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.

Wherever I may roam-Human activity alters movements of red deer (Cervus elaphus) and elk (Cervus canadensis) across two continents.

Human activity and associated landscape modifications alter the movements of animals with consequences for populations and ecosystems worldwide. Species performing long-distance movements are thought to be particularly sensitive to human impact. Despite the increasing anthropogenic pressure, it remains challenging to understand and predict animals' responses to human activity. Here we address this knowledge gap using 1206 Global Positioning System movement trajectories of 815 individuals from 14 red deer (Cervus elaphus) and 14 elk (Cervus canadensis) populations spanning wide environmental gradients, namely the latitudinal range from the Alps to Scandinavia in Europe, and the Greater Yellowstone Ecosystem in North America. We measured individual-level movements relative to the environmental context, or movement expression, using the standardized metric Intensity of Use, reflecting both the directionality and extent of movements. We expected movement expression to be affected by resource (Normalized Difference Vegetation Index, NDVI) predictability and topography, but those factors to be superseded by human impact. Red deer and elk movement expression varied along a continuum, from highly segmented trajectories over relatively small areas (high intensity of use), to directed transitions through restricted corridors (low intensity of use). Human activity (Human Footprint Index, HFI) was the strongest driver of movement expression, with a steep increase in Intensity of Use as HFI increased, but only until a threshold was reached. After exceeding this level of impact, the Intensity of Use remained unchanged. These results indicate the overall sensitivity of Cervus movement expression to human activity and suggest a limitation of plastic responses under high human pressure, despite the species also occurring in human-dominated landscapes. Our work represents the first comparison of metric-based movement expression across widely distributed populations of a deer genus, contributing to the understanding and prediction of animals' responses to human activity.

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.

Case Report: When Two Is Worse Than One-Stereo Imbalance in a Case of Wavefront-guided Scleral Lenses.

SIGNIFICANCE: Wavefront-guided scleral lenses (WGSLs) reduce visually debilitating residual higher-order aberrations. Although reduced higher-order aberrations lead to improvement in monocular high-contrast visual acuity (VA), the success of the lenses in everyday life depends on additional factors such as retinal contrast, binocular balance, and stereoacuity. PURPOSE: This report describes a case where WGSLs provided improved monocular vision compared with scleral lenses (SLs) but reduced binocularity and stereoacuity. CASE REPORT: A 48-year-old woman with moderate keratoconus right eye (OD) and severe left eye (OS) was fitted with SLs and WGSLs. Visual acuity with best SLs was 20/20 -2 OD and 20/25 -2 OS. Residual higher-order root-mean-square (HORMS) wavefront error (6 mm pupil) was 0.56 µm OD and 1.38 µm OS. Visual acuity with WGSLs was 20/16 -2 OD and 20/25 +2 OS, and residual HORMS was 0.41 µm OD and 0.98 µm OS. Monocularly, WGSLs were reported to provide better VA. However, binocularly, the patient reported an "imbalanced feeling" and preferred the SLs over WGSLs. Binocular VA at distance was 20/25 with SLs and 20/25 -2 with WGSL. To investigate, the Worth Four-Dot test was performed, and the outcomes reported fusion with SLs but suppression OS at distance with WGSLs. Stereoacuity was 160 arc seconds at near and 120 arc seconds at distance with SLs and 400 arc seconds at near and >1200 arc seconds at distance with WGSLs. Dichoptic contrast balancing showed a balance point of 0.48 with SLs and 0.17 with WGSLs, indicating a strong preference toward OD. Simulation of the patient's retinal image revealed a greater difference in image contrast between the two eyes with WGSLs. CONCLUSIONS: Wavefront-guided scleral lenses reduced HORMS and improved VA compared with SLs. However, in this case, it inadvertently caused binocular imbalance. As WGSLs become more widely available, future work should include methods to optimize binocular balance to maximize overall patient satisfaction.

Wind-energy development alters pronghorn migration at multiple scales.

Migration is a critical behavioral strategy necessary for population persistence and ecosystem functioning, but migration routes have been increasingly disrupted by anthropogenic activities, including energy development. Wind energy is the world's fastest growing source of electricity and represents an important alternative to hydrocarbon extraction, but its effects on migratory species beyond birds and bats are not well understood. We evaluated the effects of wind-energy development on pronghorn migration, including behavior and habitat selection, to assess potential effects on connectivity and other functional benefits including stopovers. We monitored GPS-collared female pronghorn from 2010 to 2012 and 2018 to 2020 in south-central Wyoming, USA, an area with multiple wind-energy facilities in various stages of development and operation. Across all time periods, we collected 286 migration sequences from 117 individuals, including 121 spring migrations, 123 fall migrations, and 42 facultative winter migrations. While individuals continued to migrate through wind-energy facilities, pronghorn made important behavioral adjustments relative to turbines during migration. These included avoiding turbines when selecting stopover sites in spring and winter, selecting areas farther from turbines at a small scale in spring and winter, moving more quickly near turbines in spring (although pronghorn moved more slowly near turbines in the fall), and reducing fidelity to migration routes relative to wind turbines under construction in both spring and fall. For example, an increase in distance to turbine from 0 to 1 km translated to a 33% and 300% increase in the relative probability of selection for stopover sites in spring and winter, respectively. The behavioral adjustments pronghorn made relative to wind turbines could reduce the functional benefits of their migration, such as foraging success or the availability of specific routes, over the long term.

Industrial energy development decouples ungulate migration from the green wave.

The ability to freely move across the landscape to track the emergence of nutritious spring green-up (termed 'green-wave surfing') is key to the foraging strategy of migratory ungulates. Across the vast landscapes traversed by many migratory herds, habitats are being altered by development with unknown consequences for surfing. Using a unique long-term tracking dataset, we found that when energy development occurs within mule deer (Odocoileus hemionus) migration corridors, migrating animals become decoupled from the green wave. During the early phases of a coalbed natural gas development, deer synchronized their movements with peak green-up. But faced with increasing disturbance as development expanded, deer altered their movements by holding up at the edge of the gas field and letting the green wave pass them by. Development often modified only a small portion of the migration corridor but had far-reaching effects on behaviour before and after migrating deer encountered it, thus reducing surfing along the entire route by 38.65% over the 14-year study period. Our study suggests that industrial development within migratory corridors can change the behaviour of migrating ungulates and diminish the benefits of migration. Such disruptions to migratory behaviour present a common mechanism whereby corridors become unprofitable and could ultimately be lost on highly developed landscapes.

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.

Migration distance and maternal resource allocation determine timing of birth in a large herbivore.

Birth timing is a key life-history characteristic that influences fitness and population performance. For migratory animals, however, appropriately timing birth on one seasonal range may be constrained by events occurring during other parts of the migratory cycle. We investigated how the use of capital and income resources may facilitate flexibility in reproductive phenology of migratory mule deer in western Wyoming, USA, over a 5-yr period (2015-2019). Specifically, we examined how seasonal interactions affected three interrelated life-history characteristics: fetal development, birth mass, and birth timing. Females in good nutritional condition at the onset of winter and those that migrated short distances had more developed fetuses (measured as fetal eye diameter in March). Variation in parturition date was explained largely by fetal development; however, there were up to 16 d of plasticity in expected birth date. Plasticity in expected birth date was shaped by income resources in the form of exposure to spring green-up. Although individuals that experienced greater exposure to spring green-up were able to advance expected birth date, being born early or late with respect to fetal development had no effect on birth mass of offspring. Furthermore, we investigated the trade-offs migrating mule deer face by evaluating support for existing theory that predicts that births should be matched to local peaks in resource availability at the birth site. In contrast to this prediction, only long-distance migrants that paced migration with the flush of spring green-up, giving birth shortly after ending migration, were able to match birth with spring green-up. Shorter-distance migrants completed migration sooner and gave birth earlier, seemingly trading off more time for offspring to grow and develop over greater access to resources. Thus, movement tactic had profound downstream effects on birth timing. These findings highlight a need to reconsider classical theory on optimal birth timing, which has focused solely on conditions at the birth site.

Life-history theory provides a framework for detecting resource limitation: a test of the Nutritional Buffer Hypothesis.

For ungulates and other long-lived species, life-history theory predicts that nutritional reserves are allocated to reproduction in a state-dependent manner because survival is highly conserved. Further, as per capita food abundance and nutritional reserves decline (i.e., density dependence intensifies), reproduction and recruitment become increasingly sensitive to weather. Thus, the degree to which weather influences vital rates should be associated with proximity to nutritional carrying capacity-a notion that we refer to as the Nutritional Buffer Hypothesis. We tested the Nutritional Buffer Hypothesis using six moose (Alces alces) populations that varied in calf recruitment (33-69 calves/100 cows). We predicted that populations with high calf recruitment were nutritionally buffered against the effects of unfavorable weather, and thus were below nutritional carrying capacity. We applied a suite of tools to quantify habitat and nutritional condition of each population and found that increased browse condition, forage quality, and body fat were associated with increased pregnancy and calf recruitment, thereby providing multiple lines of evidence that declines in calf recruitment were underpinned by resource limitation. From 2001 to 2015, recruitment was more sensitive to interannual variation in weather (e.g., winter severity, drought) and plant phenology (e.g., duration of spring) for populations with reduced browse condition, forage quality, and body fat, suggesting these populations lacked the nutritional reserves necessary to buffer demographic performance against the effects of unfavorable weather. Further, average within-population calf recruitment was determined by regional climatic variation, suggesting that the pattern of reduced recruitment near the southern range boundary of moose stems from an interaction between climate and resource limitation. When coupled with information on habitat, nutrition, weather, and climate, life-history theory provides a framework to estimate nutritional limitation, proximity to nutritional carrying capacity, and impacts of climate change for ungulates.

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.

Quantifying the Optical and Physical Consequences of Daily Cleaning on Conventional and Wavefront-guided Scleral Lenses.

SIGNIFICANCE: An equivalent 12 months of cleaning did not induce significant changes in the optical aberrations or base curves of scleral lenses. PURPOSE: This study aimed to test whether an equivalent of 12 months of manual cleaning alters the optical and physical properties of conventional and wavefront-guided scleral lenses. METHODS: Twelve scleral lenses (four repeats of three designs, termed A, B, and C) were manufactured in Boston XO material: design A, -5.00 D defocus; design B, -5.00 D defocus with -0.153-µm vertical coma; and design C, -5.00 D defocus with a full custom wavefront-guided correction (second to fifth Zernike radial orders) of an eye with severe keratoconus. One lens of each design group served as a control and was not cleaned. To simulate a year of cleaning, seven individuals cleaned nine lenses (three from each group) twice a day for 27 days using the palm technique and commercially available cleaners, resulting in 378 cleanings of each lens. Lens aberrations were optically profiled and base curve radii were measured at baseline and after every 42nd cleaning. Differences in higher-order root mean square (HORMS) wavefront error and base curve radii associated with cleaning were compared with clinical benchmarks and using sign tests. RESULTS: For the experimental lenses, median change in Seidel spherical dioptric power was +0.01 D (maximum, +0.025 D). Median change in HORMS wavefront error was 0.013 µm (maximum, 0.019 µm). All lenses exhibited HORMS changes less than one-eighth equivalent diopters (P = .002). Median percentage change in HORMS wavefront error in the three wavefront-guided lenses was 0.96% (maximum, 1.25%). Median change in base curve radii was 0.00 mm, with all lenses exhibiting changes (P = .002), less than the American National Standards Institute tolerance of 0.05 mm. CONCLUSIONS: Cleaning over an equivalent 12-month period did not induce clinically significant changes in the optical or base curve properties of conventional or wavefront-guided scleral lenses.

Case Report: What Are We Doing for Our "20/20 Unhappy" Scleral Lens Patients?

SIGNIFICANCE: Scleral lenses (SLs) partially mask higher-order aberrations (HOAs) in highly aberrated eyes. Although visual acuity (VA) may show satisfactory quantitative clinical outcomes during SL wear, residual (uncorrected) HOAs can leave subjective visual quality goals unmet. PURPOSE: The purpose of this study was to report a case where a "20/20 unhappy" patient with SLs was able to meet visual goals with wavefront-guided SLs. CASE REPORT: A 40-year-old male with bilateral keratoconus, whose Snellen VA with SLs was 20/20 right eye (OD) 20/16 left eye (OS), reported halos and glare at night and perceptual smearing. When viewing a point of light, a "Ferris wheel" shadowing was observed OD and a U-shaped shadowing OS. Residual higher-order root mean square wavefront error was 0.49 µm OD and 0.39 µm OS; visual image quality measured by visual Strehl ratio was 0.067 OD and 0.092 OS (pupil size, 4.00 mm). Wavefront-guided SLs reduced residual higher-order root mean square to 0.19 µm OD and 0.25 µm OS, VA improved to 20/10 OD and 20/13 OS, and visual Strehl improved to 0.150 OD and 0.121 OS. The patient reported reduced smearing, shadowing, and night vision concerns, meeting his visual expectations and goals. CONCLUSIONS: Wavefront sensing quantifies both lower and HOAs, which can cause visual dissatisfaction in individuals with highly aberrated eyes, despite sometimes reaching typical levels of VA. As wavefront-guided SLs targeting these residual aberrations to improve visual image quality become more available, they should be considered for 20/20 unhappy patients when conventional clinical options are unsatisfactory.

The Impact of Misaligned Wavefront-guided Correction in a Scleral Lens for the Highly Aberrated Eye.

SIGNIFICANCE: To achieve maximum visual benefit, wavefront-guided scleral lens corrections (WGCs) are aligned with the underlying wavefront error of each individual eye. This requirement adds complexity to the fitting process. With a view toward simplification in lens fitting, this study quantified the consequences of placing WGCs at two pre-defined locations. PURPOSE: This study aimed to quantify performance reduction accompanying the placement of the WGC at two locations: (1) the average decentered location (ADL; average decentration observed across individuals wearing scleral lenses) and (2) the geometric center (GC) of the lens. METHODS: Deidentified residual aberration and lens translation data from 36 conventional scleral lens-wearing eyes with corneal ectasia were used to simulate WGC correction in silico. The WGCs were decentered from the eye-specific pupil position to both the ADL and GC locations. The impact of these misalignments was assessed in terms of change (from the aligned, eye-specific pupil position) in higher-order root mean square (HORMS) wavefront error, change in log of the visual Strehl ratio (logVSX), and predicted change in logMAR visual acuity (VA). RESULTS: As expected, HORMS increased, logVSX decreased, and predicted VA was poorer at both ADL and GC compared with the aligned condition (P < .001). Thirty-four of 36 eyes had greater residual HORMS, and 33 of 36 eyes had worse logVSX values at the GC than at the ADL. In clinical terms, 19 of 36 eyes at the ADL and 35 of 36 eyes at the GC had a predicted loss in VA of three letters or greater. CONCLUSIONS: The placement of the WGC at either ADL or GC is predicted to lead to a noticeable reduction in VA for more than half of the eyes studied, suggesting the simplification of the fitting process is not worth the cost in performance.

A test of the Niche Variation Hypothesis in a ruminant herbivore.

Despite the shared prediction that the width of a population's dietary niche expands as food becomes limiting, the Niche Variation Hypothesis (NVH) and Optimal Foraging Theory (OFT) offer contrasting views about how individuals alter diet selection when food is limited. Classical OFT predicts that dietary preferences do not change as food becomes limiting, so individuals expand their diets as they compensate for a lack of preferred foods. In contrast, the NVH predicts that among-individual variation in cognition, physiology or morphology create functional trade-offs in foraging efficiency, thereby causing individuals to specialize on different subsets of food as food becomes limiting. To evaluate (a) the predictions of the NVH and OFT and (b) evidence for physiological and cognitive-based functional trade-offs, we used DNA microsatellites and metabarcoding to quantify the diet, microbiome and genetic relatedness (a proxy for social learning) of 218 moose Alces alces across six populations that varied in their degree of food limitation. Consistent with both the NVH and OFT, dietary niche breadth increased with food limitation. Increased diet breadth of individuals-rather than increased diet specialization-was strongly correlated with both food limitation and dietary niche breadth of populations, indicating that moose foraged in accordance with OFT. Diets were not constrained by inheritance of the microbiome or inheritance of diet selection, offering support for the little-tested hypothesis that functional trade-offs in food use (or lack thereof) determine whether populations adhere to the predictions of the NVH or OFT. Our results indicate that both the absence of strong functional trade-offs and the digestive physiology of ruminants provide contexts under which populations should forage in accordance with OFT rather than the NVH. Also, because dietary niche width increased with increased food limitation, OFT and the NVH provide theoretical support for the notion that plant-herbivore interaction networks are plastic rather than static, which has important implications for understanding interspecific niche partitioning. Lastly, because population-level dietary niche breadth and calf recruitment are correlated, and because calf recruitment can be a proxy for food limitation, our work demonstrates how diet data can be employed to understand a populations' proximity to carrying capacity.

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.