Supplemental feeding of wild ungulates has long been and remains a common practice across Europe and North America. Yet by drawing animals together, supplemental feeding can have unintended, negative effects on individual species and broader ecological processes. These include increased risk of disease transmission, intraspecific and interspecific competition, and predation, which are of management concern for white-tailed deer (Odocoileus virginianus) in the southeastern United States given the arrival of nonnative wild pigs (Sus scrofa) and coyotes (Canis latrans). We conducted a field experiment between March and July of 2021 to assess the effects of supplemental feeding on spatiotemporal activity patterns of deer and wild pigs at wildlife feeders, and space use of coyotes in the Piedmont region of South Carolina, USA. We observed support for our hypothesis that interspecific competition through increased visitation by larger groups of competitor species reduces use of foraging sites by other subordinate ungulates, where feeders highly visited by wild pigs were rarely visited by deer. While adult deer and wild pigs generally did not shift their temporal activity patterns at feeders, juvenile temporal activity shifted to more frequent visits of feeders during the night, supporting our hypothesis that supplemental feed could increase risk to predator exposure, as coyotes tend to be active during crepuscular hours. Our findings suggest that supplemental feed put out to encourage deer activity could actually deter deer if wild pigs occupy that area, and has potential negative demographic effects if juveniles are at increased risk of predation. Collectively, based on our data, we do not recommend supplemental feeding in the southeastern United States where white-tailed deer, coyotes, and wild pigs co-occur. More broadly, given how widespread the legal use of supplemental feed remains across the United States, we encourage landowners and policymakers to consider the full suite of potential direct and indirect, short-term and long-term negative impacts supplemental feeding can have on both target and nontarget wildlife populations.

Estimating survival and cause-specific mortality of male eastern wild turkeys (Meleagris gallopavo silvestris) is important for understanding population dynamics and implementing appropriate harvest management. To better understand age-specific estimates of annual survival and harvest rates, we captured and marked male wild turkeys with leg bands (n = 311) or bands and transmitters (n = 549) in Georgia, Louisiana, North Carolina, and South Carolina, USA, during 2014-2022. We fitted time to event models to data from radio-marked birds to estimate cause-specific mortality and annual survival. We used band recovery models incorporating both band recovery and telemetry data to further investigate harvest rates and survival. Annual survival from known-fate models in hunted populations was 0.54 (95% CI = 0.49-0.59) for adults and 0.86 (95% CI = 0.81-0.92) for juveniles. Cause-specific mortality analysis produced an annual harvest estimate of 0.29 (95% CI = 0.24-0.33) for adults and 0.02 (95% CI = 0.01-0.03) for juveniles, whereas predation was 0.15 (95% CI = 0.10-0.20) and 0.12 (95% CI = 0.08-0.17), respectively. Annual survival for adult males in a non-hunted population was 0.83 (95% CI = 0.72-0.97). Survival rate was negatively correlated with harvest rate, indicating harvest was an additive mortality source. Annual survival from band recovery models was 0.40 (95% CI = 0.37-0.44) for adults and 0.88 (95% CI = 0.81- 0.93) for juveniles, whereas annual harvest estimates were 0.24 (95% CI = 0.23-0.25) for adults and 0.04 (95% CI = 0.03-0.05) for juveniles. Both models suggested no differences in annual survival across years or among study areas, which included privately owned and public properties. Harvest was an additive mortality source for male wild turkeys, suggesting that managers interested in increasing annual survival of adult males could consider ways of reducing harvest rates.

Hunting causes direct mortality and potentially disrupts normal activities of game and non-game species. As spatial (i.e., selection of hunting areas) and temporal (i.e., only diurnally present) patterns of hunters can become predictable, hunted species may respond accordingly. Risk becomes more dynamic and complex for species that are hunted concurrent with their breeding cycle, and growing literature has noted that wild turkey (Meleagris gallopavo spp.) behaviors can be altered by hunting activity. We allocated global positioning system (GPS) units to 1,500 wild turkey hunters and affixed GPS transmitters to 175 wild turkeys during 2014-2018 on the Webb Wildlife Management Area Complex in South Carolina, USA. We evaluated whether wild turkeys shifted resource selection as a function of hunter resource selection during the progression of hunting seasons. Male wild turkeys avoided areas where stationary hunting bouts occurred during the early hunting season (and selected for these areas before hunting began), whereas females were more likely to select those areas avoided by males by the end of the hunting season. For every 15% increase in predicted probability of an area being hunted, male wild turkeys were 4.16 times less likely to select that area, whereas female wild turkeys were 1.08 times more likely to select that area relative to pre-season periods when hunters were not on the landscape. Hunting activity induced immediate responses by male wild turkeys as they sought refuge away from hunted areas. Coupled with recent research suggesting hunters are more influential than natural predators in reducing the frequency of male vocalizations and eliciting fleeing and avoidance behavior, our results indicate hunting activity could affect distribution and courtship behaviors of male wild turkeys during their breeding season. Risk of harvest is complex for species that are hunted concurrent with their breeding cycle, and growing literature has noted that wild turkey behavior can be altered by hunting activity. Our work shows that hunting activity induced immediate responses by male wild turkeys as they sought refuge away from hunted areas; however, female response was muted at best. Coupled with recent research suggesting hunters are more influential than natural predators in reducing the frequency of male vocalizations and eliciting fleeing and avoidance behavior, our results indicate hunting activity could affect distribution and courtship behaviors of male wild turkeys during their breeding season. image

Canis latrans (Coyote) is a highly mobile, generalist species that occupies a wide variety of habitats and ecoregions across North and Central America. Assessments of population genetic diversity and structure among the recently colonized populations of Coyotes in the eastern US have been conducted at broad spatial scales, revealing genetic structure concordant with immigration routes and high genetic diversity, or at local spatial scales, usually finding genetic panmixia. However, few studies have assessed eastern Coyote genetics at an intermediate spatial scale (i.e., state-level), which may be commensurate with the population dynamics of this mobile animal. We sampled 10 study areas across South Carolina during summer 2019-2020 using noninvasive genetic sampling and 10 microsatellite loci to identify individuals. We assessed genetic diversity and pairwise relatedness as well as genetic structure across our populations to determine whether there were landscape-level differences in genetic differentiation and the scale of Coyote population structure across the state. We found high levels of observed heterozygosity (0.599-0.872) and allelic richness (3.99-5.12) across our sites but low levels of relatedness. We observed low genetic differentiation (pairwise F-ST = 0.010-0.066) and insignificant isolation by distance with limited evidence of genetic structure. Coyote populations seem to be operating at a spatial extent greater than South Carolina, and gene flow is likely maintained by unrelated, transient individuals. Future research and management of Coyotes in the heterogeneous landscapes of the southeastern US should account for the geographic scale that populations encompass.

Predators impose top-down forces on prey populations, with the strength of those effects often varying over space and time and among demographic groups. In ungulates, predation risk is typically greatest for neonatal offspring, with some suggesting that predators can key in on adult activity to locate hidden neonates. However, few field studies to date have been able to directly assess the influence of maternal care on ungulate neonate survival. Using a population of white-tailed deer under heavy coyote predation pressure, we tested the maternal dispersion hypothesis, which suggests the dispersion of maternal activity temporally and spatially attenuates risk of predation for ungulate neonates during this vulnerable altricial phase. We compared support for this hypothesis with more commonly tested hypotheses regarding the influence of habitat conditions and intrinsic factors on neonatal survival. Fawn survival to 16 weeks was 27.7%, with coyotes accounting for 59% of fawn mortalities. In support of our maternal temporal diffusion hypothesis, we found that neonatal survival decreased as more maternal visits (proportionally) occurred at night. The only other significant (p < .1) predictor of fawn survival was birth timing, with fawn survival decreasing as the season progressed. Given that fawn survival declined as the proportion of nighttime visits increased, and that wild pig presence and human disturbance can push doe and fawn activity toward nocturnal hours, additional research is needed to determine whether managing pig and human disturbance can decrease fawn mortality. More broadly, given the importance of recruitment in ungulate population dynamics, our finding opens a potentially important new line of inquiry on how maternal behaviors influence predation risk in large animal predator-prey ecology.

Abstract Resource pulses are ecologically important phenomenon that occur in most ecosystems globally. Following optimal foraging theory, many consumers switch to pulsatile foods when available, examples of which include fruit mast and vulnerable young prey. Yet how the availability of resource pulses shapes the ecology of predators is still an emerging area of research; and how much individual variation there is in response to pulses is not well understood. We hypothesized that resource pulses would lead to dietary convergence in our population, which we tested by tracking both population‐level and individual coyote diets for 3 years in South Carolina, USA. We (1) described seasonal dietary shifts in relation to resource pulses; (2) compared male and female diets across seasons; and (3) tested this dietary convergence hypothesis by quantifying individual dietary variation both across and within periods when resource pulses were available. We found that pulses of white‐tailed deer fawns and blackberries composed over half of coyote diet in summer, and persimmon fruits were an important component in fall. Male and female coyotes generally had similar diets, but males consumed more deer in fall, perhaps driven by scavenging more. We found support for our dietary convergence hypothesis, where individuals had more similar diets during resource pulses compared to a non‐pulse period. We also found that this convergence happened before peak availability, suggesting a non‐symmetric response to pulse availability. We show that nearly all coyotes eat fawns, suggesting that targeted efforts to remove “fawn killers” would be in vain. Instead, given how quickly coyotes collectively converge on resource pulses, our findings show that resource pulses could potentially be used by managers to alter the behavior of apex predators. More broadly, we open a new line of inquiry into how variation in individual foraging decisions scales up to shape the effects of resource pulses on ecological communities.

Wild pigs (Sus scrofa), which are invasive in many regions globally, can alter ecosystems and compete with native species through interference competition and resource exploitation. Wild pig impacts on other species may increase with greater niche overlap, which could vary over time based on environmental conditions, resource availability, or biological traits like diet, especially as seasonal variation in wild pig diet has been widely documented. A limited number of studies have assessed spatial or temporal overlap between native species and invasive wild pigs, with only a handful simultaneously assessing overlap in these niche dimensions. We investigated the potential for interspecific interactions involving invasive wild pigs in the Piedmont region of South Carolina, USA, by examining seasonal spatiotemporal overlap with other wildlife using N-mixture models and diel activity overlap analyses. Site use by white-tailed deer (Odocoileus virginianus) and coyote (Canis latrans) was negatively associated with wild pig activity in the fall, when the species had high diel activity overlap, indicating spatial partitioning could reduce interference competition with wild pigs in this season. Conversely, white-tailed deer site use was positively associated with wild pig activity in the winter, suggesting higher spatial overlap may be necessary if resources are limited. Site use by bobcat (Lynx rufus) and nine-banded armadillo (Dasypus novemcinctus) in the spring, along with raccoon (Procyon lotor) and wild turkey (Meleagris gallopavo) site use in the summer, was positively associated with wild pig activity. With the exception of diurnal wild turkey, diel activity overlap between these species and wild pigs was high, although temporal partitioning could have occurred at finer spatiotemporal scales than we examined. Our results collectively emphasize the importance of accounting for seasonal spatial and temporal responses by individual species to invasive wild pigs, with special consideration given to species in seasons where high niche overlap with wild pigs is anticipated.

Coyotes (Canis latrans) colonized the eastern United States over the last century and formed a 3-species predator guild with bobcats (Lynx rufus) and gray foxes (Urocyon cinereoargenteus) across much of the southeastern United States. Diets among the three species vary along with respective impacts on game species such as white-tailed deer (Odocoileus virginianus) and wild turkeys (Meleagris gallopavo). To determine predation impacts on vertebrate prey and dietary overlap in consumption of prey items, we assessed diets of coyote, bobcat, and gray fox during spring, coinciding with white-tailed deer fawning and wild turkey nesting and brood rearing. We sampled across three sites along the Savannah River in South Carolina from mid-May through mid-June of 2020-2021. We collected 180 scat samples along 295.9 kilometers (71.1-122.4 km/site) of unpaved secondary roads and used DNA metabarcoding to determine vertebrate diet items. We identified predator species of scat using DNA metabarcoding and species-specific mtDNA fragment analysis (153 were coyote, 20 bobcat, and seven gray fox). Overall, we found evidence that two species, coyote and bobcat, consumed deer while all three consumed turkeys. Frequency of deer in the diet varied across sites for coyotes from 62-86% and wild turkey was present with a frequency of occurrence of 9% for coyotes, 5% for bobcats, and 14% for gray fox. Vertebrate diet specialization was evident across predator species with high frequency of deer in coyote diets, rabbits and small mammals in bobcat diets, and herpetofauna in gray fox diets. During deer fawning and wild turkey nesting and brood rearing, dietary overlap appears to be mediated by disparate selection of prey items, which reduced competition among coyotes, bobcats, and gray foxes. Use of DNA metabarcoding may augment our understanding of dietary preferences within this predator guild by providing increased resolution of diet composition among important game species.

Environmental disturbances driven by global climate variability continue to increase in both frequency and intensity. Recent studies on wild turkeys (Meleagris gallopavo) have documented both direct and indirect demographic effects of short-term, climatically driven events; however, only limited information is available on the effects of extreme environmental disturbance events on turkey demography. Thus, we evaluated space use and movements of eastern wild turkeys (M. g. silvestris; n = 20) and Rio Grande wild turkeys (M. g. intermedia; n = 22) during hurricane events in South Carolina and Texas, respectively. Hurricane Matthew occurred over a 3-day period in October 2016 impacting ongoing turkey research on the Webb Wildlife Management Area Complex in South Carolina, whereas Hurricane Harvey occurred over a 5-day period in August 2017 impacting an ongoing 6-county study region in Texas. We documented 1 and 2 direct mortalities during Hurricanes Matthew and Harvey, respectively. Mean core area size used by turkeys decreased by 75% during both hurricane events. We did not document evidence of changes in roost fidelity or distance between daily roost clusters after either hurricane. We documented direct mortality of 5 and 8% of the study population during Matthew and Harvey, respectively, and turkeys exhibited a gradient of behavioral responses dependent on hurricane intensity and duration. Because hurricanes can produce substantive modifications to areas used by wild turkeys, we suggest that managers consider initiatives to document legacy effects of extreme environmental disturbance events on local wild turkey demographics, including loss of nesting, brooding and roosting sites.

ABSTRACTWild turkeys (Meleagris gallopavo spp., hereafter turkey) are the second most pursued big game species in the United States. Turkey hunting occurs primarily during spring, and on publicly owned lands managers often monitor hunter numbers and harvest as components of managing hunter opportunity and satisfaction. Contemporary research has shown that hunting activity on public lands can influence male turkey behavior; hence, research detailing hunter behaviors is needed to better support informed management. We allocated 1,500 Global Positioning System units to hunters pursuing male turkeys during the spring hunting seasons of 2014–2018 on the Webb Wildlife Management Area Complex in South Carolina, USA. Mean number of unique hunting bouts per hunter per day was 2 (SE = 0.03, range = 1–8), whereas average time spent hunting was 230.1 minutes (SE = 4.6, range = 11.3–872.6)/hunter/day. Hunting effort was focused during the first half of hunting season, with 70.2% of the total time spent hunting occurring during the first 2 weeks of the season (~1–15 Apr). Mean distance a hunter traveled was 2,171 m (SE = 38.1, range = 10–20,685)/hunting bout. Almost 90% of hunting activity occurred between 0500 and 1200, 76% occurred before 1000. Access to roads appeared to be the primary driver of hunter movements, with 40.1% of all hunter locations <25 m from the nearest main or secondary road. On average, hunters were 480 m (SE = 0.5, range = 0–2,132) from main roads that allowed vehicle traffic, but only 123 m (SE = 0.3, range = 0–1,990) from secondary roads allowing only foot traffic. Hunters spent 54.3% of hunting bouts actively hunting and 45.7% stationary. The amount of time spent active decreased as the hunting season progressed. Hunter effort decreased significantly as the hunting season progressed, indicated by a decline in numbers of hunters and total time spent hunting. Understanding how and when turkey hunters move about the landscape is important for effectively managing hunting access and opportunity on public hunting lands. © 2020 The Wildlife Society.

ABSTRACTEastern wild turkeys (Meleagris gallopavo silvestris) use a polygynous‐promiscuous mating system, wherein males compete for mating opportunities and communicate with females via courtship behaviors. One courtship behavior is vocalization (gobbling), which attracts females and signals dominance to other males. However, temporal variation in gobbling activity may be influenced by external stimuli, environmental variation, and hunter activity. Gobbling activity is a key determinant of hunter satisfaction, and gobbling chronology is often used by state agencies to inform regulatory processes. To identify factors influencing gobbling activity, we evaluated daily gobbling chronology on 3 sites in South Carolina, USA (Webb Wildlife Management Area [WMA] Complex, Savannah River Site, Crackerneck WMA) with different levels of hunter activity. We used autonomous recording units (ARUs; n = 45) across 8,280 days to collect 53,937 hours of ambient sound recordings and identified 68,426 gobbles. Gobbling activity varied daily and site interacting with minutes since sunrise best predicted daily gobbling activity. We noted distinct differences in predicted numbers of gobbles between hunted sites and an unhunted site, suggesting that hunting may be an important determinant of gobbling activity. Across our study sites, we observed that ≥72% of gobbling activity occurred between 30 minutes before and 60 minutes after sunrise. We found no clear evidence of well‐defined unimodal or bimodal peaks in daily or weekly gobbling activity. Across sites, <44% of gobbling activity occurred during legal hunting seasons in South Carolina, with between 30% and 48% of gobbling activity occurring after legal hunting seasons. Because hunter satisfaction is primarily influenced by gobbling activity, wildlife managers in South Carolina may consider adjusting dates of turkey hunting seasons to correspond hunting with periods when most gobbling occurs. © 2018 The Wildlife Society.

Phenology is an important dimension of vertebrate reproductive strategies. Characterizing a species' reproductive phenology is a fundamental step in directing studies of influences on reproduction within and among populations throughout its range. Eastern Diamond-Backed Rattlesnakes (Crotalus adamanteus; EDBs) are endemic predators of the imperiled longleaf pine (Pinus palustris) ecosystem of the southeastern United States, and the species currently is under review for listing under the Endangered Species Act. Conservation and management of this imperiled species will greatly benefit from research on its basic biology, and particularly from information that will assist management for adult survival and population viability. Here, we present long-term data on EDB breeding and reproductive phenology from several populations in the South Carolina Coastal Plain. We summarized the phenology of EDB breeding and reproduction using observations of free-ranging, radiotelemetered rattlesnakes in South Carolina from 1997 to 2011. Breeding activity occurred from 10 July to 14 October, with a mean of 28 August. There was one outlying observation of spring breeding (18 March). Females moved to birthing sites as early as 26 June and as late as 28 August, with a mean of 15 August. Neonates were born around 31 August, with birth dates between 10 August and 14 September. We also noted six unique observations of apparent clutch size: 8, 10, 11, 11, 12, and 13 neonates. This information will contribute both to our current knowledge and conservation efforts of EDBs, and also facilitate future long-term comparative investigations throughout their geographic range.