Functional responses in habitat selection are density dependent in a large herbivore

Habitat selection is complex due to density dependence and functional responses, defined as variation in relative habitat use depending on availability. In this study we unite these concepts by empirically testing for density‐dependent functional responses in habitat selection using a large herbivore, elk Cervus canadensis manitobensis, as a model species. Theory on density‐dependent habitat selection predicts specialised behaviour when densities are low with a gradual switch to generalist behaviour (more even selection of habitat) as competition intensifies. This suggests that functional responses in habitat selection should be positive when competition is low, but that density may have a negative effect on the functional response due to constraining effects of competition on habitat use and availability. We tested this prediction using data from Global Positioning System (GPS) collared elk (n = 33) and empirical data on spatiotemporal variation in local density during winter in Riding Mountain National Park, Manitoba, Canada (2002–2011). As local density increased, winter home range size decreased and the proportion of mixed forest (providing shelter and forage to elk) used and available within the home range also decreased. Our resource selection function (RSF) revealed clear density‐dependent effects in selection, being strongest (or weakest) for high quality (or low quality) habitat types at lowest observed density leading to more even selection as densities increased. The functional response in mixed forest selection was negatively affected by local density. Increasing availability of mixed forest in the home range was associated with higher selection at low density (positive functional response); no effect of availability on selection at moderate density (no functional response); and lower selection as availability increased at high density (negative functional response). Our study demonstrates that this process is largely driven by the negative effect of density on home range size as it constrains use and availability of habitat.     

Passive external transport of freshwater invertebrates by elephant and other mud‐wallowing mammals in an African savannah habitat

1. Recent findings hint at the potential importance of mammals affecting the spatial dynamics of aquatic organisms in areas where mammals live in close association with water. Perhaps the most iconic example of such an environment is the African savannah. 2. We investigated dispersal patterns of freshwater organisms among a set of temporary ponds in SE Zimbabwe to test the hypothesis that large mammals, and particularly African elephants (Loxodonta africana), can be important vectors of aquatic organisms. Dispersal kernels were reconstructed by hatching mud collected from ‘rubbing’ trees located at increasing distances from a set of isolated ponds. To assess the relative importance of other mammalian vectors, the vertical distribution of mud on rubbing trees was mapped and related to the body size of candidate vector species. 3. Laboratory hatching of mud samples revealed large numbers of propagules of 22 invertebrate taxa as well as some aquatic macrophytes. Dispersing communities reflected source communities and diverged with increasing distance from the source. Both dispersal rates and richness of transported taxa decreased significantly with dispersal distance. No indications for differences in dispersal capacity among propagule types were detected. Instead, common propagules were more likely to travel greater distances. Most mud was attached to trees at heights >1.5 m, implicating elephants as the dominant vector. Vertical distributions of tree mud, however, also revealed clustering at heights up to 50 cm and 90–120 cm corresponding to the height of warthog, rhinoceros and buffalo, respectively. Finally, variation in the vertical distribution of mud on trees in combination with differences in vector vagility suggests that local differences in vector species composition may affect passive dispersal dynamics of aquatic organisms. 4. Based on vagility and vector load, mud‐wallowing mammals emerge as highly effective vectors that, in some areas, may be more important in transporting aquatic organisms than traditionally recognised vectors such as waterbirds. Since most large‐ and medium‐sized mammals currently have restricted geographic distributions, it is likely that mammal‐mediated dispersal was more important in the past.     

Temporal scales of density‐dependent habitat selection in a large grazing herbivore

Habitat selection is a density‐dependent process, but little is known regarding how this relationship may vary across different temporal scales. Over long time scales, grazing shapes the structure, diversity and functioning of terrestrial ecosystems, and grazing‐induced changes in forage production over time are likely to affect the level of density dependence in habitat selection. In this fully‐replicated, landscape‐scale experiment, we investigated how density‐dependent habitat selection by a large grazing herbivore, sheep Ovis aries, develops over the time scale of a decade. We also address an often‐neglected challenge in habitat selection studies; namely, whether there is variation in use within a particular habitat or vegetation type and why. We found clear evidence of density dependence in habitat selection, with a wider use of habitats at high density. Despite a change in the standing biomass of high‐productivity vegetation at high herbivore density over the years, with herb biomass declining and graminoid biomass increasing, there was no clear evidence that these grazing‐induced changes in habitat over the years were strong enough to affect the level of density‐dependent habitat selection. The difference in selection for high versus low‐productivity habitats remained similar, despite annual fluctuations in the strength of selection. We found strong variation in selection within each vegetation type, even when vegetation types were mapped at a fine‐resolution scale. Our study shows that despite the interactive effects of herbivores and habitats, they are not always sufficiently strong enough to affect the level of density‐dependent habitat selection.     

Functional responses in polar bear habitat selection

Habitat selection may occur in situations in which animals experience a trade-off, e.g. between the use of habitats with abundant forage and the use of safer retreat habitats with little forage. Such trade-offs may yield relative habitat use conditional on the relative availability of the different habitat types, as proportional use of foraging habitat may exceed proportional availability when foraging habitat is scarce, but be less than availability when foraging habitat is abundant. Hence, trade-offs in habitat use may result in functional responses in habitat use (i.e. change in relative use with changing availability). We used logistic and log-linear models to model functional responses in female polar bear habitat use based on satellite telemetry data from two contiguous populations; one near shore inhabiting sea ice within fjords, and one inhabiting pelagic drift ice. Open ice, near the ice edge, is a highly dynamic habitat hypothesised to be important polar bear habitat due to high prey availability. In open ice-polar bears may experience a high energetic cost of movements and risk drifting away from the main ice field (i.e. trade off between feeding and energy saving or safety). If polar bears were constrained by ice dynamics we therefore predicted use of retreat habitats with greater ice coverage relative to habitats used for hunting. The polar bears demonstrated season and population specific functional responses in habitat use, likely reflecting seasonal and regional variation in use of retreat and foraging habitats. We suggest that in seasons with functional responses in habitat use, polar bear space use and population distribution may not be a mere reflection of prey availability but rather reflect the alternate allocation of time in hunting and retreat habitats.     

Functional diversity of macroinvertebrates on abandoned cattle enclosures in a semi‐arid African savannah

Zimbabwe is experiencing a shift in land‐use, away from livestock farming and towards wildlife conservation. The abandonment of livestock farming may have unforeseen consequences on biodiversity and ecosystem functioning, as cattle kraals create valuable nutrient‐rich patches across the semi‐arid savannah. It is unclear how macroinvertebrates functionally respond to such nutrient‐rich patches in semi‐arid savannahs. We analysed functional diversity of both aboveground and belowground taxa on abandoned cattle kraals and savannah control plots in Save valley Conservancy (SVC). We used distance‐based multivariate techniques to estimate indices of functional diversity. Our results indicated that after two decades of abandonment, kraals had higher functional richness (FRic), functional divergence (FDiv) and functional dispersion (FDis) of macroinvertebrates when both aboveground and belowground species are combined. When aboveground macroinvertebrates were considered alone, no difference was observed for all the considered functional indices. However, only FRic was higher on kraals when belowground macroinvertebrates were separately considered. Our results suggest that two‐decade‐old abandoned kraals may have recovered enough for aboveground species to match the surrounding savannah plot and even surpassed the savannah control for belowground species functional diversity.     

Functional response in habitat selection and the tradeoffs between foraging niche components in a large herbivore

How herbivore behaviour is influenced by changes in resource levels is central for understanding trophic interactions. We examined whether foraging tradeoffs change with food levels by comparing habitat selection and space use within and between two neighbouring, predator‐free Svalbard reindeer populations. The populations faced different food levels due to contrasting grazing history. Summer resource selection in radiocollared females was assessed by a multi‐dimensional niche approach based on habitat variables obtained from a satellite image (e.g. the normalised difference vegetation index, NDVI) and a digital terrain model. The population at the overgrazed Brøggerhalvøya faced overall lower plant cover, biomass and primary productivity (i.e. lower NDVI) than the population at Sarsøyra. At Brøggerhalvøya, most reindeer selected for productive habitat when choosing home range and patches within the home range. In contrast, habitat selection at Sarsøyra was more affected by abiotic conditions such as moisture, which may influence plant quality. Here, reindeer used patches with even less biomass than the average reindeer at the poorer Brøggerhalvøya. Such a difference in habitat preference with different habitat availability (a functional response in habitat selection) probably reflected increased selection for high‐quality forage at the expense of high forage quantity at Sarsøyra. Accordingly, a negative relationship between habitat productivity and home range size was only present across individuals within Brøggerhalvøya, where forage quantity was the important foraging niche component. Individuals having poor (and large) home ranges apparently could not compensate for this by higher patch selectivity compared to individuals with richer home ranges. The results indicate changes in foraging tradeoffs at contrasting resource levels and that strong interactions occur between habitat selection, space use and the foraging niche structure in the absence of predation.     

A high‐quality chromosome‐level genome assembly of a generalist herbivore,Trichoplusia ni

The cabbage looper, Trichoplusia ni, is a globally distributed highly polyphagous herbivore and an important agricultural pest. T. ni has evolved resistance to various chemical insecticides, and is one of the only two insect species that have evolved resistance to the biopesticide Bacillus thuringiensis (Bt) in agricultural systems and has been selected for resistance to baculovirus infections. We report a 333‐Mb high‐quality T. ni genome assembly, which has N50 lengths of scaffolds and contigs of 4.6 Mb and 140 Kb, respectively, and contains 14,384 protein‐coding genes. High‐density genetic maps were constructed to anchor 305 Mb (91.7%) of the assembly to 31 chromosomes. Comparative genomic analysis of T. ni with Bombyx mori showed enrichment of tandemly duplicated genes in T. ni in families involved in detoxification and digestion, consistent with the broad host range of T. ni. High levels of genome synteny were found between T. ni and other sequenced lepidopterans. However, genome synteny analysis of T. ni and the T. ni derived cell line High Five (Hi5) indicated extensive genome rearrangements in the cell line. These results provided the first genomic evidence revealing the high instability of chromosomes in lepidopteran cell lines known from karyotypic observations. The high‐quality T. ni genome sequence provides a valuable resource for research in a broad range of areas including fundamental insect biology, insect‐plant interactions and co‐evolution, mechanisms and evolution of insect resistance to chemical and biological pesticides, and technology development for insect pest management.     

Habitat quality influences population distribution, individual space use and functional responses in habitat selection by a large herbivore

Identifying factors shaping variation in resource selection is central for our understanding of the behaviour and distribution of animals. We examined summer habitat selection and space use by 108 Global Positioning System (GPS)-collared moose in Norway in relation to sex, reproductive status, habitat quality, and availability. Moose selected habitat types based on a combination of forage quality and availability of suitable habitat types. Selection of protective cover was strongest for reproducing females, likely reflecting the need to protect young. Males showed strong selection for habitat types with high quality forage, possibly due to higher energy requirements. Selection for preferred habitat types providing food and cover was a positive function of their availability within home ranges (i.e. not proportional use) indicating functional response in habitat selection. This relationship was not found for unproductive habitat types. Moreover, home ranges with high cover of unproductive habitat types were larger, and smaller home ranges contained higher proportions of the most preferred habitat type. The distribution of moose within the study area was partly related to the distribution of different habitat types. Our study shows how distribution and availability of habitat types providing cover and high-quality food shape ungulate habitat selection and space use.     

Seasonal habitat selection and space use by a semi‐free range herbivore in a heterogeneous savanna landscape

Understanding factors that influence habitat selection in heterogeneous landscapes is fundamental for establishing realistic models on animal distribution to inform rangeland management. In this study, we tested whether seasonal variation in habitat selection within the home range of a large herbivore was influenced by constraints such as, distances from water and central place using semi‐free range cattle (Bos taurus) as a case study. We also tested whether shifts in space use over time were dependent on spatial scale and on the overall abundance of resources. We predicted that distance from water significantly influenced dry season habitat selection while the influence of the central place on habitat selection was season‐independent. We also predicted that shifts in space use over time were spatial scale‐dependent, and that large herbivores would include more diverse habitats in their home ranges during the dry season, when water and food resources are less abundant. Multinomial logit models were used to construct habitat selection models with distances from water and central place as habitat‐specific constraints. Results showed significant variations in habitat selection between the dry and wet season. As predicted, the effect of distance from central place was season‐independent, while the effect of water was not included in the top dry season models contrary to expectation. A diverse range of habitats were also selected during the dry season including agricultural fields. Results also indicated that shifts in space use were spatial scale dependent, with core areas being more sensitive to changes than the home range. In addition, shifts in space use responded to temporal changes in habitat composition. Overall, our results suggest that semi‐free range herbivores adopt different foraging strategies in response to spatial‐temporal changes in habitat availability.     

Disparate determinants of summer and winter diet selection of a generalist herbivore,Ochotona princeps

The North American pika, Ochotona princeps, is a generalist herbivore that simultaneously selects two distinct diets: one consumed immediately (summer diet), the other harvested, transported, and stored for later consumption (winter diet). I investigated factors influencing diet selection at two sites on the West Knoll of Niwot Ridge, Boulder County, Colorado during 1991 and 1992. The composition of summer and winter diets differed significantly from each other as well as from the relative abundance of food items in the environment. Thus, pikas were not foraging randomly for either diet. To explore winter and summer diet selection, I tested two existing hypotheses: (1) that plant morphology restricts the winter diet breadth to plants that are easily harvested and large, and thereby maximizes the amount collected per foraging effort, or (2) to compensate for nutrients lost during storage, pikas bias their winter diet with high-nutrient species. I also tested the hypothesis that plant secondary compounds may be higher in the winter diet either because they function as preservatives or because pikas delay consumption of these species until the toxins degrade. For individual dietary items, I measured energy, nitrogen, water, fiber, total phenolic, condensed tannin, and astringency contents. There was little evidence to suggest that morphology excluded plants from the winter diet. Plant size was not a good predictor of abundance in the winter diet. Even after harvesting costs had been experimentally removed, cushion plants were still not included in the winter diet. There was weak support for an effect of nutrients on winter diet selection; in three of four cases, the winter diet was significantly lower in water and higher in total energy content as predicted by the nutrient compensation hypothesis. However, other nutrients exhibited no consistent pattern. Nutrients were not reliable predictors of the winter diet in multiple regression analyses. There was strong support for the hypothesis of manipulation of secondary compounds. The winter diet was significantly higher in total phenolics and astringency. Total phenolics were consistent predictors of the winter diet in multiple regression analyses. The winter diets of six additional pika populations contained plant species high in secondary compounds. The results suggest that pikas preferentially select plants with high levels of secondary compounds for their winter diet, possibly because the presence of such compounds promotes preservation of the cache. This behavior may also enable the exploitation of an otherwise unusable food resource, i.e., toxic plants.     

Interspecific variation in the resprouting responses of Acacia species following simulated herbivory in a semi‐arid southern African savannah

Plants have evolved a diverse suite of tolerance traits against herbivory, including compensatory growth, increased photosynthesis and activation of dormant meristems. We studied the responses of five Acacia species to simulated herbivory in a semi‐arid southern African savannah. We clipped terminal shoots of five juvenile Acacia species (Acacia rehmanniana, A. nilotica, A. karroo, A. arenaria and A. gerarrdii) to simulate herbivory. We then determined biomass change after 5 months and also counted the number of resprouts and measured their length and diameter. All clipped shoots produced resprouts, with all the Acacia species compensating for the lost biomass. We found considerable interspecific variation in the compensation for biomass lost to herbivory in the five Acacia species. Resprouts biomass ranged from two times in A. arenaria to four times that removed in A. karroo. Acacia karroo produced many resprouts, while A. arenaria produced very few resprouts (4 vs 15 resprouts). The relationship between the number of resprouts and their growth also varied among the different Acacia species. We conclude that the response of Acacias to herbivory ranges from prolific resprouters (such as A. karroo) to poor resprouters (e.g. A. arenaria).     

Trade-offs in responses to host plants within a population of a generalist herbivore,Christoneura rosaceana

Evolutionary constraints on the ability of herbivores to efficiently use a set of phytochemically similar hosts, while maintaining a high performance on phytochemically different hosts, are central in explaining the predominance of host specialization in phytophagous insects. Such feeding trade-offs could be manifested within insect populations as negative genetic correlations in fitness on different host species. We tested the hypothesis that feeding trade-offs were present within a population of the obliquebanded leafroller,Choristoneura rosaceana (Harris). Components of fitness were measured in families originating from an apple orchard that were fed on four host-plant species in the laboratory. Under the conditions of this experiment, all across-host genetic correlations were strongly positive, suggesting that this population comprised true generalists. With the exception of diapausing propensity, the heritability of the fitness components tended to be lower in caterpillars fed on apple leaves than in insects fed other hosts. This suggests a constraint on the selective response of the fitness components in the orchard environment.     

Fire frequency drives habitat selection by a diverse herbivore guild impacting top–down control of plant communities in an African savanna

In areas with diverse herbivore communities such as African savannas, the frequency of disturbance by fire may alter the top–down role of different herbivore species on plant community dynamics. In a seven year experiment in the Kruger National Park, South Africa, we examined the habitat use of nine common herbivore species across annually burned, triennially burned and unburned areas. We also used two types of exclosures (plus open access controls) to examine the impacts of different herbivores on plant community dynamics across fire disturbance regimes. Full exclosures excluded all herbivores > 0.5 kg (e.g. elephant, zebra, impala) while partial exclosures allowed access only to animals with shoulder heights ≤ 0.85 m (e.g. impala, steenbok). Annual burns attracted a diverse suite of herbivores, and exclusion of larger herbivores (e.g. elephant, zebra, wildebeest) increased plant abundance. When smaller species, mainly impala, were also excluded there were declines in plant diversity, likely mediated by a decline in open space available for colonization of uncommon plant species. Unburned areas attracted the least diverse suite of herbivores, dominated by impala. Here, herbivore exclusion, especially of impala, led to strong declines in plant richness and diversity. With no fire disturbance, herbivore exclusion led to competitive exclusion via increases in plant dominance and light limitation. In contrast, on triennial burns, herbivore exclusion had no effect on plant richness or diversity, potentially due to relatively little open space for colonization across exclosure treatments but also little competitive exclusion due to the intermediate fire disturbance. Further, the diverse suite of grazers and browsers on triennial burns may have had a compensating effect of on the diversity of grasses and forbs. Ultimately, our work shows that differential disturbance regimes can result in differential consumer pressure across a landscape and result in heterogeneous patterns in top–down control of community dynamics.     

Density dependent and temporal variability in habitat selection by a large herbivore; an experimental approach

Both density dependent and density independent processes such as climate affect population dynamics in large herbivores. Understanding herbivore foraging patterns is essential to identify the underlying mechanisms behind variation in vital rates. However, very little is known about how animals vary their selection of habitat temporally, alone or in interaction with density during summer. At the foraging scale, we tested using a fully replicated experiment whether domestic sheep Ovis aries stocked at high (80  per  km²) and low (25  per  km²) densities (spatial contrasts) varied their habitat selection temporally over a four year period. We predicted reduced selection of high productivity vegetation types with increasing density, and that seasonal and annual variation in climate would affect this density dependent selection pattern by increasing competition for high quality habitats in late grazing season and in years with poor vegetation development and over time related to vegetation responses to grazing. As predicted from the Ideal free distribution model, selection of high productivity habitat decreased at high density. There was also a marked temporal variation in habitat selection. Selection of the most productive vegetation types declined towards the end of each grazing season, but increased over years both at low and high sheep density. There was only weak evidence for interactions, as selection ratio of highly productive habitats tended to increase more over years at low density as compared to high density. Limited interactive effects of density and annual variation on habitat selection during summer may explain why similar interactions in vital rates have rarely been reported for summer seasons. Our results are consistent with the view that variation in habitat selection is a central mechanism for climate and density related variation in vital rates.     

Seasonal shifts in habitat selection of a large herbivore and the influence of human activity

Failure to recognize factors contributing to variation in habitat models like resource selection functions (RSFs) can affect their application for projecting probabilities of occurrence, and thereby limit their relevance for conservation and management. We compared seasonal RSFs (2006–2008) for 16 adult female moose (Alces alces) with home ranges located in western Algonquin Provincial Park (APP), Ontario, Canada, to those of 14 adult females located in provincial Wildlife Management Unit (WMU) 49, 40 km west of the protected area. Wildlife and habitat management practices differed between regions: hunting was higher in WMU 49 compared to APP, and APP preserved large tracts of old growth forest rarely found in WMU 49. Seasonal RSFs projected expected similarities in moose resource use between regions (e.g., responses to wetlands and stands of eastern hemlock, Tsuga canadensis [in winter]); however, we also observed differences consistent with the hypothesis that animals, through effects of hunting, would shift habitat use seasonally and in response to roads. We further observed evidence of functional responses in habitat selection due to underlying differences in forestry practices (e.g., responses to stands of old-growth hemlock forest). Given the close proximity and shared biogeographic region between study areas, we believe that observed spatial dynamics in RSFs were ultimately reflective of divergent management strategies between areas and ensuing differences in predation and hunting mortality risk, and functional habitat.     

Host–plant quality alters herbivore responses to temperature: a case study using the generalist Hyphantria cunea

The nutritional quality of host plants is an important determinant of fitness in insect herbivores. However, it remains an open question whether the ingestion of a particular plant will have the same effects on an herbivore under differing thermal conditions. We measured the performance of the generalist‐feeding caterpillars of Hyphantria cunea Drury (Lepidoptera: Arctiidae) raised on one of five natural host plants to determine their nutritional quality: Platanus occidentalis L. (Platanaceae), Sophora japonica (L.) Schott (Fabaceae), Prunus × yedoensis Matsum. (Rosaceae), Cornus kousa Hance (Cornaceae), or Betula platyphylla Sukaczev (Betulaceae). Caterpillars performed well on P. occidentalis, S. japonica, and P. × yedoensis, but poorly on C. kousa and B. platyphylla. The nutritional phenotype of caterpillars varied among host–plant groups, with the proportion of lipid‐free body mass to lipid content being higher for caterpillars raised on P. occidentalis and S. japonica (3.8–4.2:1) than for caterpillars raised on P. × yedoensis (1.6–2.1:1). A multi‐factorial experimental design was employed to investigate the interactive effects of host–plant quality and temperature on the performance of H. cunea caterpillars raised on either P. occidentalis or P. × yedoensis at three rearing temperatures (20, 25, or 30 °C). Caterpillars raised on P. occidentalis displayed a monotonic decrease in development time with increasing temperature, but the development time of those on P. × yedoensis decreased rapidly as temperature rose from 20 to 25 °C and then stayed unchanged despite further increase in temperature. The rate at which body size increased with decreasing temperature was much steeper for caterpillars raised on P. occidentalis than for those on P. × yedoensis. Collectively, these results indicate that host plant can alter the thermal reaction norms for the key life‐history traits of herbivores. This study has implications for understanding the impacts of climate change on herbivore–plant interactions.     

Selectivity in large generalist herbivores: feeding patterns of African ungulates in a semi-arid habitat

Feeding habits of free-ranging wildebeest and zebra were monitored in a semi-arid nature reserve, bordering the southwestern part of Kruger National Park, South Africa. The purpose of study was to distinguish and define the feeding niches of two roughage grazers that occur in similar habitat types. The monthly compositions of diets were evaluated by direct observations of feeding bouts over a period of two years when rainfall patterns were average and animal populations were stable. Other analyses evaluated the standing biomass of grass species in the reserve during the wet summer and dry winter seasons. A considerable overlap of grass species composition was found in the diets of wildebeest and zebra. Ordination of bi-monthly records of the diet composition showed greater variations in scores of grasses in zebra diet in comparison to wildebeest. Seasonal patterns were more apparent in the wildebeest diet. Preference ranking of grass species indicated that zebra diet remained constant in winter and summer. Wildebeest diet however, alternated with seasons, showing high preferences during the winter months for grass species which were rejected during summer. The combined assessment of results from three separate statistical methods analysing temporal patterns and preferences in diet composition revealed contradictory trends. The solution, however, relied on the initial assumptions posed. Hence, wildebeest and zebra are essentially generalist feeders which show a limited amount of preference in their choice of diet.     

Generalist trophic ecology in a changing habitat: The case of the four‐striped mouse in a woody‐encroached savannah

Trophic generalists tolerate greater habitat change than specialists; however, few studies explore how generalist trophic ecology is affected. We established how the trophic ecology of an extreme generalist, Rhabdomys pumilio, changed in relation to a directionally changing woody‐encroached savannah in Eastern Cape, South Africa by investigating (a) foraging behaviour, (b) trophic niche and (c) feedback effects. (a) Giving‐up densities showed that R. pumilio preferred foraging in subcanopy microhabitat during the night as a result of lower thermoregulatory costs, but had similar preferences for sub‐ and intercanopy microhabitats during the day. (b) An isotope analysis revealed that the dietary composition and trophic niche occupied by R. pumilio differed among tree canopy cover levels (0%, 30% and 80%), which appeared to be related to changes in C₄ grass material and invertebrate availability. (c) Artificial seed patches suggested that R. pumilio was a potentially important postdispersal seed predator of the woody‐encroaching species, Vachellia karroo. Thus, an increase in tree canopy cover altered the trophic niche of R. pumilio by reducing foraging costs at night and providing alternative food resources in terms of availability and source. These findings demonstrate how an extreme generalist adapted to human‐induced habitat change through changes in its trophic ecology.