Seasonal trends in body size of adult male mosquitofish,Gambusia affinis,with evidence for their social control

Synopsis Collections from a natural population of mosquitofish,Gambusia affinis, in southern Indiana showed that males born early in the breeding season mature rapidly at a small body size, while males born later in the breeding season delay maturation and achieve larger body sizes. A field experiment, involving removal of males from pond populations, was conducted to test the hypothesis that delayed maturation by late summer males is, at least in part, under social control. Mean total length (TL) of adult and maturing males in late summer in ponds from which large numbers of males had previously been removed was significantly lower than mean TL of males in control ponds. These data support the hypothesis of social control and support Sohn's competition-predation hypothesis for control of adult male body size inGambusia.     

Emergent impacts of cannibalism and size refuges in prey on intraguild predation systems

Many organisms undergo ontogenetic niche shifts due to considerable changes in size during their development. These ontogenetic shifts can alter the trophic position of individuals, the type and strength of ecological interactions across species, and allow for cannibalism within species. In this study we ask if and how the interaction of a size refuge and cannibalism in the prey alters the dynamics of intraguild predation (IGP) systems. By manipulating the composition of large cannibalistic (Aeshna umbrosa) and predatory (Anax junius) dragonfly larvae in mesocosms we show that the interaction of cannibals and predators was non-linear and increased the survival of prey. The structure of the final resource community shared by prey and predator differed between small and large dragonfly treatments but not within size classes across species. In general, the small prey stage showed similar shifts in microhabitat use and refuge use when exposed to either conspecific cannibals or predators, while large cannibals showed no clear anti-predator response. However, further behavioral experiments revealed that specific behavioral components, such as distances between individuals or number of movements, differed when individuals were exposed to either cannibals or predators. This indicates that individuals discriminated between conspecific or heterospecific predators. Furthermore, in similar experiments large cannibals and predators showed different behaviors when exposed to conspecifics rather than to each other. These changes in behavior are consistent with the observed increase in prey survival. In general, the results indicate that cannibalism and ontogenetic niche shifts can result in behavior-mediated indirect interactions that reduce the impact of the predator on the mortality of its prey and alter the interactions of IGP systems. However, they also indicate that size is not the sole determinant and that we also need to account for the species identity when predicting the dynamics of communities.     

A review of the ecological parameters and implications of subsociality in Parastrachia japonensis (Hemiptera: Cydnidae), a semelparous species that specializes on a poor resource

Subsocial behavior or postovipositional parental care in insects has evolved in response to a variety of environmental stresses and ranges from briefly guarding eggs after oviposition to elaborate nidification and provisioning behaviors. Investment in parental care bears various costs, and should not continue beyond the point at which the costs to future reproductive success exceed the benefits to current reproductive effort. Progressive provisioning is a rare form of subsociality in insects. Females of the subsocial shield bug Parastrachia japonensis progressively provision their nymph-containing nests with drupes of the single resource, Schoepfia jasminodora, and this provisioning drastically enhances offspring survival. A female rears only one brood throughout her lifetime and continues provisioning the brood until about the third larval stadium, when the female dies. Thus, the female's entire reproductive effort is expressed in the success of that one brood, which suggests a reproductive strategy with enormous costs and risks. Why has such an extreme life history evolved in this species? Here, to answer that question and to contribute to our understanding of the evolutionary implications of subsocial behavior, in particular, progressive provisioning, we review what we have discovered about the ecological parameters of subsociality in this species during a long-term field study. We also discuss these parameters in P. japonensis in reference to other subsocial insects and related species and conclude with a suggestion that semelparity and progressive provisioning in this species are extreme adaptations to evolving complete dependency on an unreliable resource.     

Feeding habits of the Magellan skate: effects of sex, maturity stage, and body size on diet

The aim of this study was to evaluate the effects of sex, maturity stage, and body size on the diet of the Magellan skate, Bathyraja magellanica, in the Southwest Atlantic off Argentina, by examining stomach contents using a multiple hypothesis modeling approach. Relationships between the number of prey and sex, maturity stage, and total length (TL) were assessed by built generalized linear models (GLM). Furthermore, we tested whether there was a threshold size at which B. magellanica started or quit consuming a given prey. The overall diet of B. magellanica was mainly consisted of teleosts, followed by amphipods, isopods, and decapods. Ontogenetic diet shifts were independent of sex and maturity stage. However, discrete shifts in diet with TL were found, with individuals larger than 554 and 623 mm TL ceasing to consume amphipods and isopods, respectively. The consumption of teleosts progressively increased with increasing predator size. Likewise, ontogenetic shifts in foraging behavior were also observed with smaller individuals showing specialization on amphipods with larger specimens consuming teleosts. These results confirm that ontogenetic shifts in diet of B. magellanica are more a function of predator size rather than any other life-history traits. We propose that these food shifts are probably related to morphological limitations and abilities associated with feeding habits of skate, so when specimens of B. magellanica reach an optimum body size, they may have access to higher quality trophic resources. Our results suggest that evaluating the importance of life-history stages on the feeding habits of a species is essential for understanding how that species exploits food resources, which, in turn, is an important factor in developing a suitable plan of marine ecosystem conservation.     

Optimal age and size at maturity in annuals and perennials with determinate growth

Summary A model predicting optimal age and size at maturity is presented, exploring the conflict between growth and energy allocation to reproduction. According to the model, the factors promoting delayed maturity and large adult body size are as follows: (1) high rate of somatic growth, (2) high percentage increase in reproductive rate with body size increase, (3) long life expectancy at maturity for annuals or large number of expected productive days (when either growth or reproduction is possible) for perennials with growth ceasing at maturity, (4) life expectancy increasing with body size. All these factors are combined in the mathematical formula predicting optimal age and size at maturity, which allows for quantitative predictions. The optimal schedule of growth and reproduction may be achieved by natural selection, developmental plasticity, or when one species replaces another. Sexual size dimorphism is also discussed, resulting from different optimal age at maturity for either sex.     

Validated age and growth of the leopard shark,Triakis semifasciata,with comments on reproduction

Synopsis The age, growth, and sexual maturation of the leopard shark, Triakis semifasciata, from central California were studied. Growth band counts in vertebral centra of 162 leopard sharks produced von Bertalanffy growth curves with L, K. and t_o parameters of 1536 mm. 0.082, and -2.31, respectively, for both sexes combined. The L₈ value for females (1602 mm TL) was slightly but insignificantly higher than for males (1499 mm TL), but the K and t_o values were almost identical. Seasonal changes in size modes of young-of-the-year leopard sharks, centrum edge characteristics, and growth and tetracycline mark-recapture from the field were used to validate annual deposition of vertebral centrum band pairs. Sexual maturity was evaluated by the gonads and presence of sperm and eggs; males mature at 7 yr and at about 63% of asymptotic length, and females mature at 10 yr, and at about 72% of asymptotic length. This slow growth, late maturity, and relatively low fecundity may increase their susceptibility to over-exploitation.     

Context-dependent aggregation in Common Frog Rana temporaria tadpoles: influence of developmental stage, predation risk and social environment

1. This study examines the aggregation behaviour and activity of larvae of the Common Frog Rana temporaria in relation to the early social environment, ontogeny and the presence of chemical cues from a predatory fish to address three main questions: (i) Does previous social interaction influence aggregation behaviour in later developmental stages? (ii) To what extent does aggregation behaviour depend upon the risk level perceived by the individual? (iii) Does aggregation behaviour change through ontogeny?
2. Tadpoles were reared from eggs either in small groups or in complete isolation. In relatively early stages of development, tadpoles showed no preference for the side of the test container containing siblings over the side containing no larvae regardless of the social context experienced (isolation or contact with siblings).
3. The presence of chemical cues from a potential predator did not trigger the aggregation behaviour of tadpoles, but it had a remarkable effect on their activity and spatial distribution. Tadpoles exposed to water preconditioned by a predator spent significantly less time swimming and avoided the central area of the test container more frequently than did controls exposed to unconditioned water.
4. Tadpoles were more active, avoided the central section and associated preferentially with conspecifics (siblings) at later stages of development regardless of the social conditioning they had previously experienced.
5. Individuals reared in groups were twice as active as individuals reared in isolation. This suggests that the early social environment experienced by larvae can influence future behaviour and thereby growth and development rates.
6. The expression of conspecific attraction is probably linked to an ontogenetic shift in larval behaviour. However, reduced activity, rather than aggregation, appears to be the basic antipredator mechanism in larval Common Frog.     

Effects of diet, body size, age and temperature on growth rates in the amphipod Gammarus pulex

SUMMARY. Increase in body wet weight of Gammarus pulex fed on decaying elm leaves was followed to senescence and death. Growth in juveniles was approximately exponential; from birth to death it conformed to a logistic growth curve, with maximum absolute increments in weight about half-way through a life span of 350–450 days at 15°C. Some individuals lived longer, for up to 640–700 days. The instantaneous or specific growth rate was maximal near birth, at c. 5–6% wet wt day^?1, and declined exponentially with increasing size and age. Over the range 4.7–14.8°C there was a log-log relationship between temperature and specific growth rate. Growth was maximal at 20°C in newborn animals and at 15°C in 6–9-mg animals. The specific growth rate of young individuals was fastest on decaying leaves of elm with a well developed flora of fungi and other microorganisms. Leached elm leaves without this flora supported growth at a lower rate. The latter diet was sufficient for survival and growth of newborn individuals; detritus, faeces or other food items were not needed. Isolated specimens grew as fast as those kept in groups. Growth was generally slower on leached leaves of oak and sycamore. In newborn animals fed on the fine roots of aquatic plants (Veronica, Rorippa and Glyceria), growth was as fast as on decaying elm leaves; growth on the green living leaves of the plants was slower, as on detritus from two streams and on a pure culture of an aquatic fungus. Consumption of leached elm leaves was related to leaf thickness. In a full gut the wet weight (1.34–1.37 mg) and volume (3.8–4.1 mm³) (for 20-mg animals) was independent of leaf thickness but dependent on animal size, increasing 4-fold over the range 2–50 mg body wt. Daily consumption (dry wt) was approximately equivalent to 50% body dry wt at 5 mg and 20% at 50 mg body wet wt. Individuals fed on thick leaves ingested 50% more dry weight per day and absorbed more in the gut than when fed on thin leaves, but the relative efficiency of absorption was the same at 36–59% for 10–20-mg animals. Weight-specific absorption in the gut was highest in juveniles and decreased with increasing body weight; relative efficiency of absorption was generally lower in the larger individuals. Assuming an energy value of 5 cal mg^?1 dry wt for elm leaves, daily mean energy intake by absorption in thegutof G. pu/ex was2.2 cal mg^?1 animaldry wt (9.2 J mg^?1) in individuals of 0.4 mgdry wt (2 mg wet wt), decreasing to 0.3 cal mg^?1 (1.3 J mg^?1) at 10 mg dry wt (50 mg wet wt). Growth in Gammarus is briefly reviewed in the hght of work on other animals and it is emphasized that all aspects of feeding, growth and metabol-ism should be specifically related to size and age of the individuals, using well defined diets.     

Energetics in Liolaemini lizards: implications of a small body size and ecological conservatism

Liolaemini lizards occur in southern South America in a variety of dietary habits across a broad latitudinal and altitudinal distribution. We studied standard metabolic rates of 19 Liolaemini species and analyzed these data using both conventional and phylogenetically informed statistics. Oxygen consumption showed a significant and positive relationship with body mass (SMR = 0.109 × body mass^0.876±0.023), with a higher slope than that expected on the basis of the three-quarter power law model. After phylogenetically informed and conventional analyses, no significant differences in metabolic rates were found to be related to diet or elevation. We hypothesize that small body size, ecological conservatism and physiological compensation may explain the lack of differences in metabolic rates observed among these lizards.     

Spatial and temporal patterns in the diet of the Andean condor: ecological replacement of native fauna by exotic species

The development of conservation strategies to protect viable populations of scavenging birds requires the existence of adequate and safe food supplies in the wild. Early reports on Andean condors Vultur gryphus diet recorded guanacos and rheas, the dominant herbivores since the Pleistocene, as their main food in Patagonia. However, in the past century, guanaco and rhea populations have notably decreased as a consequence of introduced livestock, and other exotic mammals have colonized the region. We study the spatial and temporal variation of the condors' diet to determine which species are being consumed by condors, and to test whether native herbivores still have a role as a food source. We analysed 371 pellets (517 prey items), collected along 500 km in northwestern Patagonia, Argentina. Our study shows that the Andean condor depends heavily (98.5%) on exotic herbivores. Their diet was made up of c . 51% sheep/goat, 24% hare/rabbit, 17% red deer and 6% cow/horse, with only 2% other mammal species. Samples from locations surveyed after 12–15 years showed a diet shift coincident with the local tendencies in the food source. The diet composition of condors using roosts within the same zone was very similar, which suggests that they may be feeding from the same area. Thus, unhealthy carcasses could impact the entire local populations. Our results show the abundance of the invasive species in northwestern Patagonia and support the idea that native mega-herbivores are ecologically extinct in this area. Exotic species management can have a decisive impact on scavenger's survival. It is necessary to apply a strategy that includes public environmental education about the problems of scavengers (e.g. use of poison, veterinary medicines and lead bullets), and a serious productive plan, including native species as a suitable source of economic development.     

Invasion success depends on invader body size in a size-structured mixed predation–competition community

1.  The size of an individual is an important determinant of its trophic position and the type of interactions it engages in with other heterospecific and conspecific individuals. Consequently an individual's ecological role in a community changes with its body size over ontogeny, leading to that trophic interactions between individuals are a size-dependent and ontogenetically variable mixture of competition and predation.
2.  Because differently sized individuals thus experience different biotic environments, invasion success may be determined by the body size of the invaders. Invasion outcome may also depend on the productivity of the system as productivity influences the biotic environment.
3.  In a laboratory experiment with two poeciliid fishes the body size of the invading individuals and the daily amount of food supplied were manipulated.
4.  Large invaders established persistent populations and drove the resident population to extinction in 10 out of 12 cases, while small invaders failed in 10 out of 12 trials. Stable coexistence was virtually absent. Invasion outcome was independent of productivity.
5.  Further analyses suggest that small invaders experienced a competitive recruitment bottleneck imposed on them by the resident population. In contrast, large invaders preyed on the juveniles of the resident population. This predation allowed the large invaders to establish successfully by decreasing the resident population densities and thus breaking the bottleneck.
6.  The results strongly suggest that the size distribution of invaders affects their ability to invade, an implication so far neglected in life-history omnivory systems. The findings are further in agreement with predictions of life-history omnivory theory, that size-structured interactions demote coexistence along a productivity gradient.     

Species richness, species range size and ecological specialisation among African primates: geographical patterns and conservation implications

The geographical distribution of species richness and species range size of African anthropoid primates (catarrhines) is investigated and related to patterns of habitat and dietary niche breadth. Catarrhine species richness is concentrated in the equatorial regions of central and west Africa; areas that are also characterised by low average species range sizes and increased ecological specificity. Species richness declines with increasing latitude north and south of the equator, while average species range size, habitat and dietary breadth increase. Relationships between species richness, species range size and niche breadth remain once latitudinal and longitudinal effects have been removed. Among areas of lowest species richness, however, there is increased variation in terms of average species range size and niche breadth, and two trends are identified. While most such areas are occupied by a few wide-ranging generalists, others are occupied by range-restricted specialist species. That conservation efforts increasingly focus on regions of high species richness may be appropriate if these regions are also characterised by species that are more restricted in both their range size and their ecological versatility, although special consideration may be required for some areas of low species richness.     

Evolutionary influences on body size in free-living and parasitic isopods

The influence of mode of life and habitat characteristics on the evolution of body size in isopods was investigated in a comparative analysis based on data from 746 free-living and parasitic species. The phylogeny of isopods allowed 24 independent comparisons to be made between higher taxa (families or superfamilies), each corresponding to a separate branching event. The evolution of parasitism was consistently associated with reductions in body size. On the contrary, invasion of freshwater habitats was consistently coupled with increases in body size. Lineages moving to higher latitudes were significantly more likely to evolve larger body sizes than those shifting toward the equator. In addition, colonizing deeper water resulted in a weak tendency to evolve larger body size. The analysis suggests that the large size of some isopod groups parasitic on fish (e.g. Cymothoidae) may have been inherited from a free-living ancestor and is not the product of directional selection toward large size and greater fecundity.     

The influence of maternal age and mating frequency on egg size and offspring performance in Callosobruchus maculatus (Coleoptera: Bruchidae)

Maternal age influences offspring quality of many species of insects. This observed maternal age influence on offspring performance may be mediated through maternal age effects on egg size, which in turn may be directly influenced by the female's nutritional state. Thus, behaviors that influence a female's nutritional status will indirectly influence egg size, and possibly offspring life histories. Because males provide nutrients to females in their ejaculate, female mating frequency is one behavior which may influence her nutritional status, and thus the size of her eggs and the performance of her offspring. In this paper, I first quantify the influences of maternal age on egg size and offspring performance of the bruchid beetle, Callosobruchus maculatus. I then examine whether nutrients transferred during copulation reduce the magnitude of maternal age effects on egg size and larval performance when mothers are nutrient-stressed. Egg size and egg hatchability decreased, and development time increased, with increasing maternal age. Multiple mating and adult feeding by females both resulted in increased egg size. This increase in egg size of females mated multiply did not translate into reduced development time or increased body size and egg hatchability, but did correlate with improved survivorship of offspring produced by old mothers. Thus, it appears that because the influence of mating frequency on egg size is small relative to the influence of maternal age, the influence of nutrients derived from multiple mating on offspring life history is almost undetectable (detected only as a small influence on survivorship). For C. maculatus, female multiple mating has been demonstrated to increase adult female survivorship (Fox 1993a), egg production (Credland and Wright 1989; Fox 1993a), egg size, and larval survivorship, but, contrary to the suggestion of Wasserman and Asami (1985), multiple mating had no detectable influence on offspring development time or body size.     

Understanding ecological community succession: Causal models and theories,a review

Critical review of explanations for patterns of natural succession suggests a strong, common basis for theoretical understanding, but also suggests that several well known models are incomplete as explanations of succession. A universal, general cause for succession is unlikely, since numerous aspects of historical and environmental circumstances will impinge on the process in a unique manner. However, after disturbance, occupation of a site by any species causes changes in the conditions at the site. Sorting of species may result, since different species are adapted to different regions of environmental gradients. Such sorting can generate several patterns of species abundance in time, but commonly results in sequential replacements of species adapted to the varying conditions. This may be due to constraints on species' strategies, or life history traits, placed by the limited resources available to the organism. These constraints often result in inverse correlation between traits which confer success during early and late stages of succession. Facilitatory or inhibitory effects of species on each other are best understood in terms of these life history interactions, perhaps as restrictions on, or as moderation of, these processes.Strong support for the importance of correlations in life history traits stems from comparisons of simulated succession with and without these correlations. These simulations are reviewed in some detail, and followed by brief reviews of other prominent models for succession. Several aspects of the confusion and controversies in the successional literature are then discussed, with a view to a more optimistic synthesis and direction for successional ecology.     

The evolution of reaction norms: simple models for age and size at maturity

This paper presents a simple model for the evolution of reaction norms for age and size at maturity that predicts reaction norms with a variety of shapes. Using realistic parameter values the model predicts reaction norms close to those observed in Drosophila. The major assumptions of the model are: 1) that net reproductive rate is maximized, 2) that growth is determinate, and 3) that mortality rates are independent of age and size at maturity. If, additionally, juvenile mortality is uncorrelated with a growth coefficient, k, the model predicts that selection favors maturation later at a smaller size when k is reduced by environmental factors and that decreased juvenile mortality leads to delayed maturity. These two predictions conform with those found by previous models using other measures of fitness. Correlations between k and juvenile mortality can change the shape of the predicted reaction norm. Depending on the precise form of the correlation, the model can predict done- or bowl-shaped reaction norms and can predict delayed or earlier maturity as k decreases. These shapes are qualitatively different from those predicted by previous models that used different fitness measures. Systematic estimates of the parameter values for this and for related models are required to determine the appropriate fitness measure for models of reaction norms.     

Pikeperch Sander lucioperca trapped between niches: foraging performance and prey selection in a piscivore on a planktivore diet

The foraging behaviour of planktivorous pikeperch Sander lucioperca during their first growing season was analysed. Field data showed that S. lucioperca feed on extremely rare prey at the end of the summer, suggesting the presence of a bottleneck. In experiments, foraging ability of planktivorous S. lucioperca was determined when fish were feeding on different prey types (Daphnia magna or Chaoborus spp.) and sizes (D. magna of lengths 1 or 2·5 mm) when they occurred alone. From these results, the minimum density requirement of each prey type was analysed. The energy gain for three different foraging strategies was estimated; a specialized diet based on either large D. magna or Chaoborus spp. or a generalist diet combining both prey types. Prey value estimates showed that Chaoborus spp. should be the preferred prey, assuming an energy maximizing principle. In prey choice experiments, S. lucioperca largely followed this principle, including D. magna in the diet only when the density of the Chaoborus spp. was below a threshold value. Splitting the foraging bout into different sequences, however, resulted in a somewhat different pattern. During an initial phase, S. lucioperca captured both prey as encountered and then switched to Chaoborus spp. if prey density was above the threshold level. The prey selection observed was mainly explained by sampling behaviour and incomplete information about environmental quality, whereas satiation only had marginal effects. It was concluded that the observed diet based on rare prey items was in accordance with an optimal foraging strategy and may generate positive growth in the absence of prey fish in suitable sizes.     

Persistent directional selection on body size and a resolution to the paradox of stasis

Directional selection on size is common but often fails to result in microevolution in the wild. Similarly, macroevolutionary rates in size are low relative to the observed strength of selection in nature. We show that many estimates of selection on size have been measured on juveniles, not adults. Further, parents influence juvenile size by adjusting investment per offspring. In light of these observations, we help resolve this paradox by suggesting that the observed upward selection on size is balanced by selection against investment per offspring, resulting in little or no net selection gradient on size. We find that trade‐offs between fecundity and juvenile size are common, consistent with the notion of selection against investment per offspring. We also find that median directional selection on size is positive for juveniles but no net directional selection exists for adult size. This is expected because parent–offspring conflict exists over size, and juvenile size is more strongly affected by investment per offspring than adult size. These findings provide qualitative support for the hypothesis that upward selection on size is balanced by selection against investment per offspring, where parent–offspring conflict over size is embodied in the opposing signs of the two selection gradients.     

Resolving the roles of body size and species identity in driving functional diversity

Efforts to characterize food webs have generated two influential approaches that reduce the complexity of natural communities. The traditional approach groups individuals based on their species identity, while recently developed approaches group individuals based on their body size. While each approach has provided important insights, they have largely been used in parallel in different systems. Consequently, it remains unclear how body size and species identity interact, hampering our ability to develop a more holistic framework that integrates both approaches. We address this conceptual gap by developing a framework which describes how both approaches are related to each other, revealing that both approaches share common but untested assumptions about how variation across size classes or species influences differences in ecological interactions among consumers. Using freshwater mesocosms with dragonfly larvae as predators, we then experimentally demonstrate that while body size strongly determined how predators affected communities, these size effects were species specific and frequently nonlinear, violating a key assumption underlying both size- and species-based approaches. Consequently, neither purely species- nor size-based approaches were adequate to predict functional differences among predators. Instead, functional differences emerged from the synergistic effects of body size and species identity. This clearly demonstrates the need to integrate size- and species-based approaches to predict functional diversity within communities.