Allocation in reproduction is not tailored to the probable number of matings in common toad (<Emphasis Type="Italic">Bufo bufo</Emphasis>) males

The theory of life history evolution assumes trade-offs between competing fitness traits such as reproduction, somatic growth, and maintenance. One prediction of this theory is that if large individuals have a higher reproductive success, small/young individuals should invest less in reproduction and allocate more resources in growth than large/old individuals. We tested this prediction using the common toad (Bufo bufo), a species where mating success of males is positively related to their body size. We measured testes mass, soma mass, and sperm stock size in males of varying sizes that were either (1) re-hibernated at the start of the breeding season, (2) kept without females throughout the breeding season, or (3) repeatedly provided with gravid females. In the latter group, we also estimated fertilization success and readiness to re-mate. Contrary to our predictions, the relationship between testes mass and soma mass was isometric, sperm stock size relative to testes mass was unrelated to male size, fertilization success was not higher in matings with larger males, and smaller males were not less likely to engage in repeated matings than larger males. These results consistently suggest that smaller males did not invest less in reproduction to be able to allocate more in growth than larger males. Causes for this unexpected result may include relatively low year-to-year survival, unpredictable between-year variation in the strength of sexual selection and low return rates of lowered reproductive investment.     

Bigger is not always better: offspring size does not predict growth or survival for seven ascidian species

The presumed trade-off between offspring size and quality predicted by life history theory is often invoked to explain the wide range of propagule sizes observed in animals and plants. This trade-off is broadly supported by intraspecific studies but has been difficult to test in an interspecific context, particularly in animals. We tested the fitness consequences of offspring size both intra- and interspecifically for seven species of ascidians (sessile, suspension-feeding, marine invertebrates) whose offspring volumes varied over three orders of magnitude. We measured two major components of fitness, juvenile growth rates and survival, in laboratory and field experiments encompassing several food conditions. Contrary to the predictions of life history theory, larger offspring size did not result in higher rates of growth or survival, and large offspring did not perform better under nutritional stress, either intraspecifically or interspecifically. In fact, two of the four species with small offspring grew rapidly enough to catch up in size to the species with large offspring in as little as eight weeks, under wild-type food conditions. Trade-offs between growth potential and defense may overwhelm and obscure any trade-offs between offspring size and survival or growth rate. While large initial size may still confer a competitive advantage, we failed to detect any consequences of interspecific variation in initial size. This implies that larger offspring in these species, far from being inherently superior in growth or survival, require compensation in other aspects of life history if reproductive effort is to be efficient. Our results suggest that the importance of initial offspring size is context dependent and often overestimated relative to other life history traits.     

Life history trade-offs in tropical trees and lianas

It has been hypothesized that tropical trees partition forest light environments through a life history trade-off between juvenile growth and survival; however, the generality of this trade-off across life stages and functional groups has been questioned. We quantified trade-offs between growth and survival for trees and lianas on Barro Colorado Island (BCI), Panama using first-year seedlings of 22 liana and 31 tree species and saplings (10 mm < dbh < 39 mm) of 30 tree species. Lianas showed trade-offs similar to those of trees, with both groups exhibiting broadly overlapping ranges in survival and relative growth rates as seedlings. Life history strategies at the seedling stage were highly correlated with those at the sapling stage among tree species, with all species showing an increase in survival with size. Only one of 30 tree species demonstrated a statistically significant ontogenetic shift, having a relatively lower survival rate at the sapling stage than expected. Our results indicate that similar life history trade-offs apply across two functional groups (lianas and trees), and that life history strategies are largely conserved across seedling and sapling life-stages for most tropical tree species.     

Extremes, plasticity, and invariance in vertebrate life history traits: insights from coral reef fishes

Life history theory predicts a range of directional generic responses in life history traits with increasing organism size. Among these are the relationships between size and longevity, mortality, growth rate, timing of maturity, and lifetime reproductive output. Spanning three orders of magnitude in size, coral reef fishes provide an ecologically diverse and species-rich vertebrate assemblage in which to test these generic responses. Here we examined these relationships by quantifying the life cycles of three miniature species of coral reef fish from the genus Eviota (Gobiidae) and compared their life history characteristics with other reef fish species. We found that all three species of Eviota have life spans of < 100 days, suffer high daily mortality rates of 7-8%, exhibit rapid linear growth, and matured at an earlier than expected size. Although lifetime reproductive output was low, consistent with their small body sizes, short generation times of 47-74 days help overcome low individual fecundity and appear to be a critical feature in maintaining Eviota populations. Comparisons with other coral reef fish species showed that Eviota species live on the evolutionary margins of life history possibilities for vertebrate animals. This addition of demographic information on these smallest size classes of coral reef fishes greatly extends our knowledge to encompass the full size spectrum and highlights the potential for coral reef fishes to contribute to vertebrate life history studies.     

The role of life histories and trophic interactions in population recovery

Factors affecting population recovery from depletion are at the focus of wildlife management. Particularly, it has been debated how life‐history characteristics might affect population recovery ability and productivity. Many exploited fish stocks have shown temporal changes towards earlier maturation and reduced adult body size, potentially owing to evolutionary responses to fishing. Whereas such life‐history changes have been widely documented, their potential role on stock's ability to recover from exploitation often remains ignored by traditional fisheries management. We used a marine ecosystem model parameterized for Southeastern Australian ecosystem to explore how changes towards “faster” life histories might affect population per capita growth rate r. We show that for most species changes towards earlier maturation during fishing have a negative effect (3–40% decrease) on r during the recovery phase. Faster juvenile growth and earlier maturation were beneficial early in life, but smaller adult body sizes reduced the lifetime reproductive output and increased adult natural mortality. However, both at intra‐ and inter‐specific level natural mortality and trophic position of the species were as important in determining r as species longevity and age of maturation, suggesting that r cannot be predicted from life‐history traits alone. Our study highlights that factors affecting population recovery ability and productivity should be explored in a multi‐species context, where both age‐specific fecundity and survival schedules are addressed simultaneously. It also suggests that contemporary life‐history changes in harvested species are unlikely to increase their resilience and recovery ability.     

Size-selective harvesting alters life histories of a temperate sex-changing fish.

Selective mortality, whether caused naturally by predation or through the influence of harvest practices, initiates changes within populations when individuals possessing certain heritable traits have increased fitness. Theory predicts that increased mortality rates will select for changes in a number of different life history characteristics. For example, fishing often targets larger individuals and has been shown repeatedly to alter population size structure and growth rates, and the timing of maturation. For sex-changing species, selective fishing practices can affect additional traits such as the mature population sex ratio and the timing of sexual transformation. Using historical comparisons, we examined the effects of exploitation on life history characteristics of California sheephead, Semicossyphus pulcher, a temperate protogynous (female-male sex changer) labrid that inhabits nearshore rocky environments from central California, USA, to southern Baja California, Mexico. Recreational fishing intensified and an unregulated commercial live-fish fishery developed rapidly in southern California between the historical and current studies. Collections of S. pulcher from three locations (Bahia Tortugas, Catalina Island, and San Nicolas Island) in 1998 were compared with data collected 20-30 years previously to ascertain fishery-induced changes in life history traits. At Bahia Tortugas, where fishing by the artisanal community remained light and annual survivorship stayed high, we observed no changes in size structure or shifts in the timing of maturation or the timing of sex change. In contrast, where recreational (Catalina) and commercial (San Nicolas) fishing intensified and annual survivorship correspondingly declined, males and females shifted significantly to smaller body sizes, females matured earlier and changed sex into males at both smaller sizes and younger ages and appeared to have a reduced maximum lifespan. Mature sex ratios (female:male) increased at San Nicolas, despite a twofold reduction in the mean time spent as a mature female. Proper fisheries management requires measures to prevent sex ratio skew, sperm limitation, and reproductive failure because populations of sequential hermaphrodites are more sensitive to size-selective harvest than separate-sex species. This is especially true for S. pulcher, where different segments of the fishery (commercial vs. recreational) selectively target distinct sizes and therefore sexes in different locations.     

Population maintenance of the short-lived shrub Sambucus in a deciduous forest

This study quantitatively clarifies the life history of a shrub, Sambucus racemosa ssp. sieboldiana, in an old-growth forest, the Ogawa Forest Reserve, Japan, by a demographic approach using a projection matrix model that incorporates interactions between demographic parameters and canopy height dynamics. S. racemosa is a common deciduous shrub in central Japan and is known to grow predominantly at forest edges or roadsides. This indicates that it is a highly light-demanding species, and occurrence in gaps in old-growth stands suggests its "fugitive," gap-dependent life history in old-growth forests. We found that one distinctive feature of this species was that its seedlings can survive well in shaded conditions by alternating stems every year like perennial herb species. Matrix model analyses demonstrated that S. racemosa can continuously regenerate under the present disturbance regime of this forest and is highly adaptable to the structural dynamics of the old-growth forest. The maturity of S. racemosa shrubs depends on their size, and nearly all (>90%) of the mature (reproducing) individuals were found in gaps or near gaps. But wide seed dispersal by birds and the ability to form both seed banks and seedling banks, the latter of which has been regarded as a common characteristic of shade-tolerant climax species, probably increase the species' chances to encounter canopy gaps. Dynamic-canopied matrix models showed that the greatest elasticity is with shaded seedling survival. The frequent stem alternation of shaded seedlings often makes the growth rate negative, but the survival rate of seedlings in low light awaiting new gap creation is remarkably high (0.93 yr(-1)). The lower survival rate of the larger individuals and smaller minimum size to start reproduction than other canopy or subcanopy shade-tolerant species indicate that S. racemosa has the potential to reproduce before the closure of the encountered gaps and to complete its life history rapidly.     

Minimum area requirements for an at‐risk butterfly based on movement and demography

Determining the minimum area required to sustain populations has a long history in theoretical and conservation biology. Correlative approaches are often used to estimate minimum area requirements (MARs) based on relationships between area and the population size required for persistence or between species’ traits and distribution patterns across landscapes. Mechanistic approaches to estimating MAR facilitate prediction across space and time but are few. We used a mechanistic MAR model to determine the critical minimum patch size (CMP) for the Baltimore checkerspot butterfly (Euphydryas phaeton), a locally abundant species in decline along its southern range, and sister to several federally listed species. Our CMP is based on principles of diffusion, where individuals in smaller patches encounter edges and leave with higher probability than those in larger patches, potentially before reproducing. We estimated a CMP for the Baltimore checkerspot of 0.7–1.5 ha, in accordance with trait‐based MAR estimates. The diffusion rate on which we based this CMP was broadly similar when estimated at the landscape scale (comparing flight path vs. capture‐mark‐recapture data), and the estimated population growth rate was consistent with observed site trends. Our mechanistic approach to estimating MAR is appropriate for species whose movement follows a correlated random walk and may be useful where landscape‐scale distributions are difficult to assess, but demographic and movement data are obtainable from a single site or the literature. Just as simple estimates of lambda are often used to assess population viability, the principles of diffusion and CMP could provide a starting place for estimating MAR for conservation.     

Complex social structure of southern flying squirrels is related to spatial proximity but not kinship

Social individuals have organized relationships that affect fitness and so a species' tendency to be social has important implications for its population ecology, gene flow, and its distribution in space and time. We quantitatively examined the social structure of southern flying squirrels (Glaucomys volans) and tested for a role of kinship and prior familiarity in predicting social structure. To quantify social structure, we monitored nest group composition of southern flying squirrels. All squirrels at the study site were marked with passive integrated transponder (PIT) tags and nest cavity entrances were monitored with automated PIT tag recorders for a period of 28 months. Squirrels were genotyped at eight microsatellite loci. Permutation tests of associations suggested that individuals nested with other specific individuals more often than expected by chance. The lagged association rate indicated that relationships were stable and persisted across seasons and years. Multiple summer nest associates came together in winter to form larger nest groups which were likely important for social thermoregulation. A measure of prior familiarity, but not kinship, was related to the proportion of time individuals nested together during winter. We suggest that the evolution of sociality in southern flying squirrels is driven largely by mutually beneficial behaviors related to social thermoregulation although other, as of yet, unidentified mechanisms are needed to explain sociality in the warm season. We hypothesize that minimum group size requirements associated with social thermoregulation could explain the absence of this species in patchy landscapes and aspects of range boundary dynamics near their northern range boundary.     

Effects of symbiont elimination and reinfection on the life processes of the planktonic foraminifer Globigerinoides sacculifer

Laboratory experiments were carried out to determine the influence of symbiotic dinoflagellates (zooxanthellae) on the shell growth, longevity, and reproductive potential of Globigerinoides sacculifer (Brady). Its symbionts were eliminated by 72-h treatment with a photosynthetic inhibitor (DCMU). Symbiont elimination resulted in earlier gametogenesis (shortened survival time) and smaller shell sizes of G. sacculifer when compared to untreated foraminifera grown in sea water. Individuals kept in continuous darkness in untreated sea water also exhibited early gametogenesis, short survival times and small shell sizes. Aposymbiotic foraminifera formed on the average one or two chambers fewer per individual and their rate of shell size increase is slower than symbiont-bearing foraminifera. Symbionts were lysed within perialgal vacuoles of G. sacculifer when subjected to DCMU treatment or kept in continuous darkness. One DCMU-treated group was reinfected with symbionts from crushed G. sacculifer donors. Soon after reinfection, these foraminifera resumed a shell growth rate and exhibited developmental stages that were nearly equivalent to those of untreated individuals, as deduced from their shell size, frequency of sac-like chambers, rate of gametogenesis, and survival time. Our experiments indicate that the symbionts aid in calcification and that elimination of symbionts triggers gametogenesis, thus shortening the life span of the foraminiferal host. The results imply that shell growth in symbiont-bearing planktonic foraminifera occurs mainly in the euphotic zone and that they do not survive for long periods below it.     

Movement patterns of young Caribbean reef sharks, Carcharhinus perezi, at Fernando de Noronha Archipelago, Brazil: the potential of marine protected areas for conservation of a nursery ground

The movement patterns and long-term site-fidelity of primarily juvenile Caribbean reef sharks, Carcharhinus perezi, were investigated using tag-recapture and automated telemetry at an insular nursery area, the Fernando de Noronha Archipelago, Brazil. Of the 143 externally tagged juvenile sharks (<110 cm), 22 (15.3%) were recaptured between 0 and 5 km from the site of tagging after 5–800 days at liberty, suggesting some site-fidelity in young individuals of this species. Site-fidelity and movement patterns of ten juvenile sharks ranging from 78 to 110 cm total length (TL) and one opportunistically captured adult female (224 cm TL) were also investigated for periods of up to 2 years with an array of automated telemetry receivers. Tagging and telemetry data from both inside and outside a marine protected area (MPA) show that shark abundance and activity is greatest along the part of the archipelago’s coastline least disturbed by human activity. Telemetry tracking also showed that juvenile reef sharks demonstrated a high degree of site-fidelity and occupied specific locations along the coast throughout the year, with some evidence of an increase in activity space with ontogeny. Sharks appeared to range more widely at night and there were no seasonal variations in habitat use. Our results suggest that MPAs may be a useful conservation tool to protect young C. perezi and potentially other reef-dwelling carcharhinid sharks during their early life history.     

From metamorphosis to maturity in complex life cycles: equal performance of different juvenile life history pathways

Performance in one stage of a complex life cycle may affect performance in the subsequent stage. Animals that start a new stage at a smaller size than conspecifics may either always remain smaller or they may be able to "catch up" through plasticity, usually elevated growth rates. We study how size at and date of metamorphosis affected subsequent performance in the terrestrial juvenile stage and lifetime fitness of spadefoot toads (Pelobates fuscus). We analyzed capture-recapture data of > 3000 individuals sampled during nine years with mark-recapture models to estimate first-year juvenile survival probabilities and age-specific first-time breeding probabilities of toads, followed by model selection to assess whether these probabilities were correlated with size at and date of metamorphosis. Males attained maturity after two years, whereas females reached maturity 2-4 years after metamorphosis. Age at maturity was weakly correlated with metamorphic traits. In both sexes, first-year juvenile survival depended positively on date of metamorphosis and, in males, also negatively on size at metamorphosis. In males, toads that metamorphosed early at a small size had the highest probability to reach maturity. However, because very few toadlets metamorphosed early, the vast majority of male metamorphs had a very similar probability to reach maturity. A matrix projection model constructed for females showed that different juvenile life history pathways resulted in similar lifetime fitness. We found that the effects of date of and size at metamorphosis on different juvenile traits cancelled each other out such that toads that were small or large at metamorphosis had equal performance. Because the costs and benefits of juvenile life history pathways may also depend on population fluctuations, ample phenotypic variation in life history traits may be maintained.     

Survival and Recruitment of Captive-Reared and Wild-Reared Takahe in Fiordland, New Zealand

Captive rearing for release back into the wild is considered a useful management tool for endangered species because it can potentially increase the rate of recruitment by bypassing the early, high-risk stages in an individual's life history. In evaluating the benefits of captive rearing to conservation, it is important to monitor the survival rate of animals after release, to be sure that they have the skills necessary for survival in the wild. Using radio telemetry, we compared the movement and survival of captive-reared Takahe (  Porphyrio mantelli), a large flightless rail endemic to New Zealand, to wild-reared Takahe in the rugged mountains of Fiordland over a 5–year period. The results indicated that captive-reared birds survived at least as well as wild-reared birds. Survival of wild-reared Takahe up to 1 year of age, which is prior to the release of captive-reared birds, was poor over two winters marked by particularly cold temperatures, which made the benefits of captive rearing more pronounced. Differences in post-release movements and habitat selection of the two groups did not have a detrimental effect on survival rate of captive-reared birds. Although there was no difference in the survival rate of captive-reared females versus males, eight out of nine (89%) surviving females have formed pairs since their release compared with only two of eight (25%) males. This unexpected result suggests there may be a shortage of females in the wild population. We conclude that captive rearing for release back into the wild increases the adult Takahe population in Fiordland.     

Age-size plasticity for reproduction in monocarpic plants

Empirical and theoretical investigations of monocarpy have usually addressed the question of minimum or threshold sizes for reproduction. However, the range of flowering sizes observed in many monocarpic species is extraordinarily large (well beyond what can be called a "threshold"), and the sizes of flowering and nonflowering plants may overlap greatly. We attempt to explain these reproductive patterns in terms of optimal reaction norms predicted by simple deterministic life history models. We assume that individuals differ in their growth trajectories due to the heterogeneous quality of microsites and ask how the optimal age and size at flowering varies with environmental variation in growth and for different assumptions about fecundity and mortality. Under two very different growth functions (one with no age- or size-related decline in growth rate and another with such a decline as size approaches an asymptote), the optimal reaction norms imply considerable plasticity for size at reproduction, particularly when poor growth is associated with higher mortality or lower asymptotic size. Deterministic models such as these may be more applicable to long-lived than to short-lived monocarps, because fitness potential should be less affected by stochastic variability in yearly growing condition in the former than in the latter. We consider the case of a tropical monocarpic and masting tree species, Cerberiopsis candelabra (Apocynaceae), and show that our model results can account for wide ranges of reproductive size and overlap in size of flowering and nonflowering plants, in accord with observation. We suggest that empirical attention to norms of reaction across growth environments will be a more profitable approach than investigation of size thresholds per se.     

Using simulation to explore the functional relationships of terrestrial carnivore population indices

Population indices based on visits to detection stations commonly are used to monitor wildlife populations. Inferences about populations are based on 1 of 2 measures: (1) change in the proportion of stations visited at least once or (2) change in the cumulative number of visits by unique individuals. The functional relationships between index responses and population density is poorly understood and can lead to misinterpretation of index data when an incorrect functional relationship (e.g. linear) is assumed. We created a flexible simulation environment to study the response of detection-based population indices under a wide variety of conditions meant to reflect species life history and study design. Proportional indices exhibited non-linear saturating responses to changes in population density while cumulative indices responded linearly. Shapes of responses were functions of home range sizes, individual detection probabilities, and spatial arrangement of animals and sampling stations. Non-linear relationships of proportional indices lead to under-estimation of mean population density when data are aggregated from multiple detection stations deployed in a heterogeneous landscape. Cumulative indices have significant statistical advantages over proportional indices including smaller sample sizes required to detect density change, linearity, consistent index responses across a wide range of densities, and ability to aggregate data to meet minimum sample size requirements. Our simulation provides a flexible tool for the interpretation of station-based population indices.     

Inbreeding Depression Accumulation across Life‐History Stages of the Endangered Takahe

Abstract: Studies evaluating the impact of inbreeding depression on population viability of threatened species tend to focus on the effects of inbreeding at a single life‐history stage (e.g., juvenile survival). We examined the effects of inbreeding across the full life‐history continuum, from survival up to adulthood, to subsequent reproductive success, and to the recruitment of second‐generation offspring, in wild Takahe ( Porphyrio hochstetteri ) by analyzing pedigree and fitness data collected over 21 breeding seasons. Although the effect size of inbreeding at individual life‐history stages was small, inbreeding depression accumulated across multiple life‐history stages and ultimately reduced long‐term fitness (i.e., successful recruitment of second‐generation offspring). The estimated total lethal equivalents (2B) summed across all life‐history stages were substantial (16.05, 95% CI 0.08–90.8) and equivalent to an 88% reduction in recruitment of second‐generation offspring for closely related pairs (e.g., sib–sib pairings) relative to unrelated pairs (according to the pedigree). A history of small population size in the Takahe could have contributed to partial purging of the genetic load and the low level of inbreeding depression detected at each single life‐history stage. Nevertheless, our results indicate that such “purged” populations can still exhibit substantial inbreeding depression, especially when small but negative fitness effects accumulate across the species’ life history. Because inbreeding depression can ultimately affect population viability of small, isolated populations, our results illustrate the importance of measuring the effects of inbreeding across the full life‐history continuum.     

Elasticity analyses of size-based red and white abalone matrix models: management and conservation.

Prospective elasticity analyses have been used to aid in the management of fished species and the conservation of endangered species. Elasticities were examined for deterministic size-based matrix models of red abalone, Haliotis rufescens, and white abalone, H. sorenseni, to evaluate which size classes influenced population growth (lambda) the most. In the red abalone matrix, growth transitions were determined from a tag recapture study and grouped into nine size classes. In the white abalone matrix, abalone growth was determined from a laboratory study and grouped into five size classes. Survivorship was estimated from tag recapture data for red abalone using a Jolly-Seber model with size as a covariate and used for both red and white abalone. Reproduction estimates for both models used averages of the number of mature eggs produced by female red and white abalone in each size class from four-year reproduction studies. Population growth rate (lambda) was set to 1.0, and the first-year survival (larval survival through to the first size class) was estimated by iteration. Survival elasticities were higher than fecundity elasticities in both the red and white matrix models. The sizes classes with the greatest survival elasticities, and therefore the most influence on population growth in the model, were the sublegal red abalone (150-178 mm) and the largest white abalone size class (140-175 mm). For red abalone, the existing minimum legal size (178 mm) protects the size class the model suggests is critical to population growth. Implementation of education programs for novice divers coupled with renewed enforcement may serve to minimize incidental mortality of the critical size class. For white abalone, conservation efforts directed at restoring adults may have more of an impact on population growth than efforts focusing on juveniles. Our work is an example of how prospective elasticity analyses of size-structured matrix models can be used to quantitatively evaluate research priorities, fishery management strategies, and conservation options.     

Threshold to maturity in a long-lived reptile: interactions of age, size, and growth

Thresholds to sexual maturity—either age or size—are critical life history parameters. Usually investigated in short-lived organisms, these thresholds and interactions among age, size, and growth are poorly known for long-lived species. A 34-year study of captive green turtles (Chelonia mydas) that followed individuals from hatching to beyond maturity provided an opportunity to evaluate these parameters in a long-lived species with late maturity. Age and size at maturity are best predicted by linear growth rate and mass growth rate, respectively. At maturity, resource allocation shifts from growth to reproductive output, regardless of nutrient availability or size at maturity. Although captive turtles reach maturity at younger ages than wild turtles, the extensive variation in captive turtles under similar conditions provides important insights into the variation that would exist in wild populations experiencing stochastic conditions. Variation in age/size at maturity should be incorporated into population models for conservation and management planning.     

Age structure,growth rates,movement patterns and feeding in an estuarine population of the cardinalfish Apogon rueppellii

The biology of a population of the cardinalfish Apogon rueppellii has been studied over several years (1977–1983) in the Swan Estuary in south-western Australia, using ramples collected monthly from the shallows by beach seine and from various depths by otter trawl. While the life cycle of this species typically lasts for one year, at the end of which time the mean length is 50 to 60 mm, some individuals survive for a further year and attain lengths up to 104 mm. A. rueppellii shows a marked tendency to move offshore into deeper water during the winter months. This tendency is more pronounced in the 1+ than in the 0+ year class and in larger than smaller 0+ individuals. An inshore movement of A. rueppellii in the spring is followed by spawning and by oral brooding by the males, which leads to the recruitment of large numbers of a new 0+ year class on to the banks during the summer. The offshore movement is correlated with changes in salinity and temperature. The larger catches taken by otter trawl during the day than at night indicate that A. rueppellii exhibits a diel pattern of activity. Mean fecundity ranged from 70 in the 45 to 49 mm size class to 345 in the 90 to 94 mm size class. Measurements of fecundity and the number of oral-brooded eggs demonstrated that the majority of the eggs released by the female are collected and incubated by the males. Copepods are ingested in relatively greater amounts by small than by large A. rueppellii, whereas the reverse situation occurs with larger crustaceans, polychaetes and small fish. The presence of greater amounts of copepods in the diet during the day and of amphipods at night probably reflects the diel activity patterns of the prey.