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Dharmarajan, Guha

Publications and source records attributed to Dharmarajan, Guha.

Raccoon density estimation from camera traps for raccoon rabies management

Abstract Density estimation for unmarked animals is particularly challenging, yet density estimates are often necessary for effective wildlife management. Raccoons ( Procyon lotor ) are the primary terrestrial wildlife reservoir for Lyssavirus rabies within the United States. The raccoon rabies variant (RRVV) is actively managed at landscape scales using oral rabies vaccination (ORV) within the eastern United States. To effectively manage RRVV, it is important to know the density of raccoons to appropriately scale the density of ORV baits distributed on the landscape. We compared methods to estimate raccoon densities from camera‐trap data versus more intensive capture‐mark‐recapture (CMR) estimates across 2 land cover types (upland pine and bottomland hardwood) in the southeastern United States during 2019 and 2020. We evaluated the effect of alternative camera configurations and durations of camera trapping on density estimates and used an N‐mixture model to estimate raccoon densities, including covariates on abundance and detection. We further compared different methods of scaling camera‐based counts, with the maximum number of raccoons seen on any given image within a day best explaining density. Camera‐trap density estimates were moderately correlated with CMR estimates ( r = 0.56). However, densities from camera‐trap data were more reliable when classifying category of density as an index used to inform management (83% correct when compared to CMR estimates), although the densities in our study fell into the 2 lowest density classes only. Using more cameras reduced bias and uncertainty around density estimates; however, if ≤6 camera traps were used at a site, a line transect approach proved less biased than a grid design. Camera trapping should be conducted for at least 3 weeks for more accurate estimates of raccoon population density in our study area (<5% bias). We show that camera‐trap data can be used to assign raccoon densities to management‐relevant density index bins, but more studies are needed to ensure reliability across a greater range of environmental conditions and raccoon densities.

Davis, Amy J.↗

Home range and resource selection of Virginia opossums in the rural southeastern United States

The Virginia opossum (Didelphis virginiana) has a rapidly expanding distribution in North America, but many aspects of its ecology remain relatively understudied, particularly in rural areas of its core range. We collected GPS telemetry data from 93 opossums in a rural, non-agricultural landscape in South Carolina, USA (2018–2019) to examine factors influencing space use and resource selection. Estimated male home ranges (99% utilization distributions) were on average 50% larger than those of females (mean home range 115.9 ± 103.7 ha vs 76.7 ± 75.0 ha). The home range size decreased on average by 20% with each 20% increase in deciduous land cover but was not affected by season or other landscape factors. Core area sizes (65% utilization distributions) were not influenced by sex (mean core area size 29.1 ± 23.7 ha and 22.4 ha ± 13.8 for males and females, respectively) or season, but the core area size decreased by 14% with each 400 m increase in distance from a permanent water source. Resource selection by opossums primarily occurred at the landscape level. Both males and females generally selected for wetlands while avoiding pine forests and developed/open areas, likely the result of differences in resource availability and predation risk between habitats. Opossums also tended to select for linear features such as unpaved roads and edge habitat, which may facilitate movement across the landscape. Finally, the home ranges we documented are among the largest recorded for opossums in the USA, likely the result of the relatively low resource abundance throughout our study area due to comparatively minimal anthropogenic influence.

59 BASIC BIOLOGICAL SCIENCES↗

Raccoon spatial ecology in the rural southeastern United States

The movement ecology of raccoons varies widely across habitats with important implications for the management of zoonotic diseases such as rabies. However, the spatial ecology of raccoons remains poorly understood in many regions of the United States, particularly in the southeast. To better understand the spatial ecology of raccoons in the southeastern US, we investigated the role of sex, season, and habitat on monthly raccoon home range and core area sizes in three common rural habitats (bottomland hardwood, upland pine, and riparian forest) in South Carolina, USA. From 2018–2022, we obtained 264 monthly home ranges from 46 raccoons. Mean monthly 95% utilization distribution (UD) sizes ranged from 1.05 ± 0.48 km 2 (breeding bottomland females) to 5.69 ± 3.37 km 2 (fall riparian males) and mean monthly 60% UD sizes ranged from 0.25 ± 0.15 km 2 (breeding bottomland females) to 1.59 ± 1.02 km 2 (summer riparian males). Males maintained home range and core areas ~2–5 times larger than females in upland pine and riparian habitat throughout the year, whereas those of bottomland males were only larger than females during the breeding season. Home ranges and core areas of females did not vary across habitats, whereas male raccoons had home ranges and core areas ~2–3 times larger in upland pine and riparian compared to bottomland hardwood throughout much of the year. The home ranges of males in upland pine and riparian are among the largest recorded for raccoons in the United States. Such large and variable home ranges likely contribute to elevated risk of zoonotic disease spread by males in these habitats. These results can be used to inform disease mitigation strategies in the southeastern United States.

59 BASIC BIOLOGICAL SCIENCES↗

Racoons ( Procyon Lotor ) Show Higher Trypanosoma Cruzi Detection Rates than Virginia Opossums ( Didelphis Virginiana ) in South Carolina, USA

Chagas disease, a significant public health concern in the Americas, is caused by a protozoan parasite, Trypanosoma cruzi. The life cycle of T. cruzi involves kissing bugs (Triatoma spp.) functioning as vectors and mammalian species serving as hosts. Raccoons (Procyon lotor) and opossums (Didelphis virginiana) have been identified as important reservoir species in the life cycle of T. cruzi, but prevalence in both species in the southeastern US is currently understudied. We quantified T. cruzi prevalence in these two key reservoir species across our study area in South Carolina, US, and identified factors that may influence parasite detection. We collected whole blood from 183 raccoons and 126 opossums and used PCR to detect the presence of T. cruzi. We then used generalized linear models with parasite detection status as a binary response variable and predictor variables of land cover, distance to water, sex, season, and species. Our analysis indicated that raccoons experienced significantly higher parasite detection rates than Virginia opossums, with T. cruzi prevalence found to be 26.5% (95% confidence interval [CI], 20.0–33.8) in raccoons and 10.5% (95% CI, 5.51–17.5) in opossums. Altogether, our results concur with previous studies, in that T. cruzi is established in reservoir host populations in natural areas of the southeastern US.

60 APPLIED LIFE SCIENCES↗

Raccoon densities across four land cover types in the southeastern United States

Raccoons (Procyon lotor) are the primary reservoir for rabies virus in eastern North America. Management of rabies in raccoons is achieved primarily with the use of oral rabies vaccination (ORV) and effective ORV bait densities are determined in part by the densities of raccoons. Decisions regarding ORV bait densities, however, are limited by an incomplete understanding of raccoon densities across the spectrum of landscapes they occupy. We carried out a mark-recapture study of raccoons on the Savannah River Site in South Carolina, USA, from 2017–2019, to develop sex- and landscape-specific raccoon density estimates across 4 rural land cover types in the southeastern United States: bottomland hardwood, riparian forest, isolated wetland, and upland pine (Pinus spp.). We captured 404 unique raccoons 773 times over the 3-year trapping period. Estimated densities were 5.44 ± 0.37 (SE) animals/km 2 in bottomland hardwood forest, 2.62 ± 0.32 animals/km 2 in riparian forest, 2.19 ± 0.29 animals/km 2 in isolated wetlands, and 2.14 ± 0.23 animals/km 2 in upland pine. Densities were significantly higher in bottomland hardwood than all other land cover types, whereas densities among the remaining cover types were similar. These patterns are likely the result of landscape fragmentation and configuration, with riparian forests typically embedded in a matrix of less suitable cover types, leading to low densities despite presumably high resource availability. There were higher densities of males than females in every cover type except upland pine, where the sex ratio was balanced. Densities on our site were low compared to other rural areas, which likely results from the lack of human influence in terms of agriculture or development. The financial cost of baiting for ORV distribution may be reduced by considering the comparatively low densities of raccoons in these rural landscapes in the southeastern United States.

59 BASIC BIOLOGICAL SCIENCES↗

Spatial ecology of translocated raccoons

Raccoons (Procyon lotor) are routinely translocated both legally and illegally to mitigate conflicts with humans, which has contributed to the spread of rabies virus across eastern North America. The movement behavior of translocated raccoons has important ramifications for disease transmission yet remains understudied and poorly quantified. To examine the spatial ecology of raccoons following experimental translocation, we performed reciprocal 16 km-distance translocations of 30 raccoons between habitats of high and low raccoon density (bottomland hardwood and upland pine, respectively) across the Savannah River Site (SRS) in Aiken, South Carolina, USA (2018–2019). Translocation influenced patterns of raccoon space use, with translocated animals exhibiting a 13-fold increase in 95% utilization distributions (UDs) post- compared to pre-translocation (mean 95% UD 35.8 ± 36.1 km 2 vs 1.96 ± 1.17 km 2 ). Raccoons originating from upland pine habitats consistently had greater space use and larger nightly movement distances post-translocation compared to raccoons moved from bottomland hardwood habitats, whereas these differences were generally not observed prior to translocation. Estimated home ranges of male raccoons were twice the area as estimated for female raccoons, on average, and this pattern was not affected by translocation. After a transient period lasting on average 36.5 days (SD = 30.0, range = 3.25–92.8), raccoons often resumed pre-experiment movement behavior, with 95% UD sizes not different from those prior to translocation (mean = 2.27 ± 1.63km 2 ). Most animals established new home ranges after translocation, whereas three raccoons moved > 16 km from their release point back to the original capture location. Four animals crossed a 100-m wide river within the SRS post-translocation, but this behavior was not documented among collared raccoons prior to translocation. Large increases in space use combined with the crossing of geographic barriers such as rivers may lead to elevated contact rates with conspecifics, which can heighten disease transmission risks following translocation. These results provide additional insights regarding the potential impacts of raccoon translocation towards population level risks of rabies outbreaks and underscore the need to discourage mesocarnivore translocations to prevent further spread of wildlife rabies.

59 BASIC BIOLOGICAL SCIENCES↗

Phylogeography of the blacklegged tick ( Ixodes scapularis ) throughout the USA identifies candidate loci for differences in vectorial capacity

Abstract The blacklegged tick ( Ixodes scapularis ( Journal of the Academy of Natural Sciences of Philadelphia , 1821, 2 , 59)) is a vector of Borrelia burgdorferi sensu stricto ( s.s .) ( International Journal of Systematic Bacteriology , 1984, 34 , 496), the causative bacterial agent of Lyme disease, part of a slow‐moving epidemic of Lyme borreliosis spreading across the northern hemisphere. Well‐known geographical differences in the vectorial capacity of these ticks are associated with genetic variation. Despite the need for detailed genetic information in this disease system, previous phylogeographical studies of these ticks have been restricted to relatively few populations or few genetic loci. Here we present the most comprehensive phylogeographical study of genome‐wide markers in I. scapularis , conducted by using 3RAD (triple‐enzyme restriction‐site associated sequencing) and surveying 353 ticks from 33 counties throughout the species' range. We found limited genetic variation among populations from the Northeast and Upper Midwest, where Lyme disease is most common, and higher genetic variation among populations from the South. We identify five spatially associated genetic clusters of I. scapularis . In regions where Lyme disease is increasing in frequency, the I. scapularis populations genetically group with ticks from historically highly Lyme‐endemic regions. Finally, we identify 10 variable DNA sites that contribute the most to population differentiation. These variable sites cluster on one of the chromosome‐scale scaffolds for I. scapularis and are within identified genes. Our findings illuminate the need for additional research to identify loci causing variation in the vectorial capacity of I. scapularis and where additional tick sampling would be most valuable to further understand disease trends caused by pathogens transmitted by I. scapularis .

3RAD↗

Assessment of habitat‐specific competition for oral rabies vaccine baits between raccoons and opossums

Abstract Throughout the eastern United States, the National Rabies Management Program (NRMP) distributes oral rabies vaccine (ORV) baits to manage rabies virus circulation in raccoon ( Procyon lotor ) populations. The consumption of vaccine baits by non‐target species including Virginia opossums ( Didelphis virginiana ) may reduce the effectiveness of ORV programs, but competition for baits remains poorly quantified in many areas of the southeastern United States. We distributed placebo ORV baits injected with a biomarker across 4 land cover types (bottomland hardwood, upland pine, riparian, isolated wetland) on the Savannah River Site in South Carolina, USA, 2017–2019. We then trapped and collected whiskers from 247 raccoons and 78 opossums to assess biomarker presence using fluorescent microscopy. Our data revealed greater bait uptake probability by raccoons (estimated x̅ = 0.30, 95% CI = 0.19–0.44) compared to opossums (estimated x̅ = 0.11, 95% CI = 0.05–0.23) across all cover types surveyed. Probability of bait consumption was not affected by cover type or the abundance of raccoons or opossums. Among raccoons, males were more likely to consume baits than females (estimated x̅ = 0.28, 95% CI = 0.17–0.44 for males and 0.14, 95% CI = 0.05–0.31 for females) and probability of consumption increased by 0.08 with each additional day trapped during the 10‐day trapping session. Uptake rates for raccoons were relatively low compared to other studies and not influenced by competition with opossums. These low consumption rates indicate that additional research addressing the roles of baiting season, bait density, and resource selection will be important to maximize ORV bait uptake by target species in these southeastern landscapes.

59 BASIC BIOLOGICAL SCIENCES↗

Interspecific oral rabies vaccine bait competition in the Southeast United States

Here, the United States Department of Agriculture’s National Rabies Management Program (NRMP) has coordinated the use of oral rabies vaccination (ORV) to control the spread of raccoon rabies virus variant west of the Appalachian Mountains since 1997. Working with state and local partners, the NRMP deploys ORV baits containing a rabies vaccine, primarily targeting raccoon populations (Procyon lotor). Bait competition between raccoons and non-target species may limit the effectiveness of ORV programs, but the extent of bait competition remains poorly quantified, particularly in the southeastern United States. We placed placebo ORV baits in bottomland hardwood (n = 637 baits) and upland pine (n = 681 baits) habitats in South Carolina, USA during August-December 2019 and used remote cameras to examine bait competition between raccoons and non-target species. The estimated proportion of bait consumed by raccoons was 18.8 ± 2.1% in bottomland hardwood and 11.6 ± 2.1% in upland pine habitats. Vertebrate competition appeared to have a minimal effect on raccoon uptake as estimated consumption did not exceed 5% for any species or 8% of bait uptake events cumulatively. We estimated that raccoons were the primary consumer of baits in bottomland hardwood, whereas invertebrates were the primary consumer in upland pine (26.7 ± 1.3% of baits). Our results indicate a need to closely consider the effects of invertebrates on bait consumption to minimize their potential impact on ORV bait uptake by target species. Uptake probabilities by raccoons were relatively low but not primarily driven by competition with vertebrates. As such, strategies to increase the specificity of raccoon uptake may be needed to enhance the effectiveness of ORV baiting programs.

60 APPLIED LIFE SCIENCES↗

Influence of landscape attributes on Virginia opossum density

Abstract The Virginia opossum ( Didelphis virginiana ), North America's only marsupial, has a range extending from southern Ontario, Canada, to the Yucatan Peninsula, Mexico, and from the Atlantic seaboard to the Pacific. Despite the Virginia opossum's taxonomic uniqueness in relation to other mammals in North America and rapidly expanding distribution, its ecology remains relatively understudied. Our poor understanding of the ecology of this important mesopredator is especially pronounced in the rural southeastern United States. Our goal was to estimate effects of habitat on opossum density within an extensive multi‐year spatial capture‐recapture study. Additionally, we compared the results of this spatial capture‐recapture analysis with a simple relative abundance index. Opossum densities in the relatively underdeveloped regions of the southeastern United States were lower compared to the more human‐dominated landscapes of the Northeast and Midwest. In the southeastern United States, Virginia opossums occurred at a higher density in bottomland swamp and riparian hardwood forest compared to upland pine ( Pinus spp.) plantations and isolated wetlands. These results reinforce the notion that the Virginia opossum is commonly associated with land cover types adjacent to permanent water (bottomland swamps, riparian hardwood). The relatively low density of opossums at isolated wetland sites suggests that the large spatial scale of selection demonstrated by opossums gives the species access to preferable cover types within the same landscape.

54 ENVIRONMENTAL SCIENCES↗

Evolution of pathogen tolerance and emerging infections: A missing experimental paradigm

Researchers worldwide are repeatedly warning us against future zoonotic diseases resulting from humankind’s insurgence into natural ecosystems. The same zoonotic pathogens that cause severe infections in a human host frequently fail to produce any disease outcome in their natural hosts. What precise features of the immune system enable natural reservoirs to carry these pathogens so efficiently? To understand these effects, we highlight the importance of tracing the evolutionary basis of pathogen tolerance in reservoir hosts, while drawing implications from their diverse physiological and life-history traits, and ecological contexts of host-pathogen interactions. Long-term co-evolution might allow reservoir hosts to modulate immunity and evolve tolerance to zoonotic pathogens, increasing their circulation and infectious period. Such processes can also create a genetically diverse pathogen pool by allowing more mutations and genetic exchanges between circulating strains, thereby harboring rare alive-on-arrival variants with extended infectivity to new hosts (i.e., spillover). Finally, we end by underscoring the indispensability of a large multidisciplinary empirical framework to explore the proposed link between evolved tolerance, pathogen prevalence, and spillover in the wild.

59 BASIC BIOLOGICAL SCIENCES↗

Global drivers of avian haemosporidian infections vary across zoogeographical regions

Macroecological analyses provide valuable insights into factors that influence how parasites are distributed across space and among hosts. Amid large uncertainties that arise when generalizing from local and regional findings, hierarchical approaches applied to global datasets are required to determine whether drivers of parasite infection patterns vary across scales. Here we assessed global patterns of haemosporidian infections across a broad diversity of avian host clades and zoogeographical realms to depict hotspots of prevalence and to identify possible underlying drivers.

haemosporidian prevalence↗

Solving the sample size problem for resource selection functions

Abstract Sample size sufficiency is a critical consideration for estimating resource selection functions (RSFs) from GPS‐based animal telemetry. Cited thresholds for sufficiency include a number of captured animals and as many relocations per animal N as possible. These thresholds render many RSF‐based studies misleading if large sample sizes were truly insufficient, or unpublishable if small sample sizes were sufficient but failed to meet reviewer expectations. We provide the first comprehensive solution for RSF sample size by deriving closed‐form mathematical expressions for the number of animals M and the number of relocations per animal N required for model outputs to a given degree of precision. The sample sizes needed depend on just 3 biologically meaningful quantities: habitat selection strength, variation in individual selection and a novel measure of landscape complexity, which we define rigorously. The mathematical expressions are calculable for any environmental dataset at any spatial scale and are applicable to any study involving resource selection (including sessile organisms). We validate our analytical solutions using globally relevant empirical data including 5,678,623 GPS locations from 511 animals from 10 species (omnivores, carnivores and herbivores living in boreal, temperate and tropical forests, montane woodlands, swamps and Arctic tundra). Our analytic expressions show that the required M and N must decline with increasing selection strength and increasing landscape complexity, and this decline is insensitive to the definition of availability used in the analysis. Our results demonstrate that the most biologically relevant effects on the utilization distribution (i.e. those landscape conditions with the greatest absolute magnitude of resource selection) can often be estimated with much fewer than animals. We identify several critical steps in implementing these equations, including (a) a priori selection of expected model coefficients and (b) regular sampling of background (pseudoabsence) data within a given definition of availability. We discuss possible methods to identify a priori expectations for habitat selection coefficients, effects of scale on RSF estimation and caveats for rare species applications. We argue that these equations should be a mandatory component for all future RSF studies.

Street, Garrett M.↗

Lessons Learnt From the COVID-19 Pandemic

The COVID-19 pandemic, caused by the novel coronavirus SARS-CoV-2, has been characterized by unprecedented rates of spatio-temporal spread. Here, we summarize the main events in the pandemic's timeline and evaluate what has been learnt by the public health community. We also discuss the implications for future public health policy and, specifically, the practice of epidemic control. We critically analyze this ongoing pandemic's timeline and contrast it with the 2002–2003 SARS outbreak. We identify specific areas (e.g., pathogen identification and initial reporting) wherein the international community learnt valuable lessons from the SARS outbreak. However, we also identify the key areas where international public health policy failed leading to the exponential spread of the pandemic. We outline a clear agenda for improved pandemic control in the future.

60 APPLIED LIFE SCIENCES↗