2011 SOUTHEASTERN NATURALIST 10(4):721–730
Life History and Ecology of Coyotes in the Mid-Atlantic
States: A Summary of the Scientific Literature
Lauren L. Mastro*
Abstract - Relatively little information has been published on Coyotes in the eastern
United States, particularly in the mid-Atlantic region, the last area of the contiguous US
to be colonized by Coyotes. Increases in eastern Coyote distribution and abundance have
been documented, and concerns about their impact on wildlife and livestock are growing.
Information from published and unpublished manuscripts, theses, dissertations, and state
wildlife agency records in the mid-Atlantic region were examined and synthesized. This
review provides a comprehensive summary of Coyote ecology in the mid-Atlantic for
natural resource managers and researchers.
Canis latrans Say (Coyote) have expanded their range into the eastern United
States over the last 100 years. Increases in distribution and abundance have been
recorded and concerns regarding their impact on native wildlife and domestic
livestock are growing. In 2005, 35,000 cattle and calves worth >$20 million dollars
were lost to Coyotes in the eastern US, 3 times the number of animals lost to
Coyotes 14 years earlier in 1991 (NASS 1992, 2006). Coyotes are also impacting
native wildlife by hybridizing with Canis lupus rufus Audibon and Bachman
(Red Wolves), and by preying on Odocoileus virginianus (White-tailed Deer)
fawns and endangered shorebird nests. Despite increased interest, few published
studies have investigated these impacts, particularly in the mid-Atlantic region. I
summarized the existing data on Coyotes in the mid-Atlantic region in an effort
to disseminate information, identify gaps in our knowledge, and direct future
management and research.
I defined the mid-Atlantic area as the states of Delaware, Maryland, North
Carolina, Pennsylvania, Virginia, and West Virginia. I used several search engines
to ensure a comprehensive review of the literature including AGRICOLA,
BIOSIS, Wildlife and Ecological Studies Worldwide, and WorldCat. Additional
references were gained by inspecting the literature cited section of each reference.
Due to the limited information available on the eastern Coyote, I included
theses and dissertations, unpublished manuscripts, and grey literature in the form
*US Department of Agriculture, Animal and Plant Health Inspection Service, Wildlife
Services, National Wildlife Research Center, Logan Field Station, 730 Yokum Street,
Elkins, WV 26241; Lauren.L.Mastro@aphis.usda.gov.
722 Southeastern Naturalist Vol. 10, No. 4
Summary and Discussion
Spatial and temporal distribution
Coyotes began moving eastward from their historic range around 1900 (Moore
and Parker 1992, Parker 1995). Coyotes colonized the east by moving across
Canada north of the Great Lakes and into the northeast, south of the Great Lakes
and northeast along the Ohio river, and from Mississippi into the southeastern
states (Moore and Parker 1992, Parker 1995). Colonization of the mid-Atlantic
occurred primarily from two directions, south from New York (Fener et al. 2005)
and northeast from Georgia and Tennessee and, to a lesser extent, east across the
Ohio River and along Lake Erie (Parker 1995, Williams et al. 1985). Reports of
Coyotes in the mid-Atlantic states were first made in northern Pennsylvania in the
late 1930s and early 1940s (Hayden 2003, McGinnis 1979, Williams et al. 1985).
Coyotes were first reported in West Virginia’s Tucker County in 1950 (Wykle
1999), New Jersey’s Passaic County in 1958 (Lund 1975, Moore and Parker
1992), and Virginia’s Rockingham County in 1965 (Linzey 1998). Through the
1960s, 1970s, and early 1980s, Coyotes continued to occur sporadically and in
low numbers across these states (Linzey 1998, Lund 1975, Wykle 1999). Prior to
1983, only 8 Coyotes had been positively identified in Virginia, and only 6 were
positively documented in West Virginia between 1980 and 1985 (Wykle 1999).
Coyotes were first reported in Maryland in 1972 (McGinnis 1979), western North
Carolina in the early 1980s (DeBow et al. 1998, Lee et al. 1982), and Delaware
in the early 1990s (Parker 1995). During the past 10–20 years, mid-Atlantic state
wildlife agencies have reported increases in the number of Coyotes harvested by
hunters and trappers, suggesting that populations have also increased, although
no additional demographic information is available (Fig. 1).
Many authors have noted that the eastern Coyote is larger than the western
Coyote (Parker 1995, Way 2007, Way and Proietto 2005), and sometimes displays
a melanistic coat (Adams et al. 2003b, Anderson et al. 2009, Parker 1995). These
characteristics have raised questions about the genetic origins of the eastern Coyote.
In the mid-Atlantic and southeastern US, genetic testing has indicated that as
Coyotes moved eastward, a male Coyote hybridized with a female dog and their
female hybrid offspring successfully bred with Coyotes (Adams et al. 2003b).
Of 112 Coyotes tested from North Carolina, Virginia, and West Virginia, 12 had
the same dog haplotype, indicating dog ancestry likely resulted from a single
hybridization event (Adams et al. 2003b). Recently this haplotype has also been
found in Tennessee, Kentucky, and Alabama (Dennis 2010).
Additionally, eastern Coyotes have hybridized with Canis lupus lycaon (Eastern
Wolves) and Red Wolves. Studies have documented that Eastern Wolves
have hybridized with Coyotes and that Eastern Wolf genetic material is present
in the Coyote population in the northeast (Kays et al. 2010a, b; Way et al. 2010;
Wheeldon et al. 2010; Wilson et al. 2009). Kays et al. (2010a) found Eastern Wolf
genetic material in samples from central and western New York and Pennsylvania.
However, because the locations at which specific samples were collected were not
specified, it is unclear to what extent Eastern Wolf genetic material is present in
the mid-Atlantic. Coyotes have also hybridized with Red Wolves in the Red Wolf
2011 L.L. Mastro 723
recovery area in North Carolina to the extent that the US Fish and Wildlife Service
recognizes hybridization as the primary threat to Red Wolf recovery (Adams
2003a, Roth et al. 2008). However, the extent to which Red Wolf DNA is present in
the Coyote population outside of northeastern North Carolina is unknown.
Coyotes are sexually dimorphic; males are larger and weigh more than females
(Parker 1995). Adult Coyotes (n = 70) in Virginia averaged 16.2 kg and 13.4 kg for
males and females, respectively (Houben and Mason 2004), whereas adult Coyotes
(n = 39) in West Virginia averaged 14.4 kg and 11.9 kg for males and females,
respectively (Wykle 1999). On average, male Coyotes in West Virginia were 122.6
cm long and females were 117.8 cm long (Wykle 1999). The largest published
weight for a female eastern Coyote is 25.1 kg (Massachusetts) (Way and Poietto
2005). There are no published record weights of male eastern Coyotes, although
22.7-kg animals have been regularly recorded in Pennsylvania (Kyle Van Why,
USDA Wildlife Services, Harrisburg, PA, pers. comm.). No other published information
exists on the size and weights of Coyotes in the mid-Atlantic.
Pair bonding and mating. Pair bonding and mating occurs in January and
February (Chambers 1992, Parker 1995). Coyotes are monogamous and maintain
long-term pair bonds (Harrison 1992b). An instance of polygyny (1 male, 2
females) has been recorded in Coyotes in Ohio (Hennessy 2007). A survey in
Figure 1. Coyote harvest in mid-Atlantic states as reported by state wildlife agencies 1990–
2010. Estimated number of Coyotes harvested by hunters in North Carolina 2005/2006 and
2007/2008 (▲), Pennsylvania 1990–2004 and 2005–2010 (dark solid line), and Virginia
1993–2000, 2001/2002, and 2004–2009 (×'s). Actual number of Coyotes harvested by hunters
and trappers in New Jersey 1991–2010 (short dashes). Actual number of Coyote pelts
sold by trappers in West Virginia 2001–2009 (longer dashes). Actual number of Coyotes
pelts sold by trappers and actual number of Coyotes harvested by trappers in North Carolina
2002–2009 (gray hashed line).
724 Southeastern Naturalist Vol. 10, No. 4
western Tennessee found that 42% of females successfully mated and that older
females were more likely to be pregnant than yearling females (Stephenson and
Kennedy 1993). Overall, the number of breeding females is extremely variable
and depends largely on local environmental conditions (Stephenson and Kennedy
1993). In Connecticut, only 6 of 27 (22%) female reproductive tracts examined
had placental scars (Parker 1995). In Massachusetts, females have been recorded
whelping pups up to 6 years in a row (up to age 8–9) (Way and Proietto 2005).
Denning and birth. Gestation lasts about 60 days, and young are born in
March and April (Parker 1995). Eastern Coyote den sites have been described
as steep banks, rock ledges, brush-covered slopes, thickets, hollow logs, and
above-ground depressions (Parker 1995, Way et al. 2001). In Massachusetts, 17
dens averaged 2.5 m deep, with an average minimum width of 28.9 cm, and entrances
faced no particular compass direction (Way et al. 2001). In Maine, 6 den
entrances averaged a similar 37 cm; however, these entrances were more likely
to face south (120–236°) (Harrison and Gilbert 1985).
In western Tennessee, average litter size (based on placental scars) was 3.4
(Stephenson and Kennedy 1993). Litters in Massachusetts averaged 4.5 pups (n =
16) (Way et al. 2001). The size of a litter is dependent on many factors including
food availability and the ability of females to ovulate, conceive, and give birth
(Parker 1995). Adults have been recorded moving their pups to other den sites
up to 6 times (Harrison and Gilbert 1985, Way et al. 2001). New den sites were
a mean distance of 1.6 km apart (Harrison and Gilbert 1985).
Pup development. Pups emerge from the den at 4–5 weeks of age (late May)
(Harrison and Gilbert 1985, Harrison et al. 1991, Way et al. 2001). Dens are then
abandoned at 8–10 weeks of age (June/July) for rendezvous or above-ground
resting sites (Harrison and Gilbert 1985, Harrison et al. 1991, Way et al. 2001).
Pups first begin to move short distances from den and rendezvous sites at 6–8
weeks of age, but still concentrate their activities at these sites (Harrison et al.
1991). Pups spend less time at the sites through the summer and eventually abandon
them in mid-October (Harrison et al. 1991). Pups first move long distances
accompanied by an adult at 13 weeks of age (late July; Harrison et. al 1991).
Pups are weaned by 6–8 weeks of age (Parks 1979, Silver and Silver 1969) and
begin eating regurgitated food at 3 weeks (Parker 1995). Adults supply food (either
regurgitated or whole) until July, and pups usually forage independently by August
(Harrision et al. 1991). Pups may also receive food from adults that are not their parents
(Way 2003, 2004; Way et al. 2001). A den of 4 pups in Massachusetts survived
the death of their mother at 8 weeks of age because they were fed by at least 1 adult in
addition to an animal presumed to be their surviving father (Way 2004). The genetic
relationship between this adult and the pups was unknown (Way 2004).
Dispersal. Coyote pups in Maine dispersed 4–10 months after birth in two
waves, October–November and January–February (Harrison et al. 1991). Males
and females dispersed the same distance for an average of 102.5 km (Harrison
1992a). The distance and timing of dispersal is dependent upon food availability
and population density (Bekoff 1978, Harrison et al. 1991).
Age structure and mortality. A sample of 70 adult Coyotes taken in western
Virginia from 1993–1996 indicated that 71% of Coyotes were greater than 1 year
old (Houben and Mason 2004). The age structure of Coyotes in West Virginia
2011 L.L. Mastro 725
is currently being examined (Albers 2010). The oldest documented wild eastern
Coyote was a 10–12 year old female in Massachusetts that had produced a litter
of pups in her last year of life (Way et al. 2004). The majority of mortalities
among radio-collared adult Coyotes are human related, and include trapping,
hunting, and road kills (Bogan 2004, Chamberlain and Leopold 2001, Hilton
1976, Schrecengost et al. 2009, and Van Deelen and Gosselink 2006).
Coyotes are a moderately social species, some are members of social groups
which share the same territory, others are solitary and unassociated with other
Coyotes. Eastern Coyotes typically live in groups of 2–4 (Caturano 1983, Way
2000). Observations of larger groups involve 3–4 adults and 5–7 pups (Patterson
and Messier 2001, Way 2003). Groups of eastern Coyotes have been observed
consisting of male/female, male/male, female/female pairs, as well as male/1–2
juveniles, 2 males/2 females, and male/female/1–3 juveniles (Brundige 1993,
Caturano 1983, Major 1983, Patterson and Messier 2001, Way 2003).
A common misconception is that group members that are not the mated pair
are always the offspring from the previous year. An Illinois study investigated
the genetic relationship between eastern Coyote pack members (Hennessy 2007).
Out of 116 relationships among 62 individual pack members: 49.1% were fullsiblings
or parent/offspring; 31% were grandparent/grandchild, first cousins,
aunt/nephew, etc.; 13.8% were unrelated; and 6.1% were mated pair relationships.
Of the 7 mated pairs analyzed, 6 were unrelated and 1 was related on the
level of grandparent/grandchild or first cousins. In most cases, pack members
were related on some level; however, there were two individuals that were not
related to any other member of their packs and a pack where none of the three
members were related (Hennessy 2007).
Eastern Coyote home-range size varies and is influenced by habitat, geography,
food availability, and season (Brundige 1993, Caturano 1983, Crête et al.
2001, Holzman et al. 1992, Person and Hirth 1991). Additionally, reproductive
status, sex, and social status can also affect home-range size (Babb and Kennedy
1988, Harrison and Gilbert 1985, Parker and Maxwell 1989, Pearson and Hirth
1991, Way et al. 2002). Home ranges of Coyotes in the greater mid-Atlantic area
vary greatly, and no geographic trends are evident (Table 1). Both the smallest
(5.7 km2) and largest (112.8 km2) home ranges for eastern Coyotes have been
observed in New York. There is currently no published data on home-range sizes
in the mid-Atlantic region.
There is limited information on Coyote diet in the mid-Atlantic region. Two
studies have been completed in northeastern Ohio, and one study each has been
conducted in central Kentucky, Pennsylvania, and West Virginia (Table 2). Although
these studies provide some baseline data, the availability of food due to
season, prey population, or location limit the application of these data.
The first Ohio study, conducted during the winter, found that Microtus pennsylvanicus
(Ord) (Meadow Vole) were in 14 of the 50 scats collected, but only
726 Southeastern Naturalist Vol. 10, No. 4
accounted for 18% of diet by volume. Sylvilagus floridanus (J.A. Allen) (Eastern
Cottontail) were in 10 of 50 scats and made up 13% of diet by volume. Whitetailed
Deer were in 10 of 50 scats and comprised 13% of diet by volume. Procyon
lotor (L.) (Raccoon) were found in 9 of the 50 scats and comprised 12% of diet by
volume (Cepek 2004). The second Ohio study, conducted during all seasons, also
found that small mammals accounted for a large number of Coyote food items
(490 of 944 scats contained small mammals). Small mammals were followed by
White-tailed Deer (415 of 944 scats), vegetation (305 of 944 scats), and Eastern
Cottontail (142 of 944 scats). This latter study did not examine food items by
volume (Bollin-Booth 2007).
The Pennsylvania study, conducted year round, found that vegetation occurred
in 177 of 310 scats and White-tailed Deer were observed in 171 of 310 scats. Other
Table 1: Mean annual home ranges (km2) of adult Coyotes in the eastern United States. n = the
number of Coyotes a given mean home range is for, MCP = minimum convex polygon
State(s) size (km2) n Method Source
Massachusetts 50.7 7 95% MCP (Way et al. 2002)
New York 5.7 17 95% MCP (Bogan 2004)
New York 112.8 10 95% MCP (Brundige 1993)
New York 17.4 9 95% MCP (Kendrot 1998)
New Jersey 10.4 4 95% MCP (Eastman 2000)
Indiana 9.9 2 95% adaptive kernel (Atwood and Weeks 2002)
Kentucky 18.4 6 95% adaptive kernel (Cox 2003)
Tennessee 42.6 5 Minimum area (Babb and Kennedy 1988)
South Carolina 30.5 18 95% MCPa (Schrecengost 2007)
Table 2. Prey items found in >15% of samples (scat or stomach) in Coyote diet studies conducted
in mid-Atlantic and adjacent states. n = total number of samples.
Location % of samples
(author) Season Sample type (n) containing a food item
Kentucky Winter Stomach (66) 52% small mammals
(Crosset and Elliott 1991) 26% cattle
20% Eastern Cottontail
Ohio Year-round Scat (944) 52% small mammal
(Bollin-Booth 2007) 44% White-tailed Deer
15% Eastern Cottontail
Ohio Winter Scat (50) 28% Meadow Vole
(Cepek 2004) 20% Eastern Cottontail
20% White-tailed Deer
Pennsylvania Year-round Scat (310) 55% vegetation
(Witmer et al. 1995) 55% White-tailed Deer
15% small mammals
West Virginia Winter Stomach (24) 83% White-tailed Deer
(Wykle 1999) 17% fruit
2011 L.L. Mastro 727
food items of importance were insects (56 of 310 scats), fruit (49 of 310 scats), and
small mammals (46 of 310 scats). Again, this study did not examine food items by
volume (Witmer et al. 1995). The central Kentucky study, conducted only in winter,
found that small mammals occurred in more stomachs than any other food item (34
of 66 stomachs), followed by Bos Taurus L. (Cattle) (17 of 66 stomachs) and Eastern
Cottontail (13 of 66 stomachs). Samples were collected by private fur trappers, fur
dealers, farm bureau personnel, and Kentucky Department of Fish and Wildlife Resources
personnel. Again, this study did not examine food items by volume (Crossett
and Elliott 1991). The West Virginia study, conducted primarily in winter, collected
24 Coyote stomachs from private fur trappers and USDA APHIS Wildlife Services
specialists. The most frequently occurring food item was White-tailed Deer (20 of
24 stomachs), followed by fruit (4 of 24 stomachs), small mammals (2 of 24 stomachs),
and Ovis ovies (Sheep) (2 of 24 stomachs). This study also did not examine
food items by volume (Wykle 1999). The year-round diet of Coyotes in West Virginia
is currently being examined (Albers 2010).
Coyotes have recently expanded their range into the mid-Atlantic region
raising concerns about their impacts to both wildlife and domestic livestock. A
review of the literature found that gaps in our knowledge were apparent in all
areas of the ecology of eastern Coyotes. Deficiencies in the quality and quantity
of information about the mid-Atlantic Coyotes’ population, social behavior,
home range, and foraging ecology are of particular concern, as this information is
necessary for wildlife managers to assess and address their impacts and changing
role in local ecosystems. Information on Coyote foraging ecology, population
density, home range, ecological impacts, and genetics from areas outside the
mid-Atlantic states may not be applicable due to ecological and environmental
differences. Thus, these areas should be priorities of future research.
I am grateful to Marilyn Howell and the NWRC library for their assistance obtaining
documents. Andrew Burnett (New Jersey Division of Fish and Wildlife), Mike Fies (Virginia
Department of Game and Inland Fisheries), Thomas Hardisky (Pennsylvania Game
Commission), Colleen Olfenbuttel (North Carolina Wildlife Resources Commission),
and Rich Rogers (West Virginia Division of Natural Resources), generously provided
Coyote harvest statistics. Scott C. Barras and Julie K. Young reviewed the manuscript.
Christopher K. Croson and Scott C. Barras provided logistical support.
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