Nicholas E. Federoff -- Wildlife Biologist

Free Web Hosting Provider - Web Hosting - E-commerce - High Speed Internet - Free Web Page

Introduction
The U.S. Fish and Wildlife Service's (Service) red wolf (Canis rufus) recovery program has had relatively low visibility throughout its existence. This low visibility is the result of several factors, one being that the red wolf is not as well known as its cousin in the genus, the gray wolf (C. lupus). The gray wolf is larger, has a much larger (circumpolar) range that includes Asia, Europe, and North America, and often exists in much more open habitats where it is more easily seen and studied. Moreover, the red wolf has been plagued by taxonomic controversy since it was first described, which has produced skepticism about the merits of maintaining the red wolf as a separate taxon and the need for recovery. The reduced visibility was also somewhat intentional from the beginning of the red wolf recovery program due to the controversial nature of predator reintroduction. The Service kept publicity to a minimum and confined it to the local area affected by the reintroduction, abiding by the old philosophy to "let sleeping dogs lie."

With increased ecological knowledge and improved dissemination of this knowledge, public attitudes have become more positive with regard to conservation, endangered species, and the role of predators. Improved public attitudes, coupled with the high visibility and success of other wolf reintroduction programs, have resulted in some consideration being given to improving the visibility of the red wolf recovery program. One step in this direction would be outreach efforts, as represented by this paper, summarizing the interesting history of the red wolf and the significance of the species and its recovery to other endangered species and reintroduction programs.

Evolution
Already surrounded by extensive controversy regarding its systematic status and conservation, the red wolf recently was confronted by a major new assault, based on genetic studies (Roy et al. 1996, Wayne and Gittleman 1995) purporting to show that it is not a distinct species, or even subspecies, but originated through modern hybridization between the gray wolf and coyote (C. latrans). However, this position has failed to consider early historical and prehistorical records indicating that the red wolf, in much the same form as it exists today, has been present in southeastern North America since the Pleistocene. In an attempt to complete and document these records, the Service has funded a new study to catalogue and reassess all historical, fossil, and archaeological specimens attributable to the red wolf.

Nowak (1979) had argued that C. rufus is identical or immediately related to C. edwardii, a small wolf now thought to have existed 1 to 2 million years ago in the latest Pliocene and early Pleistocene. He suggested the two species represent a line that evolved entirely in North America and that was ancestral to all other wolves. He listed records of C. edwardii from 8 sites and of C. rufus from 11 fossil and 9 archaeological sites. Rundquist (1979) reported another archaeological specimen of C. rufus from North Carolina.

Subsequently, many new fossils of C. edwardii have been discovered, mostly in Florida (Morgan and Hulbert 1995). Examination of this and other material, as part of the new Service study, has suggested that C. edwardii may not be closely related to C. rufus but may have evolved into C. armbrusteri, a relatively large wolf of the middle Pleistocene. The question of the origin of C. rufus thus remains open.

There is increasing evidence that the red wolf is part of a complex of relatively advanced wolves, including C. lupus, that developed in the Old World and then migrated to North America in the late Pleistocene. Apparently, this migration occurred in a number of waves across the Bering Strait, as glaciers advanced and receded and sea levels rose and fell. The earlier migrants may then have differentiated into distinct species or subspecies. Nowak and Federoff (1996) suggested that all available specimens indicate the original presence of a single kind of wolf in the region south of the southern borders of Wisconsin, Michigan, and New York. This wolf, in turn, appeared identical to the earliest specimens of C. rufus from Louisiana.

Additional early material (previously identified as either C. lupus or C. rufus, and, in a few cases, as C. latrans), is being analyzed as part of the new Service study. Together with reassessment of previously examined specimens, preliminary results of this study support the position that only a single species of wolf inhabited the entire region east of the prairies and from Maine to Florida. This species, in turn, seems to have differed sharply from C. lupus of the western prairies and forests of the upper Great Lakes region. The overall picture would seem to involve an initial invasion of North America by a small kind of wolf. Subsequently, larger and more highly evolved kinds of C. lupus moved into the continent from the northwest and competitively forced the earlier and smaller kinds to the east and southeast.

There remains the dilemma, recognized by all investigators since Goldman (1937, 1944), that C. rufus of the southeast shows a close morphological approach to the gray wolf subspecies C. lupus lycaon of southeastern Canada. Indeed, most of the fossil and archaeological specimens of wolves from the eastern United States are not readily distinguishable as either rufus or lupus. There have been repeated suggestions (Lawrence and Bossert 1967, Nowak and Federoff 1996) that rufus and lycaon may have intergraded, and thus that the red wolf is actually not more than subspecifically distinct from C. lupus.

Recent genetic studies of lycaon, especially from the vicinity of Algonquin Provincial Park in southeastern Ontario, indicate that it may be immediately related to rufus, and that these two taxa represent a line that developed in North America separately from C. lupus (Paul Wilson, Wildlife Forensic Laboratory, Trent University, Peterborough, Ontario, personal communication). A sharp morphological distinction between Algonquin lycaon and lupus farther to the west had been reported previously by Kolenosky and Standfield (1975). Nonetheless, Nowak (1979, 1995) and Nowak and Federoff (1996) had found lycaon to be morphologically near to C. lupus than to C. rufus. This problem remains a major stumbling point in the systematics of Canis, especially in understanding the origin of the red wolf.

Nomenclature
The gray wolf was described by Linnaeus (1758), with the type locality being Sweden. The first use of a distinctive name for the North American gray wolf was by Schreber (1775). In comparison, the red wolf was first described as Lupus niger (a technically invalid name) by Bartram (1791) based on observations made in 1774. The third North American Canis, the coyote (C. latrans), was described by Say (1823) based on observations made in 1819. The first publication of a valid scientific name for the red wolf (C. l. var. Rufus) was by Audubon and Bachman (1851). They described two varieties of wolves in the Southeast that were structurally different from other wolves and described the coyote as a full species, uniquely different from wolves. Bangs (1898) stated that the Florida wolf should be elevated to a full species, and Miller (1912) did so as C. floridanus. In the meantime, Bailey (1905) had also elevated the red Texan wolf to a full species (C. rufus).

The current classification is derived from Goldman (1937, 1944). He consigned all Southeastern wolves to one species (C. rufus) and recognized three subspecies--the Texas red wolf (C. r. rufus), the Florida red wolf (C. r. floridanus), and the Mississippi Valley red wolf (C. r. gregoryi). The Mississippi Valley red wolf was named for Tappan Gregory, a pioneer in early animal self portrait flash photography, who obtained photos of red wolves in 1934 and published his experiences in "The Black Wolf of the Tensas" (Gregory 1935). Interestingly, all three photos of red wolves obtained by Gregory were of animals of the black color phase.

Recent work by Nowak and Federoff (1996), and as part of the new Service study, suggests that the historical range of the red wolf may include the entire forested region of Eastern North America from Maine to the Gulf Coast. Therefore, specimens from this range formerly classified as lycaon might now appropriately be assigned to rufus. Recent genetic studies at Trent University (see above) support possible recognition of both lycaon and rufus as components of a distinct species, which could properly be known as C. lycaon. However, the entire issue of the relationship of lycaon and rufus, and whether they are specifically or subspecifically distinct from C. lupus, has yet to be resolved.

Endangered Status
McCarley (1959, 1962) was the first to inform the scientific community that the red wolf was in danger of extinction. Pimlott and Joslin (1968) surveyed likely habitat in Arkansas, Louisiana, and Texas and could confirm extant red wolves remaining only in coastal southeast Texas and southwest Louisiana. The American Society of Mammalogists' Committee on Conservation of Land Mammals led the charge to recognize the red wolf's precarious state in the mid-1960s. The red wolf was one of 72 species included on the original official Federal Endangered Species List that was adopted in 1967 under the Endangered Species Preservation Act of 1966, the first Federal legislation addressing endangered species. This list had been developed over the previous years as an unofficial tabulation of endangered species. The red wolf had been added to this list in 1965 (Pimlott and Joslin 1968). The situation prompted the Service to assign a biologist to study the situation beginning in 1968. It soon became obvious that interbreeding with the more numerous coyote had produced a hybrid swarm that was rapidly pushing the species to extinction (Carley 1975).

The Service decided to remove the last red wolves from the wild and place them in captivity to save the species from inevitable extinction. Morphological standards for red wolves were developed from specimens in the National Museum that had been collected around the turn of the century and, therefore, were thought to have been collected prior to extensive hybridization with coyotes. These standards may have been sufficiently stringent to eliminate some red wolves but this sacrifice was considered necessary to ensure purity of the captive population. From 1973 through 1980 over 400 canids were captured in or near the red wolf's final range in southeast Texas and southwest Louisiana; only 43 of them met the morphological standards (Parker et al. 1990). Breeding experiments were then conducted with these 43 canids. Animals producing questionable or no young, based on morphological standards, were removed from the program and others bore no young, resulting in only 14 founders to begin restoration of the species' population. Because descendants of those lines died early or failed to reproduce, two founder lines have been lost since then, so the present population is descended from only 12 founder lines. The red wolf was declared extinct in the wild in 1980. It has the distinction of being the first documented mammal species to become endangered by interbreeding with a separate species and the first species to be purposely extirpated in the wild in order to save it from extinction and secure a captive-breeding population.

Recovery
Recovery actions for endangered and threatened species are normally thought of in terms of the Endangered Species Act of 1973 (Act), which was the first legislation directing such efforts. However, as indicated earlier, the Service actually started what could be considered recovery actions in 1968, when a biologist was assigned to study the situation in southeast Texas and southwest Louisiana. In addition, the Service's newly created Office of Endangered Species began to develop a recovery plan in 1971, and implementation of this plan officially began in October of 1973. All of these actions preceded passage of the Act on December 28, 1973.

Recovery efforts intensified after passage of the Act, and especially following passage of the Experimental Population Amendment to the Act in 1982. The red wolf program was instrumental in prompting this amendment and was one of only two species (Colorado squawfish was the other) used as examples in Department of the Interior testimony before the Senate Committee on Environment and Public Works, Subcommittee on Environmental Pollution on April 19,1982. The fourth experimental population (third experimental population rule) designated after passage of the amendment was for the red wolf (the Delmarva fox squirrel, Colorado squawfish, and woundfin preceded it), and it was the first experimental population for a predatory mammal.

This first experimental population of red wolves, located in northeastern North Carolina, was designated in 1986 (Parker et al. 1986), and many of the regulations promulgated under the final rule were firsts for any experimental population. These include several regulations that were later adopted by the Rocky Mountain and Mexican wolf reintroduction programs; i.e., (1) allowed taking that is incidental to an otherwise lawful activity, provided it was unavoidable, unintentional, and not the result of negligent conduct lacking reasonable due care; (2) provided for continuous monitoring of the population; (3) provided for vaccination against diseases and parasites of all released and subsequently handled animals; and (4) provided for evaluation after a trial period. Evaluation after a trial period is also used in the reintroduction of many other nonendangered species such as elk, black bear, etc. There were also some first-time regulations that were unique to the red wolf; i.e., (1) provided for the recapture of animals that move off of Federal land and (2) allowed for the taking of animals that constitute a demonstrable but non-immediate threat to human safety or that are responsible for depredations to lawfully present domestic animals or other personal property.

A second red wolf experimental population was designated in 1991 for a reintroduction into the Great Smoky Mountains National Park (Park) in eastern Tennessee (Henry 1991). Here again several regulations were first-time efforts for any species. Harassment that is not lethal or physically injurious was allowed; this provision was later used in the Rocky Mountain and Mexican wolf reintroductions. Another regulation--allowing taking that is incidental to lawful recreational activities--was a first for wolf populations. A third regulation unique to the red wolf allowed private landowners to take animals after efforts by project personnel to capture such animals have been abandoned, provided that the Service approved such actions in writing and takings are reported to the Service within 24 hours.

A final rule (Henry 1995) to provide consistency in regulations among red wolf experimental populations also plowed new ground with regard to regulations. The Service agreed to remove animals from private land where they are not wanted, if possible; this basic provision was also later used by the Mexican Wolf program. A new regulation also allowed taking that is not intentional or willful on private land.

The red wolf experimental population in the park also has the distinction of being the first experimental population to be delisted. Unfortunately, this delisting was because the reintroduction was unsuccessful (Henry 1999).

Captive Management
Captive management has been an integral part of red wolf recovery and was responsible for rebuilding the population to sufficient numbers for reintroductions. The first red wolf was placed in captivity in 1969, and the first litter born in captivity was in 1977. A number of captive management techniques were initiated by red wolf program personnel; for example, using large salmon nets to capture (secure unrestrained animals for examination) wolves running free within sizable outdoor pens and entering dens to restrain and handle adults and pups for examination. The red wolf captive-breeding program was approved by the American Zoo and Aquarium Association (AZA) for the development of a Species Survival Plan (SSP) in 1984. The Red Wolf Recovery/Species Survival Plan completed in 1990 was the first plan to combine the two concepts into one document (Parker et al. 1990). The Point Defiance Zoo and Aquarium was recognized by AZA for long-term propagation in 1987 when it received the Edward H. Bean Award. The program has also established a genome resource bank, using frozen sperm in conjunction with noninvasive detection of estrus, with the potential use for artificial insemination of selected individuals. The first successful artificial wolf insemination from semen collected through electro-ejaculation occurred in the red wolf. Unexpended funds from a contracted study of attitudes and economic impacts in areas where red wolves were reintroduced (Rosen 1997) were used by Rosen to determine the contributions of A.A. cooperators to the red wolf recovery program; this was the first effort to recognize and estimate the financial outlays of A.A. facilities to endangered species programs.

Reintroduction
The red wolf recovery program pioneered the development and utilization of several reintroduction techniques, including (1) the use of portable acclimation pens, with a ground skirt to prevent wolves from digging out; (2) the development of acclimation periods sufficient to maximize the chances of the animals' remaining in the release area; (3) the use of soft releases to minimize immediate dispersal upon release; (4) the periodic use of supplemental feeding for such purposes as localizing the animals, easing the transition to obtaining wild prey, and/or at other stressful times, such as aiding single parents in providing for young when the other parent has died; (5) the use of abdominal transmitters as a back-up telemetry system in case radio telemetry collars fail; (6) the use of recapture collars in a reintroduction project; (7) the use of different release groups (family groups, pairs with young pups, pairs just prior to whelping) to achieve specific objectives, such as maintaining pack cohesion, localizing the animals, etc.; and (8) the release of wolves in the presence of livestock. In the last example, the Service built and maintained, in cooperation with the lessee, a nursery corral to protect cows about to calve and young calves from wolf predation. Flood damage to this barbed wire nursery corral resulted in its replacement with a high tensile electric nursery corral paid for out of the livestock lessee's fees.

The red wolf recovery program was the first reintroduction program to accomplish: (1) the successful reintroduction of a species extinct in the wild in North America (only the Arabian oryx [Oryx lecheries] reintroduction preceded the red wolf on an international scale in this regard); (2) the successful reintroduction of a large predator; (3) the successful reintroduction of wolves; (4) the successful reintroduction of a mammal from captive stock in North America (only the Arabian oryx and the golden lion tamarin [Leontopithecus rosalia] reintroductions preceded the red wolf on a global scale); and (5) incorporation of private landowners as willing participants in wolf reintroductions. The first success in reintroducing wolves anywhere is a significant achievement, considering that it was done in the Eastern United States where there is the least amount of public land and the most people.

The red wolf recovery program also pioneered another concept--the use of islands off the Atlantic and Gulf Coasts as propagation sites. Bulls Island, South Carolina, was originally used to develop reintroduction techniques in 1976 and 1978. This same island and three additional islands were later used (three are still used) as propagation sites. The idea of using islands was conceived as a way to give captive stock experience in the wild before reintroducing them into mainland sites. This would, hopefully, increase survival by providing wolves the opportunity to develop or reinforce wild behavioral traits (Henry and Lucash 1998).

Additional Contributions
The taxonomic controversy surrounding the species resulted in two petitions to delist the species. The first petition was filed in 1991 and was based on results from mitochondrial DNA studies. The second petition was filed in 1995 and referenced nuclear DNA study results as the basis for filing the petition. The Service must use the best available scientific and commercial data in making a finding on petitions. Using this criterion, it was determined that neither petition presented substantial information to indicate that the petitioned action may be warranted (Henry 1992, 1997). The Service distributed the response to the 1991 petition to all Service offices with endangered species responsibilities as an example of a proper petition response.

Other endangered species programs have benefitted from the red wolf recovery program. Personnel from the program have served in an advisory capacity to other reintroduction programs, including the black-footed ferret, the Rocky Mountain wolf, and the Mexican wolf. Experienced personnel in the program were also tapped for other wolf reintroduction projects; in particular, the project leader position for the Yellowstone wolf reintroduction project. The state of Arizona also tried to hire a former red wolf program employee to head up Mexican wolf reintroduction efforts for that agency. Unfortunately, personnel matters prevented this appointment from materializing.

Literature Cited
Audubon, J. J., and J. Bachman. 1851. The viviparous quadrupeds of North America, Volume 2, New York, New York. 334 pages.

Bailey, V. 1905. Biological survey of Texas. North American Fauna Number 25. 222 pages.

Bangs, O. 1898. The land mammals of peninsular Florida and the coast region of Georgia. Proceedings, Boston Society of Natural History 28(7):157-235.

Carley. C. J. 1975. Activities and findings of the red wolf recovery program from late 1973 to July 1, 1975. U.S. Fish and Wildlife Service, Albuquerque, New Mexico. 215 pages.

------. 1944. Classification of wolves. Pages 389-636 in S. P. Young and E. A. Goldman. The wolves of North America, Part 2. American Wildlife Institute, Washington, D.C.

Gregory, T. 1935. The Black Wolf of the Tensas. Program of Activities of the Chicago Academy of Sciences 6(3):35-68.

Henry, V. G. 1991. Endangered and threatened wildlife and plants; determination of experimental population status for an introduced population of red wolves in North Carolina and Tennessee. Federal Register 56(213):56325-56334.

------. 1992. Endangered and threatened wildlife and plants; finding on a petition to delist the red wolf (Canis rufus). Federal Register 57(8):1246-1250.

------. 1995. Endangered and threatened wildlife and plants; revision of the special rule for nonessential experimental populations of red wolves in North Carolina and Tennessee. Federal Register 60(71):18940-18948.

------. 1997. Endangered and threatened wildlife and plants; 90-day finding for a petition to delist the red wolf. Federal Register 62(236):64799-64800.

------. 1999. Endangered and threatened wildlife and plants; proposed removal of nonessential experimental population status for an introduced population of red wolves in North Carolina and Tennessee and proposed revision of the special rule for a nonessential experimental population of red wolves in North carolina. Federal Register (in prep.)

Henry, V. G., and C. F. Lucash. 1998. Red wolf status report. Presented in Session 4 - Acting Locally, at the Conservation Research Consortium, Riverbanks Zoo & Garden, Columbia, South Carolina. 10 pages.

Kolenosky, G. B., and R. O. Standfield. 1975. Morphological and ecological variation among gray wolves (Canis lupus) of Ontario, Canada. Pages 62-72 in M. W. Fox, editor. The Wild canids, their systematics, behavioral ecology and evolution. Van Nostrand Reinhold, New York.

Lawrence, B. and W. H. Bossert. 1967. Multiple character analysis of Canis lupus, latrans, and familiaris, with a discussion of the relationships of Canis niger. American Zoologist 7:223-232.

------. 1962. The taxonomic status of wild Canis (Canidae) in the South-central United States. Southwestern Naturalist 7(3-4):227-235.

Miller, G. S. 1912. List of North American land mammals in the United States National Museum, 1911. U.S. National Museum Bulletin Number 79, Washington, D.C. 455 pages.

Morgan, G. S., and R. C. Hulbert, Jr. 1995. Overview of the geology and vertebrate biochronology of the Leisey Shell Pit Local Fauna, Hillsborough County, Florida. Bulletin of the Florida Museum of Natural History 37 Part I(1):1-92.

Nowak, R. M. 1979. North American Quaternary Canis. Museum of Natural History, University of Kansas Monograph 6. 154 pages.

------. 1995. Another look at wolf taxonomy. Pages 375-397 in L. N. Carbyn, S. H. Fritts, and D. R. Seip, editors. Ecology and conservation of wolves in a changing world: proceedings of the second North American symposium on wolves. Canadian Circumpolar Institute, University of Alberta, Edmonton.

Nowak, R. M., and N. E. Federoff. 1996. Systematics of wolves of eastern North America. Pages 187-203 in N. Fascione and M. Cecil, ed., Proceedings of Defenders of Wildlife Wolves of America Conference. Washington, D.C.

Parker, W. T., M. P. Jones, and P. G. Poulos. 1986. Endangered and threatened wildlife and plants; determination of experimental population status for an introduced population of red wolves in North Carolina. Federal Register 51(223):41790-41796.

Parker, W. T., R. Smith, T. Fosse, and U. S. Seal. 1990. Red Wolf Recovery/Species Survival Plan. Southeast Region, U.S. Fish and Wildlife Service, Atlanta, Georgia. 110 pages.

Pimlott, D. H., and P. W. Joslin. 1968. The status and distribution of the red wolf. Transactions, 33rd North American Wildlife and Natural Resources Conference:373-384.

Rosen, W. E. 1997. Red wolf recovery in northeastern North Carolina and the Great Smoky Mountains National Park: public attitudes and economic impacts. Department of Policy Analysis and Management, Cornell University, Ithaca, New York. 160 pages.

Roy, M. S., E. Geffen, D. Smith, and R. K. Wayne. 1996. Molecular genetics of pre-1940 red wolves. Conservation Biology 10:1413-1424.

Rundquist, J. 1979. Analysis of the flora and fauna remains from proto-historic North Carolina Cherokee Indian sites. Ph.D. dissertation, Department of Zoology, North Carolina State University, Raleigh, North Carolina. 410 pages.

Say, T. 1823. In S. H. Long and E. James. Account of an expedition from Pittsburgh to the Rocky Mountains performed in the years 1819 and 1820. 2 Volumes. Philadelphia, Pennsylvania.

Von Schreber, J. C. D. 1775. Die saugthiere in abbildungen nach der Natur mit beschreibungen. 8 Volumes. Erlangen.

Wayne, R. K., and J. L. Gittleman. 1995. The problematic red wolf. Scientific American 273(1):36-39.