Richard J. Hall’s research while affiliated with University of Georgia and other places

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Publications (83)


Human-provided food increases aggregation but does not change activity budgets in an urban wading bird
  • Article

August 2024

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11 Reads

Journal of Urban Ecology

Cali A Wilson

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Sonia Hernandez

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Julia N Weil

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[...]

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Richard J Hall

In urban areas, animals often aggregate at higher densities, move less, and alter their diets to consume anthropogenic food, all of which can affect wildlife health and the transmission of infectious diseases. However, it is unknown whether short-term changes in behavior associated with urban resources scale up to more pervasive long-term behavioral changes across landscape types. In this study, we used observational field data to explore how food provisioning affects behaviors relevant to parasite transmission in American white ibis (Eudocimus albus), a waterbird that has recently habituated to urban habitats and anthropogenic food. We found that ibis flock densities more than doubled during short intervals when birds were actively provisioned with food. We then explored activity budgets among urban sites with different levels of provisioning, and found that foraging time decreased with flock size and provisioning levels. Lastly, we compared ibis behavior in more natural wetland sites against urban sites, and found minimal to no differences in behaviors measured here. These results suggest that urbanization and provisioning alter ibis behaviors in ways that could influence, e.g. exposure to parasites in the short-term, but this has not yet resulted in significant long-term changes in activity budgets. Further studies of how urbanization and intentional feeding influences wildlife behavior can inform management strategies to benefit both wildlife and human health.


Perceived and observed biases within scientific communities: a case study in movement ecology
  • Preprint
  • File available

August 2024

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471 Reads

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Figure 1. Conceptual figure of data collection and analysis. A) Voles were monitored via capture-mark-recapture methods and the trapped locations of individuals were recorded. B) Trapped locations were pooled across individuals by functional group (combination of sex and reproductive status), treatment, and season (summer or autumn). Space use was characterised for each group as the probability of capturing a vole of that group with increasing distance from the centroid of their trapped locations. This probability was assumed to be equal in all directions from the centroid, generating a circular kernel of space use. C) Space-use overlap between pairs of voles was estimated each month based on the overlap of the kernels for each vole's functional group drawn around the centroids of their respective trapping locations. D) The amount of pairwise space-use overlap between voles was used to inform the edge weights of spatial overlap networks for voles at each site in each month JuneOctober.
Figure 3. Seasonal space use and spatial overlap networks throughout the study period in 2021 for one of the investigated vole populations (unfed-control treatment, site "Helmipöllö"). A) Coloured circles indicate space use of voles captured at the site each month. Space use was characterised separately for each functional group across the summer breeding season (June-August) and across the autumn non-breeding season (September-October). Reproductive males (dark blue) had larger space use than reproductive females (red) in summer. Larger, darker circles in June-August indicate reproductive voles. Non-reproductive voles (smaller, lighter circles) first appear in July. Reproductive and non-reproductive vole space use was similar in autumn. B) Spatial overlap networks were constructed each month with edges weighted based on the amount of space-use overlap between pairs of voles (edge weight values scale 0 to 1). Network nodes are positioned in space to match the location of voles' monthly centroids as seen in (A). Edges of higher weight (indicating greater overlap) are thicker and darker in colour.
Ecological factors alter how spatial overlap predicts viral infection dynamics in wild rodent populations

June 2024

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144 Reads

Spatial overlap between animals in wildlife populations can have important implications for pathogen transmission. Ecological factors and animal demographic traits can influence animal space use and spatial overlap, but it is unclear how these interactions drive pathogen transmission. We experimentally manipulated wild bank vole populations via resource supplementation and anthelmintic treatment. Using network analysis, we investigated the relationship between spatial overlap and infection probability of an endemic zoonotic hantavirus, including how vole sex and reproductive status interact with spatial behaviour to affect infection probability. Spatial overlap in a previous month drove current hantavirus infection probability, and food supplementation and anthelmintic treatment altered the effects of spatial overlap on infection probability. Vole sex and reproductive status were important factors determining whether spatial overlap increased or decreased the probability of hantavirus infection and interacted with resource supplementation and anthelmintic treatment, generating different infection dynamics in each treatment. Our research provides rare empirical evidence linking previous spatial overlap to current infection status in wildlife populations, with implications for understanding zoonotic exposure risk for humans. We further highlight the importance of incorporating variation in ecological factors and host demography when studying pathogen transmission in wildlife systems.


Field study design and experimental set‐up. (Site Locations) Twelve field sites were established in southern Finland. (Treatments) Sites were assigned one of four treatment pairings: unfed‐control, unfed‐deworm, fed‐control and fed‐deworm, and each treatment was replicated at three sites. (Trapping Grid Arrangement) The trapping grid included 61 traps in 11 rows and 11 columns. Rows and columns were spaced 10 m apart and traps (dots) were placed in an offset arrangement with 20 m to the next trap in the same row and column and 14.14 m to the next trap on the diagonal. Site map by Anthony Shing. Figure created with BioRender.com.
Space use of bank voles by sex and reproductive status across treatments in the summer breeding season (June–August) and autumn nonbreeding season (September–October). Circle size represents the mean space use area (m²) where the probability of capture is >0.01.
Spatial overlap networks of vole pairwise space‐use overlap. Nodes represent individual voles, edges represent pairwise space‐use overlap, thicker edges indicate greater space‐use overlap between voles. For each treatment, networks from one site are shown. Edge weights are thresholded to a minimum edge weight of 0.05 for visualization purposes.
Population‐level network size and individual vole spatial overlap. (a) Network size (number of voles) by month. One data point shown per replicate site. (b) Weighted degree (total pairwise space‐use overlap per vole, weighted by the amount of overlap with each of its neighbours) by month. Means shown as points and lines indicate the range of values for males (blue) and females (pink) in each treatment. Data shown for reproductive voles only, trends for non‐reproductive voles were similar.
Effects of food supplementation and helminth removal on space use and spatial overlap in wild rodent populations

February 2024

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99 Reads

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2 Citations

Animal space use and spatial overlap can have important consequences for population‐level processes such as social interactions and pathogen transmission. Identifying how environmental variability and inter‐individual variation affect spatial patterns and in turn influence interactions in animal populations is a priority for the study of animal behaviour and disease ecology. Environmental food availability and macroparasite infection are common drivers of variation, but there are few experimental studies investigating how they affect spatial patterns of wildlife. Bank voles (Clethrionomys glareolus) are a tractable study system to investigate spatial patterns of wildlife and are amenable to experimental manipulations. We conducted a replicated, factorial field experiment in which we provided supplementary food and removed helminths in vole populations in natural forest habitat and monitored vole space use and spatial overlap using capture–mark–recapture methods. Using network analysis, we quantified vole space use and spatial overlap. We compared the effects of food supplementation and helminth removal and investigated the impacts of season, sex and reproductive status on space use and spatial overlap. We found that food supplementation decreased vole space use while helminth removal increased space use. Space use also varied by sex, reproductive status and season. Spatial overlap was similar between treatments despite up to threefold differences in population size. By quantifying the spatial effects of food availability and macroparasite infection on wildlife populations, we demonstrate the potential for space use and population density to trade‐off and maintain consistent spatial overlap in wildlife populations. This has important implications for spatial processes in wildlife including pathogen transmission.


Extreme heat reduces host and parasite performance in a butterfly–parasite interaction

January 2024

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119 Reads

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2 Citations

Environmental temperature fundamentally shapes insect physiology, fitness and interactions with parasites. Differential climate warming effects on host versus parasite biology could exacerbate or inhibit parasite transmission, with far-reaching implications for pollination services, biocontrol and human health. Here, we experimentally test how controlled temperatures influence multiple components of host and parasite fitness in monarch butterflies (Danaus plexippus) and their protozoan parasites Ophryocystis elektroscirrha. Using five constant-temperature treatments spanning 18–34°C, we measured monarch development, survival, size, immune function and parasite infection status and intensity. Monarch size and survival declined sharply at the hottest temperature (34°C), as did infection probability, suggesting that extreme heat decreases both host and parasite performance. The lack of infection at 34°C was not due to greater host immunity or faster host development but could instead reflect the thermal limits of parasite invasion and within-host replication. In the context of ongoing climate change, temperature increases above current thermal maxima could reduce the fitness of both monarchs and their parasites, with lower infection rates potentially balancing negative impacts of extreme heat on future monarch abundance and distribution.


An objectives hierarchy, depicting a fundamental objective (ie values that indicate importance) connected to subordinate means objectives (ie how to achieve) and performance metrics to summarize consequences. In this example, performance metrics include the probability of introducing a pathogen, the percentage of infected hosts observed through monitoring, the estimated abundance of the target population, the percentage of the host population that has reached maturity, the interannual variability in harvests, and the percentage of years when harvests are below a target reference point.
A generalized model of infection dynamics and the effects of potential management actions. The four central green rectangles represent the host stock by infection status: Susceptible, Exposed, Infectious (asymptomatic and symptomatic), and Recovered. The associated green arrows indicate transition rates between host infection status (infection, latency, recovery; for simplicity, loss of immunity is not shown). The fifth green rectangle represents alternative hosts that can also supply parasites to the system apart from the focal stock. Also shown as arrows: exposure to parasites (solid black), parasite shedding (dotted orange), mortalities (dashed gray) and stocking (blue). Blue and purple rectangles show potential management actions (blue for aquaculture or stocked fisheries, and purple for wild fisheries).
Managing the threat of infectious disease in fisheries and aquaculture using structured decision making

December 2023

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43 Reads

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2 Citations

Fisheries and aquaculture provide food and economic security, especially in the developing world, but both face challenges from infectious disease. Here, we consider management of disease issues from a structured decision‐making perspective to examine how infectious disease can threaten seafood production and influence management decisions. For both wild fisheries and aquaculture, disease‐management objectives generally aim to mitigate the severity and economic burden of outbreaks. General management strategies include manipulating host densities, reducing system connectivity, conserving or improving habitat, and implementing direct treatments or some other biological interventions. To inform decisions, mathematical models can be used to explore disease dynamics and to forecast the potential effectiveness of alternative management actions. Developing and implementing disease‐management strategies also involve considering uncertainties and balancing competing stakeholder interests and risk tolerances. We conclude by outlining several steps for applying structured decision making that are broadly useful to decision makers facing issues related to disease.


Birds are not the only ones impacted by guidance to cease bird feeding

December 2023

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136 Reads

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3 Citations

1. Humans have a particularly strong connection with birds, driving the enormous popularity of residential bird feeding in much of the world. 2. We conducted a web search to document US state wildlife management agency responses to two recent avian disease outbreaks, finding that 23 agencies made recommendations to cease feeding wild birds in 2021–2022. 3. The psychological benefits of bird feeding for humans are well-documented but often overlooked in management decisions in response to avian disease outbreaks. 4. Likewise, ecological evidence does not necessarily support ceasing bird feeding to reduce the spread of every avian disease. 5. Ecological and social science need to be applied in tandem to ensure that well-intended guidance to cease feeding of birds does not have unintended consequences.


Mixed-species assemblages and disease: the importance of differential vector and parasite attraction in transmission dynamics

April 2023

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218 Reads

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2 Citations

Individuals from multiple species often aggregate at resources, group to facilitate defense and foraging, or are brought together by human activity. While it is well-documented that host-seeking disease vectors and parasites show biases in their responses to cues from different hosts, the influence of mixed-species assemblages on disease dynamics has received limited attention. Here, we synthesize relevant research in host-specific vector and parasite bias. To better understand how vector and parasite biases influence infection, we provide a conceptual framework describing cue-oriented vector and parasite host-seeking behaviour as a two-stage process that encompasses attraction of these enemies to the assemblage and their choice of hosts once at the assemblage. We illustrate this framework, developing a case study of mixed-species frog assemblages, where frog-biting midges transmit trypanosomes. Finally, we present a mathematical model that investigates how host species composition and asymmetries in vector attraction modulate transmission dynamics in mixed-species assemblages. We argue that differential attraction of vectors by hosts can have important consequences for disease transmission within mixed-species assemblages, with implications for wildlife conservation and zoonotic disease. This article is part of the theme issue ‘Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes’.


Defining post‐migratory nonbreeding movements. In each panel, breeding sites are shown as yellow polygons, stopover/staging sites as orange squares, and post‐migratory (i.e., winter) sites as blue circles. Polygon size represents stay duration at a given site. The distinctions of interest are shown with bold lines and bold font. (a) Post‐migratory nonbreeding movements can be primarily longitudinal, whereas migratory movements are primarily latitudinal. (b) Post‐migratory nonbreeding movements can be latitudinal if the first winter site is used for a relatively long duration but (c) would be considered stopover sites if they are used for the order of days instead of weeks or months. (d) Winter sites must be used after migration and during the winter season, and outside the spatial extent of an individual's breeding range. In this example, the first stopover site would otherwise meet the criteria for a winter site but is too close to the breeding site. All panels display a boreal migrant, but the same principles apply to austral migrants.
Taxonomic, dietary, and body‐mass distributions of species reported to undertake post‐migratory nonbreeding movements, compared with all documented migratory bird species in the Global Register of Migratory Species (GROMS). (a) The taxonomic distribution of species. Black bars show the number of species from each order that undertook post‐migratory movements. Gray bars show the proportion of migrants in GROMS in each order that were documented to undertake post‐migratory movements. Empty bars indicate that some species in that order were migratory, but none had documented post‐migratory nonbreeding movements. (b) Dietary categories from the Elton Traits database for each species. As in (a), black bars show the number of species in our data set and gray bars show the proportion of migrants in GROMS from each dietary category that have documented post‐migratory movements. (c) Body‐mass distributions of migratory species documented to undertake post‐migratory nonbreeding movements (black) and of species with no documented nonbreeding movements (red). Note the log scale of the x‐axis.
Examples of species that undertake post‐migratory nonbreeding movements, illustrating the diversity of taxa and geographic locations represented. Photos are all by R. Hall, with the exception of Wahlberg's Eagle (by Mike's Birds, Wikimedia Commons: https://commons.wikimedia.org/wiki/File:Wahlberg%27s_Eagle_(6609722249).jpg) and Pallid Swift (by Bogbumper, Wikimedia Commons: https://commons.wikimedia.org/wiki/File:Apuspallidus_9043.JPG).
Causes and associations of post‐migratory nonbreeding movements. Green bars indicate positive results, i.e., that a factor was associated with post‐migratory nonbreeding movements; orange bars indicate negative results. Pale bars indicate that an effect was proposed but was not tested analytically. A single study can appear in multiple categories. (a) Reported relationships between age or sex and post‐migratory nonbreeding movements. (b) Reported ecological causes of post‐migratory nonbreeding movements. Note the different y‐axis scales across panels.
Frequency of different tracking technologies used to document post‐migratory nonbreeding movements. Methods are sorted by the potential weight and modernity of the device.
Post‐migratory nonbreeding movements of birds: A review and case study

March 2023

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305 Reads

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15 Citations

Abstract Seasonal migrations are fascinating and ecologically important, but many migratory species are declining as climate change and land‐use change alter the habitats used by migrants across the annual cycle. While some migratory birds use a single wintering site, others undertake large‐scale post‐migratory movements during the nonbreeding season. Technological advances that enable tracking individual birds are uncovering more examples of post‐migratory nonbreeding movements. Documenting these movements is important for conservation, which requires understanding when and where migrants use habitats throughout their range. Here, we reviewed existing literature and collected information on the post‐migratory nonbreeding movements of 92 migratory bird species from 18 orders across six continents. Among these records, the most commonly reported drivers of movements were resource availability and climate. This strong dependence of post‐migratory nonbreeding movements on birds' abiotic and biotic environments suggests that environmental change will impact the patterns of these movements and potentially the fitness of species that undertake them. We also reviewed post‐migratory nonbreeding movements in North American‐breeding thrushes from the genus Catharus to examine the drivers of these movements in five closely related migratory species. We find that species that are less territorial are more likely to use multiple sites during the nonbreeding season; however, there is little evidence for dietary, evolutionary, or environmental differences between thrush species that move during winter and those that are stationary. While we believe our study represents the most comprehensive list of species exhibiting post‐migratory nonbreeding movements to date, biases in sampling, a lack of common terminology for these movements, and the still‐nascent availability of inexpensive, lightweight tracking devices mean that there are probably more populations that undertake such movements. Future research into the consequences of post‐migratory nonbreeding movements for individual fitness and ecosystem services would advance our understanding of their conservation importance and their evolution.


Citations (61)


... By quantifying spatial overlap in a way that aligns with likely routes of PUUV transmission, we expected that spatial overlap would predict infection probability. We also expected that space use would be influenced by food supplementation and anthelmintic treatment (Mistrick et al., 2024) and that demographic traits would be important drivers of the relationship between spatial overlap and infection probability. We predicted that increases in spatial overlap would more strongly increase infection probability in demographic groups that are more tolerant of overlap (i.e., non-reproductive voles, reproductive males) compared to those that are less tolerant of overlap (i.e., reproductive females). ...

Reference:

Ecological factors alter how spatial overlap predicts viral infection dynamics in wild rodent populations
Effects of food supplementation and helminth removal on space use and spatial overlap in wild rodent populations

... The absence of other hosts could be due to factors such as their resistance to infection, consumption of milkweeds with high cardenolide toxicity À known to suppress parasite replication À or the effect of unfavourable environmental conditions [e.g. extreme heat is known to kill spores of OE and inhibit within-host replication (Ragonese et al., 2024;Sanchez et al., 2021)]. ...

Extreme heat reduces host and parasite performance in a butterfly–parasite interaction

... Uzaktan algılama, otomatik tuzaklar ve çevresel izleme gibi teknolojilerin kullanımı, zararlı popülasyonlarının takibi ve olası salgınların öngörülmesi yeteneğini geliştirebilir (22). İklim ve çevre verilerini kullanan erken uyarı sistemleri, su ürünleri yetiştiricilerinin zararlılarla ilgili olası sorunları önceden görmesine ve önlem almasına yardımcı olabilir (23). Su ürünleri üreticilerine sürdürülebilir zararlı yönetimi uygulamaları konusunda eğitim verilmesi büyük önem taşımaktadır. ...

Managing the threat of infectious disease in fisheries and aquaculture using structured decision making

... Anecdotally, bird-feeding is undergoing rapid changes with urban by-laws and housing association rules banning the practice in some areas of Finland, but this trend is also seen in other parts of the world in response to avian-borne pathogens (Dayer et al., 2024). Therefore, Finland provides an ideal opportunity to investigate changes in bird-feeding behaviour. ...

Birds are not the only ones impacted by guidance to cease bird feeding

... B 378: 20220093 sociality influences the colonization and extinction rates of birds on islands. Finally, Trillo et al. [25] study predator eavesdropping on mating choruses, modelling how differences among species in their attractiveness and vulnerability influence parasite attack and disease transmission. ...

Mixed-species assemblages and disease: the importance of differential vector and parasite attraction in transmission dynamics

... Space use, or the manner in which an individual uses its environment, ultimately shapes how it interacts with the entire ecological community. Rather than being fixed, space use behaviors can vary substantially across space and time as environmental (e.g., food availability, habitat quality, mate availability) or social (e.g., social status, mating status) conditions change [4,[8][9][10][11][12][13]. Even stationary strategies like territoriality [11,[14][15][16] vary substantially with a range of behaviors that might include everything from defending multiple territories to showing varying degrees of site fidelity [17][18][19]. ...

Post‐migratory nonbreeding movements of birds: A review and case study

... Additionally, we characterized the observational process through statedependent detection probabilities (i.e., altered capture probability in case of infection) and the probability of detecting the infection, given that we did not blood sample each individual. We defined seven different observation events: not detected (0), non-infected capture during one secondary sampling occasion (1), non-infected capture during both secondary sampling occasions (2), infected capture during one secondary sampling occasion (3), infected capture during both secondary sampling occasions (4), infection status unknown captured during one secondary sampling occasion (5), and infection status unknown captured during both secondary sampling occasions (6). ...

Animal behavior and parasitism: Where have we been, where are we going?
  • Citing Chapter
  • August 2022

... Such parasite-induced changes in foraging can result in indirect, cascading effects on organisms consumed by the host. However, an alternative response is behavioural tolerance, where hosts maintain fitness-enhancing behaviours such as foraging despite infestation (Adelman & Hawley, 2016;Stephenson & Adelman, 2022). ...

Animal Behavior and Parasitism
  • Citing Article
  • August 2022

... While these factors remain important, there is growing interest in the role played by parasites and pathogens (hereafter parasites) in animal migration and other movement behaviours. This shift has sparked numerous empirical and theoretical studies as well as conceptual frameworks which strive to explain the disproportionate role these small creatures play in this flagship behaviour (Poulin & de Angeli Dutra, 2021;Binning et al. 2022;Hall et al. 2022). Indeed, there is now a rich body of literature using a range of conceptual, modelling and empirical approaches attempting to understand the interactions between migration and infection (Table 1 Binning et al., 2022). ...

Animal migration and infection dynamics: Recent advances and future frontiers
  • Citing Chapter
  • August 2022

... For example, enhanced forest cover can promote biodiversity and provide vital ecosystem services that support local livelihoods; however, this may also lead to conflicts over land use, particularly in areas prioritizing agricultural expansion or urban development [50]. In contrast, ecological disturbances resulting from the geographic expansion of human landuse activities, such as deforestation, may enhance human exposure to rodents and other suspected reservoir species found in primary forests [51]. This increased interaction can elevate the risk of zoonotic spillover and the emergence of infectious diseases, including mpox [19]. ...

Land reversion and zoonotic spillover risk