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Seasonal migrations and other movements

In the past 25 years new information has been obtained on the migrations and movements of mountain sheep (bighorn [Ovis canadensis], thinhorn [Ovis dalli]). This chapter provides a comprehensive overview of mountain sheep migration and other movements across their broad distribution in western North America. Across the range of mountain sheep, migrations and other seasonal movements define a complex movement portfolio that relates to and supports all aspects of mountain sheep ecology. Bighorn and thinhorn species have examples of migratory behaviors that span the continuum between annual residents and long-distance migrants. Migratory behaviors can be characterized as low- or high-elevation residents; elevational migrations and other variants; and geographic migrants. Native populations that have been extant on the landscape without notable human intervention have greater migratory propensity and more diverse migratory portfolios due to the maintenance of migration through cultural learning and social transmission. Restored and augmented populations, where the population-level knowledge of migration has been lost or greatly reduced, are largely nonmigratory, although translocations show some ability to restore short-distance elevational migrations. Seasonal spring and fall migrations are less common in desert bighorn sheep (O. canadensis spp.) or bighorn sheep living in canyon or prairie breaks landscapes. Mineral lick visitation is important and common across the range of mountain sheep. Managing for migratory diversity can help to sustain migratory behavior in the face of climate change and other anthropogenic pressures, which can limit landscape connectivity between seasonal ranges or alter the spatiotemporal dynamics of regional phenology with cascading effects to other biotic and abiotic interactions such as the need to balance forage and predation risk or the spatial refugia with increased temperatures.

Book chapter

Behavioral flexibility as a mechanism for coping with climate change

Of the primary responses to contemporary climate change – “move, adapt, acclimate, or die” – that are available to organisms, “acclimate” may be effectively achieved through behavioral modification. Behavioral flexibility allows animals to rapidly cope with changing environmental conditions, and behavior represents an important component of a species’ adaptive capacity in the face of climate change. However, there is currently a lack of knowledge about the limits or constraints on behavioral responses to changing conditions. Here, we characterize the contexts in which organisms respond to climate variability through behavior. First, we quantify patterns in behavioral responses across taxa with respect to timescales, climatic stimuli, life-history traits, and ecology. Next, we identify existing knowledge gaps, research biases, and other challenges. Finally, we discuss how conservation practitioners and resource managers can incorporate an improved understanding of behavioral flexibility into natural resource management and policy decisions.

Frontiers in Ecology and the Environment

Evidence for density dependence in foraging and migratory behavior of a subtropical nearshore seabird

Density-dependent competition for food resources influences both foraging ecology and reproduction in a variety of animals. The relationship between colony size, local prey depletion, and reproductive output in colonial central-place foragers has been extensively studied in seabirds; however, most studies have focused on effects of intraspecific competition during the breeding season, while little is known about whether density-dependent resource depletion influences individual migratory behavior outside the breeding season. Using breeding colony size as a surrogate for intraspecific resource competition, we tested for effects of colony size on breeding home range, nestling health, and migratory patterns of a nearshore colonial seabird, the brown pelican ( Pelecanus occidentalis ), originating from seven breeding colonies of varying sizes in the subtropical northern Gulf of Mexico. We found evidence for density-dependent effects on foraging behavior during the breeding season, as individual foraging areas increased linearly with the number of breeding pairs per colony. Contrary to our predictions, however, nestlings from more numerous colonies with larger foraging ranges did not experience either decreased condition or increased stress. During nonbreeding, individuals from larger colonies were more likely to migrate, and traveled longer distances, than individuals from smaller colonies, indicating that the influence of density-dependent effects on distribution persists into the nonbreeding period. We also found significant effects of individual physical condition, particularly body size, on migratory behavior, which in combination with colony size suggesting that dominant individuals remain closer to breeding sites during winter. We conclude that density-dependent competition may be an important driver of both the extent of foraging ranges and the degree of migration exhibited by brown pelicans. However, the effects of density-dependent competition on breeding success and population regulation remain uncertain in this system.

Gulf of Mexico

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.

Wyoming

Behavioral responses of Silver Carp to underwater acoustic deterrent sounds

Objective Invasive carps continue to spread across the Mississippi River basin, posing significant ecological risk. Identifying technologies to slow their dispersal is critical. The use of sound has been proposed as a method to modify the behavior of Silver Carp Hypophthalmichthys molitrix , offering a nonstructural deterrent strategy. Methods Silver Carp implanted with acoustic transmitters were released into earthen ponds equipped with telemetry arrays. The fish were exposed to a 30-min playback of three underwater sounds (chirp saw, chirp square, and 100-hp boat motor). Movement trajectories were analyzed using a two-state hidden Markov model to estimate the effects of environmental and experimental variables on fish behavior. Results The results of the hidden Markov model supported two behavioral states. State 1 was characterized by longer step lengths (distance between positions) and greater directional persistence in turning angle (change in direction between two intervals), indicative of heighted activity. State 2 was defined by shorter step lengths and less directional persistence, suggesting reduced activity. Silver Carp that were exposed to the chirp square sound had an increased likelihood of entering state 1, whereas the 100-hp boat motor sound promoted transitions to state 2. Conclusions Underwater sounds distinctly influenced the movement of Silver Carp in earthen ponds. The chirp square sound elicited heightened activity levels, demonstrating potential for use in acoustic deterrent applications. However, the response of Silver Carp to these sounds may be influenced by the size of the study environment or the absence of natural drivers of fish behavior, such as food or reproduction. This study contributes to the development of nonstructural, species-specific deterrent systems by identifying sounds that influence the behavior of invasive carps. The application of sound-based methods may play a critical role in integrated pest management strategies for invasive carps, potentially limiting their spread while minimizing effects on native species.

Transactions of the American Fisheries Society

Mapping behavioral landscapes for animal movement: a finite mixture modeling approach

Because of its role in many ecological processes, movement of animals in response to landscape features is an important subject in ecology and conservation biology. In this paper, we develop models of animal movement in relation to objects or fields in a landscape. We take a finite mixture modeling approach in which the component densities are conceptually related to different choices for movement in response to a landscape feature, and the mixing proportions are related to the probability of selecting each response as a function of one or more covariates. We combine particle swarm optimization and an Expectation-Maximization (EM) algorithm to obtain maximum likelihood estimates of the model parameters. We use this approach to analyze data for movement of three bobcats in relation to urban areas in southern California, USA. A behavioral interpretation of the models revealed similarities and differences in bobcat movement response to urbanization. All three bobcats avoided urbanization by moving either parallel to urban boundaries or toward less urban areas as the proportion of urban land cover in the surrounding area increased. However, one bobcat, a male with a dispersal-like large-scale movement pattern, avoided urbanization at lower densities and responded strictly by moving parallel to the urban edge. The other two bobcats, which were both residents and occupied similar geographic areas, avoided urban areas using a combination of movements parallel to the urban edge and movement toward areas of less urbanization. However, the resident female appeared to exhibit greater repulsion at lower levels of urbanization than the resident male, consistent with empirical observations of bobcats in southern California. Using the parameterized finite mixture models, we mapped behavioral states to geographic space, creating a representation of a behavioral landscape. This approach can provide guidance for conservation planning based on analysis of animal movement data using statistical models, thereby linking connectivity evaluations to empirical data.

Ecological Applications

A hierarchical model for estimating the spatial distribution and abundance of animals detected by continuous-time recorders

Motivation Several spatial capture-recapture (SCR) models have been developed to estimate animal abundance by analyzing the detections of individuals in a spatial array of traps. Most of these models do not use the actual dates and times of detection, even though this information is readily available when using continuous-time recorders, such as microphones or motion-activated cameras. Instead most SCR models either partition the period of trap operation into a set of subjectively chosen discrete intervals and ignore multiple detections of the same individual within each interval, or they simply use the frequency of detections during the period of trap operation and ignore the observed times of detection. Both practices make inefficient use of potentially important information in the data. Model and data analysis We developed a hierarchical SCR model to estimate the spatial distribution and abundance of animals detected with continuous-time recorders. Our model includes two kinds of point processes: a spatial process to specify the distribution of latent activity centers of individuals within the region of sampling and a temporal process to specify temporal patterns in the detections of individuals. We illustrated this SCR model by analyzing spatial and temporal patterns evident in the camera-trap detections of tigers living in and around the Nagarahole Tiger Reserve in India. We also conducted a simulation study to examine the performance of our model when analyzing data sets of greater complexity than the tiger data. Benefits Our approach provides three important benefits: First, it exploits all of the information in SCR data obtained using continuous-time recorders. Second, it is sufficiently versatile to allow the effects of both space use and behavior of animals to be specified as functions of covariates that vary over space and time. Third, it allows both the spatial distribution and abundance of individuals to be estimated, effectively providing a species distribution model, even in cases where spatial covariates of abundance are unknown or unavailable. We illustrated these benefits in the analysis of our data, which allowed us to quantify differences between nocturnal and diurnal activities of tigers and to estimate their spatial distribution and abundance across the study area. Our continuous-time SCR model allows an analyst to specify many of the ecological processes thought to be involved in the distribution, movement, and behavior of animals detected in a spatial trapping array of continuous-time recorders. We plan to extend this model to estimate the population dynamics of animals detected during multiple years of SCR surveys.

Western Ghats, Nagarahole Tiger Reserve

Analysis of changes in water-level dynamics at selected sites in the Florida Everglades

The historical modification and regulation of the hydrologic patterns in the Florida Everglades have resulted in changes in the ecosystem of South Florida and the Florida Everglades. Since the 1970s, substantial focus has been given to the restoration of the Everglades ecosystem. The U.S. Geological Survey through its Greater Everglades Priority Ecosystem Science and National Water-Quality Assessment Programs has been providing scientific information to resource managers to assist in the Everglades restoration efforts. The current investigation included development of a simple method to identify and quantify changes in historical hydrologic behavior within the Everglades that could be used by researchers to identify responses of ecological communities to those changes. Such information then could be used by resource managers to develop appropriate water-management practices within the Everglades to promote restoration. The identification of changes in historical hydrologic behavior within the Everglades was accomplished by analyzing historical time-series water-level data from selected gages in the Everglades using (1) break-point analysis of cumulative Z-scores to identify hydrologic changes and (2) cumulative water-level frequency distribution curves to evaluate the magnitude of those changes. This analytical technique was applied to six long-term water-level gages in the Florida Everglades. The break-point analysis for the concurrent period of record (1978–2011) identified 10 common periods of changes in hydrologic behavior at the selected gages. The water-level responses at each gage for the 10 periods displayed similarity in fluctuation patterns, highlighting the interconnectedness of the Florida Everglades hydrologic system. While the patterns were similar, the analysis also showed that larger fluctuations in water levels between periods occurred in Water Conservation Areas 2 and 3 in contrast to those in Water Conservation Area 1 and the Everglades National Park. Results from the analysis indicate that the cumulative Z-score curve, in conjunction with cumulative water-level frequency distribution curves, can be a useful tool in identifying and quantifying changes in historical hydrologic behavior within the Everglades. In addition to the analysis, a spreadsheet application was developed to assist in applying these techniques to time-series water-level data at gages within the Everglades and is included with this report.

Florida

Cumulative effects of multiple stressors on marine mammals: Elephant seals as a model system

Noise exposure is a potential stressor for free-ranging marine mammals and is often studied in the absence of other environmental factors. Here, a multi-investigator, interdisciplinary effort was undertaken to examine the response of elephant seals to multiple stressors. An integrated physiological and ecological approach was taken, including immunology, stress physiology, toxicology, animal behavior, population biology, and life history theory, to examine the cumulative effects of exposure to multiple stressors in elephant seals. While we measured the response of individual animals, a population response can be predicted by incorporating these results into the long-term data on elephant seal demographics.

Book chapter

Velocity-based movement modeling for individual and population level inference

Understanding animal movement and resource selection provides important information about the ecology of the animal, but an animal's movement and behavior are not typically constant in time. We present a velocity-based approach for modeling animal movement in space and time that allows for temporal heterogeneity in an animal's response to the environment, allows for temporal irregularity in telemetry data, and accounts for the uncertainty in the location information. Population-level inference on movement patterns and resource selection can then be made through cluster analysis of the parameters related to movement and behavior. We illustrate this approach through a study of northern fur seal ( Callorhinus ursinus ) movement in the Bering Sea, Alaska, USA. Results show sex differentiation, with female northern fur seals exhibiting stronger response to environmental variables.

Alaska

Where has turtle ecology been, and where is it going?

Over 9000 articles have been published on turtles and tortoises (excluding sea turtles) since 1950 according to the Web of Science, including over 8000 contained in a personal bibliography that we analyze in this paper. Research had a slow start from 1900 to 1950, with mostly anecdotal additions to our knowledge until the contributions of F. Cagle and A. Carr took turtle research to new levels as the cofathers of turtle ecology in the middle of the last century. Books written in 1939, 1952, and 1972 that compiled existing literature on turtles in the United States and Canada set the stage for growing interest in turtles. The first global compilation of turtle species did not become available until 1961. Publication frequency increased in the 1960s and especially the 1970s as interest in turtles grew, and a wave of turtle biologists emerged from doctoral degree programs. We briefly review the contributions of scientists who published extensively on turtle ecology in those and later decades up to the present. We also review advances in our knowledge of various topics, including the global distribution of turtle research efforts; changes in our perceptions of turtle species diversity over time; turtle community ecology; sex ratios, sex-determination, and climate change; overwintering behavior; sexual size dimorphism and sexual dichromatism; analyses of population genetics; turtles and vocalization; and the emergence of turtle conservation biology efforts. We conclude with a discussion of future opportunities and challenges for working with turtles.

Herpetologica

Analysis of utilization of desert habitats with dynamic simulation

The effects of climate and herbivores on cool desert shrubs in north-western Utah were investigated with a dynamic simulation model. Cool desert shrublands are extensively managed as grazing lands, and are defoliated annually by domestic livestock. A primary production model was used to simulate harvest yields and shrub responses under a variety of climatic regimes and defoliation patterns. The model consists of six plant components, and it is based on equations of growth analysis. Plant responses were simulated under various combinations of 20 annual weather patterns and 14 defoliation strategies. Results of the simulations exhibit some unexpected linearities in model behavior, and emphasize the importance of both the pattern of climate and the level of plant vigor in determining optimal harvest strategies. Model behaviors are interpreted in terms of shrub morphology, physiology and ecology.

Journal of Environmental Management

Survival of Common Loon chicks appears unaffected by Bald Eagle recovery in northern Minnesota

Recovering species are not returning to the same environments or communities from which they disappeared. Conservation researchers and practitioners are thus faced with additional challenges in ensuring species resilience in these rapidly changing ecosystems. Assessing the resilience of species in these novel systems can still be guided by species’ ecology, including knowledge of their population size, life history traits, and behavioral adaptations, as well as the type, strength, and number of ways that they interact with other species in the community. We summarized broad trends of Common Loons ( Gavia immer ) breeding at Voyageurs National Park from 1973 to 2009, and evaluated the effects of increased risk from recovering Bald Eagles ( Haliaeetus leucocephalus ) on chick survival from 2004 to 2006. Adult Common Loons appear to have increased over time. Using Bayesian survival models that accounted for imperfect detection of unmarked individuals, we determined that chick survival of Common Loons was high from year to year and was unrelated to predation risk from Bald Eagles because chicks in territories closer to active nests did not experience greater mortality than those farther away. We suggest that Common Loon chicks were unaffected by the recovery of this top predator during the three years of sampling. Previous research indicates that Bald Eagles and other predators are an important source of egg losses, but Common Loons can compensate by re-nesting. Despite current uncertainties from anthropogenic threats, knowledge of a species’ ecology remains instrumental in determining its resilience during recovery.

Minnesota

Population ecology of Roosevelt elk: Conservation and management in Redwood National and State Parks. Butch Weckerly. 2017. University of Nevada Press, Reno, Nevada, USA. 224 pp. $54.95 hardback. ISBN 978- 1943859504.

Long-term research on large ungulate populations typically conjures perceptions of extensive (and expensive) animal capture and telemetry work, and subsequent advanced modeling of resource selection and population dynamics that inform management decisions. In contrast, studies lacking a telemetry component are often limited to animal behavior or natural history. Although compelling from a standpoint of advancing understanding of ecological and evolutionary processes, results from the latter can be unfairly labeled as esoteric because they are not easily transferable to resource managers or may not provide exceptional interest to a general public drawn to these charismatic megafuana. Such dichotomies are not predetermined, however, because study conditions exist where dedicated academic researchers on tight budgets can achieve results relevant to ecology and management.

Journal of Wildlife Management

Diel predator activity drives a dynamic landscape of fear

A “landscape of fear” (LOF) is a map that describes continuous spatial variation in an animal's perception of predation risk. The relief on this map reflects, for example, places that an animal avoids to minimize risk. Although the LOF concept is a potentially unifying theme in ecology that is often invoked to explain the ecological and conservation significance of fear, little is known about the daily dynamics of an LOF. Despite theory and data to the contrary, investigators often assume, implicitly or explicitly, that an LOF is a static consequence of a predator's mere presence within an ecosystem. We tested the prediction that an LOF in a large‐scale, free‐living system is a highly dynamic map with “peaks” and “valleys” that alternate across the diel (24‐h) cycle in response to daily lulls in predator activity. We did so with extensive data from the case study of Yellowstone elk ( Cervus elaphus ) and wolves ( Canis lupus ) that was the original basis for the LOF concept. We quantified the elk LOF, defined here as spatial allocation of time away from risky places and times, across nearly 1,000‐km 2 of northern Yellowstone National Park and found that it fluctuated with the crepuscular activity pattern of wolves, enabling elk to use risky places during wolf downtimes. This may help explain evidence that wolf predation risk has no effect on elk stress levels, body condition, pregnancy, or herbivory. The ability of free‐living animals to adaptively allocate habitat use across periods of high and low predator activity within the diel cycle is an underappreciated aspect of animal behavior that helps explain why strong antipredator responses may trigger weak ecological effects, and why an LOF may have less conceptual and practical importance than direct killing.

Yellowstone National Park

Adaptive breeding habitat selection: Is it for the birds?

The question of why animals choose particular habitats has important implications for understanding behavioral evolution and distribution of organisms in the wild and for delineating between habitats of different quality for conservation and management. Habitats chosen by animals can influence fitness outcomes via the costs (e.g., predation risk) and benefits (e.g., food availability) of habitat use. Habitat preferences should therefore be under selection to favor those that confer fitness advantages (Clark and Shutler 1999). Indeed, prevailing theory suggests that the habitat preferences of animals should be adaptive, such that fitness is higher in preferred habitats (Hildén 1965, Southwood 1977, Martin 1998). However, studies have often identified apparent mismatches between observed habitat preferences and fitness outcomes across a wide variety of taxa (Valladares and Lawton 1991, Mayhew 1997, Kolbe and Janzen 2002, Arlt and Pärt 2007, Mägi et al. 2009). Certainly, one limitation of studies may be that assessment of “fitness” is typically constrained to fitness surrogates such as nest success rather than lifetime reproductive success or classic Fisherian fitness (Endler 1986). Nevertheless, important habitat choices such as nest sites influence the probability that temporarily sedentary, dependent young are discovered by enemies such as predators and parasites. We therefore expect, on average, to see congruence between evolved habitat preferences and relevant components of fitness (e.g., nest success). Here, we (1) review the prevalence of apparent mismatches between avian breeding-habitat preferences and fitness outcomes using nest-site selection as a focus; (2) describe several potential mechanisms for such mismatches, including anthropogenic, methodological, and ecological–evolutionary; and (3) suggest a framework for understanding the contexts in which habitat preferences represent adaptive decisions, with a primary focus on ecological information theory. We largely focus on habitat selection as a behavioral process at the scale of individuals (e.g., Robertson and Hutto 2006), rather than at the scale of population-level patterns (Fretwell and Lucas 1970, Morris 2003, Johnson 2007). However, these two scales cannot be wholly divorced from one another, as we will discuss.

The Auk

Long-term apparent survival of a cold-stunned subpopulation of juveniles green turtles

Understanding the effects of extreme weather on animal populations is fundamental to ecological and conservation sciences and species management. Climate change has resulted in both warm and cold temperature extremes, including an increased frequency of severe cold snaps at middle latitudes in North America. These unusually cold air masses cause rapid declines in nearshore ocean temperatures in coastal areas, with detrimental effects on marine organisms. Acute cold-stun events (hereafter cold stuns) occur when hundreds to thousands of resident juvenile sea turtles fail to escape shallow water during cold snaps. Human intervention through rescue and recovery largely mitigates direct juvenile sea turtle mortality, but delayed effects of cold stuns on rescued individuals are not well understood. Our objective was to examine long-term juvenile green turtle ( Chelonia mydas ) survival across four cold stuns of varying severity in St. Joseph Bay, Florida, between 2010 and 2018. We used the classic Cormack–Jolly–Seber model in a hierarchical Bayesian framework to estimate apparent survival (i.e., emigration and mortality) of rescued turtles at different time intervals. Our results indicated about half of a cohort rescued during a severe cold stun in January 2010 likely remained in the population 1 year later, with 10%–20% remaining 4 years later, and as few as 5% by 2018. The results also suggested higher apparent survival for cohorts rescued during two subsequent milder cold stuns. Emigration was a more plausible ecological explanation for low apparent survival than delayed mortality. Potential ecological mechanisms underlying emigration include a reduction in food availability and a behavioral response to either the severe weather event or handling during rescue (or both). However, the typical annual turnover of juvenile green turtles, though assumed low, is not well known in St. Joseph Bay. Thus, our apparent survival estimates may be reflective of higher-than-expected emigration in the broader population. Our study provides important baseline information about long-term juvenile sea turtle survival after cold stuns in temperate regions. We also highlight the importance of strategic monitoring between cold stuns to examine additional ecological questions.

Florida

Decision-making in agent-based modeling: A current review and future prospectus

All basic processes of ecological populations involve decisions; when and where to move, when and what to eat, and whether to fight or flee. Yet decisions and the underlying principles of decision-making have been difficult to integrate into the classical population-level models of ecology. Certainly, there is a long history of modeling individuals' searching behavior, diet selection, or conflict dynamics within social interactions. When all the individuals are given certain simple rules to govern their decision-making processes, the resultant population–level models have yielded important generalizations and theory. But it is also recognized that such models do not represent the way real individuals decide on actions. Factors that influence a decision include the organism's environment with its dynamic rewards and risks, the complex internal state of the organism, and its imperfect knowledge of the environment. In the case of animals, it may also involve complex social factors, and experience and learning, which vary among individuals. The way that all factors are weighed and processed to lead to decisions is a major area of behavioral theory. While classic population-level modeling is limited in its ability to integrate decision-making in its actual complexity, the development of individual- or agent-based models (IBM/ABMs) (we use ABM throughout to designate both “agent-based modeling” and an “agent-based model”) has opened the possibility of describing the way that decisions are made, and their effects, in minute detail. Over the years, these models have increased in size and complexity. Current ABMs can simulate thousands of individuals in realistic environments, and with highly detailed internal physiology, perception and ability to process the perceptions and make decisions based on those and their internal states. The implementation of decision-making in ABMs ranges from fairly simple to highly complex; the process of an individual deciding on an action can occur through the use of logical and simple (if-then) rules to more sophisticated neural networks and genetic algorithms. The purpose of this paper is to give an overview of the ways in which decisions are integrated into a variety of ABMs and to give a prospectus on the future of modeling of decisions in ABMs.

Frontiers in Ecology and Evolution