CL (carapace length) wk^-1. Growth rates were positively influenced by water temperature and negatively influenced by prawn density. Salinity had no effect on growth rates. Prawns spent between 6 and 20 wk in the Embley River before emigrating offshore from the estuary. Weekly instantaneous rates of natural mortality (M) ranged from 0.23 to 0.94, and in general were lowest during the dry season (July to September) and highest during the pre-wet and wet seasons. Only temperature significantly influenced mortality rates, with mortality rates increasing with temperature. By projecting juvenile growth rates forward through time, we established which cohorts contributed to the offshore fishery each year. In 1987 and 1988 the April fishery consisted of prawns which had settled in the river before the end of January each year. Slow growth rates during the pre-wet season of 1988 meant that only cohorts that were settled before early December 1988 contributed to the fishery in April 1989. Whether a cohort contributes to the fishery depends on the settlement date, water temperature and prawn density.     

The importance of life stage to population connectivity in whiting (Merlangius merlangus) from the northern European shelf

Juvenile movements are rarely considered in studies of connectivity. This study investigates movements in juvenile and adult life stages of an ecologically important temperate piscivore, the whiting (Merlangius merlangus). Adult movement was assessed from tag-recapture data collected between 1958 and 1980 around the coast of Scotland. Displacement distances ranged from 53 to 123 km. Adults rarely dispersed offshore in the North Sea, and exchange with the west of Scotland was negligible. Connectivity between nursery and spawning grounds was investigated using otolith microchemistry. Five juvenile nursery areas were discriminated with an accuracy of 75–85%. Comparing isotopic signatures of adults and juveniles indicated that several nursery areas contributed to each spawning aggregation. The lack of evidence for return migrations suggested an opportunistic and non-philopatric recruitment strategy within a single population unit. Differences in the proportion contribution of juveniles to adult aggregations also indicated that the “value” of nursery areas differed and the Scottish west coast could be viewed as a net source of recruits to the North Sea. Given the extent of juvenile dispersal, this study highlights the fact that all stages of the life cycle should be considered in studies of connectivity.