Dietary overlap and seasonality in three species of mormoopid bats from a tropical dry forest

Competing hypotheses explaining species’ use of resources have been advanced. Resource limitations in habitat and/or food are factors that affect assemblages of species. These limitations could drive the evolution of morphological and/or behavioural specialization, permitting the coexistence of closely related species through resource partitioning and niche differentiation. Alternatively, when resources are unlimited, fluctuations in resources availability will cause concomitant shifts in resource use regardless of species identity. Here, we used next‐generation sequencing to test these hypotheses and characterize the diversity, overlap and seasonal variation in the diet of three species of insectivorous bats of the genus Pteronotus. We identified 465 prey (MOTUs) in the guano of 192 individuals. Lepidoptera and Diptera represented the most consumed insect orders. Diet of bats exhibited a moderate level of overlap, with the highest value between Pteronotus parnellii and Pteronotus personatus in the wet season. We found higher dietary overlap between species during the same seasons than within any single species across seasons. This suggests that diets of the three species are driven more by prey availability than by any particular predator‐specific characteristic. P. davyi and P. personatus increased their dietary breadth during the dry season, whereas P. parnellii diet was broader and had the highest effective number of prey species in all seasons. This supports the existence of dietary flexibility in generalist bats and dietary niche overlapping among groups of closely related species in highly seasonal ecosystems. Moreover, the abundance and availability of insect prey may drive the diet of insectivores.     

Pulsed resource availability changes dietary niche breadth and partitioning between generalist rodent consumers

Identifying the mechanisms that structure niche breadth and overlap between species is important for determining how species interact and assessing their functional role in an ecosystem. Without manipulative experiments, assessing the role of foraging ecology and interspecific competition in structuring diet is challenging. Systems with regular pulses of resources act as a natural experiment to investigate the factors that influence the dietary niches of consumers. We used natural pulses of mast‐fruiting of American beech (Fagus grandifolia) to test whether optimal foraging or competition structure the dietary niche breadth and overlap between two congener rodent species (Peromyscus leucopus and P. maniculatus), both of which are generalist consumers. We reconstructed diets seasonally over a 2‐year period using stable isotope analysis (δ¹³C, δ¹⁵N) of hair and of potential dietary items and measured niche dynamics using standard ellipse area calculated within a Bayesian framework. Changes in niche breadth were generally consistent with predictions of optimal foraging theory, with both species consuming more beechnuts (a high‐quality food resource) and having a narrower niche breadth during masting seasons compared to nonmasting seasons when dietary niches expanded and more fungi (a low‐quality food source) were consumed. In contrast, changes in dietary niche overlap were consistent with competition theory, with higher diet overlap during masting seasons than during nonmasting seasons. Overall, dietary niche dynamics were closely tied to beech masting, underscoring that food availability influences competition. Diet plasticity and niche partitioning between the two Peromyscus species may reflect differences in foraging strategies, thereby reducing competition when food availability is low. Such dietary shifts may have important implications for changes in ecosystem function, including the dispersal of fungal spores.     

Food niche overlap among neotropical frugivorous bats in Costa Rica

Food habits of 15 species of frugivorous bats were studied at La Selva Biological Station, Costa Rica. Eight hundred and fifty-four (854) fecal samples and 169 samples from fruit parts and seeds discarded by bats beneath feeding roosts were analyzed. During eight months of study, 47 fruit species consumed by bats were identified. Five plant genera (Cecropia, Ficus, Piper Solanum, and Vismia) constituted 85% of all plants found in fecal samples. Feeding niche breadth differed significantly among the six most common species of frugivorous bats (Artibeus jamaicensis, Carollia sowelli, C. castanea, C. perspicillata, Dermanura sp., and Glossophaga commissarisi). All species, except for Dermanura sp., showed a diet dominated by one or two plant species. This suggests a pattern of resource partitioning at a generic level, in which Carollia consumed mainly Piper, Artibeus consumed Ficus and Cecropia, and Glossophaga consumed Vismia. Cluster analysis revealed higher values of food niche overlap in congeneric species than among species of different genera. Results show that if food is a limiting factor, mechanisms other than trophic selection must reduce interspecific interference or competition for food in this frugivorous bat guild.     

Seasonal dynamics of resource partitioning among foliage-gleaning passerines in northern Finland

Summary The five most abundant species were included in a year-round study with respect to six foraging niche dimensions. Approximately full multidimensional utilization functions were used for niche metrics. During summer foraging overlaps were invariably high, but in other seasons periodically lower. Foraging site breadth was lower in winter, when fewer sites are profitable for foraging than in summer. Feeding posture versatility, by contrast, was highest in winter. Seasonal foraging shifts were very prominent, as great in fact as between-species differences. Often seasonal trends were parallel in different species. Niche axes of macrohabitat-type (e.g. tree) were more open for foraging variation and axes of microhabitat-type (tree part) more rigid. Among the resident species seasonal variation in foraging was greatest in Regulus regulus and Parus cristatus, whereas the foraging behaviour was more stable for P. montanus (an abundant species with broad niche), perhaps owing to greater intraspecific competition. In these northern forests foliage-gleaners must be versatile generalists to cope with the unpredictable resources, and thus they overlap broadly in their general resource niches which are determined by their genetically fixed cost and benefit relations to each resource type. Anyhow, presumably during periodical food shortage, the actual resource uses adjusted by resource availability and competition may overlap narrowly.     

The coexistence of fish species in streams: relationships between assemblage attributes and trophic and environmental variables

Diet overlap and niche breadth are well-known species traits from trophic ecology that can assist in explaining how species interact and coexist as well as the ecological mechanisms that influence biodiversity. In the present study, we analyzed the relationships between these trophic variables and indicators of resource availability with some attributes of fish assemblages (species richness, Shannon diversity index, evenness, density and individual body size). The physical and chemical characteristics of the biotopes (topography, water quality and conservation of slopes) were examined to identify possible patterns. Monthly sampling using electrofishing was conducted in 2003 along five streams located in the Cuiabá River watershed. The relationships between environmental variables and attributes of fish assemblages were evaluated using Spearman correlation. Species richness and abundance varied among streams, with higher values (54 and 82 species) found in low-gradient streams that drained small swampy areas discharging in Cuiabá River. Diet overlap showed significant and negative correlations with species richness, the diversity index, abundance, variation in body size and food availability and positive correlations with evenness and niche breadth. Environments that had greater food availability had a reduced variety of food items (smaller amplitude) and distinct values in terms of dietary overlap. Nevertheless, to explain resource partitioning in a fish assemblage (overlap), it is necessary to consider food availability and niche breadth (degree of trophic specialization) beyond the interaction of these variables with species richness and density. In conclusion, high diversity and abundance values were strongly associated with a high degree of trophic specialization (low amplitude of trophic niche) and a small degree of overlap in the diet.     

Seasonal variation in prey preference,diet partitioning and niche breadth in a rich large carnivore guild

Large carnivore community structure is affected by direct and indirect interactions between intra-guild members. Co-existence between different species within a carnivore guild may occur through diet, habitat or temporal partitioning. Since carnivore species are highly dependent on availability and accessibility of prey, diet partitioning is potentially one of the most important mechanisms in allowing carnivores to co-exist. Intra-guild interactions may vary over time as carnivore prey preference and diet overlap can change due to seasonal changes in resource availability. We conducted scat analysis to compare the seasonal changes in prey preference, diet partitioning and niche breadth of four large carnivore species, namely leopard Panthera pardus, spotted hyena Crocuta crocuta, brown hyena Parahyaena brunnea and wild dog Lycaon pictus in central Tuli, Botswana. Large carnivores in central Tuli display a high dietary overlap, with spotted hyena and brown hyena displaying almost complete dietary overlap and the other carnivore species displaying slightly lower but still significant dietary overlap. Dietary niche breadth for both hyena species was high possibly due to their flexible foraging strategies, including scavenging, while leopard and wild dog showed a relatively low niche breadth, suggesting a more specialised diet. High dietary overlap in central Tuli is possibly explained by the high abundance of prey species in the area thereby reducing competition pressure between carnivore species. Our research highlights the need to assess the influence of diet partitioning in structuring large carnivore communities across multiple study sites, by demonstrating that in prey rich environments, the need for diet partitioning by carnivores to avoid competition may be limited.     

Reduced-impact Logging has Little Effect on Temporal Activity of Frugivorous Bats (Chiroptera) in Lowland Amazonia

Reduced-impact logging (RIL) represents a viable option for sustainable use of Neotropical lowland forests while minimizing negative effects on local biodiversity. Many Neotropical bats of the family Phyllostomidae provide ecosystem services associated with pollination and seed dispersal that promote the regeneration of disturbed areas; therefore, effects of RIL on these species is of particular concern. We determined patterns of temporal activity, degree of temporal overlap of activity, and dispersion in peaks of activity for seven abundant species of frugivorous bat in Tapajós National Forest, Pará, Brazil. In addition, we evaluated the effects of RIL at a harvest level of 18.7 m³/ha and habitat physiognomy on temporal patterns of activity for these species. Bats were surveyed for four nights at each of 96 sites for a total sampling effort of 64,512 net-m-h. Sites were distributed among four experimental blocks, two blocks of unlogged forest and two blocks of forest subjected to RIL. Half of the sites in each management type were in forest gaps and half were in closed-canopy forest. In general, species exhibited similar patterns of activity, and greater than expected temporal overlap in activity among species. RIL and forest physiognomy had little effect on activity patterns of species. RIL in Amazonia removes fewer trees than do naturally occurring treefalls and such changes in habitat structure do not alter activity patterns of frugivorous bats. Evidence suggests that RIL does not have an appreciable adverse effect on frugivorous bats in Amazonia.     

Activity patterns of two syntopic and closely related aerial-hawking bat species during breeding season in Białowieża Primaeval Forest

Temporal and spatial activity of bats is species specific and shaped by many factors such as energy requirements, climate conditions and food distribution. Pregnancy and lactation are the most energy-demanding periods throughout the female life cycle. During these periods, females have to optimize their activity patterns to maximize foraging success; however, they simultaneously need to take care of their young. In addition, daily and seasonal fluctuations of insect availability strongly affect bat foraging activity. If females, which are under strong energy constraints, belong to closely related species, they may potentially suffer from competition. One of the mechanisms that allows them to avoid competition is temporal and spatial niche partitioning. Noctule and Leisler’s bats are closely related forest-dwelling species whose diet is similar and consists mainly of ephemeral insects. The aim of our study was to test if they exhibit similar patterns in relation to the time and duration of their nocturnal activity. In Bia?owie?a Forest, we demonstrated that female nocturnal activity of both noctule and Leisler’s bats was shaped mostly by reproductive period and ambient temperature. We did not observe significant differences in the activity patterns of the two noctule species, which suggests that physiological constraints connected with reproduction and environmental conditions affect these species in a similar way and outweigh the competition between species.     

Individual–resource networks reveal distinct fruit preferences of selective individuals from a generalist population of the Helmeted Manakin

Ecological studies traditionally assume that generalist populations are homogeneous in the use of food resources, but empirical evidence supports that intraspecific differences in morphology, physiology and behaviour affect foraging decisions and promote diet variation among individuals. Furthermore, the temporal availability of resources may shape the dynamics of population trophic niche, which ultimately depends on individual niches. In this study, we investigated the seasonal changes in individual-based networks between the Helmeted Manakin Antilophia galeata, a generalist frugivorous bird, and fruiting plants, following theoretical models of interindividual diet variation based on the Optimal Diet Theory. Selective individuals were the majority of the generalist population of the Helmeted Manakin. Our results suggest that the structure of the individual-resource networks varied seasonally. We found that modularity was higher than expected by chance in the wet season, when fruit availability was also higher. In the dry season, modules disappeared and the network became more nested. These findings are consistent with the Distinct Preference Model of diet variation. We suggest that downscaling ecological networks to the individual level may reveal emergent properties that, albeit existent, are not evident in species–resources networks.     

杉阔混交林主要种群多维生态位特征

在生态位空间分割基础上,应用两种不同类型的生态位宽度公式和生态位重叠公式测定了杉阔混交林不同资源空间中主要种群的多维生态位特征。结果表明：考虑资源利用率与否对生态位宽度的测度有较大影响。杉阔混交林主要种群在不同资源空间中的生态适应性不同。在同一资源空间中,种对间生态位宽度与生态位重叠并非成正相关关系。在不同资源空间中,杉木与同一其它树种之间的生态位重叠发生变化。生态位重叠与竞争之间关系在很大程度上受种对间的生物学特性的左右。研究杉阔混交林多维生态位特征对混交林营造具有指导作用。     

Niche dynamics of sympatric non-breeding shearwaters under varying prey availability

Variation in prey availability can cause changes in species interactions among marine predators. Foraging theory predicts that niche breadth will expand when resources become limited, possibly leading to higher niche overlap among sympatric species; however, a species’ niche can become constrained by interactions with other similar species, resulting in an inability to shift niche breadth or position. In coastal Newfoundland, Capelin Mallotus villosus is the main forage fish species and its availability (i.e. biomass) during the summer has varied considerably following a population collapse in the 1990s. During the summer, non-breeding Great and Sooty Shearwaters Ardenna gravis and A. grisea migrate and aggregate at annually persistent Capelin spawning sites. We used stable isotope ratios (δ¹³C, δ¹⁵N) of blood components (plasma, red blood cells) to investigate variation in isotopic niche breadth (95% prediction ellipse areas) and overlap of the two shearwater species during 2014, 2015 and 2016. Capelin availability varied among years, illustrated by lower peak biomass in 2015 (0.126 g/m²) and 2016 (0.027 g/m²) relative to 2014 (0.254 g/m²). The isotopic niche breadth (plasma) of both shearwater species expanded similarly from 2014 (0.65–0.66‰²) to 2015 (2.22–2.57‰²) and 2016 (1.15–1.42‰²), suggesting the incorporation of alternative prey types into their diet during years of lower prey availability. Isotopic niche overlap between Great and Sooty Shearwaters remained high across years (44–63%), however, providing little evidence for dietary niche partitioning during years of lower prey availability. Findings suggest that both shearwater species are flexible foragers and can modify their diet during the non-breeding season to accommodate fluctuations in prey availability.     

Bat frugivory in two subtropical rain forests of Northern Argentina: Testing hypotheses of fruit selection in the Neotropics

Phyllostomid bats are prominent components of mammalian assemblages in the Neotropics. With many species specialized in frugivory, phyllostomids represent major partners of fleshy-fruited plants in the mutualism of seed dispersal. Here we present dietary data from two subtropical rainforests of Argentina, where fruit diversity is low and thus offer unique opportunities to test hypotheses of diet selection originally proposed for species-rich tropical assemblages. Particularly, we tested whether frugivorous phyllostomids exhibit pronounced dietary specialization in core plant taxa where fruit offer is greatly reduced as compared to tropical rainforests. We analyzed dietary overlap and niche breadth of subtropical frugivorous bats on the basis of >1000 dietary records plus >500 samples from a previous study in the region. We show that in the subtropics, frugivores from different genera remain faithful to their respective core plant taxa with few exceptions, rather than shifting toward alternative fruit resources available in the study sites. This supports predictions of specialization, which is confirmed to have a deep historical origin. The response of phyllostomid ensembles to restricted fruit diversity is at the level of species composition: absence of species for which preferred fruits do not occur in the sites. Taken together, these data lend strong support to hypotheses that explain coexistence of frugivorous phyllostomids on the basis of dietary specialization on core plant taxa with chiropterochorous fruits.     

Dietary overlap of grasshoppers on sandhill rangeland in northeastern Colorado

Summary Overlap in the diets of 14 species of grasshoppers on mixed-grass prairie in northeastern Colorado was estimated by microscopic examination of crop contents. Food availability estimates, taken by a weight-estimate method, facilitated the determination of food preferences. Two cases of high overlap in diets and period of habitat occupancy occurred between species which consumed equal or almost equal numbers of different foods (foodniche breadth) and had almost identical foraging behavior. The first case occurred early in the season between Xanthippus corallipes and Arphia conspersa with foodniche breadths of 14 and 11, respectively. The second case occurred later in the season between Amphitornus coloradus and Trachyrhachys kiowa both of which have foodniche breadths of 6. The amount of movement between plants while feeding was equal or almost equal in both cases, thus we observed high dietary overlap between species with equal ecological efficiencies. Dietary overlap between the 14 grasshopper species decreased as foodniche dimensions increased and the grasshopper population as a whole utilized each food resource approximately proportional to its relative abundance. Factors influencing the interpretation and importance of dietary overlap in this study included 1. non-synchronous life cycles, 2. seasonal changes in diets and food preferences, 3. complexity of the food resources, 4. variations in foodniche dimensions among grasshopper species, and 5. variations in diets between males and females within a species. Our findings agree with MacArthur and Levins theory that the number of competing species which can coexist is proportional to the total range of the environment divided by the niche breadth of the species.     

Abundance responses of frugivorous bats (Stenodermatinae) to coffee cultivation and selective logging practices in mountainous central Veracruz,Mexico

Shifting and permanent cultivation, selective logging, cattle production and coffee plantations are among the most important factors in montane cloud forest conversion and disturbance. Although shaded-coffee plantations can contribute to the preservation of local species richness, abundance of organisms could be determined by habitat resource availability in agricultural landscapes. We compared abundance of Sturnira and Artibeus bats (Phyllostomidae, Stenodermatinae), in shade coffee plantations and disturbed cloud forest fragments, which represent habitats with different chiropterochorous plant density. We also investigated the relationship between bat species abundance and food plant richness, abundance and diversity. We captured 956 bats, 76% in cloud forest fragments and 24% in shaded coffee plantations. Abundance of Sturnira spp. (small bats) was greater in cloud forest than in coffee plantations, but Artibeus spp. (large bats) abundance was similar in both habitats. Chiropterochorous plant abundance was positively related with bat abundance for Sturnira spp., while chiropterochorous plant richness and diversity were negatively related for Artibeus spp. This suggests that frugivorous bats with different morphological and ecological characteristics respond differentially to anthropogenic activities. For landscape management purposes, the maintenance and augmentation of diverse food resources, for frugivorous bats with different foraging requirements in coffee plantations, will benefit the resilience of bats to modification of their natural habitat.     

Ecological Responses of Frugivorous Bats to Seasonal Fluctuation in Fruit Availability in Amazonian Forests

Lowland Amazon is climatically one of the least seasonal regions on the planet, but little is known about how this is reflected in ecological seasonality. The central objective of this study was to determine whether seasonal fluctuations in the availability of fruit resources in Neotropical forests are sufficiently marked to affect the ecology and physiology of frugivorous bats. Seasonal variations in the overall bat abundance and in the captures, body condition, and reproductive activity of the two most abundant species, Carollia perspicillata and Artibeus planirostris, were studied within a region of central Brazilian Amazonia dominated by a mosaic of nonflooded (terra firme) and seasonally flooded forests (várzea and igapó). Concurrent seasonal changes in fruit availability were measured. The abundance of fruits was markedly seasonal, with far fewer resources available during the low-water season. There was a positive correlation between fruit and bat abundance. Overall, bats did not increase the consumption of arthropods during the period of fruit shortage. In A. planirostris, the body condition declined when fruits were scarcer. In both C. perspicillata and A. planirostris, foraging and reproductive activity were positively correlated with fruit availability. Consequently, the results suggest that resource seasonality is sufficiently marked to affect frugivorous bats and force them to make important eco-physiological adjustments.     

Dietary composition,breadth, and overlap between seasonally sympatric Himalayan musk deer and livestock: Conservation implications

Livestock in high altitudes of Nepal and elsewhere, frequently and freely, use potential habitat of native wildlife for foraging. Such intrusion of ecologically similar domestic species is supposed to negatively impact the resident wildlife via ‘perceived’ and/or ‘real’ competitive interactions. Hence, assessment of dietary composition and overlap between herbivores is crucial to gain insight into the potential impacts via resource exploitation by foraging livestock. Also, evaluation of dietary composition of resident wildlife across seasons is important to decipher their seasonal resource needs. Within this context, microhistological technique, that makes use of fecal pellet for identification of plant species through comparison with reference slides of plant materials in the area, was used to assess dietary composition, breadth, and overlap between seasonally sympatric Himalayan musk deer and livestock in Nepal Himalaya. Musk deer and livestock were found to have significantly different dietary consumption and that partition was contributed by different species; meaning different plant species were associated to the diets of these two groups. Of notable, however, was a considerable ‘number’ of species (i.e., species richness) shared in diets by musk deer and livestock raising a concern of unchecked number of livestock with a potential to exploit and reduce the availability of shared plant species with musk deer. Also, seasonal dietary composition of musk deer significantly varied, with increased dietary breadth in winter, suggesting a potential for intraspecific competition for forage in winter because of limited availability of resources mediated by retarded growth and harsh conditions.     

Intersexual segregation in foraging microhabitat use by Magellanic Woodpeckers (Campephilus magellanicus): Seasonal and habitat effects at the world's southernmost forests

Animals facing seasonal food shortage and habitat degradation may adjust their foraging behaviour to reduce intraspecific competition. In the harsh environment of the world's southernmost forests in the Magellanic sub‐Antarctic ecoregion in Chile, we studied intersexual foraging differences in the largest South American woodpecker species, the Magellanic Woodpecker (Campephilus magellanicus). We assessed whether niche overlap between males and females decrease when food resources are less abundant or accessible, that is, during winter and in secondary forests, compared to summer and in old‐growth forests, respectively. We analysed 421 foraging microhabitat observations from six males and six females during 2011 and 2012. As predicted, the amount of niche overlap between males and females decreased during winter, when provisioning is more difficult. During winter, males and females (i) used trees with different diameter at breast height (DBH); (ii) fed in trunk sections with different diameters; and (iii) fed at different heights on tree trunks or branches. Vertical niche partitioning between sexes was found in both old‐growth and secondary forests. Such a niche partitioning during winter may be a seasonal strategy to avoid competition between sexes when prey resources are less abundant or accessible. Our results suggest that the conservation of this forest specialist, dimorphic and charismatic woodpecker species requires considering differences in habitat use between males and females.     

Food‐web structure and mercury dynamics in a large subarctic lake following multiple species introductions

相似文献   

The Role of Fruits and Insects in the Nutrition of Frugivorous Bats: Evaluating the Use of Stable Isotope Models1

We estimated the relative contribution of fruits and insects as sources of dietary protein in two species of Neotropical frugivorous bats (Artibeus jamaicensis and Sturnira lilium) using stable carbon and nitrogen isotope analyses. An insectivorous species (Pteronotus parnellii) was also included for comparison. We found constant patterns in stable carbon and nitrogen isotope composition in blood that separated the two species of frugivorous bats from the insectivorous bat. When we used these isotopic values (combined with those of dietary fruits and insects) to estimate the percent contribution of fruits and insects to the diet of the bats, we obtained different results, depending on assumptions and model adopted. We tested models using both 8“N and 8′³C results simultaneously and separately and further used diet‐tissue fractionation factors of 3%o for nitrogen and 1 and 3.5%o for carbon. We found that a carbon‐based model with a diet‐blood enrichment factor of 3.5%o produced the most parsimonious results. The model estimated that A. jamaicensis and S. lilium obtained most of their protein requirements from fruits, whereas P. parnellii fed mostly on insects. No sexual or seasonal variations in the diet of the two frugivorous species were detected. We found no evidence that the diet of sexually active females differed from that of nonsexually active females in the two species of frugivorous bats. We suggest that future studies better define isotopic fractionation between diet and tissues of bats using captive rearing and controlled diets.     

Temporal patterns of seed use and availability in a guild of desert ants

ABSTRACT.

• 1 Temporal patterns of seed use were studied from late winter to autumn in three species of seed-harvesting ants in the Sonoran Desert. Measures of effective foraging activity, dietary niche breadth and dietary niche overlaps were obtained each month and were tested for correlation with estimates of the available seed resource.
• 2 Seeds were the only numerically important type of food in the diets of all species.
• 3 The ants partitioned the resource according to both seed species and seed size, although there was considerable overlap.
• 4 Pheidole xerophila had the smallest forager body size and is a specialist on small seeds because it harvested them in greater proportion than their rank in the soils and expanded its diet to larger seeds only when the abundance of small seeds declined.
• 5 When the abundance of the small seeds of Bouteloua barbata decreased, the middle-sized ant, Veromessor pergandei, showed a decrease in foraging activity, increase in niche breadth, and a decrease in overlap with P.xerophila.
• 6 Seed size preferences of V.pergandei did not vary seasonally, except that during the month of highest seed abundance, V.pergandei showed no size preference.
• 7 Pogonomyrmex rugosus was the largest ant; it preferred larger seeds and was inactive when small seeds were most abundant. Seasonal foraging activity and niche parameters were random in relation to seed abundance.
• 8 We suggest that nocturnal foraging by P.rugosus during the summer months was a response to interference with diurnal foraging by either predation frorn horned lizards or competition from V.pergandei.
• 9 Seasonal abundance of small seeds explains most of the seasonal foraging patterns of P.xerophila and V.pergandei. The summertime abundance of larger seeds during years of adequate precipitation may account for the seasonal activity patterns of P.rugosus.