Biology subjects

Fleming, C. H.

Publications and source records attributed to Fleming, C. H..

14 recordsLinked to original sources

bioRxiv2025

Estimating population range distributions from animal tracking data

O_LIQuantifying the space requirements of a population is a fundamental problem in spatial ecology, particularly as it relates to the identification of important utilization areas and the designation of protected areas for conservation and wildlife management. C_LIO_LITraditionally, population space use estimation techniques scale up from the individual to the population level by aggregating individual animal tracks and then using a single pooled distribution estimator like minimum convex polygons (MCP) or kernel density estimation (KDE). These techniques fail to account for the high levels of temporal autocorrelation in modern tracking datasets, and estimates are often sensitive to the number of individuals sampled. C_LIO_LIWe introduce a new population kernel density estimator (PKDE) that accounts for temporal autocorrelation in tracking data, propagates uncertainty from the individual to the population level, accounts for inter-individual variation when scaling up to the population level, and is not highly sensitive to the number of individuals tracked. Through a combination of simulated data and empirical GPS tracking datasets from three species: (a) grizzly bear (Ursus arctos horribilis); (b) lowland tapir (Tapirus terrestris); and (c) bobcat (Lynx rufus), we demonstrate that PKDE produces minimally biased estimates of population-level space use compared to conventional methods like MCP and KDE. C_LIO_LIThe use of conventional estimators can lead to substantial underestimation of population space usage, making them unsuitable for area-based conservation planning. The statistically efficient PKDE estimator provides relatively unbiased estimates of space use with fewer individuals sampled. This method has been made available as a function in the ctmm R package. C_LI

ecology↗

bioRxiv2023

Intraspecific encounters can induce home-range shifts

Direct encounters, in which two or more individuals are physically close to one another, are a topic of increasing interest as more and better movement data become available. Recent progress, including the development of statistical tools for estimating robust measures of changes in animals space use over time, facilitates opportunities to link direct encounters between individuals with the long-term consequences of those encounters. Working with movement data for coyotes (Canis latrans) and grizzly bears (Ursus arctos horribilis), we investigate whether close intraspecific encounters were associated with spatial shifts in the animals range distributions, as might be expected if one or both of the individuals involved in an encounter were seeking to reduce or avoid conflict over space. We analyze the movement data of a pair of coyotes in detail, identifying how a shift in home range location resulting from altered movement behavior was apparently a consequence of a close intraspecific encounter. With grizzly bear movement data, we approach the problem from the perspective of a set of encounter pairs within a population. We find support for the hypotheses that 1) close intraspecific encounters between bears are, on average, associated with subsequent shifts in range distributions and 2) encounters defined at finer spatial scales are followed by greater changes in space use. Our results suggest that animals can undertake long-term, large-scale spatial shifts in response to close intraspecific encounters that have the potential for conflict. These results lend support for existing theory on the evolution of territories and space use (e.g., Maynard-Smiths bourgeois strategy regarding low-conflict coexistence). Overall, we find that analyses of movement data in a pairwise context can 1) identify distances at which individuals proximity to one another may alter behavior and 2) facilitate testing of population-level hypotheses concerning the potential for direct encounters to alter individuals space use. Open Research StatementMovement data for the coyotes and grizzly bears are posted on Movebank.org as datasets 1614661371 and 1044288582, respectively. Statistical tools for estimating, manipulating, and comparing home ranges from movement data are implemented in the open-source R package ctmm. R scripts used to carry out specific analyses for this study are openly available on GitHub at https://github.com/anagkrish/encounter_homerangeshift.

ecology↗

bioRxiv2023

movedesign: Shiny R app to evaluate sampling design for animal tracking movement studies

O_LIProjects focused on movement behavior and home range are commonplace, but beyond a focus on choosing appropriate research questions, there are no clear guidelines for such studies. Without these guidelines, designing an animal tracking study to produce reliable estimates of space-use and movement properties (necessary to answer basic movement ecology questions), is often done in an ad hoc manner. C_LIO_LIWe developed movedesign, a user-friendly Shiny application, which can be utilized to investigate the precision of three estimates regularly reported in movement and spatial ecology studies: home range area, speed, and distance traveled. Conceptually similar to statistical power analysis, this application enables users to assess the degree of estimate precision that may be achieved with a given sampling design; i.e., the choices regarding data resolution (sampling interval) and battery life (sampling duration). C_LIO_LILeveraging the ctmm R package, we utilize two methods proven to handle many common biases in animal movement datasets: autocorrelated Kernel Density Estimators (AKDE) and continuous-time speed and distance (CTSD) estimators. Longer sampling durations are required to reliably estimate home range areas via the detection of a sufficient number of home range crossings. In contrast, speed and distance estimation requires a sampling interval short enough to ensure that a statistically significant signature of the animals velocity remains in the data. C_LIO_LIThis application addresses key challenges faced by researchers when designing tracking studies, including the trade-off between long battery life and high resolution of GPS locations collected by the devices, which may result in a compromise between reliably estimating home range or speed and distance. movedesign has broad applications for researchers and decision-makers, supporting them to focus efforts and resources in achieving the optimal sampling design strategy for their research questions, prioritizing the correct deployment decisions for insightful and reliable outputs, while understanding the trade-off associated with these choices. C_LI

ecology↗

bioRxiv2023

The search behavior of terrestrial mammals

Animals moving through landscapes need to strike a balance between finding sufficient resources to grow and reproduce while minimizing encounters with predators 1,2. Because encounter rates are determined by the average distance over which directed motion persists 1,3-5, this trade-off should be apparent in individuals movement. Using GPS data from 1,396 individuals across 62 species of terrestrial mammals, we show how predators maintained directed motion ~7 times longer than for similarly-sized prey, revealing how prey species must trade off search efficiency against predator encounter rates. Individual search strategies were also modulated by resource abundance, with prey species forced to risk higher predator encounter rates when resources were scarce. These findings highlight the interplay between encounter rates and resource availability in shaping broad patterns mammalian movement strategies.

ecology↗

bioRxiv2022

Clarifying space use concepts in ecology: range vs. occurrence distributions

Quantifying animal movements is necessary for answering a wide array of research questions in ecology and conservation biology. Consequently, ecologists have made considerable efforts to identify the best way to estimate an animals home range, and many methods of estimating home ranges have arisen over the past half century. Most of these methods fall into two distinct categories of estimators that have only recently been described in statistical detail: those that measure range distributions (methods such as Kernel Density Estimation that quantify the long-run behavior of a movement process that features restricted space use) and those that measure occurrence distributions (methods such as Brownian Bridge Movement Models and the Correlated Random Walk Library that quantify uncertainty in an animal movement path during a specific period of observation). In this paper, we use theory, simulations, and empirical analysis to demonstrate the importance of applying these two classes of space use estimators appropriately and distinctly. Conflating range and occurrence distributions can have serious consequences for ecological inference and conservation practice. For example, in most situations, home-range estimates quantified using occurrence estimators are too small, and this problem is exacerbated by ongoing improvements in tracking technology that enable more frequent and more accurate data on animal movements. We encourage researchers to use range estimators to estimate the area of home ranges and occurrence estimators to answer other questions in movement ecology, such as when and where an animal crosses a linear feature, visits a location of interest, or interacts with other animals. Open Research StatementTracking data on Aepyceros melampus, Beatragus hunteri, Bycanistes bucinator, Cerdocyon thous, Eulemur rufifrons, Glyptemys insculpta, Gyps coprotheres, Madoqua guentheri, Ovis canadensis, Propithecus verreauxi, Sus scrofa, and Ursus arctos are publicly archived in the Dryad repository (Noonan et al. 2018; https://doi.org/10.5061/dryad.v5051j2), as are data from Procapra gutturosa (Fleming et al. 2014a; https://doi.org/10.5061/dryad.45157). Data on Panthera onca were taken from (Morato et al. 2018). Additional data are publicly archived in the Movebank repository under the following identifiers: Canis latrans, 8159699; Canis lupus, 8159399; Chrysocyon brachyurus, 18156143; Felis silvestris, 40386102; Gyps africanus, 2919708; Lepus europaeus, 25727477; Martes pennanti, 2964494; Panthera leo, 220229; Papio cynocephalus, 222027; Syncerus caffer, 1764627; Tapirus terrestris, 443607536; Torgos tracheliotus, 2919708; and Ursus americanus, 8170674.

ecology↗

bioRxiv2022

Mitigating pseudoreplication and bias in resource selection functions with autocorrelation-informed weighting

O_LIResource selection functions are among the most commonly used statistical tools in both basic and applied animal ecology. They are typically parameterized using animal tracking data, and advances in animal tracking technology have led to increasing levels of autocorrelation between locations in such data sets. Because resource selection functions assume that data are independent and identically distributed, such autocorrelation can cause misleadingly narrow confidence intervals and biased parameter estimates. C_LIO_LIData thinning, generalized estimating equations, and step selection functions have been suggested as techniques for mitigating the statistical problems posed by autocorrelation, but these approaches have notable limitations that include statistical inefficiency, unclear or arbitrary targets for adequate levels of statistical independence, constraints in input data, and (in the case of step selection functions) scale-dependent inference. To remedy these problems, we introduce a method for likelihood weighting of animal locations to mitigate the negative consequences of autocorrelation on resource selection functions. C_LIO_LIIn this study, we demonstrate that this method weights each observed location in an animals movement track according to its level of non-independence, expanding confidence intervals and reducing bias that can arise when there are missing data in the movement track. C_LIO_LIEcologists and conservation biologists can use this method to improve the quality of inferences derived from resource selection functions. We also provide a complete, annotated analytical workflow to help new users apply our method to their own animal tracking data using the ctmm R package. C_LI

ecology↗

bioRxiv2021

Movement ecology of vulnerable lowland tapirs across a gradient of human disturbance

Animal movement is a key ecological process that is tightly coupled to local environmental conditions. While agriculture, urbanisation, and transportation infrastructure are critical to human socio-economic improvement, these have spurred substantial changes in animal movement across the globe with potential impacts on fitness and survival. Notably, however, human disturbance can have differential effects across species, and responses to human activities are thus largely taxa and context specific. As human disturbance is only expected to worsen over the next decade it is critical to better understand how species respond to human disturbance in order to develop effective, case-specific conservation strategies. Here, we use an extensive telemetry dataset collected over 22 years to fill a critical knowledge gap in the movement ecology of lowland tapirs (Tapirus terrestris) across a gradient of human disturbance within three biomes in southern Brazil: the Pantanal, Cerrado, and Atlantic Forest. From these data we found that the mean home range size across all monitored tapirs was 8.31 km2 (95% CI: 6.53 - 10.42), with no evidence that home range sizes differed between sexes nor age groups. Interestingly, although the Atlantic Forest, Cerrado, and Pantanal vary substantially in habitat composition, levels of human disturbance, and tapir population densities, we found that lowland tapir movement behaviour and space use were consistent across all three biomes. Human disturbance also had no detectable effect on lowland tapir movement. Lowland tapirs living in the most altered habitats we monitored exhibited movement behaviour that was comparable to that of tapirs living in a near pristine environment. Contrary to our expectations, we observed very little individual variability in lowland tapir space use and movement, and human impacts on the landscape also had no measurable effect on their movement. Lowland tapir movement behaviour thus appears to exhibit very little phenotypic plasticity. Crucially, the lack of any detectable response to anthropogenic disturbance suggests that human modified habitats risk being ecological traps for tapirs and this information should be factored into conservation actions and species management aimed towards protecting lowland tapir populations.

ecology↗

bioRxiv2021

Population-level inference for home-range areas

O_LIHome-range estimates are a common product of animal tracking data, as each range informs on the area needed by a given individual. Population-level inference on home-range areas--where multiple individual home-ranges are considered to be sampled from a population--is also important to evaluate changes over time, space, or covariates, such as habitat quality or fragmentation, and for comparative analyses of species averages. Population-level home-range parameters have traditionally been estimated by first assuming that the input tracking data were sampled independently when calculating home ranges via conventional kernel density estimation (KDE) or minimal convex polygon (MCP) methods, and then assuming that those individual home ranges were measured exactly when calculating the population-level estimates. This conventional approach does not account for the temporal autocorrelation that is inherent in modern tracking data, nor for the uncertainties of each individual home-range estimate, which are often large and heterogeneous. C_LIO_LIHere, we introduce a statistically and computationally efficient framework for the population-level analysis of home-range areas, based on autocorrelated kernel density estimation (AKDE), that can account for variable temporal autocorrelation and estimation uncertainty. C_LIO_LIWe apply our method to empirical examples on lowland tapir (Tapirus terrestris), kinkajou (Potos flavus), white-nosed coati (Nasua narica), white-faced capuchin monkey (Cebus capucinus), and spider monkey (Ateles geoffroyi), and quantify differences between species, environments, and sexes. C_LIO_LIOur approach allows researchers to more accurately compare different populations with different movement behaviors or sampling schedules, while retaining statistical precision and power when individual home-range uncertainties vary. Finally, we emphasize the estimation of effect sizes when comparing populations, rather than mere significance tests. C_LI

ecology↗

bioRxiv2021

A semi-variance approach to visualising phylogenetic autocorrelation

O_LIComparing traits across species has been a hallmark of biological research for centuries. While inter-specific comparisons can be highly informative, phylogenetic inertia can bias estimates if not properly accounted for in comparative analyses. In response, researchers typically treat phylogenetic inertia as a form of autocorrelation that can be detected, modelled, and corrected for. Despite the range of methods available for quantifying the strength of phylogenetic autocorrelation, no tools exist for visualising these autocorrelation structures. C_LIO_LIHere we derive variogram methods suitable for phylogenic data, and show how they can be used to straightforwardly visualise phylogenetic autocorrelation. We then demonstrate their utility for three empirical examples: sexual size dimorphism (SSD) in the Musteloidea, maximum per capita rate of population growth, r, in the Carnivora, and brain size in the Artiodactyla. C_LIO_LIWhen modelling musteloid SSD, the empirical variogram showed a tendency for the variance in SSD to stabilise over time, a characteristic feature of Ornstein-Uhlenbeck (OU) evolution. In agreement with this visual assessment, model selection identified the OU model as the best fit to the data. In contrast, the infinitely diffusive Brownian Motion (BM) model did not capture the asymptotic behaviour of the variogram and was less supported than the OU model. Phylogenetic variograms proved equally useful in understanding why an OU model was selected when modelling r in the Carnivora, and why BM was the selected evolutionary model for brain size in the Artiodactyla. C_LIO_LIBecause the variograms of the various evolutionary processes each have different theoretical profiles, comparing fitted semi-variance functions against empirical semi-variograms can serve as a useful diagnostic tool, allowing researchers to understand why any given evolutionary model might be selected over another, which features are well captured by the model, and which are not. This allows for fitted models to be compared against the empirical variogram, facilitating model identification prior to subsequent analyses. We therefore recommend that any phylogenetic analysis begin with a non-parametric estimate of the autocorrelation structure of the data that can be visualized. The methods developed in this work are openly available in the new R package ctpm. C_LI

evolutionary biology↗

bioRxiv2020

Estimating encounter location distributions from animal tracking data

O_LIEcologists have long been interested in linking individual behavior with higher-level processes. For motile species, this upscaling is governed by how well any given movement strategy maximizes encounters with positive factors, and minimizes encounters with negative factors. Despite the importance of encounter events for a broad range of ecological processes, encounter theory has not kept pace with developments in animal tracking or movement modeling. Furthermore, existing work has focused primarily on the relationship between animal movement and encounter rates while no theoretical framework exists for directly relating individual movement with the spatial locations of encounter events in the environment. C_LIO_LIHere, we bridge this gap by introducing a new theoretical concept describing the long-term encounter location probabilities for movement within home ranges, termed the conditional distribution of encounters (CDE). We then derive this distribution, as well as confidence intervals, implement its statistical estimator into open source software, and demonstrate the broad ecological relevance of this novel concept. C_LIO_LIWe first use simulated data to show how our estimator provides asymptotically consistent estimates. We then demonstrate the general utility of this method for three simulation-based scenarios that occur routinely in biological systems: i) a population of individuals with home ranges that overlap with neighbors; ii) a pair of individuals with a hard territorial border between their home ranges; and iii) a predator with a large home range that encompassed the home ranges of multiple prey individuals. Using GPS data from white-faced capuchins (Cebus capucinus) tracked on Barro Colorado Island, Panama, and sleepy lizards (Tiliqua rugosa) tracked in Bundey, South Australia, we then show how the CDE can be used to estimate the locations of territorial borders, identify key resources, quantify the location-specific potential for competition, and/or identify any changes in behaviour that directly result from location-specific encounter probability. C_LIO_LIThis novel target distribution enables researchers to better understand the dynamics of populations of interacting individuals. Notably, the general estimation framework developed in this work builds straightforwardly off of home range estimation and requires no specialised data collection protocols. This method is now openly available via the ctmm R package. C_LI

ecology↗

bioRxiv2020

A comprehensive framework for handling location error in animal tracking data

Animal tracking data are being collected more frequently, in greater detail, and on smaller taxa than ever before. These data hold the promise to increase the relevance of animal movement for understanding ecological processes, but this potential will only be fully realized if their accompanying location error is properly addressed. Historically, coarsely-sampled movement data have proved invaluable for understanding large scale processes (e.g., home range, habitat selection, etc.), but modern fine-scale data promise to unlock far more ecological information. While GPS location error can often be ignored in coarsely sampled data, fine-scale data require more care, and tools to do this have not kept pace. Current approaches to dealing with location error largely fall into two categories--either discarding the least accurate location estimates prior to analysis or simultaneously fitting movement and error parameters in a hidden-state model. In some cases these approaches can provide a level of correction, but they have known limitations, and in some cases they can be worse than doing nothing. Here, we provide a general framework to account for location error in the analysis of triangulated and trilatcralizcd animal tracking data, which includes GPS, Argos Doppler-shift, triangulated VHF, trilatcralized acoustic and cellular location data. We apply our error-modelselection framework to 190 GPS, cellular, and acoustic devices representing 27 models from 14 manufacturers. Collectively, these devices were used to track a wide range of taxa comprising birds, fish, reptiles, and mammals of different sizes and with different behaviors, in urban, suburban, and wild settings. In almost half of the tested device models, error-model selection was necessary to obtain the best performing error model, and in almost a quarter of tested device models, the reported DOP values were actually misinformative. Then, using empirical tracking data from multiple species, we provide an overview of modern, error-informed movement analyses, including continuous-time path reconstruction, home-range distribution, home-range overlap, speed, and distance estimation. Adding to these techniques, we introduce new error-informed estimators for outlier detection and autocorrelation visualization. Because error-induced biases depend on many factors--sampling schedule, movement characteristics, tracking device, habitat, etc.--differential bias can easily confound biological inference and lead researchers to draw false conclusions. We demonstrate how error-informed analyses on calibrated tracking data can provide more accurate estimates are that are insensitive to location error, and allow researchers to use all of their data.

ecology↗

bioRxiv2020

ctmmweb: A graphical user interface for autocorrelation-informed home range estimation

Estimating animal home ranges is a primary purpose of collecting tracking data. All conventional home range estimators in widespread usage, including minimum convex polygons and kernel density estimators, assume independently sampled data. In stark contrast, modern GPS animal tracking datasets are almost always strongly autocorrelated. This incongruence between estimator assumptions and empirical reality leads to systematically underestimated home ranges. Autocorrelated kernel density estimation (AKDE) resolves this conflict by modeling the observed autocorrelation structure of tracking data during home range estimation, and has been shown to perform accurately across a broad range of tracking datasets. However, compared to conventional estimators, AKDE requires additional modeling steps and has heretofore only been accessible via the command-line ctmm R package. Here, we introduce ctmmweb, which provides a point-and-click graphical interface to ctmm, and streamlines AKDE, its prerequisite autocorrelation modeling steps, and a number of additional movement analyses. We demonstrate ctmmwebs capabilities, including AKDE home range estimation and subsequent home range overlap analysis, on a dataset of four jaguars from the Brazilian Pantanal. We intend ctmmweb to open AKDE and related autocorrelation-explicit analyses to a wider audience of wildlife and conservation professionals.

ecology↗

bioRxiv2019

The challenges of estimating the distribution of flight heights from telemetry or altimetry data

BackgroundGlobal positioning systems (GPS) and altimeters are increasingly used to monitor vertical space use by aerial species, a key aspect of their niche that we need to know to understand their ecology and conservation needs, and to manage our own use of the airspace. However, there are various sources of error in flight height data (\"height\" above ground, as opposed to \"altitude\" above a reference like the sea level): vertical error from the devices themselves, error in the ground elevation below the tracked animals, and error in the horizontal position of the animals and thus the predicted ground elevation below them.\n\nMethodsWe used controlled field trials, simulations, and the reanalysis of raptor case studies with state-space models to illustrate the effect of improper error management.\n\nResultsErrors of a magnitude of 20 meters appear in benign conditions (expected to be larger in more challenging context). These errors distort the shape of the distribution of flight heights, inflate the variance in flight height, bias behavioural state assignments, correlations with environmental covariates, and airspace management recommendations. Improper data filters such as removing all negative recorded flight height records introduce several biases in the remaining dataset, and preclude the opportunity to leverage unambiguous errors to help with model fitting. Analyses that ignore the variance around the mean flight height, e.g., those based on linear models of flight height, and those that ignore the variance inflation caused by telemetry errors, lead to incorrect inferences.\n\nConclusionThe state-space modelling framework, now in widespread use by ecologists and increasingly often automatically implemented within on-board GPS data processing algorithms, makes it possible to fit flight models directly to raw flight height records, with minimal data pre-selection, and to analyse the full distribution of flight heights, not just the mean. In addition to basic research about aerial niches, behaviour quantification, and environmental interactions, we highlight the applied relevance of our recommendations for airspace management and the conservation of aerial wildlife.

ecology↗

bioRxiv2019

How range residency and long-range perception change encounter rates

Encounter rates link movement strategies to intra- and inter-specific interactions, and therefore translate individual movement behavior into higher-level ecological processes. Indeed, a large body of interacting population theory rests on the law of mass action, which can be derived from assumptions of Brownian motion in an enclosed container with exclusively local perception. These assumptions imply completely uniform space use, individual home ranges equivalent to the population range, and encounter dependent on movement paths actually crossing. Mounting empirical evidence, however, suggests that animals use space non-uniformly, occupy home ranges substantially smaller than the population range, and are often capable of nonlocal perception. Here, we explore how these empirically supported behaviors change pairwise encounter rates. Specifically, we derive novel analytical expressions for encounter rates under Ornstein-Uhlenbeck motion, which features non-uniform space use and allows individual home ranges to differ from the population range. We compare OU-based encounter predictions to those of Reflected Brownian Motion, from which the law of mass action can be derived. For both models, we further explore how the interplay between the scale of perception and home range size affects encounter rates. We find that neglecting realistic movement and perceptual behaviors can systematically bias encounter rate predictions.

ecology↗