My friends at Coyote Yahoo Groups supplied me with a copy of Dr. Jacqueline Frair and Dr. Mark Teece's on-going Adirondack NY Eastern Coyote(Coywolf) Study outline ............The goal of their research is to understand the mechanisms by which coyotes affect biodiversity through competition with native predators and predation on white-tailed deer....Tangible products from this research will be at least two manuscripts in peer-reviewed journals, presentations at regional and national conferences, and a new collaboration among faculty of different disciplines at SUNY-ESF. Intangible produces include training for at least one graduate student (S. Warsen), outreach on carnivore ecology in the Adirondacks through interactions with trappers and land managers, and field and lab experience for the cadre of undergraduate volunteers interested in studying carnivore ecology ......We anxiously await final results from this very relevant Coywolf research

Proposal to the 2010 NSRC Competition

 Project Title:
Evolving niche of the "coy-wolf" in northeastern forests and implications for biodiversityLead Principal Investigator: Dr. Jacqueline L. Frair Department of Environmental and Forest Biology State University of New York – College of Environmental Science and Forestry 405 Illick Hall, 1 Forestry Drive, Syracuse, New York 13210 Phone: 315-470-4905, FAX: 315-470-6934, Email:

 Dr. Mark Teece Department of Chemistry, State University of New York – College of Environmental Science and Forestry 415 Jahn Laboratory, 1 Forestry Drive, Syracuse, New York 13210 Phone: 315-470-4736, FAX: 315-470-6856, Email:

 Stacy McNulty Paul Jensen Adirondack Ecological Center NYS Dept. of Environmental Conservation 6312 State Route 28N 232 Hudson Street Extension, PO Box 220 Newcomb, NY 12852 Warrensburgh, NY 12885-0220 Phone: 518-582-4551, FAX: 518-582-2181 Phone: 518-623-1240, FAX: 518-623-3603 Email:

 Coyotes

 When Europeans first settled North America, the northeastern United States was home to wolves (

 The earliest records of coyotes in the northeastern U.S. occurred in 1925 in New York State2. Over the past few decades coyotes have become the most ubiquitous large carnivore throughout the Northeast, and they are here to stay. Something unique about coyotes in the Northeast is that they share genes, morphology, and (potentially) behavior with wolves, having interbred with wolves in eastern Canada prior to colonizing the northeastern states3. The ecological role of these "coy-wolves" in the Northeast remains of great interest to scientists, resource managers, and the general public alike4.
 5-8. Native predators in the Northern Forest that compete with coyotes include red fox (Vulpes vulpes), gray fox (Urocyon cineroargenteus), and bobcat (Lynx rufus). The diet of these species overlap that of coyotes9, although not completely, and reductions in one predator population will reduce predation pressure on some prey species with cascading effects to lower trophic levels5. Sorting out the potentially cascading effects of interference competition among predators in the Northern Forest begins fundamentally with a clear indication of the dietary breadth of the coyote and its overlap with native predators.
Following the second pathway, predation by coyotes can directly affect their prey species with cascading effects to lower trophic levels
 17 compared to 88% in 1975-8018 and 94% in 1986-8919. This trend may indicate niche specialization with coyotes evolving to efficiently hunt deer, but it also may reflect changes in the abundance of snowshoe hare (Lepus americanus), which declined in the diet from 33, to 13, to 1% over the same time period.
To regulate prey numbers, predators must largely specialize on a particular species and kill a significant fraction of their population. In the central Adirondack region of New York State, the diet of coyotes has become increasingly focused on white-trailed deer, with deer comprising only 39% of the diet in 1956-61

 In more northerly (and westerly) regions, snowshoe hare is the primary component of coyote, red fox and bobcat diets9, and coyotes demonstrate prey-switching when hare populations are low20,21.

Additional years of study combined with information on deer and hare population trends are required to address the question of coyote specialization on deer. Nevertheless a large proportion of coyote diets consist of deer and coyotes are capable of killing deer19, but whether coyotes kill deer that would otherwise die from limited food availability or severe weather (compensatory mortality), or if mortality from coyotes is additional to these other sources (additive mortality)22 remains unclear. Whereas dietary preferences indicate the importance of different prey species to the predator, deeper understanding of predation processes is required to infer the importance of the predator to specific prey species.
 Project Goals and Supporting Objectives The goal of this research is to understand the mechanisms by which coyotes affect biodiversity through competition with native predators and predation on white-tailed deer. Our specific research objectives are:
1) Calculate niche overlap among coyotes, bobcats, red fox, and (if possible) gray fox in the Adirondack region of NY State by documenting the relative importance of different prey species to their annual energy intake, 2) Identify whether coyotes are specializing on deer or switching to deer when snowshoe hare abundance is low by contrasting contemporary and historical coyote diets in the central Adirondack region and comparing trends in diet to trends in deer and snowshoe hare population density, and 3) Quantify the degree to which coyote predation on deer may be compensatory by back-tracking coyotes to deer carcasses to determine the fraction of deer consumed by coyotes that were actually killed by coyotes rather than killed by other causes. This research advances informational needs for NSRC Theme Four (Biodiversity and Protected Area Management) by clarifying the potential for coyotes to positively affect biodiversity (providing an ecosystem service by reducing deer populations and their browsing pressure), and to negatively affect biodiversity (through competition with native predators) in the Northern Forest. Project Background and Justification Coyotes routinely make the daily news in the eastern US given that they became common throughout the region only in the past thirty years. There is no doubt that the addition of a large bodied predator in such abundance is influencing biodiversity – but as we have outlined previously, there remains a great deal of uncertainty about precisely how. We‟ve identified two alternative pathways, competition with native predators and predation on white-tailed deer, by which coyotes could profoundly influence ecological communities, and by extension ecological health, in the Northern Forest. Long-term studies tracking the evolving diet of coyotes in the central Adirondacks

 To compare contemporary diets to historical data requires using the same methodology (scat-based analysis). However, we will also use state-of-the-art techniques involving stable isotopes and GPS devices (worn by the coyotes themselves!) to gain both a broader and deeper understanding of the mechanisms driving coyote impacts on other species in the region.
 Figure 1. Study area centered at the Huntington Wildlife Forest in Newcomb, NY. Approach For comparison of contemporary coyote diets to historic data (Objective 2), coyote scats will be collected in winter and summer on roads and trails throughout the HWF. PI Frair has extensive experience analyzing diets from coyote scats



 A total of 100 scats per season are needed to achieve a stable diet estimate. We will record the frequency of occurrence in scats of each prey tem following the methods of Brundige (1993)19, and contemporary coyote diets will be compared to those documented in this region during three previous periods: 1956-196117, 1975-198018, and 1986-198919. Recognizing that unequal surface:volume ratios make smaller prey items less digestible than larger items, which potentially biases estimates of prey importance, we will correct estimates based on the differential digestibility of prey items following Boser (2009)25.

 Finally, to compare historic trends in coyote diets to fluctuations in the populations of their primary prey species, specifically white-tailed deer and snowshoe hare, we will use deer harvest and hare pelt-sealing records to produce indices of abundance for each study period. These records are available from the NYS Department of Environmental Conservation in printed reports organized by wildlife management unit and county. We will work with Paul Jensen, Region 5 Wildlife Biologist, to secure these data and produce population indices (see letter of support).
 15N) in living or keratinized tissue (like hair) changes in a predictable manner with trophic level, due mainly to the preferential excretion of the lighter nitrogen isotope, yielding higher values at higher trophic levels26. The ratio of carbon isotopes (δ13C) changes little between trophic levels; however differences occur between species and reflect the source of primary productivity in the food web27. When used together, stable nitrogen and carbon isotope measurements can be used to trace energy flow, understand trophic relationships, and piece together a species‟ diet28.
To compare diets among multiple predator species (Objective 1) throughout the Adirondack region, an alternative analytical approach using naturally-occurring stable nitrogen and carbon isotopes will be used (routinely measured by co-PI Teece). The ratio of nitrogen isotopes (δ
 both seasonal and annual diets. In contrast to scat-based analyses, relatively few samples are needed from a given area (n ≥ 4) to depict population-level variation in diets29.
We will measure the isotopic composition of hair samples from animals, which we can readily acquire from fur-trappers in the Adirondack region. The advantages of using hair are that it is the easiest and least expensive type of tissue to collect, and hair δ values reflect the diet over the widest range of time relative to other types of tissue. Specifically, the portion of hair closest to the body reflects the most recent diet while the distal portion of hair reflects diet for the entire period of time the hair has been growing, allowing evaluation of
 30 and mixSIR31. Mixing models quantify the percent contribution of different sources to a mixture and determine the most likely diet composition.
Identifying the relative makeup of predator diets requires that the isotopic signatures of prey items be known, that unique prey items (or isotopically similar groups) can be identified, and that prey signatures have been acquired over the same spatial and temporal domain that the predator‟s diet is measured. A pilot study conducted in western NY State by PI Frair indicates that common prey items for coyotes are isotopically distinct from each other (Figure 2), and therefore this approach has a high probability of differentiating prey contributions to predator diets. Once δ values are obtained for prey species and predators, predator diets will be pieced together using stable isotope mixing models developed for multiple prey items available through IsoSource
 19,33,34. We will deploy specialized radio-collars equipped with Global Positioning System (GPS) technology that record detailed coyote movements (locations every 0.5 hour) and relay coyote paths to researchers every 3 days. This allows us to: 1) rapidly find deer carcasses and obtain a large sample size by searching only places where coyotes spent time (where at least 2 consecutive locations occur within 25-m of each other), and 2) identify whether deer were killed by coyotes or other causes with greater certainty by getting to deer carcasses within 3 days of the coyote having fed on it. These techniques have been pioneered for coyotes elsewhere in the state (R. Holevinski, Ph.D. Candidate, SUNY ESF under direction of J. Frair), where preliminary results indicate that coyotes kill less than 10% of the deer they consume.
Finally, to evaluate the degree to which coyote predation may be compensatory in deer populations (Objective 3) we will evaluate the fraction of "known-fate" deer carcasses that were killed by coyotes. Snow-based back-tracking is an inefficient way to find deer kills, and has in the past yielded too few carcasses to make reliable inferences regarding coyote kill rates



 Coyotes may kill a higher proportion of the deer they consume in the Adirondacks because deep snow conditions give them an advantage over deer, especially in late winter32. The GPS-facilitated approach to estimating kill rates has proven efficient and even extends efforts into snow-free periods allowing us to quantify fawn kill rates as well as adult kill rates35.
 Tangible products from this research will be at least two manuscripts in peer-reviewed journals, presentations at regional and national conferences, and a new collaboration among faculty of different disciplines at SUNY-ESF. Intangible produces include training for at least one graduate student (S. Warsen), outreach on carnivore ecology in the Adirondacks through interactions with trappers and land managers, and field and lab experience for the cadre of undergraduate volunteers interested in studying carnivore ecology
Products and Outcomes 25. Fresh scat samples (< 2 weeks old based on consistency and cohesiveness), those most likely to be from coyotes based on a diameter ≥ 2 cm4, will be placed in nylon bags, labeled by date and location of collection, and washed twice to remove faecal material21,23. Scat contents will then be separated by hand, and prey items will be identified from bone fragments and hair, both macroscopically and microscopically, with the aid of reference bone and hair collections and a hair key24.
This project is important because we have likely grossly overestimated the potential impact of coyotes on deer populations given the few studies that indicate coyotes are capable of killing deer, while at the same time grossly under-appreciating the cascading trophic effects that reductions in native predator populations are likely to have had following the coyote "invasion".

 The research is timely not only because of the great interest in coyotes at present, but also because it builds on concurrent studies in central and western NY (lead by PI Frair). Taken collectively these studies provide a broad scope of inference regarding the influence of coyotes on biodiversity in the Northern Forest. These concurrent studies also provide field supplies (like GPS collars) that reduce the overall cost of this project proposal by more than $33,000. Further, M.S. student Scott Warsen is committed to take on this study and has already secured fellowships covering salary and tuition (except for summer 2011) that further reduce the total project cost by $66,246. 17,18,19 combined with our experience working in that system grounds the current study in the Huntington Wildlife Forest (HWF), a 6000-ha research forest in the central Adirondacks near Newcomb, NY (Figure 1). Coyote and deer populations have been intermittently studied in this area since the 1950s providing a rich database to build upon.
By one pathway or the other, coyotes are influencing biodiversity, restructuring ecological communities, and subsequently affecting the health of the Northern Forest. This research will provide clarity on the ecological role of coyotes in the Northern Forest.14. Of particular importance to the Northern Forest is the potential for coyote predation to reduce populations of white-tailed deer (Odocoileus virginianus), a species recognized as a „keystone herbivore‟ or „ecosystem engineer‟. In the absence of wolves, high deer numbers and their subsequent browsing pressure has driven the composition and structure of plant communities, triggered changes in nutrient and carbon cycling, and subsequently altered invertebrate and vertebrate communities15. In short, overabundance of deer is a very serious threat to biodiversity. Whether coyotes are capable of regulating deer numbers and filling the vacant niche of the wolf remains unknown.
Given that coyotes compete with and predate upon a wide array of species, they may profoundly affect the structure of ecological communities in the Northern Forest. As a large, generalist carnivore, the potential for coyotes to exert top-down influences on a broad array of organisms is great. Top-down influences follow two key pathways from an apex predator to potential prey species. First, reductions in the abundance of other predators via interference competition indirectly link increasing coyote density to increasing abundance or diversity of waterfowl, songbirds, and rodentsCanis lupus lycaon) and cougars (Felix concolor) while the smaller coyote (Canis latrans) occurred only in the Great Plains and western states1. A series of human-mediated events that culminated at the turn of the 20th Century – namely, the extirpation of large carnivores combined with large-scale forest clearing – allowed coyotes to dramatically expand their range across North America.
Introduction Canis latrans) have become the top predator throughout most of the Northern Forest in the past few decades. Given that coyotes complete with and predate upon a wide array of species, their potential to exert strong top-down influences on community structure is great. Coyotes are known to limit populations of smaller carnivores, like red fox (Vulpes vulpes), and to predate upon white-tailed deer (Odocoileus virginianus) – either of which may have cascading trophic effects that influence the Northern Forest ecosystem. This research asks three questions fundamental to understanding the ecological role of coyotes in the Northeast: 1) what is the dietary niche of coyotes and how does it overlap that of native predators in the region?, 2) have coyotes become white-tailed deer specialists?, and 3) what proportion of deer consumed by coyotes are actually killed by coyotes? Each of these questions is critical to tracing the pathways linking the growing coyote population to changes in biodiversity. The research will be the purview of a M.S. student at SUNY-ESF who will compare contemporary and historical coyote diets, calculate niche overlap among coyotes, red fox, and bobcat (Felis rufus), and will estimate the rate at which coyotes kill adult and fawn deer. The work will be conducted in the central Adirondack region of New York State, and, added to a growing body of information on coyotes elsewhere in NY, will provide a comprehensive picture of the ecological niche of the coyote and in the Northern Forest and its implications for biodiversity.
Evolving niche of the "coy-wolf" in northeastern forests and implications for biodiversity
Abstract smcnulty@esf.edu Email: pgjensen@gw.dec.state.ny.us
Relevant (non-paid) Project Cooperators: mteece@esf.edu
Co-Principal Investigator: jfrair@esf.edu