Evaluating the effect of predators on white-tailed deer: Movement and diet of coyotes

Coyotes (Canis latrans) may affect adult and neonate white-tailed deer (Odocoileus virginianus) survival and have been implicated as a contributor to the decline of deer populations. Additionally, coyote diet composition is influenced by prey availability, season, and region. Because coyote movement and diet vary by region, local data are important to understand coyote population dynamics and their impact on prey species. In southeast Minnesota, we investigated the effect of coyotes on white-tailed deer populations by documenting movement rates, distances moved, and habitats searched by coyotes during fawning and nonfawning periods. Additionally, we determined survival, cause-specific mortality, and seasonal diet composition of coyotes. From 2001 to 2003, we captured and radiocollared 30 coyotes. Per-hour rate of movement averaged 0.87 km and was greater (P = 0.046) during the fawning (1.07 km) than the nonfawning period (0.80 km); areas searched were similar (P = 0.175) between seasons. Coyote habitat use differed during both seasons; habitats were not used in proportion to their availability (P < 0.001). Croplands were used more (P < 0.001) than their proportional availability during both seasons. Use of grasslands was greater during the fawning period (P = 0.030), whereas use of cropland was greater in the nonfawning period (P < 0.001). We collected 66 fecal samples during the nonfawning period; coyote diets were primarily composed of Microtus spp. (65.2%), and consumption of deer was 9.1%. During the study, 19 coyotes died; annual survival rate range was 0.33–0.41, which was low compared with other studies. Consumption of deer was low and coyotes searched open areas (i.e., cropland) more than fawning areas with dense cover. These factors in addition to high coyote mortality suggested that coyote predation was not likely limiting white-tailed deer populations in southeast Minnesota. © 2011 The Wildlife Society.     

Predation by coyotes on white-tailed deer neonates in South Carolina

Coyotes (Canis latrans) are novel predators throughout the southeastern United States and their depredation of white-tailed deer (Odocoileus virginianus) neonates may explain observed declines in some deer populations in the region, but direct evidence for such a relationship is lacking. Our objective was to quantify neonate survival rates and causes of mortality at the United States Department of Energy's Savannah River Site (SRS), South Carolina to directly evaluate degree of predation in this deer population. From 2006 to 2009, we radio-monitored 91 neonates captured with the aid of vaginal implant transmitters in pregnant adult females and opportunistic searches. Overall Kaplan–Meier survival rate to 16 weeks of age was 0.230 (95% CI = 0.155–0.328), and it varied little among years. Our best-fitting model estimated survival at 0.220 (95% CI = 0.144–0.320). This model included a quadratic time trend variable (lowest survival rate during the first week of life and increasing to near 1.000 around week 10), and Julian date of birth (survival probability declining as date of birth increased). Predation by coyotes was the most frequent cause of death among the 70 monitored neonates that died, definitively accounting for 37% of all mortalities and potentially accounting for as much as 80% when also including probable coyote predation. Predation by bobcats (Felis rufus) accounted for 7% (definitive) to 9% (including probable bobcat predation) of mortalities. The level of coyote-induced mortality we observed is consistent with the low recruitment rates exhibited in the SRS deer population since establishment of coyotes at the site. If representative of recruitment rates across South Carolina, current harvest levels appear unsustainable. This understanding is consistent with the recent declining trend in the statewide deer population. The effects of coyote predation on recruitment should be considered when setting harvest goals, regardless of whether local deer population size is currently above or below desired levels, because coyotes can substantially reduce fawn recruitment. Published 2012. This article is a U.S. Government work and is in the public domain in the USA.     

Widespread occurrence of a domestic dog mitochondrial DNA haplotype in southeastern US coyotes

Sequence analysis of the mitochondrial DNA control region from 112 southeastern US coyotes (Canis latrans) revealed 12 individuals with a haplotype closely related to those in domestic dogs. Phylogenetic analyses grouped this new haplotype in the dog/grey wolf (Canis familiaris/Canis lupus) clade with 98% bootstrap support. These results demonstrate that a male coyote hybridized with a female dog, and female hybrid offspring successfully integrated into the coyote population. The widespread distribution of this haplotype from Florida to West Virginia suggests that the hybridization event occurred long ago before the southeastern USA was colonized by coyotes. However, it could have occurred in the southeastern USA before the main front of coyotes arrived in the area between male coyotes released for sport and a local domestic dog. The introgression of domestic dog genes into the southeastern coyote population does not appear to have substantially affected the coyote's genetic, morphological, or behavioural integrity. However, our results suggest that, contrary to previous reports, hybridization can occur between domestic and wild canids, even when the latter is relatively abundant. Therefore, hybridization may be a greater threat to the persistence of wild canid populations than previously thought.     

White‐tailed deer exploit temporal refuge from multi‐predator and human risks on roads

Although most prey have multiple predator species, few studies have quantified how prey respond to the temporal niches of multiple predators which pose different levels of danger. For example, intraspecific variation in diel activity allows white‐tailed deer (Odocoileus virginianus) to reduce fawn activity overlap with coyotes (Canis latrans) but finding safe times of day may be more difficult for fawns in a multi‐predator context. We hypothesized that within a multi‐predator system, deer would allocate antipredation behavior optimally based on combined mortality risk from multiple sources, which would vary depending on fawn presence. We measured cause‐specific mortality of 777 adult (>1‐year‐old) and juvenile (1–4‐month‐old) deer and used 300 remote cameras to estimate the activity of deer, humans, and predators including American black bears (Ursus americanus), bobcats (Lynx rufus), coyotes, and wolves (Canis lupus). Predation and vehicle collisions accounted for 5.3 times greater mortality in juveniles (16% mortality from bears, coyotes, bobcats, wolves, and vehicles) compared with adults (3% mortality from coyotes, wolves, and vehicles). Deer nursery groups (i.e., ≥1 fawn present) were more diurnal than adult deer without fawns, causing fawns to have 24–38% less overlap with carnivores and 39% greater overlap with humans. Supporting our hypothesis, deer nursery groups appeared to optimize diel activity to minimize combined mortality risk. Temporal refuge for fawns was likely the result of carnivores avoiding humans, simplifying diel risk of five species into a trade‐off between diurnal humans and nocturnal carnivores. Functional redundancy among multiple predators with shared behaviors may partially explain why white‐tailed deer fawn predation rates are often similar among single‐ and multi‐predator systems.     

Survival of White-Tailed Deer Fawns in Southern Illinois

Abstract: Survival of white-tailed deer (Odocoileus virginianus) fawns has been quantified throughout much of North America. However, few studies have assessed the influence of intrinsic factors (e.g., fawn age and birth mass) and habitat on fawn survival. During 2002-2004, we captured and radiocollared 166 fawns in southern Illinois, USA, to estimate survival rates, determine causes of mortality, and identify factors influencing fawn survival. We used a known fates model in program MARK to estimate survival rates and compare explanatory models based on Akaike's Information Criterion corrected for small sample sizes (AIC_c). We developed 2 candidate sets of a priori models to quantify factors influencing fawn survival: model set 1 included intrinsic factors and model set 2 focused on habitat variables. We recorded 64 mortalities and the overall survival rate was 0.59 (95% CI = 0.51-0.68). Predation was the leading source of mortality (64%) and coyotes (Canis latrans) were the most prominent predator. For model set 1, model {S_{age X year}} had the lowest AIC_c value suggesting that the age at mortality varied among capture years. For model set 2, model {S_{landscape+forest}} had the lowest AIC_c value and indicated that areas inhabited by surviving fawns were characterized by a few large (i.e., > 5 ha) irregular forest patches adjacent to several small nonforest patches, and survival areas also contained more edge habitat than mortality areas. Due to the magnitude of coyote predation, survival areas could have represented landscapes where coyotes were less effective at locating and capturing fawns when compared to mortality areas. This study was the first account of macrohabitat characteristics directly influencing fawn survival. Wildlife managers can use this information to determine how habitat management activities may affect deer populations.     

Using faecal DNA sampling and GIS to monitor hybridization between red wolves (Canis rufus) and coyotes (Canis latrans)

The US Fish and Wildlife Service's (USFWS) Red Wolf Recovery Program recognizes hybridization with coyotes as the primary threat to red wolf recovery. Efforts to curb or stop hybridization are hampered in two ways. First, hybrid individuals are difficult to identify based solely on morphology. Second, managers need to effectively search 6000 km(2) for the presence of coyotes and hybrids. We develop a noninvasive method to screen large geographical areas for coyotes and hybrids with maternal coyote ancestry by combining mitochondrial DNA sequence analysis of faeces (scat) and geographic information system (GIS) technology. This method was implemented on the Alligator River National Wildlife Refuge (1000 km(2)) in northeastern North Carolina. A total of 956 scats were collected in the spring of 2000 and 2001 and global positioning system (GPS) coordinates were recorded. Seventy-five percent of the scats were assigned to species and five coyote/hybrid scats were detected. Placement of scat location coordinates on a map of the experimental population area revealed that four of the coyote/hybrid scats were detected within the home ranges of sterilized hybrids. The other coyote/hybrid scat indicated the presence of a previously unknown individual. We suggest this method be expanded to include more of the experimental population area and be optimized for use with nuclear markers to improve detection of hybrid and back-crossed individuals.     

Effects of large-scale removal of coyotes on pronghorn and mule deer productivity and abundance

We tested the hypothesis that predation by coyotes (Canis latrans) impacts pronghorn (Antilocapra americana) and mule deer (Odocoileus hemionus) populations. We did so by examining the effects of coyote removal on pronghorn and mule deer populations within 12 large areas (>10,500 km²) located in Wyoming and Utah during 2007 and 2008. Pronghorn productivity (fawn to adult female ratio) and abundance were positively correlated with the number of coyotes removed and removal effort (hours spent hunting coyotes from aircraft) although the correlation between pronghorn productivity and removal effort was not statistically significant (P = 0.08). Mule deer productivity and abundance were not correlated with either the number of coyotes removed or removal effort. Coyote removal conducted during the winter and spring provided greater benefit than removals conducted during the prior fall or summer. Our results suggest that coyote removal conducted over large areas increases fawn survival and abundance of pronghorn but not mule deer. © 2011 The Wildlife Society.     

Population structure of California coyotes corresponds to habitat-specific breaks and illuminates species history

Little is known about the relationship between animal movements and the emergent structure of populations, especially for species occupying large continuous distributions. Some such mammals disperse disproportionately into habitat similar to their natal habitat, a behavioural bias that might be expected to lead to habitat-conforming genetic structure. We hypothesized that coyotes (Canis latrans) would exhibit such natal-biased dispersal, and used 13 microsatellite loci to test, correspondingly, whether genetic structure conformed to major habitat breaks. First, we used a model-based approach to assign coyote genotypes to distinct genetic clusters irrespective of geographical location. Visualization on a geographical information system revealed a strong concordance between the locations of cluster assignments and habitat bioregions, not explainable in terms of physical dispersal barriers or intervening low-quality habitat. Next, we used a multiple Mantel test, which controlled for effects of geographical distance (itself, marginally significant; P = 0.06), to statistically determine that genetic distance was indeed higher between than within bioregions (P < 0.001). Whereas previously published examples of landscape effects on gene flow have typically been explainable in terms of species-wide habitat affinities or dispersal barriers, our finding that genetic subdivisions were associated with unobstructed boundaries between contiguous habitats suggests a role for intraspecific variability in habitat affinities as a factor underlying genetic structure. In addition, our data combined with previously published data suggest a pattern of genetic isolation-by-distance throughout western North America, consistent with independent evidence that the western half of the coyote range predates European settlement.     

Can Coyotes Affect Deer Populations in Southeastern North America?

ABSTRACT The coyote (Canis latrans) is a recent addition to the fauna of eastern North America, and in many areas coyote populations have been established for only a decade or two. Although coyotes are known predators of white-tailed deer (Odocoileus virginianus) in their historic range, effects this new predator may have on eastern deer populations have received little attention. We speculated that in the southeastern United States, coyotes may be affecting deer recruitment, and we present 5 lines of evidence that suggest this possibility. First, the statewide deer population in South Carolina has declined coincident with the establishment and increase in the coyote population. Second, data sets from the Savannah River Site (SRS) in South Carolina indicate a new mortality source affecting the deer population concurrent with the increase in coyotes. Third, an index of deer recruitment at SRS declined during the period of increase in coyotes. Fourth, food habits data from SRS indicate that fawns are an important food item for coyotes during summer. Finally, recent research from Alabama documented significant coyote predation on fawns there. Although this evidence does not establish cause and effect between coyotes and observed declines in deer recruitment, we argue that additional research should proactively address this topic in the region. We identified several important questions on the nature of the deer—coyote relationship in the East.     

Resident and transient coyotes exhibit differential patterns of movement behavior across heterogeneous landscapes in the southeastern United States

Coyotes (Canis latrans) are a highly adaptable canid species whose behavioral plasticity has allowed them to persist in a wide array of habitats throughout North America. As generalists, coyotes can alter movement patterns and change territorial strategies between residency (high site fidelity) and transiency (low site fidelity) to maximize fitness. Uncertainty remains about resident and transient coyote movement patterns and habitat use because research has reached conflicting conclusions regarding patterns of habitat use by both groups. We quantified effects of habitat on resident and transient coyote movement behavior using first passage time (FPT) analysis, which assesses recursive movement along an individual''s movement path to delineate where they exhibit area‐restricted search (ARS) behaviors relative to habitat attributes. We quantified monthly movement rates for 171 coyotes (76 residents and 53 transients) and then used estimated FPT values in generalized linear mixed models to quantify monthly habitat use for resident and transient coyotes. Transients had greater movement rates than residents across all months except January. Resident FPT values were positively correlated with agricultural land cover during fall and winter, but negatively correlated with agriculture during spring. Resident FPT values were also negatively correlated with developed habitats during May–August, deciduous land cover during June–August, and wetlands during September–January except November. FPT values of transient coyotes were positively correlated with developed areas throughout much of the year and near wetlands during July–September. Transient FPT values were negatively correlated with agriculture during all months except June and July. High FPT values (ARS behavior) of residents and transients were generally correlated with greater densities of edge habitat. Although we observed high individual variation in space use, our study found substantive differences in habitat use between residents and transients, providing further evidence that complexity and plasticity of coyote habitat use is influenced by territorial strategy.     

Survival of Neonatal White-Tailed Deer in an Exurban Population

Abstract: As humans continue to move further from the urban epicenter and expand into suburban and exurban areas, problems involving coexistence of wildlife and human populations will become increasingly common. Wildlife biologists will be tasked with reducing wildlife-human conflicts, and their effectiveness will be a function of their understanding of the biology and life-history characteristics of wildlife populations residing in areas with high human density. In this study, we examined causes and timing of deaths of neonatal white-tailed deer (Odocoileus virginianus) in an exurban area of Alabama in 2004 and 2005, estimated survival rates, and determined factors that influenced survival for the initial 8 weeks of life. We found 67% mortality, with the leading causes being predation by coyotes (Canis latrans; 41.7%)and starvation due to abandonment (25%). These results suggest that coyote predation may be a significant source of natural mortality in exurban areas. Contrary to our original expectations, vehicle collisions were not an important cause of mortality.     

Interference competition between coyotes and raccoons: a test of the mesopredator release hypothesis

Some predator species appear to conform to the mesopredatorrelease hypothesis (MRH), in which larger predators help limitpopulations of smaller predators. This hypothesis has been usedto explain the possible relationship between coyotes, mesopredators,and resultant cascades involving nonpredators. However, relationshipsbetween coyotes and noncanid mesopredators are poorly understood,and predictions from the MRH have rarely been rigorously tested.We monitored sympatric raccoon and coyote populations to assess2 predictions derived from the MRH: coyote predation is an importantcause of mortality in raccoon populations or raccoons avoidareas used by coyotes. Between March 2000 and September 2001,we recorded 3553 locations for 27 radio-collared raccoons and1393 locations for 13 coyotes captured on the Max McGraw WildlifeFoundation in Illinois, USA. No raccoon mortality from coyotepredation was observed during the study, and raccoon survivalwas >0.7 each season. All raccoon 95% home ranges exhibitedoverlap with 95% coyote home ranges in each season. The meanproportion of raccoon locations within 95% coyote home rangesdid not vary by sex but did vary by season. Raccoon overlapof coyote core areas varied considerably among individuals withinseasons, ranging from 0% to 83%. However, 45% of raccoons had<10% overlap with coyote core areas, whereas only 14% ofraccoons exhibited >50% overlap. Mean overlap with core areasdid not vary by season or sex. For those raccoons with homeranges overlapping coyote core areas, mean proportion of observedraccoon locations within coyote core areas was generally greaterthan the mean proportion of random locations. Scent-stationexperiments failed to document raccoon avoidance of specificsites that had been marked with coyote urine. We did not findsupport for a mortality prediction or avoidance prediction tosupport MRH with regard to raccoons and coyotes. These resultssuggest that relationships among mammalian predators may notbe simply dictated by body size, particularly for species outsidethe Canidae.     

Fine‐scale movements and behaviors of coyotes (Canis latrans) during their reproductive period

In canids, resident breeders hold territories but require different resources than transient individuals (i.e., dispersers), which may result in differential use of space, land cover, and food by residents and transients. In the southeastern United States, coyote (Canis latrans) reproduction occurs during spring and is energetically demanding for residents, but transients do not reproduce and therefore can exhibit feeding behaviors with lower energetic rewards. Hence, how coyotes behave in their environment likely differs between resident and transient coyotes. We captured and monitored 36 coyotes in Georgia during 2018–2019 and used data from 11 resident breeders, 12 predispersing residents (i.e., offspring of resident breeders), and 11 transients to determine space use, movements, and relationships between these behaviors and landcover characteristics. Average home range size for resident breeders and predispersing offspring was 20.7 ± 2.5 km² and 50.7 ± 10.0 km², respectively. Average size of transient ranges was 241.4 ± 114.5 km². Daily distance moved was 6.3 ± 3.0 km for resident males, 5.5 ± 2.7 km for resident females, and 6.9 ± 4.2 km for transients. We estimated first‐passage time values to assess the scale at which coyotes respond to their environment, and used behavioral change‐point analysis to determine that coyotes exhibited three behavioral states. We found notable differences between resident and transient coyotes in regard to how landcover characteristics influenced their behavioral states. Resident coyotes tended to select for areas with denser vegetation while resting and foraging, but for areas with less dense vegetation and canopy cover when walking. Transient coyotes selected areas closer to roads and with lower canopy cover while resting, but for areas farther from roads when foraging and walking. Our findings suggest that behaviors of both resident and transient coyotes are influenced by varying landcover characteristics, which could have implications for prey.