Oecologia

Frugivorous insect larvae are dependent on fruiting plants for development, leading to complex host-parasite interactions that may be influenced by many factors at various scales. We compared the relative effects of factors at the individual, neighborhood, and landscape scales in forest patches. Our results suggest that in areas like upstate New York, where agricultural land uses are dominant, individual scale factors are the most influential. Specifically, parasitism increased with host fruit crop size, but was not associated with host species richness or proximity to forest edge. Notably, the most parasitized hosts were non-native species, including Frangula alnus Mill. (Glossy Buckthorn), indicating a potential role of invasive species to shape host-parasite interactions in our system. Our results underscore the importance of host-specific traits in structuring parasitism patterns and suggest management could consider both the ecological context of host traits and the influence of invasive species at multiple scales.

Repeated grazing and trampling can reduce forb populations in South African mesic grasslands by limiting photosynthesis and depleting carbohydrate reserves stored in underground organs. We examined the effects of repeated defoliation on growth, bulb starch reserves and water status in the geophyte Eucomis autumnalis over 468 days. Four intense summer and autumn defoliations reduced above-ground production, bulb mass, starch reserves and bulb water pools, with the strongest impacts occurring in late summer and autumn when plants normally replenish reserves. Repeated defoliation disrupted seasonal source sink dynamics, forcing resources into regrowth and driving bulbs towards energetic bankruptcy. These results highlight the importance of avoiding heavy autumn grazing and providing periodic full-year rests to conserve geophytic forbs in mesic grasslands.

Arctic and Boreal raptor communities will continue to be affected by borealization and other climate change related processes, providing a challenge for ecologists predicting future sates. However, by using community assembly theory and species traits, future communities may be predictable. In this study, we analyzed variation in reproduction traits as a consequence of diet specialization for 29 raptors, 2 skuas and 3 corvids. We assessed and implemented foraging traits for specialists and generalists into predator-prey models from which successful invasion conditions were derived. Specialist raptors produced larger clutch sizes, had a higher proportion of fledged per clutch and also expressed more variation compared to generalist raptors. These results suggest a relationship between diet specialization and reproductive traits which was also observed within phylogenetic orders. Specialist owls (Strigiformes) produced higher clutch sizes with a larger clutch range compared to generalist owls. The same pattern was observed for falcons (Falconiformes). No clear difference in reproduction was observed for specialist and generalist hawks, kites and eagles (Accipitriformes). Corvids expressed clutch sizes similar to that of specialist raptors while having the lowest proportion of fledged per clutch. Differences in foraging traits between specialists and generalists could be distinguished using functional response curves. A predator-prey model parameterized with foraging trait data showed that a generalist can coexist with a resident specialist if it has access to prey unavailable to the resident specialist. Otherwise, the native specialist outcompetes the invading generalist due to foraging efficiency. The combined empirical and theoretical findings in this study show how diet specialization affects both reproduction and the potential invasion success of raptors.

Understanding the relationships between soil macrofauna and decomposition is crucial for predicting how land-use change impacts ecosystem function in fragmented systems. This is because soil macrofauna affect decomposition and also respond to the changes in abiotic conditions across habitat gradients. This study investigates edge effects on the macrofauna contributions to decomposition across forest-field ecotones. We used bait lamina assay to quantify aboveground and belowground feeding activity of soil macrofauna in Autumn 2025 in three deciduous forest-old field ecotones and one coniferous forest-old field ecotone, in Southwestern Ontario, Canada. Vegetation diversity and composition, LAI and soil characteristics (i.e., soil organic matter, pH, temperature and moisture) were measured at each plot along the ecotone. Pitfall trap data collected in Summer 2025 at the same sites were used to characterize macrofauna communities. We used generalized linear mixed effects models to estimate the effect of distance to edge, site, and depth into the soil on bait lamina consumption and soil macrofauna, with transect nested within site as random effects. Consumption activity increased with distance into the forest from the field, with the edge representing an intermediate; and, decreased with increasing depth into the soil. In contrast, soil macrofauna abundance, especially isopods, decrease with distance into the forest from the field. These trends varied significantly across sites, so that consumption activity and abundance sometimes remained constant across the ecotone (i.e., site x distance interaction). The results demonstrate that macrofaunal contributions to bait consumption varied along the ecotone, shaped by interacting environmental gradients and shifts in community composition unique to each site.

Climate change is expected to alter plant populations not only through direct environmental shifts but also via changes in biotic interactions, such as with herbivores and pathogens. As plant species are also expected to differ in their responses to both climate and antagonists, plant responses to both factors are expected to be variable and species-specific. To assess whether interactive effects of climate and antagonists on plant population dynamics are common and whether the strength and direction of plant responses vary across species, we conducted a multi-year field experiment that manipulated realistic climate change and experimentally reduced insect herbivores and fungal pathogens. We measured responses of plant vital rates, such as survivorship, growth, and reproduction across six grassland species. Using Integral Projection Models (IPMs) and Life Table Response Experiments (LTREs), we quantified changes in population growth rate across experimental treatments and the contribution of each vital rate to that observed change. Two of the study species declined so drastically over the course of the experiment that demographic quantification of population growth rates was not possible. From the remaining species, Bromus erectus and Plantago lanceolata show significant interactive responses of climate and antagonist reduction on population growth rates. In contrast, Dianthus carthusianorum and Tragopogon orientalis showed limited responses to experimental treatments. Notably, our results indicate that in some species biotic interactions may amplify the effects of climate change: the presence of plant antagonists exacerbates the negative effects of the future climate treatment on plant population dynamics. Our findings highlight the complexity in predicting plant population responses to climate change and provide insights for grassland management under future environmental conditions.

1. Ruby-throated hummingbirds (Archilochus colubris) exhibit flower preferences and readily visit human subsidies like feeders. However, foraging behavior and plant-animal interactions may vary across spatial, temporal, and landscape gradients. Despite the popularity and ubiquity of Ruby-throated Hummingbirds in the eastern U.S., there has never been a quantitative assessment of their flower preferences or feeder use across broad scales. 2. We investigated how feeder visitation, flower visitation, and flower trait preferences vary by latitude, season, and land use in the northeastern United States using over 6 million occurrence records of Ruby-throated Hummingbirds and flowering plants, >2,100 annotated hummingbird-flower interactions, and >2,700 feeder visit occurrences. 3. We found that hummingbird feeder use declined over the year, increased with latitude, and was higher in developed landscapes. Flower visitation increased over the year across all latitudes, with higher visitation in developed landscapes. Finally, we found that native plant use diverged between landscapes, such that the probability of visiting a native flower increased over time in non-developed land uses but declined over time in developed ones, demonstrating that hummingbirds track the advancement of native floral phenology and use non-native, cultivated flowers as a human subsidy due to either availability or preferences. 4. Our preference and network models revealed that while hummingbird-plant network structure was similar across landscapes, the composition of important taxa shifted from native, wild species like Monarda and Impatiens to non-native, cultivated species like Salvia. 5. Using trait-based models of flower visitation, we found that hummingbirds preferred native, tubular, and red/orange flowers fitting the hummingbird pollination syndrome despite visiting >260 different plant species. Red and orange flowers were preferred across all seasons, suggesting color may be a reliable signal of nectar availability across species and contexts. Native and tubular flowers were strongly preferred during the breeding season; however, preferences relaxed during spring and fall migration. 6. These findings reveal the consistent preferences of Ruby-throated Hummingbirds for native, tubular, and red/orange flowers, and underscore how spatial and temporal factors reshape foraging behavior and trait preferences. Our results also highlight the value of community-collected data in characterizing plant-pollinator interactions across broad spatial and temporal scales.

Reproduction in species with parental care involves sustaining a brood of offspring through an energetically demanding period, when shifts in resource availability, weather, predation risk, and parental condition can strongly alter offspring survival. The most extreme outcome is complete brood failure (death of all offspring), which is relatively frequent in many bird species and may occur when conditions cross a viability threshold. Although complete brood failure is important for shaping fitness variation and population dynamics, we have limited understanding of how intra- and interspecific density dependence governs these events, or of how factors such as habitat quality and disease burden contribute to them, because deriving this requires fine-scale, individual-level data collected across generations for multiple overlapping species. Using a dataset totalling 38,509 nesting attempts from great tits (Parus major) and blue tits (Cyanistes caeruleus) in Wytham Woods, Oxford, UK, we examined how brood failure is shaped by local conspecific and heterospecific density, habitat structure, and avian malaria infection for a subset. Complete brood failure was frequent (14.75%), mostly involving chick mortality in the nest consistent with starvation, rather than brood removal by predators. Relationships between density and brood failure were strong but species-specific. Specifically, great tit failure risk was higher in neighbourhoods that remained densely populated across years, whereas blue tit failure risk was lower where annual great tit or combined density was high, but not where annual blue tit density itself was high. This suggests that local overall density reflects continuing constraint for great tits, while local annual density may partly track favourable within-year conditions and settlement patterns for blue tits. In great tits, failure was also more common where oak density was low and farther from the closest river (Thames), while habitat associations were weak in blue tits. Malaria infection was spatially heterogeneous and covaried with density and habitat, but infection status did not significantly explain complete brood failure. Together, these results show that complete brood failure is shaped by spatially structured local ecological context, and how density dependence in these events can differ in direction and timescale between sympatric species.

Artificial light at night (ALAN) can alter the movement of nocturnal pollinators such as moths, with potential consequences for pollination services. However, most studies have focused on the presence or properties of lights, while the role of spatial lighting configuration remains poorly understood. We conducted a small-scale field experiment to test how different lighting configurations affect pollination success in moth-pollinated plants, using low-intensity LED garden lights. Potted phytometer plants of three moth-pollinated species were exposed to one of three treatments: no-light control, isolated LED point lights, or 25 m linear arrays of multiple LED lights. We quantified pollination success as both the probability of seed set and reproductive output through seed and capsule production. Experimental lighting affected plant reproductive success, with light arrays increasing seed-set probability, seed and seed capsule number relative to unlit controls, whereas point lights showed no clear effect. These results provide preliminary evidence that the spatial configuration of artificial lights may influence nocturnal pollination outcomes. Future work combining phytometer assays with direct tracking of moth movement is needed to assess whether light arrays facilitate or redirect the dispersal of nocturnal pollinators.

Understanding how habitat connectivity shapes biodiversity remains a major ecological challenge. In particular, the roles of connectivity and ecological heterogeneity on co-variation in plant species diversity and intraspecific genetic diversity is not understood. We combined species distribution modelling, resistance-to-movement mapping, landscape connectivity analysis and population genomics to investigate diversity patterns in three wet meadow herbs, Scorzonera humilis, Oenanthe peucedanifolia and Lychnis flos-cuculi, and their surrounding plant communities. Genetic diversity patterns differed strongly among co-occurring species. Connectivity metrics explained genetic diversity only in O. peucedanifolia, and environmental drivers of genetic diversity were highly species specific. Importantly, genetic diversity changed with the presence of some species in the community, but it was consistently unrelated to indicators of local plant community diversity. Overall, the processes shaping within-species biodiversity may differ fundamentally from those structuring habitat connectivity and plant species communities, with important implications for conservation.

While mutualistic symbioses with nitrogen-fixing bacteria enable plants to access fixed nitrogen, they also require substantial carbon investment. Under carbon limitation, such as shading, shifts in biomass allocation can decouple symbiotic investment from leaf and root growth, potentially compromising plant nitrogen status. Because shading shifts biomass allocation toward light acquisition, it could influence nitrogen fixing symbiosis in two opposing ways. If nodulation remains coupled to leaves rather than roots, nitrogen status should be maintained despite reduced root growth. Alternatively, if root growth constrains nodulation, nitrogen status should decline. We tested these hypotheses by manipulating light availability (full sunlight vs. 50% shade) and quantifying biomass allocation and symbiotic nodulation. Under shading, plants allocated proportionally more biomass to shoots than to roots and invested less biomass in root nodules. Relationships between nodulation and leaf or root biomass differed between treatments but converged with increasing plant size, although shaded plants never attained the root biomass observed in full sunlight. Leaf nitrogen concentration was maintained under shading because nodulation remained coupled to leaf investment despite reduced root allocation. These findings highlight that, under carbon limitation, maintaining leaf and nodule coupling enables plants to reduce nodule investment without compromising the nitrogen benefits of symbiosis.

Age-dependent survival is central to understanding population dynamics and life-history evolution. We analysed carcass weight and age-at-harvest data from 6022 red foxes (Vulpes vulpes) collected across Sweden between 1967 and 1971 to evaluate latitudinal effects on body mass and age-dependent survival. Carcass weights decreased from south to north in both adults and sub-adults, contrary to Bergmann's rule, with southern foxes weighing approximately 1.27 times more than northern foxes. The latitudinal weight gradient exceeded the sex difference in both age classes, and no sex x region interaction was detected. The decrease in weight with latitude is consistent with reduced prey availability and harsher winter conditions in the north, which limit growth and body size during development. Using a Bayesian age-at-harvest model with region-specific population growth rates (lambda), we estimated age-dependent survival probabilities for four latitudinal regions and both sexes. Despite the strong latitudinal gradient in weight, survival did not show a corresponding pattern - regional differences were uncertain, with all credible intervals spanning zero. Regional population growth rates were consistent with slight decline in the north and near-stability in the south-central region, which suggests that body condition and population dynamics are coupled at the regional scale despite no survival gradient. The decoupling of body condition and survival across regions suggests that mortality patterns are similar across the latitudinal gradient. We discuss these patterns in terms of latitudinal productivity gradients, prey availability, and life-history trade-offs in a widely distributed carnivore.

Agroecosystems are dynamic ecosystems, constituted by patches of vegetation and agricultural use, where biodiversity is shaped by spatial and temporal variability. While most studies have focused on spatial composition and configuration, the role of temporal variability remains poorly understood. Yet, temporal dynamics can strongly modify species composition, abundance, and persistence in ecological communities. Temporal variability is particularly relevant in agroecosystems with rainfed agriculture where environmental conditions shift dramatically between rainy and dry seasons. In this paper, we assess the role of temporal variability on biodiversity maintenance in an agricultural matrix using a metacommunity model that simulates an agricultural landscape under rainfed conditions, that is, with abrupt seasonal changes in the agricultural patches. This model couples a local community network dynamic with a migration dynamic and is based on empirically documented features of rainfed agricultural matrices. Our results show that temporal variability provides new opportunities for species to recover from low densities. However, the effect of temporal variability is not straightforward. It depends on the initial and final conditions, the migration and mortality rates and the intensity of temporal variability. Overall, our findings highlight the need to further investigate temporal variability to better understand its role in shaping biodiversity in agricultural landscapes.

The energetic demands of courtship are central to sexual selection, but their magnitude and temporal variation remain poorly quantified in many signalling systems. We used closed-chamber respirometry and low-light video analysis to estimate courtship-associated metabolic rates in males of the bioluminescent ostracod Photeros sp. EGD. Low-activity metabolic rate varied strongly across the diel cycle: in small vessels that constrained movement, individually measured males consumed significantly more oxygen at night than during the day. We then compared oxygen consumption across vessels that differed in opportunities for movement and courtship. Metabolic rates were highest in large vessels that permitted bioluminescent courtship displays, intermediate in medium vessels that allowed swimming but not full displays, and lowest in small vessels that constrained movement. Oxygen consumption in large vessels at night was approximately 500% of small-vessel daytime rates, 280% of small-vessel nighttime rates, and 160% of medium-vessel nighttime rates. Because measurements integrated oxygen use over multi-hour intervals, these values represent time-averaged metabolic demand rather than instantaneous costs of individual light pulses or display trains. Video analyses suggested a positive association between signalling rate and oxygen consumption, although this relationship was not statistically supported in our large-vessel dataset, which had low statistical power. Together, these results show that male Photeros undergo strong diel shifts in metabolic state and that the whole-animal performance required to construct bioluminescent courtship displays may often impose substantial energetic demands.

Surges in food availability create localized but intense foraging opportunities, often attracting multi-species consumer groups. Agricultural practices can trigger these surges, prompting bird associations. However, the strength and duration of the association, as well as its drivers, remain unclear. This study examines waterbird association with harvesting and ploughing events in rice fields. The duration and magnitude of these associations were determined and three hypotheses addressed to explain them: (1) increased food availability, (2) enhanced foraging success and (3) reduced time allocated to vigilance. Waterbird counts and GPS tracking revealed strong associations with management events. Bird numbers spiked during events but declined within one to two days. Food availability (and soil penetrability) increased significantly during events - crayfish and rice during harvesting, worms and soil penetrability during ploughing - supporting Hypothesis 1. However, this did not improve foraging performance (intake rate, foraging success), rejecting Hypothesis 2. Higher competition, interference or kleptoparasitism in these large mixed-species flocks may offset increased food availability benefits. Alternatively, functional responses of target species may limit prey intake due to physiological or behavioural constraints. Hypothesis 3 was also unsupported, as birds did not reduce vigilance. It is plausible that birds may be drawn to events by the perception of a feast, not actual benefits. Gregariousness and foraging behaviour by local enhancement may explain such associations. Results highlight the complexity of bird responses to food surges while suggesting waterbirds in rice fields maintain stable foraging performance during agricultural management events and otherwise, indicating resilience to agricultural timing shifts.

Fat accumulation and use is critical for sustaining life. Most insects show a typical response to feeding where fat is accumulated when sufficient sugars and other carbohydrates are consumed. Parasitoid insects are an exception, because most species do not accumulate fat when feeding on a sugar-rich diet. Studies on fat metabolism generally measure fat content early in life without considering lipid metabolism as a dynamic process that is expected to change as life progresses. In this paper, we compared fat accumulation and use throughout the lives of adult female Drosophila melanogaster and females of 5 inbred lines of the parasitoid wasp Leptopilina heterotoma. We expected that fat accumulation would take place irrespective of teneral fat content in D. melanogaster. We found that fat D. melanogaster initially used fat reserves, while lean flies economized on fat stores. Both lean and fat flies started accumulating fat after 7 days of life, indeed showing a typical response for insects. Unlike other parasitoids, L. heterotoma populations differs in fat accumulation patterns that we expected to observe also between inbred lines. In none of the inbred lines, however, did fat accumulation take place. Our results did reveal that inbred lines differed in the rate at which fat was used mainly later during life. We further confirmed that D. melanogaster pupal size was highly correlated with adult female size for both D. melanogaster and L. heterotoma. Overall, our findings for L. heterotoma provide strong evidence that genetic background has a major impact on the rate at which fat is used over a lifetime.

The exchange of energy and organisms across habitat boundaries links aquatic and terrestrial ecosystems and sustains ecosystem functioning. Although disturbance may disrupt these linkages, the mechanisms at play remain poorly understood. Here, we investigated the extent to which flow intermittency may disrupt riparian-aquatic ecosystem linkages by altering consumer communities in the recipient ecosystem or by altering resource quality in the donor ecosystem. We ran an experiment in an intermittent river network in California, focusing on a critical forest-to-river subsidy (organic matter in the form of leaf litter), its transformation, and its reciprocal benefit (aquatic insect production). Using three riparian species (willow, cottonwood, and oak) at sites spanning a gradient of flow permanence, we quantified intraspecific plasticity in leaf traits (specific leaf area, nitrogen and phosphorus concentrations, and {delta}13C), measured decomposition rates, and estimated the secondary production of aquatic shredders (Plecoptera). Across all leaf species, decomposition rates were 16-36% lower at intermittent than perennial sites, an effect largely driven by intraspecific leaf trait plasticity rather than changes in consumer abundance. At high flow intermittency, willow experienced water stress (enriched {delta}13C) and reduced specific leaf area, while cottonwood showed primarily stoichiometric responses (reduced leaf nitrogen and phosphorus). Despite these divergent strategies, all species produced lower-quality litter at intermittent sites. Variance partitioning confirmed that initial litter quality uniquely explained 51.5% of variation in decomposition rates, more than double the contribution of invertebrate community metrics; and structural equation modeling revealed that both leaf traits and stonefly (Plecoptera) secondary production significantly predicted decomposition rates, with leaf traits exerting the stronger effect. Notably, stonefly secondary production was 37-98% lower at intermittent sites across leaf species. Because these insects later emerge as terrestrial adults, they provide a significant energy flux to riparian predators, and, thus, impoverished litter quality suppresses the reciprocal transfer of energy back to terrestrial food webs. As drought intensifies globally, the decoupling of terrestrial-aquatic linkages may begin in the riparian canopy.

Various hypotheses have been proposed to explain why some species persist or even flourish in urban areas. Yet, despite its central role in determining when and where animals encounter resources, disturbance, and risk, movement behaviour remains an overlooked mechanism of urban success. In urban areas, human activities are strongly periodic, i.e. predictable in space and time. This may favour species able to adjust their behaviour to predictable cycles of resources and risks in space and time. Here, we tested this hypothesis and tracked movement behaviour along an urbanisation gradient in three mammal species with different urban success: red fox (Vulpes vulpes), an urban dweller; raccoon (Procyon lotor), an invasive urban dweller; and wild boar (Sus scrofa), an urban utiliser. We analysed periodicity in movement behaviour and investigated whether increasing urbanisation is associated with periodic reorganisation of activity timing, space use, and further analysed alterations in habitat selection along the urbanisation gradient. Our results show that foxes aligned their movement behaviour with human activity, having stronger day-night contrasts and more repeatable space use than their rural counterparts. Urban raccoons showed a contrasting strategy; they were more active during the day, without changes in their movement routines under increasing urbanisation, suggesting a flexible strategy that explains their urban success. In contrast, wild boars reduced routine movement behaviours with increasing urbanisation, consistent with their occurrence in less predictable suburban environments and avoidance of city centres. In summary, our results suggest that movement behaviour may be a key mechanism enabling animals to persist in cities, revealing distinct behavioural strategies for coping with urban environments.

Grasslands are vulnerable to increasing drought with global warming, but process-based models lack the mechanistic knowledge required to predict the magnitude of drought impacts. While a plant hydraulics framework has been successful in advancing process understanding of drought responses in trees, and how drought responses vary across rainfall gradients, similar approaches have rarely been applied to grasses. Here, we quantified the progression of key drought response processes in sixteen dominant perennial grasses (seven C3 and nine C4) with differing climatic origins across eastern Australia. We found that stomatal closure, hydraulic impairment and leaf browning occurred concurrently, in contrast to the progressive sequence typically observed in trees. We also found that drought response traits were not correlated with species climate of origin. The early impairment of leaf hydraulic conductance and leaf browning along with the lack of correlation with climate of origin suggest that grasses may employ fundamentally different strategies to adapt to low water availability than trees. These results highlight the need for grass-specific parameterization of drought responses in process-based models.

O_LISpawning salmon transport marine-derived nutrients (MDN) into riparian forests, influencing soil, plant, and animal communities, yet their effects on fungal communities remain poorly understood. C_LIO_LIWe used DNA metabarcoding to examine fungal responses to three spatial patterns of salmon-derived nitrogen (N) in southwest Alaska: (i) patchy inputs from wildlife-deposited carcasses, (ii) a 21-year carcass relocation experiment, and (iii) natural N gradients with distance from streams. C_LIO_LIDecomposing carcasses increased saprotrophic fungal diversity, identifying taxa responsible for salmon carcass decomposition. Long-term carcass relocation reduced diversity of medium-distance fringe ectomycorrhizal fungi (EMF), whereas recent, patchy carcass inputs increased diversity of both medium-distance fringe and long-distance EMF--guilds often associated with low-nutrient environments. Along natural stream N gradients, EMF responses varied markedly within functional guilds and genera, revealing unexpected variation in N sensitivity among closely related taxa. C_LIO_LIPulsed, spatially heterogeneous nutrient inputs enhanced diversity of typically nitrophobic EMF, likely reflecting their capacity to maintain extensive mycelial networks, exploit nutrient hotspots, and mobilize organic N and phosphorus. The diversity of responses along natural N gradients suggests that mechanisms linking EMF traits to nutrient acquisition and tolerance remain unresolved. Our findings emphasize the importance of linking fungal community composition with functional attributes and nutrient dynamics. C_LI

Despite the ecological importance of ungulate migrations, we lack a complete understanding of why some ungulates migrate and others do not. Though progress has been made towards understanding differences across species and between populations, migratory behavior varies even within populations: in many populations, some individuals remain behind as residents (partial migration). Theoretical population-level work has suggested that these different migratory tactics can coexist, but such approaches stop short of providing insights into how individuals make the decision to stay or go each year. Using long-term data from three ungulate populations, we find that individuals probabilities of migrating are highly variable across years, which points to a non-trivial context-dependent decision-making process, whose underlying mechanisms must be probed via individual-level modeling. Drawing on existing knowledge, we propose a decision-making model of ungulate migration onset wherein individuals probabilistically decide to start migrating based on the local intensity of environmental and/or social cues. Residents arise as a robust collective organization phenomenon in our model. At sufficiently large population sizes, the number of residents is invariant with total population size, consistent with empirical patterns. Instead, resident numbers are influenced by the severity of the bad season, by relevant character differences among individuals, and by how individuals contribute and respond to environmental and/or social cues; for instance, when social cues contribute to decision-making in addition to environmental ones, fewer residents result, and migration is more likely to be complete. Overall, our model provides a potential mechanistic explanation for how residents might emerge within migratory ungulate populations.