Non-reproductive Individuals Research Articles

Contrasts in body size and growth suggest that high population density results in faster pace of life in Damaraland mole-rats (Fukomys damarensis)

We studied the correlates of population density and body size, growth rates, litter size, and group size in Damaraland mole-rats (Fukomys damarensis (Ogilby, 1838)) at two study sites with contrasting population densities. Group size, litter size, and the probability of recapture were independent of study site. However, body size differed between the two study sites, suggesting that population density may affect life-history traits in social mole-rats. At the low-density site (0.13 groups/ha), individuals were significantly larger and subordinate males showed higher growth rates than at the high-density site (0.41 groups/ha), which may indicate that high population density in subterranean rodents enhances pace of life. The larger size of nonreproductive individuals at the low-density site could adapt individuals at lower population densities to larger dispersal distances.

Imitating the cost of males: A hypothesis for coexistence of all-female sperm-dependent species and their sexual host.

All‐female sperm‐dependent species are particular asexual organisms that must coexist with a closely related sexual host for reproduction. However, demographic advantages of asexual over sexual species that have to produce male individuals could lead both to extinction. The unresolved question of their coexistence still challenges and fascinates evolutionary biologists. As an alternative hypothesis, we propose those asexual organisms are afflicted by a demographic cost analogous to the production of males to prevent exclusion of the host. Previously proposed hypotheses stated that asexual individuals relied on a lower fecundity than sexual females to cope with demographic advantage. In contrast, we propose that both sexual and asexual species display the same number of offspring, but half of asexual individuals imitate the cost of sex by occupying ecological niches but producing no offspring. Simulations of population growth in closed systems under different demographic scenarios revealed that only the presence of nonreproductive individuals in asexual females can result in long‐term coexistence. This hypothesis is supported by the fact that half of the females in some sperm‐dependent organisms did not reproduce clonally.

Reproductive biology of Durvillaea antarctica (Chamisso) Hariot in the sub-Antartic ecoregion of Magallanes (51–56° S)

Durvillaea antarctica (Chamisso) Hariot is a large kelp commonly found on the exposed rocky shores of the Chilean coastline. It is endemic to the Southern Hemisphere and is used for direct consumption in the human diet and for alginate extraction; it is commonly known as “cochayuyo” (word from Chilean indigenous population “Mapudungun” language). The species occurs in the sub-Antarctic region and is found from Coquimbo (30° S) to Cape Horn (56° S). This study investigates and describes the seasonal reproductive strategies of D. antarctica located in the far south of the Chilean coast, in the region of the Magallanes (51–56° S), Chile, during the period of a year. Samples were taken in quadrants of 10 m2 parallel to the coastline in the intertidal zone of three locations during consecutive seasons. Both the phenological stages and the different states of maturity in individuals were determined. These findings were used to describe the main characteristics of the reproductive cycle and the spatial heterogeneity that may exist between the reproductive processes within each studied area. The results indicate that in all three studied localities of D. antarctica reproductive individuals are present throughout the year. The largest amount of reproductive individuals tends to exist in autumn and winter, with the numbers subsequently decreasing in spring and summer. In all studied localities, it was possible to identify male and female non-reproductive individuals. The largest number of male and female plants in the studied populations was found from autumn to winter; however, there were shifts in the proportions during these periods that lead to an increase in male over female fronds. In all localities, both male and female individuals in different stages of maturity were found, with the level of maturity depending on locality and season. Mature fronds were found more frequently during autumn and winter although the measures of maturity vary between different localities. Based on these results we conclude that D. antarctica is a species with present reproductive individuals throughout the year, similar to populations of this species described for other sub-Antarctic latitude. It is possible that there could be a cryptic species in the region, presenting morphological and reproductive variability that did not agree well with the circumscription of the species. Therefore, studies of ecophysiological and molecular aspects are needed to clarify the taxonomic status of this species in the sub-Antarctic region of Magallanes.

An Efficient Antioxidant System in a Long-Lived Termite Queen.

The trade-off between reproduction and longevity is known in wide variety of animals. Social insect queens are rare organisms that can achieve a long lifespan without sacrificing fecundity. The extended longevity of social insect queens, which contradicts the trade-off, has attracted much attention because it implies the existence of an extraordinary anti-aging mechanism. Here, we show that queens of the termite Reticulitermes speratus incur significantly lower oxidative damage to DNA, protein and lipid and have higher activity of antioxidant enzymes than non-reproductive individuals (workers and soldiers). The levels of 8-hydroxy-2’-deoxyguanosine (oxidative damage marker of DNA) were lower in queens than in workers after UV irradiation. Queens also showed lower levels of protein carbonyls and malondialdehyde (oxidative damage markers of protein and lipid, respectively). The antioxidant enzymes of insects are generally composed of catalase (CAT) and peroxiredoxin (Prx). Queens showed more than two times higher CAT activity and more than seven times higher expression levels of the CAT gene RsCAT1 than workers. The CAT activity of termite queens was also markedly higher in comparison with other solitary insects and the queens of eusocial Hymenoptera. In addition, queens showed higher expression levels of the Prx gene RsPRX6. These results suggested that this efficient antioxidant system can partly explain why termite queens achieve long life. This study provides important insights into the evolutionary linkage of reproductive division of labor and the development of queens’ oxidative stress resistance in social insects.

Oxidative stress in a capital breeder (Vipera aspis) facing pregnancy and water constraints.

The physiological mechanisms underlying the 'cost of reproduction' remain under debate, though oxidative stress has emerged as a potential candidate. The 'oxidative cost of reproduction' has received considerable attention with regards to food and antioxidant availability; however, the limitation of water availability has thus far been neglected. In this study, we experimentally examined the combined effect of pregnancy and water deprivation on oxidative status in a viviparous snake (Vipera aspis), a species naturally exposed to periods of water and food deprivation. We predicted a cumulative effect of pregnancy and dehydration on oxidative stress levels. Our results support the occurrence of an oxidative cost of reproduction as we found higher oxidative damage levels in pregnant females than in non-reproductive individuals, despite an up-regulation of antioxidant defences. Surprisingly, water deprivation was associated with an up-regulation of antioxidant defences, and did not increase oxidative damage, either alone or in combination with reproduction.

Differences in cooperative behavior among Damaraland mole rats are consequences of an age-related polyethism

In many cooperative breeders, the contributions of helpers to cooperative activities change with age, resulting in age-related polyethisms. In contrast, some studies of social mole rats (including naked mole rats, Heterocephalus glaber, and Damaraland mole rats, Fukomys damarensis) suggest that individual differences in cooperative behavior are the result of divergent developmental pathways, leading to discrete and permanent functional categories of helpers that resemble the caste systems found in eusocial insects. Here we show that, in Damaraland mole rats, individual contributions to cooperative behavior increase with age and are higher in fast-growing individuals. Individual contributions to different cooperative tasks are intercorrelated and repeatability of cooperative behavior is similar to that found in other cooperatively breeding vertebrates. Our data provide no evidence that nonreproductive individuals show divergent developmental pathways or specialize in particular tasks. Instead of representing a caste system, variation in the behavior of nonreproductive individuals in Damaraland mole rats closely resembles that found in other cooperatively breeding mammals and appears to be a consequence of age-related polyethism.

Nipped in the bud: mesograzer feeding preference contributes to kelp decline.

Small invertebrate grazers can disproportionately reduce plant fitness by discriminately consuming valuable tissues, but the context and attendant consequences of this activity at higher levels of ecological organization rarely are considered. To assess the impact of a gastropod mesograzer Lacuna vincta on fecundity and potential reproductive output of the habitat-forming kelp Saccharina latissima, we measured the intensity and distribution of grazing damage on kelp blades at five sites of varying kelp density, during the annual reproductive peak (October-November) in Nova Scotia. We found most grazing damage on reproductive individuals consisted of superficial excavations, and was concentrated on the central sorus (region where sporangia develop) compared to the vegetative blade margins. Grazing intensity on sori (percent grazed) averaged 29.6% across sites and sampling periods. The distribution of grazing on non-reproductive individuals was opposite to that of reproductive ones, indicating that snails shift feeding from blade margins to the center as sori develop. Choice and no-choice feeding assays in the laboratory revealed that focused grazing on sori is likely due to an active feeding preference for sporogenous over vegetative tissue. This preference was correlated with the distribution of chemical defense between tissues (phlorotannin content was ~2.5 times higher in vegetative tissue than sori), but not nutritional quality (no difference in C/N ratio). We deduce, with support from histological observations, that consumption of sorus tissue by L. vincta reduces fecundity of S. latissima. Extrapolating our results to estimate potential reproductive output within kelp beds suggests that spore supply and recruitment limitation may be predominantly imposed by the scarcity of reproductive individuals in the most degraded kelp beds. However, loss of reproductive output to grazing could extend recruitment limitations that impede recovery of waning kelp populations in Nova Scotia.

Considering transient population dynamics in the conservation of slow life-history species: An application to the sandhill crane

The importance of transient dynamics of structured populations is increasingly recognized in ecology, yet these implications are not largely considered in conservation practices. We investigate transient and long-term population dynamics to demonstrate the process and utility of incorporating transient dynamics into conservation research and to better understand the population management of slow life-history species; these species can be theoretically highly sensitive to short- and long-term transient effects. We are specifically interested in the effects of anthropogenic removal of individuals from populations, such as caused by harvest, poaching, translocation, or incidental take. We use the sandhill crane (Grus canadensis) as an exemplar species; it is long-lived, has low reproduction, late maturity, and multiple populations are subject to sport harvest. We found sandhill cranes to have extremely high potential, but low likelihood for transient dynamics, even when the population is being harvested. The typically low population growth rate of slow life-history species appears to buffer against many perturbations causing large transient effects. Transient dynamics will dominate population trajectories of these species when stage structures are highly biased towards the younger and non-reproducing individuals, a situation that may be rare in established populations of long-lived animals. However, short-term transient population growth can be highly sensitive to vital rates that are relatively insensitive under equilibrium, suggesting that stage structure should be known if perturbation analysis is used to identify effective conservation strategies. For populations of slow life-history species that are not prone to large perturbations to their most productive individuals, population growth may be approximated by equilibrium dynamics.

Can modern technology prevent extinction?

Conservation Biology![Figure][1] The northern white rhinoceros is a critically endangered species. PHOTO: © DANITA DELIMONT CREATIVE/ALAMY STOCK PHOTO Rising temperatures, habitat loss, and increasing urbanization mean that many animal species are on the brink of extinction. However, Saragusty et al. suggest that advances in stem cell and assisted reproductive technologies might allow scientists to save severely endangered species and to bring extinct species back to life. Examining the case of the northern white rhinoceros, an instance in which only three nonreproducing individuals are known to exist, the authors outline the potential and caveats of applying current methodological advances to save this iconic species. Furthermore, the guidelines laid out for the northern white rhino are likely to be applicable to many other critically endangered species. Zoo Biol. 10.1002/zoo.21284 (2016). [1]: pending:yes

Do female newts modify thermoregulatory behavior to manipulate egg size?

Reproductive females manipulate offspring phenotypes by modifying conditions during embryogenesis. In ectotherms, the environmental control over embryogenesis is often realized by changes in maternal thermoregulation during gravidity. To determine if reproduction influences thermoregulatory behavior in species where females lay eggs shortly after fertilization (strict oviparity), we compared preferred body temperatures (Tp) between reproductive (egg-laying) and non-reproductive female newts, Ichthyosaura alpestris. Next, we exposed reproductive females to temperatures mimicking Tp ranges of reproductive and non-reproductive individuals to find out whether the maternally modified thermal regime influences ovum and jelly coat volume, and early cleavage rates at the time of oviposition. In the thermal gradient, reproductive females maintained their body temperatures within a narrower range than non-reproductive individuals. The exposure of ovipositing females to temperatures preferred during their reproductive and non-reproductive period had a negligible influence on egg size and early cleavage rates. We conclude that the modification of maternal thermoregulatory behavior provides a limited opportunity to manipulate egg traits in newts.

Population structure of Sinelobus cf. stanfordi (Crustacea, Tanaidacea) from Pinheiros River mangrove, Guaratuba Bay, southern Brazil

An analysis of the population structure of the tanaid Sinelobus cf. stanfordi was performed at Pinheiros River mangrove, Guaratuba Bay, Paraná State, southern Brazil (25°49′S 48°34′W). Fifteen samples of 12 × 12 cm were obtained bimonthly from the surface of galleries (interconnected burrow complex) constructed by the adult mangrove crab Ucides cordatus from June 2007 to April 2008. The carapace length (CL) of a total of 714 tanaids was measured. Tanaids were divided into four demographic categories: males (M), ovigerous females (OV), preparatory females (PREP) and non-reproductive individuals (NREP) with a mean CL of 0.55, 0.49, 0.48 and 0.43 mm, respectively. In the NREP group, specimens smaller than 0.32 mm were classified as juveniles. NREP predominated in all sampling months, and among sexually mature individuals, a strong predominance of females was observed throughout the study period. Temporal fluctuations in abundance were observed, with a peak in August 2007. The reproductive period of this species is continuous, with a peak in the summer, like most S. cf. stanfordi populations.

Testing the genetic determination of the soldier caste in the silver ant

Division of labor among workers is a hallmark of social insects that has largely contributed to their ecological success. In a number of species, ants in particular, environmental cues have long been recognized to determine the different phenotypes of workers. Recently, however, a genetic basis for worker polymorphism has been documented in some species. The silver ant Cataglyphis bombycina is characterized by the co-existence of two physiologically distinct castes of non-reproductive individuals: workers and soldiers. Soldiers are not a worker subcaste; they belong to a third caste, along with the queen and the worker castes. Using microsatellite DNA markers, we tested whether soldier caste determination has a genetic component, by comparing the distribution of patrilines between the soldier and the worker castes. Our data show evidence of genotypic variation in caste propensity in only 2 out of 7 colonies sampled. In addition, most patrilines produce both workers and soldiers across all colonies. These results support moderate genotypic influence to soldier caste determination and suggest that non-genetic, likely environmental factors, also influence caste fate among non-reproductive offspring in this species. We also provide new estimates of the queen mating frequency, which support biogeographic variations in mating behavior in C. bombycina.

Reproduction, aging and mortality rate in social subterranean mole voles (Ellobius talpinus Pall.).

Eusocial subterranean rodents of the Bathyergidae family have enormous longevity. The long lifespan of these species is associated with negligible senescence, that is, an absence of the signs of age-related deterioration in physical condition. The question arises as to whether these features are unique to eusocial Bathyergids or typical of other social subterranean rodents as well. In the present study, we analysed data from observations of a social subterranean Microtinae rodent, the northern mole vole (Ellobius talpinus Pall.), which, like mole-rats, has reproductive skew. Among the individuals captured in the wild and maintained in captivity, females that reproduced lived significantly longer than non-breeding females. We did not find any changes in muscle strength with age in any of the demographic groups studied. Faecal glucocorticoid concentrations before death were significantly higher in non-breeding females than in breeding females and males. Increased adrenocortical activity may be one mechanism responsible for the decreased lifespan of non-reproducing individuals of social subterranean rodents. We conclude that the patterns of aging, although different in some respects, are generally common for social subterranean rodents of different taxonomic groups.

Fertility Signaling and Partitioning of Reproduction in the Ant Neoponera apicalis.

All individuals in social insect colonies benefit from being informed about the presence and fertility state of reproducers. This allows the established reproductive individuals to maintain their reproductive monopoly without the need for physical control, and the non-reproductive individuals to make appropriate reproductive choices. Here, we studied whether fertility signaling is responsible for the partitioning of reproduction in the ant Neoponera apicalis. This species forms small colonies from one single-mated queen, with workers establishing reproductive hierarchies when hopelessly queenless. Previous studies identified putative fertility signals, particularly the hydrocarbon 13-methylpentacosane (13-MeC25), and have shown that precise status discrimination based on these signals could be involved in the regulation of reproductive activities. Here, we extend these findings and reveal that all individuals, be they queens or workers, differ in their cuticular hydrocarbon profile according to fertility state. Proportions of 13-MeC25 were a strong predictor of an individual's ovarian activity, and could, thus, advertise the established reproducer(s) in both queenright and queenless conditions. Furthermore, this compound might play a key role in the establishment of the reproductive hierarchy, since workers with low fertility at the onset of hierarchy formation already have relatively high amounts of 13-MeC25. Dyadic encounters showed that individuals with experimentally increased amounts of 13-MeC25 triggered less agonistic interactions from top rankers, in accord with them "advertising" higher status. Thus, these bioassays supported the use of 13-MeC25 by competing ants. This simple recognition system potentially allows permanent regulation of partitioning of reproduction in this species.

Main biotic drivers of tree growth in a developing Juniperus thurifera stand in central Spain

Over the last few decades, abandonment of traditional management practices in Spain has led to widespread stand densification and has favoured the expansion of some forest species that previously exhibited more restricted ranges. Spanish juniper (Juniperus thurifera L.) woodlands are experiencing this phenomenon due to agricultural land abandonment and a decrease in the livestock pressure. Yet the main drivers underlying stand structure and dynamics at this novel scenario are poorly understood. In this study, we investigate the main biotic drivers of tree growth in a high-density stand of the dioecious J. thurifera at an early developmental stage (mean tree age of 32, ca. 50 years after land abandonment). Tree growth was measured by coring 299 individuals of different reproductive classes (male, female and non-reproductive). Neighbourhood models were used to assess the relative importance of tree size and neighbourhood competition on tree growth of each reproductive class in the study plot. We found that tree size had the strongest effect on tree growth, whereas the effect of intraspecific competition was negligible. We observed differences in growth patterns among reproductive classes along trunk diameter sizes. Thus, at smaller sizes, the three reproductive classes presented identical patterns of growth. However, at bigger sizes, females were the fastest growing individuals, followed by males and non-reproductive individuals. Overall, our results suggest that in young J. thurifera monospecific forests, where self-thinning processes may have not undergone yet, tree size and the reproductive class could play a relatively more important role than competition as drivers of tree growth. These findings constitute new information which contributes to understanding growth dynamics at early developmental stages in this dioecious species. Furthermore, our results provide guidelines for silvicultural managing, suggesting that at these young juniper stands thinning would likely not translate into enhanced growth on remnant trees.

Discriminating between the effects of founding events and reproductive mode on the genetic structure of Triops populations (Branchiopoda: Notostraca).

Crustaceans that initially colonize a freshwater temporary pond can strongly bias the subsequent genetic composition of the population, causing nearby populations to be genetically distinct. In addition, these crustaceans have various reproductive modes that can influence genetic differentiation and diversity within and between populations. We report on two species of tadpole shrimp, Triops newberryi and Triops longicaudatus “short”, with different reproductive modes. Reproduction in the tadpole shrimp can occur clonally (parthenogenesis), with self fertilization (hermaphroditism), or through outcrossing of hermaphrodites with males (androdioecy). For all these reproductive modes, population genetic theory predicts decreased genetic diversity and increased population differentiation. Here we use mitochondrial control region (mtCR) sequences and nuclear microsatellite loci to determine if the difference in reproductive mode affects the high genetic structure typical of persistent founder effects. Previous authors indicated that T. newberryi is androdioecious because populations are composed of hermaphrodites and males, and T. longicaudatus “short” is hermaphroditic or parthenogenetic because males are absent. In our data, T. newberryi and T. longicaudatus “short” populations were highly structured genetically over short geographic distances for mtCR sequences and microsatellite loci (T. newberryi: ΦST = 0.644, F ST = 0.252, respectively; T. l. “short”: invariant mtCR sequences, F ST = 0.600). Differences between the two Triops species in a number of diversity measures were generally consistent with expectations from population genetic theory regarding reproductive mode; however, three of four comparisons were not statistically significant. We conclude the high genetic differentiation between populations is likely due to founder effects and results suggest both species are composed of selfing hermaphrodites with some level of outcrossing; the presence of males in T. newberryi does not appreciably reduce inbreeding. We cannot exclude the possibility that males in T. newberryi are non-reproductive individuals and the two species have the same mating system.

Geographic variation in caste ratio of trematode colonies with a division of labour reflect local adaptation.

Similarly to the division of labour in social insects, castes of morphologically distinct individuals exist within colonies of some species of parasitic trematodes. These colonies occur in the first intermediate host of the trematode's complex life cycle and are composed of clonal individuals. Individuals of the reproductive caste have significantly larger bodies while non-reproductive individuals are small and appear to be specialised for defence against co-infecting trematode colonies. In parallel with colony organisation of social insects, demographic traits such as the proportion of the small, non-reproducing individuals relative to the large, reproducing individuals and colony size are expected to vary and adjust to local conditions. In the case of colonies from geographically and potentially genetically distinct populations, this variation is hypothesised to become fixed by evolutionary divergence, as reported in social insect studies. In this study, the adaptive demography theory was further tested by looking at caste ratio and colony organisation of Philophthalmus sp. (a parasitic trematode with a recently discovered division of labour) colonies from geographically distinct populations. Results indicate that the caste ratio from geographically distinct Philophthalmus sp. colonies differs; the proportion of small, defensive individuals is higher in colonies from the location where the risk of competition is highest. This is suggestive of local adaptation, as caste ratios do not change over time under standardised laboratory conditions. This is the first evidence to support the adaptive demography theory in a species with a division of labour other than social insects.

Male and female voles do not differ in their assessments of predation risk

A forager's willingness to trade off safety for food varies with its energetic state. Animals in a low energetic state should accept higher risk than animals with larger energy reserves. In mammals, energy expenditure by females on gestation and lactation may exceed the relatively low cost of sperm production by males. It follows, if reproductive costs are indeed higher for females than for males, that reproductive females may be more likely than males to trade safety for food. Thus, we evaluated the use of safe versus risky foraging patches by male and female meadow voles using putatively safe and risky habitats. We also used behavioural trials to assess whether sexual differences in personality could account for any differences in patch use. Voles preferred to forage in safe patches over risky ones. There was no difference between male and female voles, or between reproductive and non-reproductive individuals, in their respective use of safe versus risky foraging patches. Personality also had no effect on patch choice. The results are consistent with recent studies on other species that have failed to find differences in reproductive costs between the sexes. Experiments on foraging behaviour might thus provide simple and repeatable tests for sexual differences and similarities in reproductive costs.

Influence of light conditions on the allometry and growth of the understory palm Geonoma undata subsp. edulis (Arecaceae) of neotropical cloud forests

Knowledge on the growth responses of understory palms to changing light conditions within neotropical cloud forests is limited. The low light regime of these environments, in addition to persistent cloudiness, low ambient temperatures, and slow nutrient cycles, imposes significant constraints on biomass accumulation. Here, we evaluate how changes in the understory light conditions influenced the allometry and growth of G. undata subsp. edulis in two cloud forests in Costa Rica. We examined the structural relationships between stem diameter, stem height, and crown area in reproductive and nonreproductive individuals. We related the variation in stem growth and crown area with allometry, leaf production and longevity, and light conditions that we measured using hemispherical photographs over 1 year. The allometric and growth pattern of G. undata subsp. edulis was characterized by its investment in crown area, which was strongly and positively related to increments in palm height and reproduction. Growth, measured as the increase in crown area and stem height, was not explained by the variation in the light regime spanning 1 year. However, reproductive individuals were generally taller, more slender, and had larger leaf areas than nonreproductive individuals. Our results demonstrated that stem growth responses were mostly controlled by initial crown size rather than by temporal differences in the understory light regimes of cloud forests. These results suggest that cloud forest understory palms have a limited capacity to respond to light changes and rely mostly on an opportunistic strategy for biomass accumulation and reproduction.

Factors influencing stable nitrogen isotope ratios in wing membranes of insectivorous bat species: A field study

Nitrogen stable isotope analysis studies have become a useful tool in dietary studies to quantify differences in diet composition among species or individuals. However, laboratory studies have revealed several factors that influence the nitrogen isotope ratios (δ15N) in animal tissues (e.g. habitat, amount of ingested food, and metabolic rate). The influence of these factors has not been validated for free-ranging bat species, which is important for interpreting nitrogen isotope data in field studies. In this study, we measured δ15N from wing membranes (δ15Nw) and habitat samples (δ15Nh; soil and leaf, or sediment) to test for effects of diet, age, and individual energy demand in three free-living bat species (Myotis daubentonii, Nyctalus noctula and Nyctalus leisleri). We hypothesised that based on differences in diet composition δ15Nw values should increase in the sequence M. daubentonii, N. leisleri, and highest N. noctula. Furthermore, juveniles should have significantly lower δ15Nw compared to their mothers. Thirdly, the δ15Nw values of reproductively active individuals should be significantly higher than those of non-reproductive individuals. Values of δ15Nw were significantly influenced by differences in feeding ecology among species and individual reproductive status, but not by animal age. Our results show that species feeding on primary consumers have lower δ15Nw values than species feeding on secondary and tertiary consumers. Further, independent young did not differ in their δ15Nw values from adults suggesting that their tissues already reflect the insect diet. The effect of reproductive status varied among species which might reflect intra-specific differences in foraging behaviour of generalist vs. specialist species. In this study, we demonstrated that δ15N can provide information on the dietary range of bats, but researchers should account for variation owing to reproductive status, habitat, and species.