Environment-dependent genetic correlations between development time and body mass in a scorpionfly

Development time and body mass at maturation are two important fitness traits fundamental for our understanding of life history theory. Generally, fast development is associated with small adult body mass, as it will take longer to grow large. However, the strength of this trade-off may depend on average food availability, as the potential benefit of long development will depend on the rate of food intake. Here, I report results of a food manipulation experiment during larval development of the scorpionfly Panorpa cognata (Insecta, Mecoptera). Development time showed considerable genetic variation, yet food level had no influence and there was a strong genetic correlation in development time across environments. As expected, larval and adult body weight was significantly affected by food availability. Furthermore, body mass was influenced by a highly significant genotype-by-environment interaction. The reaction norm for body mass in response to food treatment was much stronger in families with long development time compared with rapidly developing genotypes. This effect was accompanied by a shift in the genetic correlation between development time and body size when comparing the two food levels. Specifically, the genetic correlation between body mass and development time changed from being positive at high food levels to a negative genetic correlation at low food levels. These results are consistent with other empirical findings demonstrating a similar shift in genetic correlations between body mass and development time when comparing favourable and unfavourable environmental conditions.     

Seasonality, body condition, and timing of reproduction in Propithecus verreauxi verreauxi in the Kirindy Forest

Mammals that live in seasonal environments may adjust their reproductive cycles to cope with fluctuations in food availability. Because lemurs in Madagascar experience highly seasonal variation in food availability, we examined the effects of fluctuating food availability on body condition and reproduction in one of the larger living species, Verreaux's sifaka (Propithecus verreauxi verreauxi), in the Kirindy Forest of western Madagascar. Seven years of demographic data were combined with an intensive study of 25 individuals over the course of 18 months. In contrast to other populations of Verreaux's sifaka, females were found to have greater body mass than males. Both male and female sifaka exhibited significant losses of body mass and fat during the dry season. Females were more likely to give birth and successfully wean an infant when they had higher body mass during the mating season. They mated during the periods of high and declining food availability, gave birth during the lean season, and then timed mid/late lactation with the period of increasing food availability. Thus, we conclude that sifaka follow the "classic" reproductive strategy (sensu van Schaik and van Noordwijk [Journal of Zoology (London) 206:533-549, 1985]).     

Prey size, prey nutrition, and food handling by shrews of different body sizes

We tested some predictions relating metabolic constraints offoraging behavior and prey selection by comparing food handlingand utilization in four sympatric shrew species: Sorex minutus(mean body mass = 3.0 g), S. araneus (8.0 g), Neomys anomalus(10.0 g), and N. fodiens (14.4 g). Live fly larvae, mealwormlarvae, and aquatic arthropods were offered to shrews as smallprey (body mass <0.1 g). Live earthworms, snails, and smallfish were offered as large prey (>0.3 g). The larvae werethe high-nutrition food (>8 kJ/g), and the other prey werethe low-nutrition food (<4 kJ/g). The smallest shrew, S.minutus, utilized (ate + hoarded) <30% of offered food,and the other species utilized >48% of food. The largerthe shrew, the more prey it ate per capita. However, highlyenergetic insect larvae composed 75% of food utilized by S.minutus and only >40% of the food utilized by the other species. Thus, inverse relationships appeared between shrewbody mass and mass-specific food mass utilization and betweenshrew body mass and mass-specific food energy utilization:the largest shrew, N. fodiens, utilized the least food massand the least energy quantity per 1 g of its body mass. Also,the proportion of food hoarded by shrews decreased with increase in size of shrew. With the exception of S. araneus, the sizeof prey hoarded by the shrews was significantly larger thanthe size of prey eaten. Tiny S. minutus hoarded and ate smallerprey items than the other shrews, and large N. fodiens hoardedlarger prey than the other shrews.     

Maternal body size as an evolutionary constraint on egg size in a butterfly

Genetic and developmental constraints have often been invoked to explain patterns of existing morphologies. Yet, empirical tests addressing this issue directly are still scarce. We here set out to investigate the importance of maternal body size as an evolutionary constraint on egg size in the tropical butterfly Bicyclus anynana, employing an artificial two-trait selection experiment on simultaneous changes in body and egg size (synergistic and antagonistic selection). Selection on maternal body size and egg size was successful in both the synergistic and the antagonistic selection direction. Yet, responses to selection and realized heritabilities varied across selection regimes: the most extreme values for pupal mass were found in the synergistic selection directions, whereas in the antagonistic selection direction realized heritabilities were low and nonsignificant in three of four cases. In contrast, for egg size the highest values were obtained in the lines selected for low pupal mass. Thus, selection on body size yielded a stronger correlated response in egg size than vice versa, which is likely to bias (i.e., constrain), if weakly, evolutionary change in body size. However, it seems questionable whether this will prevent evolution toward novel phenotypes, given enough time and that natural selection is strong. Correlated responses to selection were overall weak. Egg and larval development times tended to be associated with changes in maternal size, whereas variation in pupal development times weakly tended to follow variation in egg size. Lifetime fecundity was similar across selection regimes, except for females simultaneously selected for large body mass and small egg size, exhibiting increased fecundity. Multiple regressions showed that lifetime fecundity and concomitantly reproductive investment were primarily determined by longevity, as expected for an income breeder, whereas egg size was primarily determined by pupal mass. Evidence for a phenotypic trade-off between egg size and number was weak.     

Sexual and seasonal variation in the diet and foraging behaviour of a sexually dimorphic carnivore, the honey badger (Mellivora capensis)

The honey badger, or ratel, Mellivora capensis has not been well studied despite its extensive distribution. As part of the first detailed study, visual observations of nine habituated free-living individuals (five females, four males) were used to investigate seasonal, annual and sexual differences in diet and foraging behaviour. Theory predicts that generalist predators 'switch' between alternative prey species depending on which prey species are currently most abundant, and diet breadth expands in response to decreased availability of preferred food types. There were significant seasonal differences in the consumption of eight prey categories related to changes in prey availability but no seasonal differences in food intake per kg of body mass. As predicted, the cold-dry season diet was characterized by low species richness and low foraging yield but high dietary diversity, while the reverse was true in the hot-dry and hot-wet seasons. In accordance with these predictions, results suggest that the honey badger maintains its intake level by food switching and by varying dietary breadth. Despite marked sexual size dimorphism, male and female honey badgers showed no intersexual differences in prey size, digging success, daily food intake per unit body weight or foraging behaviour. Results do not support the hypothesis that size dimorphism is primarily an adaptation to reduce intersexual competition for food.     

GEOGRAPHIC VARIATION IN LIFE-HISTORY TRAITS OF THE ANT LION,MYRMELEON IMMACULATUS: EVOLUTIONARY IMPLICATIONS OF BERGMANN'S RULE

In eastern North America, body size of the larval ant lion Myrmeleon immaculatus increases from south to north, following Bergmann's rule. We used a common-garden experiment and a reciprocal-transplant experiment to evaluate the effects of food and temperature on ant lion growth, body size, and survivorship. In the laboratory common-garden experiment, first-instar larvae from two southern (Georgia, South Carolina) and two northern (Connecticut, Rhode Island) populations were reared in incubators under high- and low-food and high- and low-temperature regimes. For all populations, high food increased final body mass and growth rate and decreased development time. Growth rates were higher at low temperatures, but temperature did not affect larval or adult body mass. Survivorship was highest in high-food and low-temperature treatments. Across all food and temperature treatments, northern populations exhibited a larger final body mass, shorter development time, faster growth rate, and greater survivorship than did southern populations. Results were similar for a field reciprocal-transplant experiment of third-instar larvae between populations in Connecticut and Oklahoma: Connecticut larvae grew faster than Oklahoma larvae, regardless of transplant site. Conversely, larvae transplanted to Oklahoma grew faster than larvae transplanted to Connecticut, regardless of population source. These results suggest that variation in food availability, not temperature, may account for differences in growth and body size of northern and southern ant lions. Although northern larvae grew faster and reached a larger body size in both experiments, northern environments should suppress growth because of reduced food availability and a limited growing season. This study provides the first example of countergradient selection causing Bergmann's rule in an ectotherm.     

Life history variation in a dobsonfly, Protohermes grandis (Megaloptera: Corydalidae): effects of prey availability and temperature

SUMMARY. 1. The life cycle of a predatory insect, Protohermes grandis (Megaloptera: Corydalidae), was compared in four streams in central Japan. The effects of annual temperature regime and prey availability on life history characteristics were also assessed.
2. The larval period was 2 years and small adults emerged in the Morito River, where summer water temperature was high and large prey scarce.
3. In the Nagura River, rich in large prey, the larval period was also 2 years in spite of slightly lower temperature, and the adult size was largest among the streams.
4. In Anado Fork with a low summer temperature, the larval development took 3 years, and large adults emerged. Large prey were abundant in this stream.
5. Seasonal abundance of large prey also affected the time large larvae left the stream to pupate. Larvae emigrated earlier in streams where the density of large prey sharply decreased after spring, than in streams where large prey were available throughout the year.     

Effects of spatio-temporal variation in food supply on red squirrel Sciurus vulgaris body size and body mass and its consequences for some fitness components

Food availability is likely to influence body condition and, in turn, fitness. The intensity of this response may vary between populations of the same species on a small spatial and temporal scale. We used 5 yr of data from 6 Eurasian red squirrel Sciurus vulgaris populations from the southern Alps to explore differences in body size and body mass among neighbouring populations, in relation to habitat type and variation in food supply. We also investigated sexual dimorphism in these traits and whether phenotypic variation affects local survival and female reproductive success. Mean hind foot length, a measure of body size, did not differ between sexes but differed between areas. Seasonal variation in body mass was small with no evidence for fattening in autumn. Females were slightly heavier than males, but this difference was largely explained by mass gain of females during reproduction. The size of conifer seed crops, the major food supply, varied strongly over years and between habitats, but this variation corresponded only weakly with autumn body mass. Differences in size and mass between populations were partially explained by habitat‐related differences in body size and variability of seed‐crops, suggesting differential selection for smaller squirrels in spruce‐larch forests against selection for larger and heavier animals in mixed broadleaves and conifer forests and in Scots pine forests with more stable seed production. The probability of reproduction by females increased with body mass, but varied strongly between habitats and years, with more females reproducing in years with rich seed‐crops. In both sexes, body mass positively affected probability of settlement and length of residency. Our results suggest that in temporally variable environments that differ in overall amount of food resources, individual variation in body mass is related to habitat type, and that having a relatively high body mass, within each population, positively affects male and female settlement success and local survival, and female reproductive success.     

Insects from the grazing food web favoured the evolutionary habitat shift to bright environments in araneoid spiders

The Araneoidea comprises a diverse group of web-building spiders, and part of this diversity is believed attributable to habitat expansion to bright environments. We clarified the fitness-related advantages of living in such environments by examining prey availability and the growth rates of 10 species in three families inhabiting grassland (bright) and forest understory (dim) habitats. Spiders in the grassland habitat captured more prey, derived mainly from the grazing food web, than those in the forest-floor environment, and this difference was manifested in their growth rate. Independent contrasts indicated that increased utilization of insects from the grazing food web led to an evolutionary increase in adult body size. These results suggest that the shift to bright environments enabled araneoid spiders to evolve diverse life-history traits, including rapid growth and large size, which were not possible in dim environments.     

Abundance–body mass relationships in size-structured food webs

In communities sharing a common energy source, the energetic equivalence hypothesis predicts that numerical abundance (N) scales with body mass (M) as M^?0.75. However, in size‐structured food webs all individuals do not share a common energy source, and the energy available (E) to larger individuals is constrained by inefficient energy transfer through the food chains that support them. This is expected to lead to steeper scalings of N with M. Here, we formalize and test an existing model for predicting abundance–body mass scaling, where the decline in E with M is calculated from the mean predator–prey body mass ratio (from size‐based nitrogen stable isotope analysis) and trophic transfer efficiency. We show that the steep predicted scalings of abundance and body mass (N scales as M^?1.2, B scales as M^?0.2) in a marine food web are consistent with empirical estimates and can be attributed to the small predator–prey body mass ratio (106 : 1). As a previous study has shown that environmental stability may favour low predator–prey mass ratios and long food chains, we predict that steeper abundance–body mass relationships will be found in more stable environments.     

Correlated responses to artificial body size selection in growth, development, phenotypic plasticity and juvenile viability in yellow dung flies

Most life history traits are positively influenced by body size, whereas disadvantages of large body size are poorly documented. To investigate presumed intrinsic costs of large size in the yellow dung fly (Scathophaga stercoraria; Diptera: Scathophagidae), we established two replicates each of three body size laboratory selection lines (small, control and large; selection on males only), and subjected flies of the resulting extended body size range to various abiotic stresses. Response to selection was symmetrical in the small and large lines (realized h(2) = 0.16-0.18). After 24 generations of selection body size had changed by roughly 10%. Female size showed a correlated response to selection on male size, whereas sexual size dimorphism did not change. Development time also showed a correlated response as, similar to food limited flies, small line flies emerged earlier at smaller body size. At the lowest larval food limit possible, flies of all lines emerged at the same small body size after roughly the same development time; so overall phenotypic plasticity in body size and development time strongly increased following selection. Juvenile mortality increased markedly when food was extremely limited, large line flies showing highest mortality. Winter frost disproportionately killed large (line) flies because of their longer development times. Mortality at high temperatures was high but size-selective effects were inconsistent. In all environments the larger males suffered more. Initial growth rate was higher for males and at unlimited food. Small line individuals of both sexes grew slowest at unlimited larval food but fastest at limited larval food, suggesting a physiological cost of fast growth. Overall, extension of the natural body size range by artificial selection revealed some otherwise cryptic intrinsic juvenile viability costs of large size, mediated by longer development or faster growth, but only in stressful environments.     

Differential physiological responses to prey availability by the great egret and white Ibis

In long-lived species, the balance between the benefits of reproduction and the costs from reduced survival or productivity is particularly challenging in dynamic environments like wetlands, where food levels vary greatly year to year. Some wetland species exhibit changes in reproductive strategies in response to food availability but whether physiological responses function in a similar manner is unclear. We compared the pre-breeding physiological responses (fecal corticosterone [FCORT], heat shock protein 60 [HSP60], and mass) of 2 species of wading birds with contrasting foraging strategies (great egret [Ardea alba], an exploiter, and white ibis [Eudocimus albus], a searcher) during years with contrasting levels of prey availability. Both species were in good physiological condition, with low levels of HSP60 and FCORT, during a year with high prey availability (2006). In a contrasting year with lesser prey availability (2007), HSP60 and FCORT concentrations indicated that ibis physiological condition was reduced, whereas egrets showed little change. Egrets and male ibis increased body mass, whereas female ibis decreased mass, in the year with low prey availability. Although poorly understood, we hypothesize that the differential response between female ibis and the others is associated with differential investment strategies based on long-term costs of reproduction. Model results identified prey availability and the 2-week water recession rate as the primary habitat variables that were associated with the physiological condition of white ibises, whereas great egret physiological condition was influenced mostly by 2-week water recession rate. Our results support the hypothesis that prey availability and hydrological factors play crucial roles in regulating populations of wading birds in the Florida Everglades. The results of this study show a more complete pathway by which hydrologic patterns affect wading birds, and it suggests that ibis are more sensitive to habitat conditions than are egrets. This information can be used to refine species models designed to evaluate water management scenarios and will improve our ability to manage and restore wetland ecosystems © 2012 The Wildlife Society.     

Physiological responses to variable environments: storage and respiration in starving rotifers

1. Zooplankton exist in environments where food availability varies greatly over time, and success depends in part on the ability to store resources when food is abundant and to conserve them when food is scarce. This paper reports on interspecific differences in the size of stored reserves, and in respiration rate during food deprivation, of four species of planktonic rotifers.
2. The size of reserves varied from 42 to 71% of initial (well-fed) body mass. Interspecific differences in reserve size explained some of the previously observed differences in starvation time.
3. The initial response of respiration rate to food deprivation was quite variable between species. Brachionus calyciflorus was the only species to conserve energy by decreasing respiration rate in response to food deprivation. In contrast, the respiration rate of starved Asplanchna priodonta increased, while that of A. silvestrii and Synchaeta pectinata did not change, during food deprivation.
4. Theory predicts that temporal variation in resource level may facilitate the coexistence of competing species. This theory depends upon trade-offs between traits that confer competitive success in different environments. Although rotifers show a trade-off between competitive ability and maximum population growth rate, we found no evidence for trade-offs between either of those two traits and the size of reserves.     

Evolution of prey holding behaviour and large male body size in Ninox owls (Strigidae)

The hawk owl genus Ninox is unique among raptorial birds in that it includes three species in which males are substantially larger than females. This is a reversal of the normal pattern observed in both diurnal and nocturnal raptorial birds in which females are larger. Interestingly, these three Ninox species also are both the largest of the 22 species in the genus and the only species that exhibit the striking behaviour of ‘prey holding’ in which large (> 600 g) mammalian or avian prey is captured at night and held with body parts intact, and draped below a roost for the entire day without being consumed. Because explanations of the evolution of large male size suggest that it results from competition among males, the adaptive significance of prey holding was studied in a wild population of powerful owl Ninox strenua. Prey holding is largely confined to breeding males and its occurrence varies significantly across the breeding cycle, being most frequent during incubation and brooding. The study did not clearly resolve whether prey holding is a form of food storage or territorial display; however, both functions can select for large male body size and therefore play a significant role in the evolution of nonreversed size dimorphism. Although female‐only incubation and brooding is typical of Ninox owls and other owl species, prey holding appears to occur only in the large Ninox species because of the unique combination of large body size, large prey size, separate sex roles, and obligate cavity nesting. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 284–292.     

Natural selection on body size is mediated by multiple interacting factors: a comparison of beetle populations varying naturally and experimentally in body size

Body size varies considerably among species and among populations within species, exhibiting many repeatable patterns. However, which sources of selection generate geographic patterns, and which components of fitness mediate evolution of body size, are not well understood. For many animals, resource quality and intraspecific competition may mediate selection on body size producing large-scale geographic patterns. In two sequential experiments, we examine how variation in larval competition and resource quality (seed size) affects the fitness consequences of variation in body size in a scramble-competing seed-feeding beetle, Stator limbatus. Specifically, we compared fitness components among three natural populations of S. limbatus that vary in body size, and then among three lineages of beetles derived from a single base population artificially selected to vary in size, all reared on three sizes of seeds at variable larval density. The effects of larval competition and seed size on larval survival and development time were similar for larger versus smaller beetles. However, larger-bodied beetles suffered a greater reduction in adult body mass with decreasing seed size and increasing larval density; the relative advantage of being large decreased with decreasing seed size and increasing larval density. There were highly significant interactions between the effects of seed size and larval density on body size, and a significant three-way interaction (population-by-density-by-seed size), indicating that environmental effects on the fitness consequences of being large are nonadditive. Our study demonstrates how multiple ecological variables (resource availability and resource competition) interact to affect organismal fitness components, and that such interactions can mediate natural selection on body size. Studying individual factors influencing selection on body size may lead to misleading results given the potential for nonlinear interactions among selective agents.     

Behavioural and life history effects of predator diet cues during ontogeny in damselfly larvae

A central issue in predator–prey interactions is how predator associated chemical cues affect the behaviour and life history of prey. In this study, we investigated how growth and behaviour during ontogeny of a damselfly larva (Coenagrion hastulatum) in high and low food environments was affected by the diet of a predator (Aeshna juncea). We reared larvae in three different predator treatments; no predator, predator feeding on conspecifics and predator feeding on heterospecifics. We found that, independent of food availability, larvae displayed the strongest anti-predator behaviours where predators consumed prey conspecifics. Interestingly, the effect of predator diet on prey activity was only present early in ontogeny, whereas late in ontogeny no difference in prey activity between treatments could be found. In contrast, the significant effect of predator diet on prey spatial distribution was unaffected by time. Larval size was affected by both food availability and predator diet. Larvae reared in the high food treatment grew larger than larvae in the low food treatment. Mean larval size was smallest in the treatment where predators consumed prey conspecifics, intermediate where predators consumed heterospecifics and largest in the treatment without predators. The difference in mean larval size between treatments is probably an effect of reduced larval feeding, due to behavioural responses to chemical cues associated with predator diet. Our study suggests that anti-predator responses can be specific for certain stages in ontogeny. This finding shows the importance of considering where in its ontogeny a study organism is before results are interpreted and generalisations are made. Furthermore, this finding accentuates the importance of long-term studies and may have implications for how results generated by short-term studies can be used.     

Prey availability affects territory size,but not territorial display behavior,in green anole lizards

The availability of food resources can affect the size and shape of territories, as well as the behaviors used to defend territories, in a variety of animal taxa. However, individuals within a population may respond differently to variation in food availability if the benefits of territoriality vary among those individuals. For example, benefits to territoriality may differ for animals of differing sizes, because larger individuals may require greater territory size to acquire required resources, or territorial behavior may differ between the sexes if males and females defend different resources in their territories. In this study, we tested whether arthropod abundance and biomass were associated with natural variation in territory size and defense in insectivorous green anole lizards, Anolis carolinensis. Our results showed that both male and female lizards had smaller territories in a habitat with greater prey biomass than lizards in habitats with less available prey, but the rates of aggressive behaviors used to defend territories did not differ among these habitats. Further, we did not find a relationship between body size and territory size, and the sexes did not differ in their relationships between food availability and territory size or behavioral defense. Together, these results suggest that differences in food availability influenced male and female territorial strategies similarly, and that territory size may be more strongly associated with variation in food resources than social display behavior. Thus, anole investment in the behavioral defense of a territory may not vary with territory quality.     

Mass gained during breeding positively correlates with adult survival because both reflect life history adaptation to seasonal food availability

Both mass (as a measure of body reserves) during breeding and adult survival should reflect variation in food availability. Those species that are adapted to less seasonally variable foraging niches and so where competition dominates during breeding, will tend to have a higher mass increase via an interrupted foraging response, because their foraging demands increase and so become more unpredictable. They will then produce few offspring per breeding attempt, but trade this off with higher adult survival. In contrast, those species that occupy a more seasonal niche will not gain mass because foraging remains predictable, as resources become superabundant during breeding. They can also produce more offspring per breeding attempt, but with a trade-off with reduced adult survival. We tested whether the then predicted positive correlation between levels of mass gained during seasonal breeding and adult survival was present across 40 species of tropical bird measured over a 10-year period in a West African savannah. We showed that species with a greater seasonal mass increase had higher adult survival, controlling for annual mass variation (i.e. annual variation in absolute food availability) and variation in the timing of peak mass (i.e. annual predictability of food availability), clutch size, body size, migratory status and phylogeny. Our results support the hypothesis that the degree of seasonal mass variation in birds is probably an indication of life history adaptation: across tropical bird species it may therefore be possible to use mass gain during breeding as an index of adult survival.     

Prey size spectra and prey availability of larval and small juvenile cod

The aim of the present study is to describe the prey preference characteristics of cod larvae and assess preference variability in relation to species and size composition of copepod prey. A further aim is to examine the hypothesis that dietary prey size spectra remain the same during the larval stage when viewed on a relative predator/prey size scale. The study is based on stomach analysis of larval/juvenile cod in the size range 10–35 mm from nursery grounds in the North Sea. Stomach contents (species, size) were compared to environmental composition and preference indices were calculated. Prey size spectra had the expected relationship to larval cod size, and preference for given copepod species could be ascribed to their relative size. Additional species-specific preferences were evident, for example the larger Pseudocalanus and the larger Calanus spp. were highly preferred. Available prey biomass was highest in the areas of a hydrographic front, where larvae have been shown to concentrate. Changes in prey availability with larval growth depend strongly on specific prey biomass spectra at a given location. Both increasing and decreasing prey availability at increasing larval size were indicated, dependent on location. The findings illustrate the usefulness of coupling dietary prey size spectra and biomass spectra of available prey sizes during studies of ichthyoplankton feeding ecology.     

Smaller predator-prey body size ratios in longer food chains

Maximum food-chain length has been correlated with resource availability, ecosystem size, environmental stability and colonization history. Some of these correlations may result from environmental effects on predator-prey body size ratios. We investigate relationships between maximum food-chain length, predator-prey mass ratios, primary production and environmental stability in marine food webs with a natural history of community assembly. Our analyses provide empirical evidence that smaller mean predator-prey body size ratios are characteristic of more stable environments and that food chains are longer when mean predator-prey body size ratios are small. We conclude that environmental effects on predator-prey body size ratios contribute to observed differences in maximum food-chain length.