Maternal Reproductive Investment Research Articles

Interplay of cooperative breeding and predation risk on egg allocation and reproductive output.

Predation risk can influence behavior, reproductive investment, and, ultimately, individuals' fitness. In high-risk environments, females often reduce allocation to reproduction, which can affect offspring phenotype and breeding success. In cooperative breeders, helpers contribute to feed the offspring, and groups often live and forage together. Helpers can, therefore, improve reproductive success, but also influence breeders' condition, stress levels and predation risk. Yet, whether helper presence can buffer the effects of predation risk on maternal reproductive allocation remains unstudied. Here, we used the cooperatively breeding sociable weaver Philetairus socius to test the interactive effects of predation risk and breeding group size on maternal allocation to clutch size, egg mass, yolk mass, and yolk corticosterone. We increased perceived predation risk before egg laying using playbacks of the adults' main predator, gabar goshawk (Micronisus gabar). We also tested the interactive effects of group size and prenatal predator playbacks on offspring hatching and fledging probability. Predator-exposed females laid eggs with 4% lighter yolks, but predator-calls' exposure did not clearly affect clutch size, egg mass, or egg corticosterone levels. Playback-treatment effects on yolk mass were independent of group size, suggesting that helpers' presence did not mitigate predation risk effects on maternal allocation. Although predator-induced reductions in yolk mass may decrease nutrient availability to offspring, potentially affecting their survival, playback-treatment effects on hatching and fledging success were not evident. The interplay between helper presence and predator effects on maternal reproductive investment is still an overlooked area of life history and physiological evolutionary trade-offs that requires further studies.

Do non-direct heterospecific cues of avian predator activity alter reproductive modes of a passerine bird?

Predators affect prey by killing them or inducing changes in their physiology and behaviour through a fear effect associated with predation risk. In birds, perceived predation risk influences reproductive decisions, such as the reduction of parental investment in offspring during both egg production and nestling rearing. Visual and vocal cues of predator presence have been widely used to test the direct effects of predation risk. However, few studies have examined the indirect cues of predator activity such as dead avian prey or their remains. In this study, for the first time, we experimentally studied whether piles of feathers, simulating the remains of avian prey, induce changes in the reproductive decisions of adult birds. Before and during egg laying, great tit, Parus major, pairs were exposed to piles of bright down and cover feathers from domestic goose (treatment), woodchips (procedural control), or were not exposed (control). Our experiment affected maternal investment in individual eggs, but did not influence other reproductive parameters. Females from the treatment group laid larger and more asymmetrical (pointed) eggs than control females. Moreover, females from the procedural control group laid larger eggs than those from the control group, but without differences in egg shape. However, the eggs from the treatment and procedural control groups did not differ. This indicates that great tit females can perceive feathers and woodchips as informative cues, such as potential predation risk or habitat suitability, or as novel items in the environment. Importantly, females respond to such cues by changing their maternal investment in eggs, which may result from an adaptive mechanism aimed at increasing offspring fitness in the face of specific environmental conditions experienced by a female. Our study contributes to the understanding of how female songbirds adjust their maternal reproductive investment in response to publicly available social and environmental cues.

Experimental reduction of haemosporidian infection affects maternal reproductive investment, parental behaviour and offspring condition.

When hosts have a long coevolutionary history with their parasites, fitness costs of chronic infection have often been assumed to be negligible. Yet, experimental manipulation of infections sometimes reveals effects of parasites on their hosts, particularly during reproduction. Whether these effects translate into fitness costs remains unclear. Here, we present the results of an experimental study conducted in a free-ranging population of red-winged blackbirds (Agelaius phoeniceus) naturally experiencing a high prevalence of haemosporidian infections, with more than 95% of breeding adults infected with parasites from one or more haemosporidian genus. To assess effects of infection during reproduction, we manipulated adult red-winged blackbird females' parasite burden by administering an anti-haemosporidian medication before onset of egg-laying. Experimental reduction of infection resulted in significant benefits to mothers and their offspring. Medicated females laid heavier clutches, invested more in incubation and provisioning behaviour, and produced more fledglings than control females. Nestlings of medicated females had higher haematocrit, higher blood glucose, and lower reactive oxygen metabolites than nestlings of control females. Overall, our results provide evidence that, even in a species with high prevalence of infection, parasites can lead to decreased maternal investment and offspring quality, substantially reducing fitness.

Multiple aspects of the maternal reproductive investment in a polytocous species: What do mothers really control?

One of the factors facilitating the expansion and proliferation of wild boar Sus scrofa is the plasticity of its reproductive biology. Nevertheless, the real influence of maternal and environmental factors on number and sex of the offspring is still controversial. While the litter size was shown to be related with the maternal condition, the strength of this relation remains to be understood, together with the possible role played by environmental conditions. Analogously, it is unclear whether wild boar females can adjust their offspring sex. We investigated multiple aspects of wild boar maternal investment by means of a 10-year dataset of female reproductive traits and a set of biologically meaningful environmental variables. The maternal condition slightly affected the litter size but not the offspring sex, and environment did not affect the litter size or the offspring sex. Moreover, mothers did not cope with the higher costs entailed by producing sons by placing them in the most advantageous intrauterine position, nor by allocating less resources on daughters. Our set of results showed that the female reproductive investment is quite rigid in comparison with other aspects of wild boar reproductive biology. Wild boar females seem to adopt a typical r-strategy, producing constantly large litters and allocating resources on both sexes regardless of internal and external conditions. Such strategy may be adaptive to cope with environmental unpredictability and an intense human harvest, contributing to explain the extreme success of wild boar within human-dominated landscapes.

Cold shock induces a terminal investment reproductive response in C. elegans

Challenges from environmental stressors have a profound impact on many life-history traits of an organism, including reproductive strategy. Examples across multiple taxa have demonstrated that maternal reproductive investment resulting from stress can improve offspring survival; a form of matricidal provisioning when death appears imminent is known as terminal investment. Here we report a reproductive response in the nematode Caenorhabditis elegans upon exposure to acute cold shock at 2 °C, whereby vitellogenic lipid movement from the soma to the germline appears to be massively upregulated at the expense of parental survival. This response is dependent on functional TAX-2; TAX-4 cGMP-gated channels that are part of canonical thermosensory mechanisms in worms and can be prevented in the presence of activated SKN-1/Nrf2, the master stress regulator. Increased maternal provisioning promotes improved embryonic cold shock survival, which is notably suppressed in animals with impaired vitellogenesis. These findings suggest that cold shock in C. elegans triggers terminal investment to promote progeny fitness at the expense of parental survival and may serve as a tractable model for future studies of stress-induced progeny plasticity.

Effects of Maternal Reproductive Investment on Sex-Specific Pollutant Accumulation in Seabirds: A Meta-Analysis.

Avian egg production demands resources such as lipids and proteins. Relative egg size and mass varies across species, reflecting differences in maternal investment. This variability may affect the maternal transfer of anthropogenic pollutants including lipophilic polychlorinated biphenyls (PCBs) and protein-associated per- and polyfluoroalkyl substances (PFASs) and mercury (Hg). We conducted a meta-analysis on seabirds and investigated whether interspecies variation in maternal investment contributes toward skewed pollutant concentration ratios between males and females, as Cmale/Cfemale (80 studies). Overall concentrations of PCBs and perfluorooctanesulfonic acid (PFOS) were 1.6 and 1.3 times higher, respectively, in males than females, whereas mercury was similar between sexes. Few studies compared females and eggs ( n = 6), highlighting a knowledge gap. We found that an increasing maternal investment as a clutch-to-female mass ratio resulted in lower PCB concentrations in females than in males during the incubation period, but no sex-specific differences were observed for mercury and PFOS. Egg production is both a lipid dominated and protein-limited process. Females transfer lipophilic pollutants more easily to eggs, and to a higher degree with increasing maternal investment, but feeding ecology may be more important. Interspecies variation in maternal pollutant transfer may lead to negative effects scaling from an offspring to population level.

How many cubs can a mum nurse? Maternal age and size influence litter size in polar bears.

Life-history theory predicts that females' age and size affect the level of maternal investment in current reproduction, balanced against the future reproductive effort, maintenance and survival. Using long-term (30 years) individual data on 193 female polar bears ( Ursus maritimus), we assessed age- and size-specific variation on litter size. Litter size varied with maternal age, younger females had higher chances of losing a cub during their first months of life. Results suggest an improvement in reproductive abilities early in life due to experience with subsequent reproductive senescence. Litter size increased with maternal size, indicating that size may reflect individual quality. We also found an optimum in the probability of having twins, suggesting stabilizing selection on female body size. Heterogeneity was observed among the largest females, suggesting that large size comes at a cost.

Age-Specific Reproductive Investment and Offspring Performance in an Orb-web Spider, Argiope radon

Temporal variation in reproductive investment, e.g. maternal egg provisioning, has a substantial effect on offspring fitness therefore has received great attention by evolutionary biologists. Maternal allocation into egg size and egg content directly influences performance of offspring in many taxa, but spiders have rarely been investigated in this regard. In this study, we investigate the temporal changes in maternal reproductive investment and offspring performance in an orb-web, Argiope radon. A group of male and female spiders were mated randomly in the laboratory. Female spiders were kept under standard condition until they laid egg sacs. For each egg sac, egg sac mass, egg size and egg protein content were measured across all egg sacs. Once the spiderlings emerged, emerging time and toleration to starvation of the spiderlings were recorded. Egg sacs laid early in life were heavier, have larger eggs and offspring emerge sooner than the egg sacs laid by the old females. The spiderlings from the early egg sacs were more likely to emerge, however, these offspring had less toleration to starvation than the egg sacs laid by old females. There was no significant correlation between egg size and egg protein content with offspring performance. The results showed how the female spiders strategically allocate resources to egg sacs and its consequences on the offspring emerging time and toleration to starvation. This study suggests an adaptive maternal reproductive investment strategy which allows the female spiders to gain maximum fitness in each reproductive bout.

Sex-Specific Differences in Essential Lipid Requirements of Daphnia magna

Sex-specific differences in nutritional requirements may crucially influence the performances of the sexes, which may have implications for sexual reproduction and thus is of great ecological and evolutionary interest. In the freshwater model species Daphnia magna, essential lipid requirements have been extensively studied. Dietary deficiencies in sterols and polyunsaturated fatty acids (PUFA) have been shown to constrain somatic growth and parthenogenetic reproduction of female Daphnia. In contrast, nutrient requirements of male Daphnia have not been studied yet. Supplementation experiments were conducted to investigate differences in sterol (cholesterol) and PUFA (eicosapentaenoic acid, EPA) requirements between female and male D. magna. Thresholds for sterol-limited juvenile growth were higher in females than in males, suggesting that females are more susceptible to dietary sterol deficiencies than males. Sex-specific differences in maximum somatic growth rates were evident primarily in the presence of dietary EPA; females could not exploit their generally higher growth potential in the absence of dietary PUFA. However, the thresholds for EPA-limited growth did not differ between sexes, suggesting that both sexes have similar dietary EPA requirements during juvenile growth. During a life history experiment, the gain in body dry mass was higher in females than in males, irrespective of food treatment. In both sexes, the gain in body dry mass increased significantly upon EPA supplementation, indicating that both sexes benefited from dietary EPA supply also later in life. However, the positive effects of EPA supplementation were most pronounced for female reproduction-related traits (i.e. clutch sizes, egg dry masses, and total dry mass investment in reproduction). The high maternal investment in reproduction resulted in a depletion of nutrients in female somata. In contrast, the comparatively low paternal investment in reproduction allowed for the accumulation of nutrients in male somata. We conclude that males are generally less susceptible to dietary nutrient deficiencies than females, because they can rely more on internal body stores. Our data suggest that the performances of the sexes are differentially influenced by lipid-mediated food quality, which may have consequences for sexual reproduction and thus the production of resting eggs and the maintenance of Daphnia populations.

Maternal modulation of paternal effects on offspring development.

The paternal transmission of environmentally induced phenotypes across generations has been reported to occur following a number of qualitatively different exposures and appear to be driven, at least in part, by epigenetic factors that are inherited via the sperm. However, previous studies of paternal germline transmission have not addressed the role of mothers in the propagation of paternal effects to offspring. We hypothesized that paternal exposure to nutritional restriction would impact male mate quality and subsequent maternal reproductive investment with consequences for the transmission of paternal germline effects. In the current report, using embryo transfer in mice, we demonstrate that sperm factors in adult food restricted males can influence growth rate, hypothalamic gene expression and behaviour in female offspring. However, under natural mating conditions females mated with food restricted males show increased pre- and postnatal care, and phenotypic outcomes observed during embryo transfer conditions are absent or reversed. We demonstrate that these compensatory changes in maternal investment are associated with a reduced mate preference for food restricted males and elevated gene expression within the maternal hypothalamus. Therefore, paternal experience can influence offspring development via germline inheritance, but mothers can serve as a modulating factor in determining the impact of paternal influences on offspring development.

Dietary antioxidants in life-history trade-offs: differential effects of a-tocopherol supplementation on blue tit Cyanistes caeruleus mothers and offspring during reproduction

α-Tocopherol is assumed to be the most biologically active dietary antioxidant in vivo, but despite its potential importance little is known about its impacts on wild birds. Reproduction is presumed to be costly for parents through several routes, including increased oxidative stress, particularly for bird species producing large clutches. If dietary antioxidants can ameliorate oxidative stress associated with reproduction, mothers supplemented with dietary antioxidants are predicted to be in improved condition and/or invest more resources in reproduction than controls. We provided adult blue tit pairs with an α-tocopherol-enriched or control food supplement during nest building and egg laying, then cross-fostered half broods between treatment groups to test the theory that α-tocopherol-supplemented mothers would invest more in self-maintenance or reproduction than controls. We found that α-tocopherol supplementation had no effect on the maternal condition or reproductive investment. However, effects on nestlings were evident: nestlings from α-tocopherol-supplemented mothers were smaller at hatching. There was no effect on chick fledging mass, fledging success or lipid peroxidation, but the catch-up growth exhibited by chicks from α-tocopherol-supplemented parents may be considered costly. Thus, our results do not provide evidence for a benefit of maternal α-tocopherol supplementation at a biologically relevant dose on either themselves or their offspring. We discuss our findings in terms of ongoing research on the multifaceted roles that dietary ‘antioxidants’ can have in vivo, and the issues of disentangling their impacts on physiology and behaviour in the wild.

Nest-box temperature affects clutch size, incubation initiation, and nestling health in great tits

Prenatal maternal effects can be a source of phenotypic plasticity and may play a role in adaptation to climate change. However, we do not know how far temperature could influence such effects, if at all. We studied the influence of temperature during egg laying on maternal reproductive investment and on the phenotype of adult females, adult males, and nestlings. We expected temperature to have an effect, as it influences maintenance costs for females, who can also use it as a cue of the advancement of the breeding season. We experimentally increased night-time nest-box temperatures by approximately 1 °C throughout the entire laying period in great tits (Parus major). Clutch size was negatively correlated with laying date in heated females. Heated females did not delay incubation after clutch completion as frequently as control females did. Finally, blood sedimentation rate, which is an indicator of acute infections and inflammatory diseases, was positively correlated with hatching date in control broods. This suggests that nestlings were of lower quality in late-hatched broods than in early-hatched broods. This seasonal effect was not detected in heated nests. Our results show that a small increase in temperature during laying can influence breeding strategy and nestling characteristics. These results suggest that birds used temperature as a cue of seasonal advancement to adjust breeding phenology, with beneficial effects on nestling health. To better understand the consequences of maternal adjustments during egg laying, it would be interesting to combine studies with heating treatment during different periods of the breeding cycle.

Age‐related prenatal maternal effects and postnatal breeding experience have different influences on nestling development in an altricial passerine

Reproductive success and nestling performance are related to the age of parents across several vertebrate taxa. However, because breeding experience and prenatal maternal investment in reproduction often covary, the source of these age‐related differences can be difficult to determine. In this study, we evaluated the influence of prenatal maternal effects and postnatal breeding experience on the performance of nestling tree swallows Tachycineta bicolor by conducting a carefully controlled partial cross‐fostering experiment. We swapped half‐broods of nestlings between the nest of a young first‐time breeding female and the nest of a female known to have previously raised and fledged young. Our manipulation did not influence the within‐brood nestling hierarchies, and controlled for the effects of egg laying order. We found that nestlings of older females were heavier just prior to fledging regardless of the breeding experience of the attending female. In addition, fledglings raised by experienced females grew their flight feathers faster, and had greater probability of fledging. Our study demonstrates that prenatal investment in reproduction by older females can have long‐term consequences on nestling mass, and suggests limited potential for compensatory mass gains prior to fledging. Because our analyses controlled for feeding rates, our results also suggest that foraging quantity and quality are not the only benefits nestlings gain by being raised by an experienced female.

Effects of parasitism on host reproductive investment in a rodent-flea system: host litter size matters.

Parents may alter offspring phenotype depending on the type of environment they encounter. Parasitism is a common stressor; therefore, maternal reproductive investment could change in response to parasitic infection. However, few experiments have investigated the relationship between parasitism and maternal investment, whereas earlier field studies provided contradictory evidence. We investigated number, sex ratio, and growth of offspring in two rodent species, solitary altricial Meriones crassus and social precocial Acomys cahirinus, exposed to parasitism by fleas Xenopsylla ramesis and Parapulex chephrenis. No effect of treatment on litter size or sex ratio of a litter was found in either rodent species. Flea parasitism was found to affect pre-weaning body mass gain in M. crassus, but not in A. cahirinus pups. Furthermore, it appeared that female M. crassus invested resources into their offspring differently in dependence of litter size. In small litters (1-3 offspring), pups from infested females gained more body mass before weaning than pups from uninfested mothers. However, this trend was reversed in females with large litters indicating that parasitized females have a finite amount of resources with which to provision their young. Thus, M. crassus mothers parasitized by fleas seemed to receive some sort of external cues (e.g., stress caused by infestation) that prompted them to alter offspring provisioning, depending on species-specific possibilities and constraints. Therefore, parasites could be a mediator of environmentally induced maternal effects and offspring provisioning may have adaptive value against parasitism.

Heritable variation in maternally derived yolk androgens, thyroid hormones and immune factors.

Maternal reproductive investment can critically influence offspring phenotype, and thus these maternal effects are expected to be under strong natural selection. Knowledge on the extent of heritable variation in the physiological mechanisms underlying maternal effects is however limited. In birds, resource allocation to eggs is a key mechanism for mothers to affect their offspring and different components of the egg may or may not be independently adjusted. We studied the heritability of egg components and their genetic and phenotypic covariation in great tits (Parus major), using captive-bred full siblings of wild origin. Egg mass, testosterone (T) and androstenedione (A4) hormone concentrations showed moderate heritability, in agreement with earlier findings. Interestingly, yolk triiodothyronine hormone (T3), but not its precursor, thyroxine hormone (T4), concentration was heritable. An immune factor, albumen lysozyme, showed moderate heritability, but yolk immunoglobulins (IgY) did not. The genetic correlation estimates were moderate but statistically nonsignificant; a trend for a positive genetic correlation was found between A4 and egg mass, T and lysozyme and IgY and lysozyme, respectively. Interestingly, phenotypic correlations were found only between A4 and T, and T4 and T3, respectively. Given that these egg components are associated with fitness-related traits in the offspring (and mother), and that we show that some components are heritable, it opens the possibility that natural selection may shape the rate and direction of phenotypic change via egg composition.

Costly sons do not lead to adaptive sex ratio adjustment in pilot whales, Globicephala melas

Maternal investment in reproduction and parental care is an important determinant of both offspring and maternal fitness. However, optimal investment strategies may differ depending on offspring sex, potentially resulting in a sex-biased distribution of maternal resources or adaptive variation in offspring sex ratio. We used morphometric and genetic data collected from over 3400 long-finned pilot whales in 40 pods to investigate whether females experience differential costs of producing sons and daughters and whether they differentially invest in male and female offspring. We found that male calves grew faster than female calves during the first 5 years of life, suggesting that sons may require greater investment from lactating mothers. This appeared to result in mothers experiencing a higher cost to future reproductive opportunities when producing male offspring as the presence of dependent sons (but not daughters) reduced the probability that a female would be pregnant. Despite these costs, we found no evidence that mothers adaptively adjusted their investment in sons and daughters according to their body condition or their social and physical environment. These results suggest that mothers may be constrained from biasing investment in the sexes, or that additional benefits may be masking such costs.

The evolution of the patterning of human lactation: A comparative perspective

Some four decades ago, Jeanne Altmann started her detailed field studies of baboon mothers and their infants with a focus on the behavioral ecology of maternal reproductive investment. Around the same time, Sarah Hrdy studied langur mothers and their infants, focusing on the influence of the social environment on a female's reproductive options and decisions. Their pioneering work has inspired many subsequent studies of female primate reproduction in its natural context and shaped our own work on primate mothers.

No evidence for sex biases in milk macronutrients, energy, or breastfeeding frequency in a sample of filipino mothers

Maternal reproductive investment includes both the energetic costs of gestation and lactation. For most humans, the metabolic costs of lactation will exceed those of gestation. Mothers must balance reproductive investment in any single offspring against future reproductive potential. Among mammals broadly, mothers may differentially invest in offspring based on sex and maternal condition provided such differences investment influence future offspring reproductive success. For humans, there has been considerable debate if there are physiological differences in maternal investment by offspring sex. Two recent studies have suggested that milk composition differs by infant sex, with male infants receiving milk containing higher fat and energy; prior human studies have not reported sex-based differences in milk composition. This study investigates offspring sex-based differences in milk macronutrients, milk energy, and nursing frequency (per 24 h) in a sample of 103 Filipino mothers nursing infants less than 18 months of age. We found no differences in milk composition by infant sex. There were no significant differences in milk composition of mothers nursing first-born versus later-born sons or daughters or between high- and low-income mothers nursing daughters or sons. Nursing frequency also showed no significant differences by offspring sex, sex by birth order, or sex by maternal economic status. In the Cebu sample, there is no support for sex-based differences in reproductive investment during lactation as indexed by milk composition or nursing frequency. Further investigation in other populations is necessary to evaluate the potential for sex-based differences in milk composition among humans.

Experience affects mating behavior, but does not impact parental reproductive allocation in a lizard

Investment into reproduction is influenced by multiple factors and varies substantially between males and females. Theory predicts that males should adjust their ejaculate size or quality in response to variation in female experience or phenotypic quality. In addition, sperm investment by males may also be influenced by their own status and experience. Although such adjustments of male ejaculate size can impact reproductive success (via fertilization success), fitness returns from male sperm investment may be influenced (either limited or facilitated) by the level of maternal investment. To test this hypothesis, we conducted an experiment that simultaneously evaluated the effect of paternal and maternal experience (which incorporates mating status, age, body size, and other related variables) on paternal sperm investment and maternal reproductive allocation in the lizard Anolis sagrei. During staged mating trials, experienced males were more likely to copulate with females, but these individuals were less likely to transfer sperm during mating than were naive individuals. Maternal experience had no impact on these mating behaviors. In contrast to expectations, experience and phenotypic quality (of both sexes) had no impact on male ejaculate size or quality (proportion of live sperm) or on maternal reproductive investment (in terms of egg size and yolk steroids). These findings were intriguing given the mating system and past evidence for differential maternal investment in relation to sire quality in A. sagrei. The results found in this study highlight the complexity of reproductive investment patterns, and we urge caution when applying general conclusions across populations or taxa.

Bathymetric and interspecific variability in maternal reproductive investment and diet of eurybathic echinoderms

While conditions in shallow-water and deep-sea environments differ markedly, it remains unclear how eurybathic species adapt their life histories to cope with these changes. The present study compared indicators of maternal reproductive investment of three common echinoderms collected shallower than 20m and deeper than 850m: Cucumaria frondosa (Holothuroidea), Solaster endeca and Henricia sanguinolenta (Asteroidea). Depth-specific and species-specific differences were found in gonad indices (GI), potential fecundity, oocyte size frequency, as well as lipid classes and fatty acids measured in gonad tissue and oocytes. The asteroids, S. endeca and H. sanguinolenta, exemplified the interspecific trade-off between size and number of oocytes: the former had fewer larger oocytes than the latter, with higher total lipid proportions in them. However, intraspecifically, larger oocytes found in deep specimens of S. endeca did not translate into lower fecundity but rather into a seemingly higher GI, indicating greater investment per oocyte without reducing fecundity. Oocytes were absent in specimens of C. frondosa sampled in deep water, suggesting delayed or impaired maturation at the limit of their depth tolerance. Analysis of S. endeca sterol proportions in gonads and oocytes across depths showed higher sterol input into oocytes in females from the deep. Gonads of S. endeca and H. sanguinolenta contained similar essential fatty acids, but showed significant differences in major fatty acids and some of the less dominant ones, indicating diet specificities. Analyses within S. endeca showed evidence of similar feeding mode (carnivory) at both depths, but suggested shifts in the diet or synthesis of fatty acids, presumably reflecting differences in available food sources and/or adaptations to their respective environments.