Chronic wasting disease alters the movement behavior and habitat use of mule deer during clinical stages of infection.

Integrating host movement and pathogen data is a central issue in wildlife disease ecology that will allow for a better understanding of disease transmission. We examined how adult female mule deer (Odocoileus hemionus) responded behaviorally to infection with chronic wasting disease (CWD). We compared movement and habitat use of CWD-infected deer (n = 18) to those that succumbed to starvation (and were CWD-negative by ELISA and IHC; n = 8) and others in which CWD was not detected (n = 111, including animals that survived the duration of the study) using GPS collar data from two distinct populations collared in central Wyoming, USA during 2018-2022. CWD and predation were the leading causes of mortality during our study (32/91 deaths attributed to CWD and 27/91 deaths attributed to predation). Deer infected with CWD moved slower and used lower elevation areas closer to rivers in the months preceding death compared with uninfected deer that did not succumb to starvation. Although CWD-infected deer and those that died of starvation moved at similar speeds during the final months of life, CWD-infected deer used areas closer to streams with less herbaceous biomass than starved deer. These behavioral differences may allow for the development of predictive models of disease status from movement data, which will be useful to supplement field and laboratory diagnostics or when mortalities cannot be quickly retrieved to assess cause-specific mortality. Furthermore, identifying individuals who are sick before predation events could help to assess the extent to which disease mortality is compensatory with predation. Finally, infected animals began to slow down around 4 months prior to death from CWD. Our approach for detecting the timing of infection-induced shifts in movement behavior may be useful in application to other disease systems to better understand the response of wildlife to infectious disease.

A test of the frost wave hypothesis in a temperate ungulate.

Growing evidence supports the hypothesis that temperate herbivores surf the green wave of emerging plants during spring migration. Despite the importance of autumn migration, few studies have conceptualized resource tracking of temperate herbivores during this critical season. We adapted the frost wave hypothesis (FWH), which posits that animals pace their autumn migration to reduce exposure to snow but increase acquisition of forage. We tested the FWH in a population of mule deer in Wyoming, USA by tracking the autumn migrations of n = 163 mule deer that moved 15-288 km from summer to winter range. Migrating deer experienced similar amounts of snow but 1.4-2.1 times more residual forage than if they had naïve knowledge of when or how fast to migrate. Importantly, deer balanced exposure to snow and forage in a spatial manner. At the fine scale, deer avoided snow near their mountainous summer ranges and became more risk prone to snow near winter range. Aligning with their higher tolerance of snow and lingering behavior to acquire residual forage, deer increased stopover use by 1 ± 1 day (95% CI) day for every 10% of their migration completed. Our findings support the prediction that mule deer pace their autumn migration with the onset of snow and residual forage, but refine the FWH to include movement behavior en route that is spatially dynamic.

Dynamic balancing of risks and rewards in a large herbivore: Further extending predator-prey concepts to road ecology.

Animal behaviour is shaped by the ability to identify risks and profitably balance the levels of risks encountered with the payoffs experienced. Anthropogenic disturbances like roads generate novel risks and opportunities that wildlife must accurately perceive and respond to. Basic concepts in predator-prey ecology are often used to understand responses of animals to roads (e.g. increased vigilance, selection for cover in their vicinity). However, prey often display complex behaviours such as modulating space use given varying risks and rewards, and it is unclear if such dynamic balancing is used by animals in the context of road crossings. We tested whether animals dynamically balance risks and rewards relative to roads using extensive field-based and GPS collar data from elk in Yoho National Park (British Columbia, Canada), where a major highway completely bisects their range during most of the year. We analysed elk behaviour by combining hidden Markov movement models with a step-selection function framework. Rewards were indexed by a dynamic map of available forage biomass, and risks were indexed by road crossings and traffic volumes. We found that elk generally selected intermediate and high forage biomass, and avoided crossing the road. Most of the time, elk modulated their behaviour given varying risks and rewards. When crossing the highway compared with not crossing, elk selected for greater forage biomass and this selection was stronger as the number of highway crossings increased. However, with traffic volume, elk only balanced foraging rewards when they crossed a single time during a travel sequence. Using a road ecology system, we empirically tested an important component of predator-prey ecology-the ability to dynamically modulate behaviour in response to varying levels of risks and rewards. Such a test articulates how decision-making processes that consider the spatiotemporal variation in risks and rewards allow animals to successfully and profitably navigate busy roads. Applying well-developed concepts in predator-prey theory helps understand how animals respond to anthropogenic disturbances and anticipate the adaptive capacity for individuals and populations to adjust to rapidly changing environments.

Le comportement animal est influencé par la capacité des animaux à identifier et minimiser les risques rencontrés, tout en maximisant les gains obtenus. Les perturbations anthropiques, telles que les routes, engendrent de nouveaux risques et opportunités pour la faune. Les concepts de l'écologie prédateur-proie sont fréquemment utilisés pour comprendre les réactions des animaux aux routes (e.g. vigilance accrue, choix de couvert à proximité des routes). Cependant, même s'il est connu que les proies ajustent fréquemment leur utilisation de l'espace de façon à minimiser les risques et maximiser les récompenses, il n'est pas clair si une telle optimisation est utilisée par les animaux lorsqu'ils traversent des routes. Ici, nous avons évalué comment les animaux ajustent leur sélection d'habitat par rapport aux routes en fonction des risques et des récompenses disponibles. Nous avons examiné cette question chez les wapitis du parc national Yoho (Colombie-Britannique, Canada), où une autoroute majeure divise complètement leur domaine vital pendant une majeure partie de l'année. À l'aide d'une analyse de sélection d'habitat à fine échelle, nous avons testé si les wapitis optimisent les risques liés aux traversées d'autoroute et les récompenses alimentaires obtenues lorsqu'ils se déplacent entre des zones d'alimentation. Les récompenses ont été estimées à l'aide d'une carte dynamique de la biomasse végétale disponible pour les wapitis, et les risques ont été estimés en fonction des traversées de route et du trafic automobile rencontré. Nos résultats indiquent que les wapitis sélectionnaient généralement des zones d'alimentation avec une biomasse intermédiaire à élevée, et évitaient de traverser l'autoroute. La plupart du temps, les wapitis ajustaient leur sélection d'habitat en fonction des risques et des récompenses. Les wapitis sélectionnaient des zones d'alimentation avec une biomasse plus élevée lorsqu'ils traversaient l'autoroute, comparé à lorsqu'ils ne traversaient pas. Ils optimisaient également la biomasse végétale obtenue en fonction du nombre de traversées de l'autoroute effectuées durant une séquence de déplacement. Cependant, les wapitis optimisaient uniquement les récompenses alimentaires avec le trafic automobile durant les séquences de déplacement avec une seule traversée. Nous avons testé empiriquement un élément essentiel de l'écologie prédateur-proie, soit la capacité d'ajuster de façon dynamique un comportement en réponse à des niveaux variables de risques et de récompenses, dans le contexte de l'écologie routière. Notre étude permet d'illustrer comment les processus décisionnels considérant à la fois les risques, les récompenses et leur variation spatiotemporelle, permettent aux animaux de naviguer de façon optimale les routes très fréquentées. L'utilisation de concepts bien établis de l'écologie prédateur-proie aide à comprendre comment les animaux réagissent aux perturbations anthropiques, et contribue à anticiper la capacité d'adaptation des individus et des populations face à des environnements en transformation rapide.

Migrating mule deer compensate en route for phenological mismatches.

Billions of animals migrate to track seasonal pulses in resources. Optimally timing migration is a key strategy, yet the ability of animals to compensate for phenological mismatches en route is largely unknown. Using GPS movement data collected from 72 adult female deer over a 10-year duration, we study a population of mule deer (Odocoileus hemionus) in Wyoming that lack reliable cues on their desert winter range, causing them to start migration 70 days ahead to 52 days behind the wave of spring green-up. We show that individual deer arrive at their summer range within an average 6-day window by adjusting movement speed and stopover use. Late migrants move 2.5 times faster and spend 72% less time on stopovers than early migrants, which allows them to catch the green wave. Our findings suggest that ungulates, and potentially other migratory species, possess cognitive abilities to recognize where they are in space and time relative to key resources. Such behavioral capacity may allow migratory taxa to maintain foraging benefits amid rapidly changing phenology.

Defining null expectations for animal site fidelity.

Site fidelity-the tendency to return to previously visited locations-is widespread across taxa. Returns may be driven by several mechanisms, including memory, habitat selection, or chance; however, pattern-based definitions group different generating mechanisms under the same label of 'site fidelity', often assuming memory as the main driver. We propose an operational definition of site fidelity as patterns of return that deviate from a null expectation derived from a memory-free movement model. First, using agent-based simulations, we show that without memory, intrinsic movement characteristics and extrinsic landscape characteristics are key determinants of return patterns and that even random movements may generate substantial probabilities of return. Second, we illustrate how to implement our framework empirically to establish ecologically meaningful, system-specific null expectations for site fidelity. Our approach provides a conceptual and operational framework to test hypotheses on site fidelity across systems and scales.

Multi-mode movement decisions across widely ranging behavioral processes.

Movement of organisms plays a fundamental role in the evolution and diversity of life. Animals typically move at an irregular pace over time and space, alternating among movement states. Understanding movement decisions and developing mechanistic models of animal distribution dynamics can thus be contingent to adequate discrimination of behavioral phases. Existing methods to disentangle movement states typically require a follow-up analysis to identify state-dependent drivers of animal movement, which overlooks statistical uncertainty that comes with the state delineation process. Here, we developed population-level, multi-state step selection functions (HMM-SSF) that can identify simultaneously the different behavioral bouts and the specific underlying behavior-habitat relationship. Using simulated data and relocation data from mule deer (Odocoileus hemionus), plains bison (Bison bison bison) and plains zebra (Equus quagga), we illustrated the HMM-SSF robustness, versatility, and predictive ability for animals involved in distinct behavioral processes: foraging, migrating and avoiding a nearby predator. Individuals displayed different habitat selection pattern during the encamped and the travelling phase. Some landscape attributes switched from being selected to avoided, depending on the movement phase. We further showed that HMM-SSF can detect multi-modes of movement triggered by predators, with prey switching to the travelling phase when predators are in close vicinity. HMM-SSFs thus can be used to gain a mechanistic understanding of how animals use their environment in relation to the complex interplay between their needs to move, their knowledge of the environment and navigation capacity, their motion capacity and the external factors related to landscape heterogeneity.

Responses to natural gas development differ by season for two migratory ungulates.

While migrating, animals make directionally persistent movements and may only respond to human-induced rapid environmental change (HIREC), such as climate and land-use change, once a threshold of HIREC is surpassed. In contrast, animals on other seasonal ranges (e.g., winter range) make more localized and tortuous movements while foraging and may have the flexibility to adjust the location of their range and the intensity of use within it to minimize interactions with HIREC. Because of these seasonal differences in movement, animals on seasonal ranges should avoid areas that contain any level of HIREC, however, during migration, animals should use areas that contain low levels of HIREC, avoiding it only once a threshold of HIREC has been surpassed. We tested this hypothesis using a decade of GPS collar data collected from migratory mule deer (Odocoileus hemionus; n = 56 migration, 143 winter) and pronghorn (Antilocapra americana; n = 70 migration, 89 winter) that winter on and migrate through a natural gas field in western Wyoming. Using surface disturbance caused by well pads and roads as an index of HIREC, we evaluated behavioral responses across three spatial scales during winter and migration seasons. During migration, both species tolerated low levels of disturbance. Once a disturbance threshold was surpassed, however, they avoided HIREC. For mule deer, thresholds were consistently ~3%, whereas thresholds for pronghorn ranged from 1% to 9.25% surface disturbance. In contrast to migration, both species generally avoided all levels of HIREC while on winter range. Our study suggests that animal responses to HIREC are mediated by season-specific movement patterns. Our results provide further evidence of ungulates avoiding human disturbance on winter range and reveal disturbance thresholds that trigger mule deer and pronghorn responses during migration: information that managers can use to maintain the ecological function of migration routes and winter ranges.

Body size and digestive system shape resource selection by ungulates: A cross-taxa test of the forage maturation hypothesis.

The forage maturation hypothesis (FMH) states that energy intake for ungulates is maximised when forage biomass is at intermediate levels. Nevertheless, metabolic allometry and different digestive systems suggest that resource selection should vary across ungulate species. By combining GPS relocations with remotely sensed data on forage characteristics and surface water, we quantified the effect of body size and digestive system in determining movements of 30 populations of hindgut fermenters (equids) and ruminants across biomes. Selection for intermediate forage biomass was negatively related to body size, regardless of digestive system. Selection for proximity to surface water was stronger for equids relative to ruminants, regardless of body size. To be more generalisable, we suggest that the FMH explicitly incorporate contingencies in body size and digestive system, with small-bodied ruminants selecting more strongly for potential energy intake, and hindgut fermenters selecting more strongly for surface water.

Elk migration influences the risk of disease spillover in the Greater Yellowstone Ecosystem.

Wildlife migrations provide important ecosystem services, but they are declining. Within the Greater Yellowstone Ecosystem (GYE), some elk Cervus canadensis herds are losing migratory tendencies, which may increase spatiotemporal overlap between elk and livestock (domestic bison Bison bison and cattle Bos taurus), potentially exacerbating pathogen transmission risk. We combined disease, movement, demographic and environmental data from eight elk herds in the GYE to examine the differential risk of brucellosis transmission (through aborted foetuses) from migrant and resident elk to livestock. For both migrants and residents, we found that transmission risk from elk to livestock occurred almost exclusively on private ranchlands as opposed to state or federal grazing allotments. Weather variability affected the estimated distribution of spillover risk from migrant elk to livestock, with a 7%-12% increase in migrant abortions on private ranchlands during years with heavier snowfall. In contrast, weather variability did not affect spillover risk from resident elk. Migrant elk were responsible for the majority (68%) of disease spillover risk to livestock because they occurred in greater numbers than resident elk. On a per-capita basis, however, our analyses suggested that resident elk disproportionately contributed to spillover risk. In five of seven herds, we estimated that the per-capita spillover risk was greater from residents than from migrants. Averaged across herds, an individual resident elk was 23% more likely than an individual migrant elk to abort on private ranchlands. Our results demonstrate links between migration behaviour, spillover risk and environmental variability, and highlight the utility of integrating models of pathogen transmission and host movement to generate new insights about the role of migration in disease spillover risk. Furthermore, they add to the accumulating body of evidence across taxa that suggests that migrants and residents should be considered separately during investigations of wildlife disease ecology. Finally, our findings have applied implications for elk and brucellosis in the GYE. They suggest that managers should prioritize actions that maintain spatial separation of elk and livestock on private ranchlands during years when snowpack persists into the risk period.

Drivers of site fidelity in ungulates.

While the tendency to return to previously visited locations-termed 'site fidelity'-is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals' recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity. We compared inter-year site fidelity in 669 individuals across eight ungulate species fitted with GPS collars and occupying a range of environmental conditions in North America and Africa. We used a distance-based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren-ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a 'win-stay, lose-switch' strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter-annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species-specific differences in attraction to known sites, contribute to variation in the expression of this behaviour. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change.

Emerging Perspectives on Resource Tracking and Animal Movement Ecology.

Resource tracking, where animals increase energy gain by moving to track phenological variation in resources across space, is emerging as a fundamental attribute of animal movement ecology. However, a theoretical framework to understand when and where resource tracking should occur, and how resource tracking should lead to emergent ecological patterns, is lacking. We present a framework that unites concepts from optimal foraging theory and landscape ecology, which can be used to generate and test predictions on how resource dynamics shape animal movement across taxa, systems, and scales. Consideration of the interplay between animal movement and resource dynamics not only advances ecological understanding but can also guide biodiversity conservation in an era of global change.

Wave-like Patterns of Plant Phenology Determine Ungulate Movement Tactics.

Animals exhibit a diversity of movement tactics [1]. Tracking resources that change across space and time is predicted to be a fundamental driver of animal movement [2]. For example, some migratory ungulates (i.e., hooved mammals) closely track the progression of highly nutritious plant green-up, a phenomenon called "green-wave surfing" [3-5]. Yet general principles describing how the dynamic nature of resources determine movement tactics are lacking [6]. We tested an emerging theory that predicts surfing and the existence of migratory behavior will be favored in environments where green-up is fleeting and moves sequentially across large landscapes (i.e., wave-like green-up) [7]. Landscapes exhibiting wave-like patterns of green-up facilitated surfing and explained the existence of migratory behavior across 61 populations of four ungulate species on two continents (n = 1,696 individuals). At the species level, foraging benefits were equivalent between tactics, suggesting that each movement tactic is fine-tuned to local patterns of plant phenology. For decades, ecologists have sought to understand how animals move to select habitat, commonly defining habitat as a set of static patches [8, 9]. Our findings indicate that animal movement tactics emerge as a function of the flux of resources across space and time, underscoring the need to redefine habitat to include its dynamic attributes. As global habitats continue to be modified by anthropogenic disturbance and climate change [10], our synthesis provides a generalizable framework to understand how animal movement will be influenced by altered patterns of resource phenology.

Drought reshuffles plant phenology and reduces the foraging benefit of green-wave surfing for a migratory ungulate.

To increase resource gain, many herbivores pace their migration with the flush of nutritious plant green-up that progresses across the landscape (termed "green-wave surfing"). Despite concerns about the effects of climate change on migratory species and the critical role of plant phenology in mediating the ability of ungulates to surf, little is known about how drought shapes the green wave and influences the foraging benefits of migration. With a 19 year dataset on drought and plant phenology across 99 unique migratory routes of mule deer (Odocoileus hemionus) in western Wyoming, United States, we show that drought shortened the duration of spring green-up by approximately twofold (2.5 weeks) and resulted in less sequential green-up along migratory routes. We investigated the possibility that some routes were buffered from the effects of drought (i.e., routes that maintained long green-up duration irrespective of drought intensity). We found no evidence of drought-buffered routes. Instead, routes with the longest green-up in non-drought years also were the most affected by drought. Despite phenological changes along the migratory route, mule deer closely followed drought-altered green waves during migration. Migrating deer did not experience a trophic mismatch with the green wave during drought. Instead, the shorter window of green-up caused by drought reduced the opportunity to accumulate forage resources during rapid spring migrations. Our work highlights the synchronization of phenological events as an important mechanism by which climate change can negatively affect migratory species by reducing the temporal availability of key food resources. For migratory herbivores, climate change poses a new and growing threat by altering resource phenology and diminishing the foraging benefit of migration.

Chronic wasting disease undermines efforts to control the spread of brucellosis in the Greater Yellowstone Ecosystem.

Wildlife diseases pose a substantial threat to the provisioning of ecosystem services. We use a novel modeling approach to study the potential loss of these services through the imminent introduction of chronic wasting disease (CWD) to elk populations in the Greater Yellowstone Ecosystem (GYE). A specific concern is that concentrating elk at feedgrounds may exacerbate the spread of CWD, whereas eliminating feedgrounds may increase the number of elk on private ranchlands and the transmission of a second disease, brucellosis, from elk to cattle. To evaluate the consequences of management strategies given the threat of two concurrent wildlife diseases, we develop a spatiotemporal bioeconomic model. GPS data from elk and landscape attributes are used to predict migratory behavior and population densities with and without supplementary feeding. We use a 4,800 km2 area around Pinedale, Wyoming containing four existing feedgrounds as a case study. For this area, we simulate welfare estimates under a variety of management strategies. Our results indicate that continuing to feed elk could result in substantial welfare losses for the case-study region. Therefore, to maximize the present value of economic net benefits generated by the local elk population upon CWD's arrival in the region, wildlife managers may wish to consider discontinuing elk feedgrounds while simultaneously developing new methods to mitigate the financial impact to ranchers of possible brucellosis transmission to livestock. More generally, our methods can be used to weigh the costs and benefits of human-wildlife interactions in the presence of multiple disease risks.

Migrating bison engineer the green wave.

Newly emerging plants provide the best forage for herbivores. To exploit this fleeting resource, migrating herbivores align their movements to surf the wave of spring green-up. With new technology to track migrating animals, the Green Wave Hypothesis has steadily gained empirical support across a diversity of migratory taxa. This hypothesis assumes the green wave is controlled by variation in climate, weather, and topography, and its progression dictates the timing, pace, and extent of migrations. However, aggregate grazers that are also capable of engineering grassland ecosystems make some of the world's most impressive migrations, and it is unclear how the green wave determines their movements. Here we show that Yellowstone's bison (Bison bison) do not choreograph their migratory movements to the wave of spring green-up. Instead, bison modify the green wave as they migrate and graze. While most bison surfed during early spring, they eventually slowed and let the green wave pass them by. However, small-scale experiments indicated that feedback from grazing sustained forage quality. Most importantly, a 6-fold decadal shift in bison density revealed that intense grazing caused grasslands to green up faster, more intensely, and for a longer duration. Our finding broadens our understanding of the ways in which animal movements underpin the foraging benefit of migration. The widely accepted Green Wave Hypothesis needs to be revised to include large aggregate grazers that not only move to find forage, but also engineer plant phenology through grazing, thereby shaping their own migratory movements.

Managing Genetic Diversity and Extinction Risk for a Rare Plains Bison (Bison bison bison) Population.

Unfenced plains bison are rare and only occur in a small number of locations throughout Canada and the United States. We examined management guidelines for maintenance of genetic health and population persistence for a small and isolated population of plains bison that occupy the interface between a protected national park and private agricultural lands. To address genetic health concerns, we measured genetic diversity relative to other populations and assessed the potential effects of genetic augmentation. We then used individual-based population viability analyses (PVA) to determine the minimum abundance likely to prevent genetic diversity declines. We assessed this minimum relative to a proposed maximum social carrying capacity related to bison use of human agricultural lands. We also used the PVA to assess the probability of population persistence given the limiting factors of predation, hunting, and disease. Our results indicate that genetic augmentation will likely be required to achieve genetic diversity similar to that of other plains bison populations. We also found that a minimum population of 420 bison yields low probability of additional genetic loss while staying within society-based maxima. Population estimates based on aerial surveys indicated that the population has been below this minimum since 2007. Our PVA simulations indicate that current hunting practices will result in undesirable levels of population extinction risk and further declines in genetic variability. Our study demonstrates that PVA can be used to evaluate potential management scenarios as they relate to long-term genetic conservation and population persistence for rare species.

Spatial memory shapes migration and its benefits: evidence from a large herbivore.

From fine-scale foraging to broad-scale migration, animal movement is shaped by the distribution of resources. There is mounting evidence, however, that learning and memory also guide movement. Although migratory mammals commonly track resource waves, how resource tracking and memory guide long-distance migration has not been reconciled. We examined these hypotheses using movement data from four populations of migratory mule deer (n = 91). Spatial memory had an extraordinary influence on migration, affecting movement 2-28 times more strongly than tracking spring green-up or autumn snow depth. Importantly, with only an ability to track resources, simulated deer were unable to recreate empirical migratory routes. In contrast, simulated deer with memory of empirical routes used those routes and obtained higher foraging benefits. For migratory terrestrial mammals, spatial memory provides knowledge of where seasonal ranges and migratory routes exist, whereas resource tracking determines when to beneficially move within those areas.

Determinants of elephant foraging behaviour in a coupled human-natural system: Is brown the new green?

Crop raiding by wildlife poses major threats to both wildlife conservation and human well-being in agroecosystems worldwide. These threats are particularly acute in many parts of Africa, where crop raiders include globally threatened megafauna such as elephants, and where smallholder agriculture is a primary source of human livelihood. One framework for understanding herbivore feeding behaviour, the forage-maturation hypothesis, predicts that herbivores should align their movements with intermediate forage biomass (i.e., peak green-up); this phenomenon is known as "surfing the green wave." Crop-raiding elephants, however, often consume not just foliage, but also fruits and tubers (e.g., maize and potatoes), which generally mature after seasonal peaks in photosynthetic activity. Thus, although elephants have been reported to surf the green wave in natural habitats, they may utilize a different strategy in cultivated landscapes by selecting crops that are "browning down." We sought to understand the factors that underpin movement of elephants into agricultural landscapes. In Mozambique's Gorongosa National Park, we used movement data from GPS-collared elephants, together with precipitation records, remotely sensed estimates of landscape greenness (NDVI), DNA-based diet analysis, measurements of plant nutritional quality and survey-based metrics of crop availability to understand spatiotemporal variation in elephant crop-raiding behaviour. Elephants tracked peak NDVI while foraging inside the Park. During the dry season, however, when NDVI within the Park declined and availability of mature crops was high, crop raiding increased dramatically, and elephants consistently selected crop plants that were browning down while foraging in cultivated landscapes. Crops contained significantly higher digestible energy than wild food plants, but comparable (and sometimes lower) levels of digestible protein, suggesting that this foraging strategy maximized energy rather than protein intake. Our study is the first to combine GPS tracking data with high-resolution diet analysis and community-based reporting of crop availability to reveal fine-scale plasticity in foraging behaviour of elephants at the human-wildlife interface. Our results extend the forage-maturation hypothesis by showing that elephants surf waves of plant brown-down in cultivated landscapes. These findings can aid efforts to reduce human-elephant conflict by enabling wildlife managers to prioritize mitigation actions in time and space with limited resources.