Effects of food supplementation on the physiological ecology of female Western diamond-backed rattlesnakes (<Emphasis Type="Italic">Crotalus atrox</Emphasis>)

Food availability is an important factor in the life histories of organisms because it is often limiting and thus can affect growth, mass change, reproduction, and behaviors such as thermoregulation, locomotion, and mating. Experimental studies in natural settings allow researchers to examine the effects of food on these parameters while animals are free to behave naturally. The wide variation among organisms in energy demands and among environmental food resources suggest that responses to changes in food availability may vary among organisms. Since most supplemental feeding field experiments have been conducted on species with high energy demands, we conducted a supplemental feeding study on free-ranging, female Western diamond-backed rattlesnakes (Crotalus atrox), a species with low energy demands and infrequent reproductive investment. Snakes were offered thawed rodents 1–4 times per week. Over two active seasons, we collected data on surface activity, home range size, growth, mass change, and reproduction of supplementally fed and control snakes. Fed and control snakes did not differ in surface activity levels (proportion of time encountered above versus below ground) or home range size. Fed snakes grew and gained mass faster, and had a dramatically higher occurrence of reproduction than control snakes. Also, fed snakes were in better body condition following reproduction than snakes that were not fed. However, litter characteristics such as offspring number and size were not increased by feeding, suggesting that these characteristics may be fixed. These data experimentally demonstrate that food availability can directly impact some life history traits (i.e., growth and reproduction for C. atrox), but not others (i.e., surface activity and home range size for C. atrox). The relationship between food availability and life history traits is affected in a complex way by ecological traits and physiological constraints, and thus interspecific variation in this relationship is likely to be high.     

Analysis of variance of first-year growth in captive garter snakes (Thamnophis elegans) by family and sex

Although body size in reptiles has important consequences for their ecology, it is quite variable in most species, presumably because growth in these animals is influenced by many factors. In this study, we analyse the effects of family membership and sex on first-year growth of garter snakes (Thamnophis elegans) held under identical conditions in the laboratory. Litters, but not the two sexes, differed significantly in mean size at birth. Larger snakes ate more food and had a higher absolute (but not relative) growth rate in length than smaller snakes. Thus, there was significant variation in length increase among litters, but not between the sexes. All else being equal, variance in size at birth could contribute significantly to variance in adult body size.     

Which proximate factor determines sexual size dimorphism in tiger snakes?

Diverse interactions between factors that influence body size complicate the identification of the primary determinants of sexual size dimorphism. Using data from a long‐term field study (1997–2009), we examined the contributions of the main proximate factors potentially influencing sexual size dimorphism from birth to adulthood in tiger snakes (Notechis scutatus). Data on body size, body mass and body condition of neonates, juveniles and adults were obtained by mark–recapture. Frequent recaptures allowed us to monitor reproductive status, diet and food intake, and to estimate survival and growth rates in age and sex classes. Additional data from females held briefly in captivity enabled us to assess reproductive output and the body mass lost at parturition (proxies for reproductive effort). From birth to maturity, individuals of both sexes experienced similar growth and mortality rates. We found no difference in diet, feeding and survival rates between the sexes, nor between juveniles and adults. On maturity, despite comparable diet and food intake by both sexes, the high energy requirements of vitellogenesis and gestation were responsible for a depletion of body reserves and probably resulted in a marked decrease in growth rates. Males were largely exempt from such costs of reproduction, and so could grow faster than females and attain larger body sizes. The absence of niche divergence between the sexes (uniformity of habitat, lack of predators) suggests that the impact of differential energetic investment for reproduction on growth rate is probably the main proximate factor influencing sexual size dimorphism in this species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 668–680.     

Vertebral number in adders, Vipera berus: direct and indirect effects on growth

Given the importance of body size, and thereby growth rate, for many reproductive parameters in snakes, morphological traits conferring an advantage in terms of growth may be important targets of selection. Studies have demonstrated effects of vertebral number of growth rate in garter snakes. In this study effects of total number of body vertebrae and of number of abnormal body vertebrae (obtained by counting number of ventral scutes and number of abnormal scutes) on growth rate in free-ranging male and female adders, Vipera berus (L.), are examined by calculating directional performance gradients (estimating linear effects) and stabilizing performance gradients (estimating curvilinear effects). After controlling for body size (SVL) female adders demonstrated a significant positive directional gradient for vertebral number, and a significant interaction between body size and vertebral number, showing that females with more vertebrae have higher size-specific growth rates, and that this effect is strongest among small, fast growing individuals. Females also showed a weak stabilizing effect of abnormal vertebrae. Males, on the other hand, showed a positive directional gradient for number of abnormal vertebrae, whereas no effect of vertebral number was observed. Indirect effects of the same variables were evaluated by use of path analysis. Generally, indirect effects were weak and did not substantially increase the amount of explained variance in growth rate. Field data showed that the correlation between vertebral number and growth rate in females was stronger in years with higher overall growth rate. To evaluate whether vertebral number and food availability show an interactive effect I used captive born juvenile adders in an experiment with two different food levels. The experiment confirmed the field data. No relationship between vertebral number and growth was observed in the low food level group, whereas in the high food level group a significant positive correlation was demonstrated. Finally, the heritability of vertebral number was examined using a mother-offspring regression and a full-sib analysis. The estimated heritabilities were 0.30 and 0.39, respectively. From these results it is concluded that both vertebral number and abnormal vertebral number may significantly affect growth in adders, but that this effect may differ between sexes and among years.     

Short and chubby or long and slim? Food intake,growth and body condition in free‐ranging pythons

Abstract An animal's sex and body size can influence not only its rate of food consumption, but also the way in which it allocates the resultant energy among the competing demands of maintenance, growth, reproduction and storage. A 13‐year mark–recapture study of pythons (Liasis fuscus) in tropical Australia provides extensive data on these topics. Rates of food intake and growth were highest in small pythons, and decreased more rapidly with body size in males than in females. Allocation to storage (as measured by the snake's mass relative to its body length) showed a more complex pattern. Body condition was high at hatching, but dropped rapidly as energy was allocated to growth rather than storage. Condition then increased through juvenile life, was at a maximum close to maturation, and was higher in females than in conspecific males. Body condition thereafter decreased with increasing body length. These allocation ‘decisions’ reflect the relative advantages of growth versus energy storage at different body sizes. Hatchling snakes grow rapidly (and hence become thin) because greater body size enables the snake to ingest larger prey items. Adult females amass larger energy reserves than males, because they need reserves to produce the clutch. Large snakes become thinner because their feeding rates are low, and they cannot compensate with increased prey size because large‐bodied mammalian prey are rare in our study area.     

Sexual size dimorphism and growth plasticity in snakes: an experiment on the Western Diamond-backed Rattlesnake (Crotalus atrox)

We conducted an experiment to examine the effects of sex and food intake on growth, mass gain, and attainment of sexual maturity in Western Diamond-backed Rattlesnakes (Crotalus atrox). We also measured testosterone levels to determine whether testosterone might be involved in the male-biased sexual size dimorphism observed in this species. We collected neonate rattlesnakes and raised them in the laboratory for 2 years on either a high-intake diet (fed one mouse per week) or a low-intake diet (fed one mouse every 3 weeks). High-intake snakes grew and gained mass more rapidly than low-intake snakes, but males did not grow or gain mass more rapidly than females in either treatment group. High-intake snakes attained reproductive maturity earlier than low-intake snakes, indicating that size, not age, is the critical determinant of reproductive maturity. Males had higher levels of testosterone than females but did not grow more quickly, suggesting that testosterone may not affect growth in this species and may therefore not be the proximate determinant of sexual size dimorphism.     

Effects of various environmental conditions on growth and reproduction of the sea hareAplysia oculifera (Adams and Reeve, 1850)

We studied the influences of food type, food quantity, water currents, starvation and light on growth and reproduction of the sea hareaplysia oculifera (Adams and Reeve, 1850) under laboratory conditions. Out of five species of algae served as food,Enteromorpha intestinalis promoted the fastest growth ofA. oculifera, Ulva spp. slower growth,Cladophora sp. allowed maintenance spp. slower growth,Cladophora sp. allowed maintenance of steady body mass, and the brown algaeColpomenia sp. andPadina pavonia were rejected by the sea hares. When sea hares were exposed to four levels of water currents, growth rates decreased as water currents increased. Sea hares fed on 50% ration grew slower than those fed on 100% ration (ad libitum). During 10 days of starvation sea hares lost weight, but when subsequently fed 100% ration they recovered and grew at a rate similar to those fed continuously with 100% ration. Under shade and under natural sunlight sea hares grew at the same rates. Whenever growth rates decreased, sea hares began to spawn at a smaller body size.A. oculifera demonstrated physiological plasticity that adapted them to varied and unpredictable environmental conditions. At different conditions of food availability they applied different tactics of resource allocation between growth and reproduction.     

Is fecundity in plaice (Pleuronectes platessa L.) down‐regulated in response to reduced food intake during autumn?

The effect of controlling food intake during the autumn, which is the time of late vitellogenesis, on fecundity, atresia and follicle and ovary growth was examined for plaice Pleuronectes platessa . Eighteen fish were kept in individual pens and either fed on a high or low ration diet. Fish which increased in whole body condition exhibited an increase in carcass condition which means that when food intake is sufficient to maintain whole body condition some resources are used as storage. Follicle growth rate was positively correlated with change in Fulton's condition and total atresia was negatively correlated with change in Fulton's condition. Thus, the rate of vitellogenesis was dependent on the availability of an exogenous food source. Fecundity at the end of the experiment was positively correlated with mass and total length. Food intake had no effect on relative fecundity; however, fish which had a lower food intake lost mass and had a greater intensity of atresia, lowering their absolute fecundity. One fish in a very low condition at the start of the experiment skipped spawning and one fish exhibited a decrease in average follicle diameter during the experiment which is hypothesized to be a prelude to mass atresia.     

Length-mass relationships of macro-invertebrates in a freshwater stream in Japan

Relationships between weight (W; dry weight) and length (L; head capsule width or total body length) were examined in 25 freshwater macro-invertebrates at species level and four at order level using the formula W = aL^b. Our estimates at the order level dropped into the range of those in North America, Europe, New Zealand, and previous studies in Japan with the exception of Trichoptera. Differences encountered in Trichoptera may be due to high interspecific variation in length-mass relationships and size-range of component taxa. In addition, our results show that the relationships could differ at the genus level. Intrageneric morphological variations related to microhabitat differences have been reported; careful estimates may thus be needed for the length-mass relationships even in the same genus if the targets have diverse ecological traits.     

Food consumption and growth of larvae and juveniles of three cyprinid species at different food levels

Synopsis The relationships between food availability, consumption and growth were analyzed from the onset of feeding to an age of 90 days in three cyprinid species. Fish were held at 20 ± 0.5° C and given two (three) constant rations of approximately 30, (40) or 100% dry body weight (dbw) ind^-1 day^-1. Food consisted of living zooplankton, the size of which correlated with fish size. At high food densities consumption rates decreased rapidly with fish size in all three species. At reduced rations, fish consumed most of the food offered until they were larger than 10 mg dbw. In all species and at each feeding level daily rations consumed increased allometrically with body size. Respiration rate, expressed as routine metabolic rate differed insignificantly between the three species. At high ration levels, growth rates of small bleak, Alburnus alburnus, were distinctly lower than those of roach, Rutilus rutilus, and blue bream, Abramis ballerus. At low food supply all three species grew at similar rates. Assimilation efficiency at low food conditions was approximately twice that of the well-fed groups. If the caloric equivalents of oxygen consumption as measured in well-fed fish are applied to fish fed at low rations their energy budgets do not balance. This indicates the limitations of fish larvae in the partitioning of energy for growth or activity at such conditions.     

Environmental and genetic variation in the haematocrit of fledgling pied flycatchers Ficedula hypoleuca

We exposed females of a highly placentotrophic viviparous scincid lizard (Pseudemoia pagenstecheri) to various environmental factors during pregnancy, and quantified the effects of these treatments on their offspring. The clear result was that the phenotypes of neonatal lizards can be substantially modified by the environment that their mother experiences during gestation. Restricting prey availability to the females reduced the size of their offspring. Limiting the females' basking opportunities delayed their seasonal timing of parturition, and modified body proportions (tail length relative to snout-vent length) of the neonates. More surprisingly, female lizards that were regularly exposed to the scent of sympatric lizard-eating snakes gave birth to offspring that were heavier, had unusually long tails relative to body length, and were highly sensitive to the odour of those snakes (as measured by tongue-flick responses). The neonates' antipredator responses were also modified by the experimental treatment to which their mother was exposed. The modifications in body mass, tail length and response to snake scent plausibly reduce the offspring's vulnerability to predatory snakes, and hence may constitute adaptive maternal manipulations of the neonatal phenotype. Received: 6 July 1998 / Accepted: 5 December 1998     

Resource limitation,predation risk and compensatory growth in a damselfly

Periods of poor nutrition during early development may have negative fitness consequences in subsequent periods of ontogeny. In insects, suppression of growth and developmental rate during the larval stage are likely to affect size and timing of maturity, which in turn may lead to reduced reproductive success or survivorship. In light of these costs, individuals may achieve compensatory growth via behavioural or physiological mechanisms following food limitation. In this study, we examined the effects of a temporary period of food restriction on subsequent growth and age and size at maturity in the larval damselfly Ischnura verticalis (Odonata: Coenagrionidae). We also asked whether this temporary period of reduced nutrition affected subsequent foraging behaviour under predation risk. I. verticalis larvae exposed to a temporary food shortage suffered from a reduced growth rate during this period relative to a control group that was fed ad libitum. However, increased growth rates later in development ensured that adult body size measurements (head and pronotum widths) did not differ between the treatments upon emergence. In contrast, adult dry mass did not catch up to that of the controls, indicating that the increased growth rates for size dimensions occur at the cost of similar gains in mass. Predators reduced foraging effort of larvae, but this reduction did not differ between control larvae and those previously exposed to poor nutrition.     

Male mating success in a North American pitviper: influence of body size,testosterone, and spatial metrics

Males with enhanced traits relative to conspecifics often show increased mating and reproductive success and thus have a fitness advantage. The opportunity or potential for sexual selection is predicted to occur under these conditions. Here, we investigated proximate determinants of mating success in male copperhead snakes (Agkistrodon contortrix), a medium‐sized pitviper of North America. Specifically, we investigated the relationships of body size (snout‐vent length, body mass), body condition index, spatial metrics (total distance moved, home range size), and plasma testosterone concentration on mating success in males. The single mating season lasts from August through September. We compared a set of candidate linear mixed models and selected the best‐fitting one using the adjusted Akaike Information Criterion (AICc). The AICc‐selected model (model 2), with testosterone, body condition index, and home range size as predictor variables, showed that male mating success was positively correlated with testosterone. To our knowledge, this is the first report to show the relationship of testosterone and individual mating success in any snake species. A parallel study conducted on male fitness in A. contortrix of the same population used microsatellite markers to assign parentage of fathers (known mothers). Unlike our study, they found that snout‐vent length was positively correlated with reproductive success and that males were experiencing greater sexual selection. This relationship has been detected under natural conditions in other species of snakes. Although behavioural data are important in any mating system analysis, they should not stand alone to infer parentage, relationships or selection metrics (e.g. Bateman gradients). Long‐term sperm storage by females, female cryptic choice, and other factors contribute to the complexity of mating success of males. Accordingly, we thus conclude that estimates of reproductive success and fitness in cryptic species, such as copperheads and other snakes, require robust molecular methods to draw accurate conclusions regarding proximate and evolutionary responses. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 185–194.     

Trends in body condition of native piscivores following invasion of Lakes Erie and Ontario by the round goby

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Food supply controls the body condition of a migrant bird wintering in the tropics

The hypothesis that migratory bird populations are limited during the non-breeding season is increasingly supported by empirical studies that also suggest consequences that carry-over into subsequent seasons. Although variation in food supply is the best supported explanation for non-breeding season limitation, the ecological mechanisms and physiological consequences are not well understood. We both supplemented and reduced Ovenbird (Seiurus aurocapilla) food availability on replicate plots in Jamaica in each of 3 years to determine the direct role of food in limiting the physical condition of Ovenbirds. Annual variation in rainfall and food supply created a natural experiment in parallel with manipulations. Sex and age-classes of Ovenbirds did not respond differently in terms of body condition to either food manipulation or natural variation in environmental conditions, suggesting that this population is not structured by strong dominance relationships. Ovenbird body mass, fat, and pectoralis muscle shape were positively and predictably related to manipulated food availability. Feather regrowth rate also responded positively to food supplementation and negatively to food reduction in the drier of 2 years. Prior to manipulation, annual variation in body mass corresponded to annual variation in food supply and rainfall, providing additional, correlational evidence of food limitation. Since multiple intercorrelated body condition indices of Ovenbirds responded directly to food supply, and since food supply influenced body condition independently of other habitat features, we argue that food is a primary driver of non-breeding season population limitation. Moreover, since these effects were observed during the late non-breeding period, when individuals are preparing to migrate, we infer that food availability likely initiates carry-over effects.     

The effects of prey species on food conversion efficiency and growth of an insectivorous lizard

Little is known about the effects of different prey species on lizard growth. We conducted a 6‐week study to determine the relative effects of prey species on growth parameters of hatchling western fence lizards, Sceloporus occidentalis. Lizards were fed house cricket nymphs, Acheta domesticus, or mealworm larvae, Tenebrio molitor. The effects of prey species on growth were determined by measuring prey consumption, gross conversion efficiency of food [gain in mass (g)/food consumed (g)], gain in mass, and gain in snout–vent length. Lizards grew well on both the prey species. However, lizards that fed on crickets consumed a significantly higher percentage of their body mass per day than those fed mealworms. Nevertheless, lizards that consumed mealworms ingested significantly more metabolizable energy, had significantly higher food conversion efficiencies, significantly higher daily gains in mass, and significantly greater total growth in mass than lizards that fed on crickets. Zoo Biol 27:181–187, 2008. © 2008 Wiley‐Liss, Inc.     

Validating growth and development of a seabird as an indicator of food availability: captive‐reared Caspian Tern chicks fed ad libitum and restricted diets

ABSTRACT For seabirds raising young under conditions of limited food availability, reducing chick provisioning and chick growth rates are the primary means available to avoid abandonment of a breeding effort. For most seabirds, however, baseline data characterizing chick growth and development under known feeding conditions are unavailable, so it is difficult to evaluate chick nutritional status as it relates to foraging conditions near breeding colonies. To address this need, we examined the growth and development of young Caspian Terns (Hydroprogne caspia), a cosmopolitan, generalist piscivore, reared in captivity and fed ad libitum and restricted (ca. one‐third lower caloric intake) diets. Ad libitum‐fed chicks grew at similar rates and achieved a similar size at fledging as previously documented for chicks in the wild and had energetic demands that closely matched allometric predictions. We identified three general characteristics of food‐restricted Caspian Tern chicks compared to ad libitum chicks: (1) lower age‐specific body mass, (2) lower age‐specific skeletal and feather size, such as wing chord length, and (3) heightened levels of corticosterone in blood, both for baseline levels and in response to acute stress. Effects of diet restriction on feather growth (10–11% slower growth in diet‐restricted chicks) were less pronounced than effects on structural growth (37–52% slower growth) and body mass (24% lower at fledging age), apparently due to preferential allocation of food resources to maintain plumage growth. Our results suggest that measurements of chick body mass and feather development (e.g., wing chord or primary length) or measurement of corticosterone levels in the blood would allow useful evaluation of the nutritional status of chicks reared in the wild and of food availability in the foraging range of adults. Such evaluations could also inform demography studies (e.g., predict future recruitment) and assist in evaluating designated piscivorous waterbird conservation (colony) sites.     

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.     

Effects of induced variation in anuran larval development on postmetamorphic energy reserves and locomotion

Anuran larvae exhibit high levels of phenotypic plasticity in growth and developmental rates in response to variation in temperature and food availability. We tested the hypothesis that alteration of developmental pathways during the aquatic larval stage should affect the postmetamorphic performance of the Iberian painted frog (Discoglossus galganoi). We exposed tadpoles to different temperatures and food types (animal- vs. plant-based diets) to induce variation in the length of the larval period and body size at metamorphosis. In this species, larval period varied with temperature but was unaffected by diet composition. In contrast, size at metamorphosis was shaped by the interaction between food quality and temperature; tadpoles fed on an animal-based diet became bulkier metamorphs than those fed on plant-based food at high (22°C) but not at low (12°C) temperature. Body condition of newly metamorphosed frogs was unrelated to the temperature or food type experienced during the premetamorphic stage. Frogs maintained at high temperature during the larval period showed reduced jumping ability, especially when fed on the plant-based diet. However, when considering size-independent jumping ability, cold-reared individuals exhibited the lowest performance, and herbivores reared at 17°C the highest. Cold-reared (12°C) frogs accumulated larger amounts of energy reserves than individuals raised at 17°C or 22°C. This was still the case after correction for differences in body mass, thus indicating some size-independent effect of developmental temperature. Despite the higher lipid content of the carnivorous diet, the differences in energy reserves between herbivores and carnivores were relatively weak and associated with differences in body size. These results suggest that the consequences of environmental variation in the larval habitat can extend to the terrestrial phase and influence juvenile growth and survival.     

Ambient temperature does not affect fuelling rate in absence of digestive constraints in long-distance migrant shorebird fuelling up in captivity

Pre-flight fuelling rates in free-living red knots Calidris canutus, a specialized long-distance migrating shorebird species, are positively correlated with latitude and negatively with temperature. The single published hypothesis to explain these relationships is the heat load hypothesis that states that in warm climates red knots may overheat during fuelling. To limit endogenous heat production (measurable as basal metabolic rate BMR), birds would minimize the growth of digestive organs at a time they need. This hypothesis makes the implicit assumption that BMR is mainly driven by digestive organ size variation during pre-flight fuelling. To test the validity of this assumption, we fed captive knots with trout pellet food, a diet previously shown to quickly lead to atrophied digestive organs, during a fuelling episode. Birds were exposed to two thermal treatments (6 and 24°C) previously shown to generate different fuelling rates in knots. We made two predictions. First, easily digested trout pellet food rather than hard-shelled prey removes the heat contribution of the gut and would therefore eliminate an ambient temperature effect on fuelling rate. Second, if digestive organs were the main contributors to variations in BMR but did not change in size during fuelling, we would expect no or little change in BMR in birds fed ad libitum with trout pellets. We show that cold-acclimated birds maintained higher body mass and food intake (8 and 51%) than warm-acclimated birds. Air temperature had no effect on fuelling rate, timing of fuelling, timing of peak body mass or BMR. During fuelling, average body mass increased by 32% while average BMR increased by 15% at peak of mass and 26% by the end of the experiment. Our results show that the small digestive organs characteristic of a trout pellet diet did not prevent BMR from increasing during premigratory fuelling. Our results are not consistent with the heat load hypothesis as currently formulated.