Abstract

Competitions between carnivores are complex and common between species

of canids. Apex predators, which are predators that rely on meat for 100% of their

diet and mesocarnivores that has a diet of 75% encompassed by meat are known to

have competition that leads to one affecting the other through, predating on one

another, breeding, habitat usage, and feeding habits. This study looks into the

possible affects that coyote (Canis latrans) have on the populations of red wolves

(Canis rufus) in eastern, North Carolina. From review of earlier studies of sympatric,

coexisting predators in North America and Africa it is concluded that coyotes have

an effect on red wolves in eastern, North Carolina.

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Introduction

Just as people affect each other so do animals. With animals that are

carnivorous the issue of one affecting the other happens to be in depth, and for some

that are endangered the outcomes can be detrimental. Biologists have looked into

the behavior of how carnivores affect one another in multiple places around the

world such as Africa and North America. For example, in Africa researcher and

biologist Dr. Richard W. Yarnell studies the influence of large predators on their

feeding habits and feeding ecology by looking at the two mesocarnivores, the brown

hyena (Hyaena brunnea) and the black backed jackal (Canis mesomelas).

Further research in North America by Dr. Lynda Randa has looked at prey

switching of sympatric canids in response to variable prey abundance between red

foxes and coyotes. Meaning that with the affects and pressures that coyote had on

red fox in the same environment the red fox swapped what they typically feed on in

order to make room for the coyote in the area. By looking at these carnivores in

these varying environments we can translate what was found and use this

information to apply it North American predators such as the red wolf (Canis rufus)

and coyote (Canis latrans). The purpose of this study is to look at the competition

between red wolves and coyote by answering the following question: Does the

coyote (Canis latrans) effect the populations of red wolves (Canis rufus) in Eastern,

North Carolina, and if so, how?

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Background Information

Identification and Behavior

Red wolves (Canis rufus) are a species of wolf that are native to the

southeastern, United States. The characteristics of the red wolf are of two origins

within the rufus species. Those characteristics are of red variety (Appendix A, Figure

1, p. 21) and black variety (Appendix A, Figure 2, p.22) (Mech, 2003). The red

variety of red wolves are found today in eastern, North Carolina and parts of

northern, South Carolina (Mech, 2003). The lesser known the black variety of red

wolves are found in Florida and are fewer in number than the red variety (Mech,

2003). The red variety, which this research focuses on, is characterized as canids

that are morphologically similar to a gray wolf (Canis lupus) and a coyote.

At first glance looking at the red wolf and the coyote (Appendix A, Figure 3, p.

23) both canids look similar and are hard to distinguish. However, there are features

that help to identify the two easily. The similar features between the red wolf and

coyote are as follows, their facial features, their long legs, their long muzzle, large

ears, and to some their coat or pelt color. Red wolves may have these similarities

with coyotes, but these characteristics, when looked at more closely, also set them

apart (Mech, 2003).

For instance, when you look at the face of a red wolf in between the ears and

the muzzle side by side with a coyote and a gray wolf (Appendix A, Figure 4, p. 24)

there is a more wolfish appearance than the appearance of a coyote. When it comes

to coat or pelt color the red wolf has a reddish grey topcoat and markings with a

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light tan under coat, whereas the coyote has the markings of a tawny brown under

coat and blonde under coat and facial markings. Looking at the legs of each animal

the red wolf has a bigger paw and distal region that the coyote, and the legs are also

thicker and more muscled than the coyote.

Another indicator to help distinguish between red wolves and coyote is by

their behavior. Coyotes are more bold and territorial in their actions whereas red

wolves are more reclusive and the packs show more human avoidance and urban

avoidance (Beckoff, 2001). Coyotes seem to thrive in urban environments as habitat

generalists; however, their dens during pup rearing season are hard to find. Red

wolves focus more on the art of being unseen with their packs being hard to trace or

find and during the pup-rearing season they are made even more rare to find

(Hinton, 2013) Furthermore, even as habitat generalists red wolves prefer conifer

forests over human modified boreal forests, urban areas (Mech, 2003).

The diet of the red wolf mainly consists of deer, nutria, voles, mice, and other

small mammals. Red wolves do not go after big game such as elk, bison, or moose

like their grey wolf counter parts. Therefore, the farmer and rancher on the east

coast has nothing to fear from these predators when it comes to their livestock.

Furthermore, as with all wolves, red wolves will not attack people. There are no

documented or known wolf attacks from any species of wolf in the United States.

Historical Range and Modern Ranges

As previously stated red wolves are found in the southeastern, United States

in eastern, North Carolina, northern South Carolina, very northern parts of Georgia,

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and a few packs reaching to Florida, of the black variety (Mech, 2003). Fossil records

of the red wolf and the gray wolf show their historical range consisted of all of the

southeastern, United States reaching to parts of Ohio and going through the lower

parts of Texas (Appendix B, Figure 7, p. 27; Hinton, 2013).

Some documents state that the red wolf is a variation of the grey wolf (Canis

lupus). This information is inaccurate in that the red wolf is morphologically

different and followed a different evolutionary track. In comparison with the grey

wolf the red wolf is smaller, but still larger than a coyote. Grey wolves evolved in

North America in parts of Canada and the red wolf evolved in the southern United

states and branched out into their historical range reaching as far north as New

York, as far south as Florida, and as far east as eastern Texas (Mech, 2003). For

visuals on the historical range refer to Appendix B Figure 7 on page 27.

Likewise the coyote evolved in North America and grew in number and

ranged throughout the western states such as Nevada, California, New Mexico,

Texas, and even reaching as Far East as Louisiana. Texas and Louisiana are where

the red wolves and coyote have historically always shared the environment. Coyote

came to North Carolina from the western parts of U.S. in 1980 due to the killing of

grey wolves (Canis lupus). Grey wolves are main predators of the coyote and with

the killing of the wolves it allowed for the coyote to become more bold and

aggressive, whereas with the wolf present coyote were more focused in avoidance

from the grey wolf (Hinton, 2013).

Naturalist William Bartram has an account with wolves in his travels to

Florida that he mentions in his book that he published in 1791, Travels. For years

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many believed this to be an encounter with red wolves, but it was later discovered

that they were another species of wolf later to be known as the Florida wolf also

known as the black variation of the red wolf. These discoveries lead to the beginning

interests in the red wolf recovery project.

Red Wolf Recovery Project

With the decline in population in the red wolf population the United States

declared the red wolf as an endangered species in 1967. What lead to this

declaration were numerous factors consisting of human interference and a parasite

infestation as a result of interbreeding with coyote (Mech, 2003). The Endangered

species act began a recovery project in 1973 upon the US Congress passing the ESA.

Essentially, the program was set to calculate and document the remaining wolves

found in the wild. The research yielded results that showed the red wolf as being

rare, free ranging in the wild and that a good number of the pups documented were

of mixed species. The pups DNA revealed that there was a mix with coyote

(Mech,2003; Beckoff, 2001).

The United States Fish and Wildlife Service further concluded and stated that

the recovery of the red wolf needed to be accomplished through captive breeding.

Thus, the red wolf captive breeding program began in November of 1973 (Mech,

2003; Hinton, 2013). In 1980 the red wolf species was declared biologically extinct

(Hinton, 2013; Mech, 2003). This was the same year in which the coyote had

entered North Carolina and began to rapidly populate the state (Mech, 2003;

Beckoff, 2001). The breeding program initially began with the capturing of 400

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wolves from Louisiana and southeastern, Texas. During the project it was

discovered that from the canids they had captured there had been interbreeding

between red wolves and coyote. Because the ESA and USWS wanted to preserve the

red wolf as a species only approximately 43 wolves were used (Mech, 2003).

Amongst those 43 wolves only 14 were absolutely of pure strain. Red wolves are the

only known species of wolf that have been recorded to interbreed with coyote.

Today there is a new species of canid known as the coywolf. The coywolf is a hybrid

between coyotes and red wolves whose range is from Florida to New York

(Appendix A, Figure 5, p. 25). Their size is smaller than a red wolf but larger than a

coyote, and stockier build than a coyote (Appendix A, Figure 6, p. 26).

Due to morphological and evolutionary discrepancies in the scientific

community about the red wolf being considered a full separate species of wolf the

USFWS conducted a review of the red wolf species using William Bartram and

Audubon’s notes on the red wolf characteristics as well as DNA testing. It was shown

that the red wolf was in fact its own distinct wolf species not being a subspecies of

grey wolf that bred with coyote, as it was once believed (Mech, 2003; Hinton, 2013).

Reintroduction of the red wolf began preparations in 1982 and in 1984 another

recovery of 330 wolves began. The actual release program began in 1986 followed

soon after with the release of 63 wolves in 1987 (Mech, 2003).

The red wolf recovery program was highly active throughout the years

between 1987 through 1994, and by June of 2002 had implemented 100 red wolves

into the wild (Mech, 2003). The red wolves were released in Alligator River, North

Carolina, Bulls Island State Park, South Carolina, and Cape Romain National Wildlife

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Refuge, South Carolina. Ultimately these packs listed were and are still successful. A

pair of red wolves were released in the Great Smoky Mountains National Park but

were unsuccessful and the program in the Great Smoky Mountains was disbanded.

As of today the red wolf recovery program is still on going, however, as of

August 2014 North Carolina called for a review of the Red Wolf Recovery Program

to be reviewed in October 2014 in order to discuss the appropriateness of the red

wolf species and to decide the overall success of the recovery program. Today there

are approximately 265 wolves in captivity, with that number steadily increasing,

and approximately 100 wolves in the wild, with that number steadily decreasing.

According to varying sources the red wolf recovery program is not either a

success or a failure. The program itself has yielded some successful red wolf packs

in areas, but has failed overall in bringing the red wolf back to its historic range and

numbers. As an example the release in the Great Smoky Mountains is considered

puzzling and the results as to why the pack did not take in the area are still

unknown. According to Dr. L. David Mech who is considered to be the authority on

the biology and ecology of wolves due to his publishing of Wolves: Behavior, Ecology,

and Conservation which he published in 2003 in partnership with Luigi Boitani, has

said that he views the program has served as an example of what occurs with over

relaxation of regulations under experimental implication and design (Mech, 2003).

He states that a few of the reasons that the red wolf recovery program has

ultimately been a moot project neither yielding positives or negatives are due to

regulations that provide leniency and flexible land ownership (Mech, 2003).

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The statements made by Dr. L. David Mech raise questions with regards to

the North Carolina government and their wishes to review the overall program.

Such as: How strict should regulations be for landowners who live in areas where

endangered or threatened species live and should they be allowed to become

lenient? Is the red wolf recovery program a solid program as is or should we revise

it or be rid of it? Should we restore endangered species to historical ranges or do we

work to preserve them in the environment that is available to them? Even with the

relevance of these questions, for the sake of this research the main question that

everyone is asking and one that no one has even thought about answering is, does

the coyote (Canis latrans) effect the populations of red wolves (Canis rufus) in

Eastern, North Carolina, and if so, how?

Methods

There have been limited studies done on red wolves and coyotes, though

there have been no studies on the two species together at all. The methods that

were used in this research were adopted from, and influenced by, previous, relevant

research on other carnivores. The conclusions drawn, and the information

presented within the previous research offers parallels to the situation of the red

wolf and coyote. A, separate study, conducted in North America by researcher and

biologist Dr. Lynda Randa, analyzed the affects coyotes had on the red fox (Vulpes

vulpes) within the red fox environment. She observed that within the environments

inhabited by both the red fox and the coyote, the two predated on the same prey

species: voles, rabbits, and mice. They conducted their study by analyzing behavior

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and prey availability. What she found was in order for the two carnivores to coexist

they would feed on varying prey and avoid competition. Furthermore, if the

abundance of prey were scarce the canids would expand their home ranges, in

which coyote would push out the red foxes within that range (Randa, 2009).

Interference competition between grey wolves (Canis lupus) and coyotes in

Yellowstone National Park researched by J. A. Merkle, D.R. Stahler, and D.W. Smith

to better understand interactions and coexistence between both species. Merkle

studied the behavior and interspecific interactions between the two canids from

1995 to 2007 (Merkle, 2009). Their study found that the gray wolves showed

tolerance to a mesocarnivore within their territory. However, if coyote interfered

with left over kills, kills, or were around a den the wolves would predate on coyote.

It was further shown that with scarcity of prey wolves would hunt coyote pairs.

Interbreeding among coyote and grey wolves was not shown, like it is in the red

wolf and Mexican grey wolf species. Coyote unless starved or inexperienced with

wolves showed avoidance of wolves, but were opportunistic about stealing left over

food cashes that packs were known to travel back too.

Research by Dr. Richard Yarnell conducted in Africa, studied the influence

large predators had on the feeding habits and feeding ecology of mesocarnivores;

specifically looking at the brown hyena (Hyaena brunnea) and the black backed

jackal (Canis mesomelas). Both predators feed on the same prey, mainly consisting

of the carrion of sea birds, cape fur seals, and other means. In the presence of an

apex predator their diet of living animals consists of young antelope, small

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mammals, and rodents (Yarnell, 2013). Yarnell looked into how these two

mesocarnivores competed for their food resources.

What Yarnell found was that these two meso-carnivores would switch

dietary habits in the presence of, or lack of, an apex predator within their

environments. The apex predator for both the brown hyena and the black backed

jackal is the African Lion (Panthera leo). In the areas where lions were present the

two meso-carnivores continued to eat carrion and would co-exist sympatrically.

However, it was observed that in the event the lion was absent from the

environment, the habits of the two meso carnivores changed; they began to directly

compete against each other, for resources, fresh kills, and carrion finds (Yarnell,

2013). Black-backed Jackals are submissive to brown hyena in the absence of an

apex predator, whereas both meso-carnivores are submissive to the apex predator

when present.

In a study conducted by Jan F. Kamler in 2007 the predation of a

mesocarnivores by an apex predator was observed. Kamler looked into the factors

that lead to the predation of black backed jackals by African Wild Dogs (Lycoan

pictus). These canids were observed for this research because interspecific

predation among African canids is rare whereas interspecific predation is common

amongst northern temperate canids, as observed in wolves and coyote (Kamler,

2007; Mech, 2003). The behavior of each canid was observed as well as aggression

levels between the two species. Results showed that when ranges were vast and

abundance of prey was high the two canids coexisted, sharing the same territory.

However, it was further observed that if another apex predator, such as spotted

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hyena or African lions, was present aggression level between the two species of

canids was high and predation of the mesocarnivores was present. Some

observations showed the black backed jackals eating the carrion left over from

African wild dog kills, when usually African wild dogs guard previous kills as a

source for food when hunts are unsuccessful (Kamler, 2007). In one observation it

was noted that a pack of African wild dogs, in an environment with low prey

abundance, were chasing pairs of black backed jackals for sport (Kamler, 2007).

Kamler suggested that further research among the 13 species of African canines be

conducted for confirmation of common nature among African canines (Kamler,

2007).

Another study by Heather E. Watts and K. E. Holekamp in 2009 looked at

interspecific competition between spotted hyena (Crocuta crocuta) and African lions

and the affects the competition had on reproduction among the spotted hyena. The

relevance of this journal to this project can be related to the affects coyote have on

red wolf pup rearing seasons and overall reproduction. Furthermore, it must be

noted that predation among carnivores is not limited to apex predators predating

on mesocarnivores; apex predators will also predate on each other in rare

circumstances when prey abundance is low (Watts; Holekamp, 2009; Mech, 2003)

(Mech; Boitani, 2001). For their research Watts and Holekamp looked at two

populations of spotted hyena, one with African lions present in the home range and

one without African lions in the home and only having spotted hyena as the apex

predator within the home range (Watts; Holekamp, 2009). Aggression levels,

behavior, predation, prey abundance, and reproduction were observed amongst the

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two populations of spotted hyena, as well as aggression levels between spotted

hyena and African lions. What the results showed were in the population with lions

present the reproduction of the spotted hyena females was affected by 24%,

showing a decrease in pups born compared to the population of spotted hyena that

lived in an environment without African lions (Watts; Holekamp, 2009). It was again

suggested that further research into the 13 species of African canines be conducted

for more conclusive evidence.

Results

The articles conducted in North America and in Africa are of different

predators, but these articles shows an affect on prey abundance, habitat size, and

reproduction are affected by other carnivores. In Holekamp’s study, on the

reproduction levels in spotted hyenas, the study can be related to the affects that

coyote have on red wolf pup rearing seasons and overall reproduction.

Each individual study found that, if prey abundance was inadequate that

competition would be present. The separate studies by Randa, Yarnell, and Merkle

show that low prey meant that predation or coexistence between the predators

within that environment.

With this related information I feel confident in concluding that coyote do

affect the red wolf species. Mech in 2003 wrote a book, Wolves: Biology, Ecology,

and Conservation which he states how it was difficult for the biologists to find pure

strains of red wolves for the recovery and captive breeding programs (Mech, 2003).

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Furthermore, red wolves began to drastically decline the same year, 1980, that

coyote populated all 100 counties of North Carolina.

The Wildlife and Fisheries of North Carolina has been observed this year that

in areas heavily populated by coyotes, red wolves will not have as many pups as in

areas that have low level coyote interference, though know study has been done to

confirm this observation (Hinton, 2013).

Another impact on the red wolf species, by the coyote, is that the two species

are nearly identical in appearance making it difficult for farmers and hunters to

discern the canid they are hunting. As of 2013 four packs of wild red wolves were

shot and killed because of misidentification (Dalesio, 2014).

Discussion

The driving force behind this study was the knowledge that red wolves began a

rapid decline in 1980, which was the same year that coyotes populated North

Carolina; to me that was no coincidence. The other overarching issue was how

closely the two species resembled each other to the unaided eye. Those two factors

brought me to my question, Does the coyote (Canis latrans) effect the populations of

red wolves (Canis rufus) in Eastern, North Carolina, and if so, how?

When I gathered my information I specifically chose David Mech’s book as

my main source of background information since I could compare the red wolves to

other species of wolves found in other parts of the world, and because David Mech is

considered the biological authority on wolves. In using Mech’s book I specifically

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researched articles and peer reviewed journals that related to behavior and

biodiversity relatedness on canids. From those journals I could then relate that

information back to articles and issues brought up by Mech and J.W. Hinton.

Although, extinction and replacement is a natural cycle in the Earth’s history,

these cycles greatly impact ecosystems when keystone species go extinct. As a

keystone species red wolves are important to North Carolina’s ecosystem as a whole

and currently there are no other keystone species that could replace the role that

red wolves have on the environment.

The parallels shown in the related research should aid in further study of the

coyote and red wolf issue in North Carolina, and further research should be done to

better understand the impact that coyote have on the red wolf species. Preservation

of not only the species but the environment should be a priority.

Conclusion

In conclusion red wolves are a native apex predator to eastern, North

Carolina and are critically endangered. Red wolves began a rapid decline in 1980

and were declared biologically extinct. Also, in 1980 coyote, due to declining

numbers of their natural predator the grey wolf began to populate all 100 counties

of North Carolina. The issues facing the relations between these two species are how

closely they resembling each other, and how unlike other species of wolves, red

wolves will interbreed with coyote.

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Studies on other separate species of canines aided in drawing indirect

evidence on the issues facing the red wolf and coyote competition in eastern, North

Carolina. Randa’s study on red foxes and coyotes showed how two species of

predators were able to coexist in the same environment, and Watt’s study on

African lions impact on reproduction levels of spotted hyena further showed how

another apex predator, impact pup rearing and births.

Through looking at the evidence presented in related research and the current

events on red wolves it is concluded that coyotes do impact red wolves. However,

further research should be conducted to fully understand the competition between

red wolves and coyote in order to enact methods to help protect red wolves from

extinction.

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Acknowledgements

I would like to thank Dr. Jeff Llewellyn, Professor of Biology at Brevard

College, who was my senior project advisor. I would also like to thank Dr. Bob Cabin,

Associate Professor of Environmental Studies and Ecology at Brevard College, who

was the SCI 494/495 class professor, and my academic advisor. Both Dr. Llewellyn

and Dr. Cabin greatly helped me with forming my study to what it now is! I would

also like to thank the Wolf Conservation Center in Salem, New York for sending me

current events in Red Wolf news, as well as allowing me to adopt Nikai! I would

further like to thank Dr. Edward Schwartzman of the North Carolina Heritage

Program for Transylvania County, North Carolina, who sent me the range maps for

the red wolf! Finally, I would like to thank my parents for supporting me through all

of my academics and being encouraging when I needed it most!

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Literature Cited

BARTRAM: 1791. Travels. Norton Anthology. 2002. The Norton Book on Nature Writing. W. W. Norton & Company; College Edition. 1135 pp.

KAMLER, J. F. and DAVIES-MOSTERT, H. T. 2007. Predation on Black Backed Jackals (Canis mesomelas) by African Wild Dogs (Lycoan pictus). African Journal of Ecology. Vol. 45 (4): 667-668

MECH, D. L. and BOITANI, L. 2003. Wolves Behavior, Ecology, and Conservation. University of Chicago Press. 448 pp.

MECH, D. L., CHRISTENSEN, B. W., ASA, C. S., CALLAHAN, M., and YOUNG, J. K. 2014. Production of hybrids between western grey wolves and western coyote. PloS ONE. Vol. 9 (2): 1-7

MERKLE, J. A., STAHLER, D. R., and SMITH, D. W. 2009. Interference competition between gray wolves and coyote in Yellowstone National Park. Canadian Journal of Zoology. Vol. 87 (1): 56-63

RANDA, L. A., COOPER, D. M., MESERVE, P. L., and YUNGER, J. A. 2009. Prey switching of sympatric canids in response to variable prey abundance. Journal of Mammology. Vol. 90 (3): 594-603

WATTS, H. E. and HOLEKAMP, K. E. 2008. Interspecific competition influences reproduction in spotted hyenas. Journal of Zoology. Vol. 276 (4): 402-410

YARNELL, R. W., PHIPPS, W. L., BURGESS, L. P., ELISS, J. A., HARRISON, S. W. R., DELL, S., MAC TAVISH, D., MAC TAVISH, L. M., and SCOTT, D. M. 2013. Influence of large predators on the feeding ecology of two African mesocarnivores the black backed jackal and brown hyena. South African Journal of Wildlife Research. Vol. 43 (2): 155-156

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Appendix A

Figure 1: Red variety of Red Wolf The image is by the Wolf Conservation Center in Salem, New York.

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Appendix A Continued

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Figure 2: Black Variety of Red Wolf (found in Florida.)William Bartram documented this variation in his book, Travels, in 1791. This image was obtained through World Wildlife.

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Appendix A Continued

Figure 3: Both Red Wolves and Coyote Look Similar (Top is red wolf and Bottom is coyote). The red wolf image is by the Wolf Conservation Center in Salem, New York and the Coyote image is by Nature.

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Appendix A Continued

Figure 4: Grey Wolf:Wolf Conservations Center’s 2014 Pic of my wolf, Nikai. Nikai is a Canadian Rocky Mountain Grey Wolf that is found historically in the U.S. in the lower 48. I adopted Nikai on 9/20/14.

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Appendix A Continued

Figure 5: Coyote Hybrid Known As the Coywolf

Coywolves are hybrids of red wolf and coyote and range up and down the east coast of the United States. They pose a threat since they are considered a new species because they are feral unlike other hybrids that are sterile. This image was obtained through the Nature Documentary on PBS.

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Appendix A Continued

Figure 6: Skull Comparison to Show Size

PBS Nature Documentary Image of Skulls to show size of new species. Left to Right: Red Wolf, Coywolf, and Coyote. The coywolf is a red wolf coyote hybrid that has become an entirely new species.

Appendix B

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Figure 7: Historical Range of Red Wolves

Dr. Edward Schwartzman of North Carolina Heritage for Transylvania

County provided this image.

Appendix B Continued

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Figure 8: The Modern Range of Red Wolves found in North Carolina

Appendix B Continued

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Figure 9: Historical Range of Coyotes

This image was found on an information site dedicated to coyote facts.

Appendix B Continued

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Figure 10: Modern Ranges of Coyotes

This is an image by National Geographic.

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