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Re-evaluation of the evidence for the importation of red foxes from Europe to colonial America: Origins of the southeastern red fox (Vulpes vulpes fulva)
Biological Conservation 158 (2013) 74–79
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Re-evaluation of the evidence for the importation of red foxes from Europe to colonial America: Origins of the southeastern red fox (Vulpes vulpes fulva) Jennifer K. Frey ⇑ New Mexico State University, Department of Fish, Wildlife and Conservation Ecology, P.O. Box 30003, MSC 4901, Las Cruces, NM 88003-8003, USA
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Article history: Received 10 March 2012 Received in revised form 19 July 2012 Accepted 26 July 2012 Available online 28 November 2012 Keywords: Red fox Vulpes vulpes fulva Exotic species Geographic range shifts Scientific misconception United States
a b s t r a c t It is widely accepted that the red fox subspecies of southeastern North America (Vulpes vulpes fulva) is descended from red foxes imported from Europe to the American colonies for sport hunting. Thus, V. v. fulva is considered an exotic invasive organism that has apparently expanded its range to occupy much of the contiguous United States, with exception of the range of relict native populations in the western mountains and Sacramento Valley. I examined the evidence for importations and found that all claims of introduction stemmed to just two literature sources that were based on vague second-hand information. Together with results of morphological and genetic studies, this information indicates that V. v. fulva is native to North America. Thus, historical shifts in distribution and abundance of V. v. fulva and current management approaches should be re-evaluated in light of native status. Evaluating original sources of information can curtail the proliferation of inaccurate information and conclusions that influence conservation decisions. Ó 2012 Elsevier Ltd. All rights reserved.
1. Introduction The red fox (Vulpes vulpes) occurs throughout most of the Holarctic, except some desert and tundra areas, and it has been introduced to Australia and some other islands (Lariviere and Pasitschniak-Arts, 1996). The International Union for Conservation of Nature considers it one of the world’s worst 100 exotic invasive species due to negative impacts on native fauna where it has been introduced, including Australia and North America (GISD, 2012). It is thought that both native and exotic populations of red foxes occur in North America, with native foxes restricted to Alaska, Canada, and relict populations in the western United States (US; Churcher, 1959; GISD, 2012; Kamler and Ballard, 2002). Most treatments of the red fox in North America state that red foxes from Europe were introduced by the American colonists to southeastern North America for equestrian sport hunting with hounds – a claim that essentially has become ‘‘common knowledge.’’ Consequently, many authors have concluded that the southeastern subspecies in North America, Vulpes vulpes fulva, is directly descended from those introductions or a hybrid between European and native North American foxes (e.g. Ables, 1975; Leopold and Chamberlain, 2001; Kamler and Ballard, 2002; Rhoads, 1903; but see Aubry et al., 2009; Sacks et al., 2010; Statham et al., 2012). The distribution of V. v. fulva is thought to have expanded since colonial times (Kamler and Ballard, 2002). Its distribution currently ⇑ Tel.: +1 575 640 9782; fax: +1 575 646 1281 E-mail address: [email protected] 0006-3207/$ - see front matter Ó 2012 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.biocon.2012.07.027
extends from the Great Lakes south to the Gulf of Mexico and from the eastern seaboard of the US, westward across the southern Great Plains to Colorado and New Mexico (Frey, 2004; Hall, 1981), and Kamler and Ballard (2002) concluded that V. v. fulva replaced the native V. v. regalis in the northern Great Plains. It also is thought to have been introduced and established in low elevation regions within various areas of western North America (e.g. Buskirk and Zielinski, 2003; Kamler and Ballard, 2002; Lewis et al., 1999), although specific records detailing the source population and the date, location, and numbers of animals introduced are generally lacking (but see Aubry, 1983, 1984; Whitlow and Hall, 1933). If these scenarios are true, then as suggested by Kamler and Ballard (2002), all current populations of the red fox in the contiguous US, with exception of relict populations occurring in the western mountains (V. v. macroura, V. v. cascadensis, V. v. necator) and Sacramento Valley (V. v. patwin), are not native to North America and, hence, present a threat to native species. This interpretation has practical management implications as the attitudes and approaches towards native and exotic species are usually diametrically opposed (e.g. Clout and Williams, 2009). Despite the important ramifications, the exotic status of V. v. fulva usually has been taken as fact without any reasonable attempt to validate the claim of introduction based on original sources of information (e.g. Kamler and Ballard, 2002). Most modern treatments cite relatively recent works, especially Churcher (1959), that are themselves based on earlier works that cite even earlier works. Some treatments go so far as to provide no citations at all (e.g. Hamilon and Whitaker, 1979; Presnall, 1958), which are
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subsequently cited as evidence by other works (e.g. Fields, 2007; Long, 2003). A notable exception is the recent work by Statham et al. (2012). They built upon a prior phylogeographic analysis of native red foxes in North America (Aubry et al., 2009) to explicitly test the origins of red foxes throughout the eastern contiguous US, which they considered to have been established since the mid-1800s (Statham et al., 2012). They found no evidence of European haplotypes anywhere in North America and concluded that ‘‘recently established’’ populations in the eastern US were primarily the result of a southward range expansion from eastern Canada and northeastern US, and were not derived from introductions from Europe (Statham et al., 2012). However, their sampling included only 18 individuals from populations considered recently established in the eastern US, and did not include key areas where colonial introductions are frequently said to have occurred (e.g. eastern Virginia, Maryland). In addition, the methods they employed were based on mtDNA and, hence, were only able to track matrilineal ancestry. To better understand the origins of the southeastern red fox, I compiled original information concerning the importation of red foxes to the American colonies, such as dates, source locations, release locations, and numbers of animals. In large part, this was made possible by the availability of electronic copies of rare old texts, such as through the Biodiversity Heritage Library, American Journeys, Internet Archive, Google Books, Proquest American Periodicals Services Historical database, and other library resources.
2. Evidence for importation The chain of evidence for all references to the importation of red foxes from Europe to colonial America appears to extend back to only two references. The first was a travel journal written by the Swedish naturalist Pehr Kalm describing his experiences in eastern North America from 1748 to 1751 (Egerton, 2006). In Philadelphia on 7 November 1748, Kalm (1772: pp. 221–222) wrote: ‘‘Mr. Bartram [i.e. American botanist John Bartram who lived in New Jersey near Philadelphia] and several others assured me, that, according to the unanimous testimony of the Indians, this kind of foxes never was in the country, before the Europeans settled in it. But of the manner of their coming over I have two different accounts: Mr. Bartram and several other people were told by the Indians, that these foxes came into America soon after the arrival of the Europeans, after an extraordinary cold winter, when all the sea to the northward was frozen: from hence they would infer, that they could perhaps get over to America upon the ice, from Greenland or the northern parts of Europe and Asia. But Mr. Evans [i.e. Lewis Evans, an engineer from vicinity of Philadelphia], and some some [sic] others, assured me, that the following account was still known by the people. A gentleman of fortune in New England, who had a great inclination for hunting, brought over a great number of foxes from Europe, and let them loose in his territories, that he might be able to indulge his passion for hunting.⁄This is said to have happened almost at the very beginning of New England’s being peopled with European inhabitants. These foxes were believed to have so multiplied, that all the red foxes in the country were their offspring.’’ [The asterisk is to a footnote added by John Reinhold Forster who translated Kalm’s work into English. Forster did not consider either of Kalm’s accounts satisfactory; he believed that the red fox immigrated to North America ‘‘in remote times’’ from eastern Asia and then spread across the continent.] The second source of information is from the first volume (1829) of the American Turf Register and Sporting Magazine, which primarily served as a venue to register the performance and pedi-
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grees of purebred horses. However, it also contained information on rural sports including articles on the natural history of American game species. The editor, John Stuart Skinner, solicited readers to submit letters describing anecdotes about wildlife. The most widely cited ‘‘records’’ of importations of the red fox are based on an editorial note and two letters to the editor in this volume. Skinner’s editorial note (Skinner, 1829: pp. 9) follows an article on the manner in which George Washington hunted foxes: ‘‘The red fox is supposed to have been imported from England, to the Eastern Shore of Maryland, by a Mr. Smith, and to have emigrated across the ice to Virginia, in the hard winter of 1779–80, when the Chesapeake was frozen over.’’ In response, one letter to the editor from Fredericksburg, Virginia (Skinner, 1829: pp. 74) stated: ‘‘This I do not doubt; but I think it probable they were brought over and turned out at other places, and at very early periods. In 1789, when quite a boy, I was at the death of the first red fox killed in Perry county, Pa. . . . Not a person present, or any one who saw it for some days had ever seen or heard of an animal of the kind. At last it was shown to a Mr. Lenarton, an old Jersey-man, who pronounced it an English fox. He said the red fox was imported into New York from England, by one of the first English governors, who was said to be a great sportsman, and turned out on Long island, where they remained for many years, but at last made their way, on the ice, to the main land, and spread over the country.’’ In another letter to the editor written from Richmond, Virginia (Skinner, 1829: pp. 197): ‘‘The tradition of this part of the country. . .differs from the conjecture which seems to prevail farther north, in this; that . . . he is supposed to have been brought from the continent—Germany, I think—and not from the island of Great Britain.’’ In an Outing magazine article on the origins of fox hunting in America, Hiss (1897: pp. 14–15) provided a greatly embellished version of the story told in Skinner’s (1829) editorial comment, which he described as a ‘‘tradition’’ and ‘‘old tale.’’ Cogent details of Hiss’ story are that in 1730 eight wealthy farmers from Talbot County, Maryland, instructed the captain of the tobacco schooner ‘‘Monocacy,’’ which was owned by one of them, to bring back eight pairs of red foxes on the next trip to Liverpool, England. The animals were released in Maryland on the eastern shore of the Chesapeake Bay and a festival celebrated the event. The foxes multiplied and expanded into Virginia across the ice during the hard winter of 1779–80. In his book on fox hunting in America Potts (1912: pp. 5–7), dismissed the two stories written as letters to the editor of American Turf Register and Sporting Magazine as interesting legends and he considered Hiss (1897) story as true. Potts (1912: pp. 7) stated that the story had been verified by Skinner. Thus, stories told by Skinner (1829) and Hiss (1897) are the same. Potts (1912:7) changed some of the details in Hiss (1897), calling the name of the schooner ‘‘Monaccasy,’’ and stating that the foxes were consigned to a Mr. Smith. These stories were frequently recounted in periodicals and natural history literature, sometimes word-for-word (although sometimes lacking citations) and sometimes with misconstrued information (e.g. Pennant, 1784; Doughty and Doughty, 1830; Hunter, 1876; Rhoads, 1903). Reports of introductions of red foxes in 1650 (e.g. Gilmore, 1946; Long, 2003) are apparently misconstrued based on reports of the first importation of fox hounds to North America by Robert Brooke, an event that has been interpreted as the beginnings of fox hunting in America (Hiss, 1897; Brooke, 1910). 3. Supporting evidence for native status I found no direct evidence or firsthand accounts of any importation of red foxes to colonial America. The total ‘‘evidence’’ consists
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of a vague second-hand report given to Kalm by Evans, the vague story told by Skinner, which might be the same as the one told by Kalm, a vague second-hand story by Lenarton given by another person in a letter to the editor, and a rumor of a different source population given in a letter to the editor. Hiss’ more detailed version of the one given by Skinner, was written ca. 250 years after the supposed event and likely represents literary license. Even the earliest story occurred a century or so after the alleged importation. A recurring theme in these stories is that the red fox expanded its range over ice during a hard winter, which might ultimately derive from the supposed comments by the Native Americans that were relayed to Kalm by Bartram. By applying evidentiary standards used for interpreting species occurrence records, these scant, second-hand reports are deemed highly questionable (Frey, 2006; McKelvey et al., 2008). In contrast, introductions of foxes to Alaskan islands and Australia are relatively well documented (Bailey, 1993; Long, 2003). Given the popularity of fox hunting in colonial America, importations might have been reported in local newspapers, at least if introduction occurred during the last few decades of the colonial period when there were more media outlets. I found no such articles in my searches of the Proquest American Periodicals Series database, which includes >1000 sources dating back to 1741. If such an importation occurred, the paucity of records suggests it would have been an exceedingly rare event, probably because foxes (including the gray fox [Urocyon cinereoargenteus]) that could be used for sport were already present in the eastern US. There is a common modern misperception that gray foxes were not considered suitable targets for equestrian fox hunts with hounds because they climb trees (e.g. Voigt, 1999; Peddicord, 2010). However, the early literature is replete with accounts extolling the virtues of gray foxes as a quarry for such hunts (e.g. Audubon and Bachman, 1846; Godman, 1831; Page, 1892). In Australia where foxes were entirely absent, the desire to establish a population for hunting would have been more keen. However, even in Australia most of the early attempts to introduce the red fox failed (Long, 2003). It seems unlikely that any rare release of imported red foxes into North America would have established a population, especially given that any such importation would have been for hunting purposes. Thus, I found no compelling evidence that red foxes from Europe were successfully introduced in colonial America. Morphological and genetic data support the interpretation that V. v. fulva is native to North America. If the red fox of the eastern US is the result of importations, then it would be referable to the subspecies native to Great Britain and other areas of Europe, V. v. crucigera, as it is in Australia (Ellerman and Morrison-Scott, 1951). However, Doughty and Doughty (1830) noted that none of the cabinets (i.e. Victorian natural history collections) in Philadelphia contained specimens of the European red fox, and all known specimens from the US at that time were referable to V. v. fulva (also recorded in the literature as V. fulvous, V. fulvus, V. fulva, and V. pennsylvanica). I used the Mammal Networked Information System (http://manisnet.org/) to search for specimens of red foxes in the eastern US; none were referred to V. v. crucigera (all were referred to V. v. fulva). All studies that have compared morphology of red foxes from eastern North America and Eurasia have found consistent differences, which have been recognized at either the species or subspecies level (Baird, 1857; Bangs, 1897; Churcher, 1959; Merriam, 1900; Paradiso, 1969). Molecular genetic data based on mtDNA have shown that red foxes from Eurasia are in a subclade different from populations in North America (Aubry et al., 2009). With that phylogeography as a framework, Statham et al. (2012) examined genetic signals of red foxes from throughout the eastern US and found no Eurasian haplotypes. Similarly, specimens of red foxes taken at low elevations in California that were hypothesized by Kamler and Ballard (2002) to represent
introduced V. v. fulva, had haplotypes within North American clades rather than in the Eurasian subclade (Sacks et al., 2010, 2011; Statham et al., 2012). Further, molecular data (Aubry et al., 2009; Statham et al., 2012) have not supported the views of Kamler and Ballard (2002) that ‘‘exotic’’ V. v. fulva spread like a wave across the US replacing native red foxes. Thus, Statham et al. (2012) concluded that ‘‘recently established’’ populations in the eastern US were the result of a natural range expansion from southern Canada.
4. Distribution of the red fox in the eastern US Today the red fox is thought to occur throughout the US east of the Rocky Mountains, with exception of coastal Florida, the southern Great Plains, and southern Texas (Cypher, 2003). Aubry et al. (2009) and Statham et al. (2012) each provided a map of the geographic range of red foxes that were considered to be unequivocally indigenous (i.e. no introductions have been proposed as influencing distribution) in North America prior to European settlement. Areas south of that range where red foxes are currently known to occur, were denoted ‘‘recently established populations’’ and an explicit goal of Statham et al. (2012) was to evaluate ancestry of those populations. In the US east of the Rocky Mountains, the only region considered within the unequivocal range of indigenous red foxes was northern New England (i.e. Maine and the northern portions of New York, Vermont, and New Hampshire); rationale for this interpretation was detailed in Statham et al. (2012), and was based primarily on information contained in Audubon and Bachman (1851); Churcher (1959); Newberry (1857), and Rhoads (1903). However, the pre-settlement geographic range of the red fox in the eastern US requires further evaluation. There is very little reliable information about wildlife in America prior to 1800. The early 1800s was the period during which most of the initial scientific exploration of the country occurred, and also was a period of proliferation of news periodicals and natural history publications. It is important to recognize that lack of a record is not evidence that a species was absent (Keay, 1901) or that the first documentation of a species in a region is not necessarily the first occurrence of that species in the area (Frey, 2009). For instance, the red fox was reported in the Great Plains during several of the first scientific expeditions to the region, including what is now North Dakota in 1805 by Meriwether Lewis and William Clark (http://lewisandclarkjournals.unl.edu/index.html), what is now Minnesota in 1805 by Zebulon Pike (Pike, 1810), and what is now Nebraska in 1819 by Edwin James (Genoways and Ratcliffe, 2008; James, 1823). Given that these observations preceded European settlement of those regions, there is no reason to presume that they represent anything other than native occurrence of the red fox on the Great Plains. There also are credible reports of red foxes in the American colonies. For instance, Thomas Morton and John Josselyn reported both red foxes and gray foxes in Massachusetts in the early 1600s at the beginning of European settlement (Josselyn, 1865a,b; Morton, 1883). Kalm (1772), who primarily visited Pennsylvania, stated that both the red fox and gray fox occurred in the colonies, but that the red fox was scarce. On basis of Kalm’s report, Pennant (1784) stated that red foxes decreased in abundance to the south and that none were found south of Pennsylvania. Further, red foxes, but rarely gray foxes, were viewed as an important depredator of livestock and poultry (e.g. Thompson, 1853; Warren, 1897), and hence bounties were instituted for their control. Bounties on red foxes were some of the earliest wildlife laws enacted, including 1635 in Massachusetts and 1641 in Rhode Island (Rabushka, 2008). In 1714 a bounty on wolves in New Jersey was expanded to include red foxes (Rhoads,
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1903). In Pennsylvania, a bounty on foxes was established in 1718, and records from Chester County, show bounties paid on more than 453 foxes during 11 years prior to 1747 (Barclay, 2002). Kalm (1772) stated that there were bounties for red fox in Pennsylvania and all other provinces. The idea that red foxes were introduced to the eastern US was fueled, in part, by studies of vertebrate remains at archeological sites in southern Pennsylvania that lacked evidence of red foxes (Gilmore, 1946; Miller, 1940). While the red fox may have been absent from some local faunas, it is now possible to evaluate late Quaternary mammal distributions across thousands of sites in the US using the Faunmap electronic database (http://www.ucmp. berkeley.edu/faunmap/about/index.html). Although Faunmap is a powerful tool for estimating historical mammal distributions, some records may not be verified. However, errors in this case are likely to be minimized because there are many skeletal characteristics that distinguish between the red fox and gray fox (Gilmore, 1946), and other North American members of the genus Vulpes (i.e. the arid-land foxes V. velox and V. macrotis) and Urocyon (i.e. U. littoralis) are unlikely to have been present east of the Great Plains during this epoch. Using Faunmap, I examined 2,861 sites from the late Holocene (450–4500 B.P.) for records of red foxes and gray foxes in order to reconstruct their distribution east of the Rocky Mountains in the US. Sites of this age appeared best represented in the Ozark-Quachita Interior Highlands and the Interior Low Plateaus and Central Lowlands; sites were sparse in the Appalachian Highlands and the Gulf and Atlantic Coastal Plain (except Florida; physiographic regions follow http://tapestry. usgs.gov/physiogr/physio.html). I found 112 records of red foxes and 178 records of gray foxes. Records of both species were dispersed across the eastern US and exhibited similar distribution patterns. Records of red foxes occurred from Maine to the Missouri Plateau (e.g. North Dakota and South Dakota) and as far south as southern Georgia, Arkansas, and Oklahoma. Records of red foxes were sparse in the Appalachian Plateau and Interior Low Plateaus regions; there were no records from Florida and the High Plains. Records of gray foxes were comparatively more dense throughout the eastern US, but were absent from New England (i.e. northeast of New York). The northernmost records of gray foxes were from southern Ontario, central Wisconsin, and southern Minnesota; to the west records were sparse or absent in most of the Great Plains and Osage Plains. Thus, historical records and other evidence support a preEuropean native occurrence of red foxes in the US east of the Rocky Mountains. Consistent with its boreal affinities, the evidence presented herein (see also Allen, 1876; Rhoads, 1903) suggests that the red fox was primarily associated with the Canadian and Transition life zones (sensu Merriam, 1898), which in the eastern US extends from Maine south along the Appalachian Mountains and westward from New York through the Great Lakes region and the Missouri Plateau of the Great Plains. The red fox was likely rare or absent in much of the Austral zone, especially in southern deciduous forests. However, given the species’ ecological plasticity and capacity for long distance movements (Lariviere and Pasitschniak-Arts, 1996), it also might have been sparsely distributed in some regions of the Austral zone where an open habitat structure was present, such as prairies, beaches, and perhaps some pine forests. However, based on repeated testimony of the appearance of red foxes in places where it had not previously been noticed, it appears that the red fox historically expanded its distribution and abundance within the Austral zone (e.g. Audubon and Bachman, 1851; Hahn, 1909; Rhoads, 1903; Schmidly, 2002). Several interrelated factors may account for the apparent increase in distribution and abundance of red foxes within the Austral zone of the eastern US. Prior to European settlement, various types of forests dominated most of the eastern US. Foster et al.
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(2002) demonstrated widespread changes to wildlife assemblages in response to changes in abundance of forests as a result of postsettlement land-use. For instance, red foxes were included in a group of open land species that experienced historical increases in abundance that peaked in the mid-1800s as land was cleared for settlements, timber, and farms, but have recently declined as forests have regenerated. In contrast, gray foxes were in a group of woodland species that experienced concomitant historical declines in abundance, but recent increases (Foster et al., 2002). Indeed, many early historical accounts of foxes noted apparent increases in abundance and distribution of red foxes, and concomitant decreases in abundance of gray fox, as areas were settled and cleared of forests in the Austral zone (e.g. Allen, 1876; Hahn, 1909; Newberry, 1857). These changes in abundance are consistent with the habitat selection, predator escape strategies, locomotion, and other adaptations of each species (Cypher, 2003; Feeney, 1999). The gray fox is a true omnivore that prefers plants and eats a high proportion of insects, in addition to small mammals and birds; in contrast, the red fox is primarily carnivorous with a diet dominated by mammals (Cypher, 2003; Hockman and Chapman, 1983; Trapp and Hallberg, 1975). The most important prey for red foxes in the eastern US are cottontails (Sylvilagus spp.), although diets vary by region, and may also contain large proportions of other mammals such as voles (Microtus spp.) and woodchucks (Marmota monax). Most of the important prey species for red foxes are associated with early successional habitats and their populations are thought to have increased with deforestation and settlement, peaking in the late 1800s and early 1900s (Litvaitis, 2001). This increase in abundance and distribution of favored prey would have preferentially benefited the red fox. Lastly, intraspecific competition is thought to have an important influence on distribution and abundance of canids. Historically, wolves (Canis spp.) occurred throughout all of the eastern US, but were extirpated from most of the region in the wake of settlement due to control efforts. Mesopredator release due to extirpation of wolves could have benefited red foxes (Elmhagen and Rushton, 2007; Ritchie and Johnson, 2009). However, following the loss of wolves, coyotes (C. latrans) subsequently expanded their range from the plains region east to the Atlantic seaboard, which may result in renewed suppression of red fox populations by coyotes (Cypher, 2003).
5. Conclusions The belief by some early naturalists that V. v. fulva was an exotic was influenced by observations of apparent increases in abundance and range expansions of the red fox into areas where the species originally had been scarce or absent in southeastern North America (e.g. Allen, 1876; Audubon and Bachman, 1851; Bailey, 1896; Baird, 1857; Gilmore, 1946; Hahn, 1909). More recently, apparent expansion of V. v. fulva into western North America likewise has been attributed to these foxes being ‘‘descendants from European stock and therefore . . . better adapted to human activities, urban areas, and human-induced changes to the environment’’ (Kamler and Ballard, 2002: pp. 375). My study, which is based on independent lines of evidence, provides broad support for the central conclusions of Statham et al. (2012) – that red foxes in the eastern US are not the descendants of importations from Europe. However, my results also suggest a need to reevaluate origins of populations of red foxes, particularly in the central US, given evidence that the species was historically present in that region. Further, native status of V. v. fulva in North America calls for a reinterpretation of mechanisms responsible for changes in its distribution and abundance. As with most game species, there is evidence of isolated translocations of red foxes within the US and there are reported instances of animals escaping from fur farms (e.g. Aubry, 1983, 1984;
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Kellogg, 1939). Thus, conservation issues related to introduced populations of V. v. fulva in some areas of the western US where red foxes did not occur historically will remain (e.g. Perrine et al., 2007; Sacks et al., 2010). In these regions, conservation and management decisions should be predicated on native status of the red fox in North America. Finally, as with the origins of V. v. fulva, the proliferation of inaccurate information can be averted by verifying the original sources of cited ‘‘facts.’’ The ready access to electronic copies of rare old text can provide new perspectives on old problems. References Ables, E.D., 1975. Ecology of the red fox in North America. In: Fox, M.W. (Ed.), The Wild Canids: their Systematics Behavioral Ecology and Evolution. Van Nostrand Reinhold, New York, 508p (pp. 216–236). Allen, J.A., 1876. The former range of some New England carnivore mammals. Am. Nat. 10, 708–715. Aubry, K.B., 1983. 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Invasive Species Management: a Handbook of Principles and Techniques. Oxford Univ. Press, Oxford, United Kingdom, pp. 308. Cypher, B.L., 2003. Foxes. In: Feldhame,r G.A., Thompson, B.C., Chapman, J.A. (Eds.), Wild mammals of North America: Biology, Management, and Conservation. John Hopkins Univ., Baltimore, Maryland, pp. 511–546. Doughty, J., Doughty, T., 1830. The Cabinet of Natural History and American Rural Sports, vol. 1. J. and T. Doughty, Philadelphia. Egerton, F.N., 2006. A history of the ecological sciences, Part 22: Early European naturalists in eastern North America. Bull. Ecol. Soc. Am. 87, 341–356. Ellerman, J.R., Morrison-Scott, T.C.S., 1951. Checklist of palaearctic and Indian mammals 1758–1946. Brit. Mus. Nat. Hist. Lond., 810. Elmhagen, B., Rushton, S.P., 2007. Trophic control of mesopredators in terrestrial ecosystems: top-down or bottom-up. Ecol. Lett. 10, 197–206. Feeney, S., 1999. Comparative Osteology, Myology, and Locomotor Specializations of the Fore and Hind Limbs of the North American Foxes Vulpes Vulpes and Urocyon Cinereoargenteus. Unpublished Ph.D. Dissertation, Univ. Massachusetts, Amherst. Fields, S.E., 2007. Mammals of the north-central piedmont of South Carolina. Southeastern Nat. 6, 577–596. Foster, D.R., Motzkin, G., Bernardos, D., Cardoza, J., 2002. Wildlife dynamics in the changing New England landscape. J. Biogeogr. 29, 1337–1357. Frey, J.K., 2004. Taxonomy and distribution of the mammals of New Mexico: an annotated checklist. Occas. Papers Mus. Texas Technol. Univ. 240, 1–32. Frey, J.K., 2006. Inferring species distributions in the absence of occurrence records: an example considering wolverine (Gulo gulo) and Canada lynx (Lynx canadensis) in New Mexico. Biol. Conserv. 130, 16–24. Frey, J.K., 2009. Distinguishing range expansions from previously undocumented populations using background data from museum records. Divers. Distrib. 15, 183–187. Genoways, H.H., Ratcliffe, B.C., 2008. Engineer cantonment, missouri territory, 1819–1820: America’s first biodiversity inventory. Great Plains Res. 18, 3–31. Gilmore, R.M., 1946. Mammals in archeological collections from southwestern Pennsylvania. J. Mammal. 27, 227–234.
Global Invasive Species Database [GISD]. 2012. GISD; Managed by the Invasive Species Specialist Group, IUCN Species Survival Commission. 19.07.12. Pennant, T., 1784. Arctic zoology, Vol. 1. Henry Hughs, London. Perrine, J.D., Pollinger, J.P., Sacks, B.N., Barrett, R.H., Wayne, R.K., 2007. Genetic evidence for the persistence of the critically endangered Sierra Nevada red fox in California. Conserv. Genet. 8, 1083–1095. Pike, Z.M., 1810. An account of expeditions to the sources of the Mississippi and through the western parts of Louisiana, to the sources of the Arkansaw, Kans, La Platte, and Pierre Jaun, Rivers performed by order of the government of the United States during the years 1805, 1806, and 1807, and a tour through the interior parts of New Spain, when conducted through these provinces, by order of the captain-general, in the year 1807. C. & A. Conrad, Philadelphia. Potts, A., 1912. Fox Hunting in America. Carnahan Press, Washington. Presnall, C.C., 1958. The present status of exotic mammals in the United States. J. Wildl. Manage. 22, 49–50. Rabushka, A., 2008. Taxation in Colonial America. Princeton Univ. Press, New Jersey. Rhoads, S.N., 1903. The Mammals of Pennsylvania and New Jersey. Privately Published, Philadelphia. Ritchie, E.G., Johnson, C.N., 2009. Predator interactions, mesopredator release and biodiversity conservation. Ecol. Lett. 12, 982–998. Sacks, B.N., Statham, M.J., Perrine, J.D., Wisely, S.M., Aubry, K.B., 2010. North American montane red foxes: expansion, fragmentation, and the origins of the Sacramento Valley red fox. Conserv. Genet. 11, 1523–1539.
J.K. Frey / Biological Conservation 158 (2013) 74–79 Sacks, B.N., Moore, M., Statham, M.J., Wittmer, H.U., 2011. A restricted hybrid zone between native and introduced red fox (Vulpes vulpes) populations suggests reproductive barriers and competitive exclusion. Mol. Ecol. 20, 326– 341. Schmidly, D.J., 2002. Texas Natural History: a Century of Change. Texas Tech Univ. Press, Lubbock. Skinner, J.S., 1829. American Turf Register and Sporting Magazine, vol. 1. J.S. Skinner, Baltimore. Statham, M.J., Sacks, B.N., Aubry, K.B., Perrine, J.D., Wisely, S.M., 2012. The origins of recently established red fox populations in the United States: translocations or natural range expansions? J. Mammal. 93, 52–65.
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Thompson, Z., 1853. Natural History of Vermont, with Numerous Engravings, and an Appendix. Published by the author, Burlington. Trapp, G.R., Hallberg, D.L., 1975. Ecology of the gray fox (Urocyon cinereoargenteus): a review. In: Fox, M.W. (Ed.), The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution. Van Nostrand Reinhold, New York, pp. 164–178. Voigt, D.R., 1999. Red fox. In: Novak, M., Baker, J.A., Obbard, M.E., Malloch, B. (Eds.), Wild Furbearer Management and Conservation in North America, Ontario Fur Managers Federation, Toronto, pp. 379–392. Warren, M.D., 1897. Enemies of Poultry. C.M. Busch, Harrisburg, Pennsylvania. Whitlow, W.B., Hall, E.R., 1933. Mammals of the Pocatello region of southeastern Idaho. Univ. Calif. Publ. Zool. 40, 235–275.
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Short communication
Re-evaluation of the evidence for the importation of red foxes from Europe to colonial America: Origins of the southeastern red fox (Vulpes vulpes fulva) Jennifer K. Frey ⇑ New Mexico State University, Department of Fish, Wildlife and Conservation Ecology, P.O. Box 30003, MSC 4901, Las Cruces, NM 88003-8003, USA
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Article history: Received 10 March 2012 Received in revised form 19 July 2012 Accepted 26 July 2012 Available online 28 November 2012 Keywords: Red fox Vulpes vulpes fulva Exotic species Geographic range shifts Scientific misconception United States
a b s t r a c t It is widely accepted that the red fox subspecies of southeastern North America (Vulpes vulpes fulva) is descended from red foxes imported from Europe to the American colonies for sport hunting. Thus, V. v. fulva is considered an exotic invasive organism that has apparently expanded its range to occupy much of the contiguous United States, with exception of the range of relict native populations in the western mountains and Sacramento Valley. I examined the evidence for importations and found that all claims of introduction stemmed to just two literature sources that were based on vague second-hand information. Together with results of morphological and genetic studies, this information indicates that V. v. fulva is native to North America. Thus, historical shifts in distribution and abundance of V. v. fulva and current management approaches should be re-evaluated in light of native status. Evaluating original sources of information can curtail the proliferation of inaccurate information and conclusions that influence conservation decisions. Ó 2012 Elsevier Ltd. All rights reserved.
1. Introduction The red fox (Vulpes vulpes) occurs throughout most of the Holarctic, except some desert and tundra areas, and it has been introduced to Australia and some other islands (Lariviere and Pasitschniak-Arts, 1996). The International Union for Conservation of Nature considers it one of the world’s worst 100 exotic invasive species due to negative impacts on native fauna where it has been introduced, including Australia and North America (GISD, 2012). It is thought that both native and exotic populations of red foxes occur in North America, with native foxes restricted to Alaska, Canada, and relict populations in the western United States (US; Churcher, 1959; GISD, 2012; Kamler and Ballard, 2002). Most treatments of the red fox in North America state that red foxes from Europe were introduced by the American colonists to southeastern North America for equestrian sport hunting with hounds – a claim that essentially has become ‘‘common knowledge.’’ Consequently, many authors have concluded that the southeastern subspecies in North America, Vulpes vulpes fulva, is directly descended from those introductions or a hybrid between European and native North American foxes (e.g. Ables, 1975; Leopold and Chamberlain, 2001; Kamler and Ballard, 2002; Rhoads, 1903; but see Aubry et al., 2009; Sacks et al., 2010; Statham et al., 2012). The distribution of V. v. fulva is thought to have expanded since colonial times (Kamler and Ballard, 2002). Its distribution currently ⇑ Tel.: +1 575 640 9782; fax: +1 575 646 1281 E-mail address: [email protected] 0006-3207/$ - see front matter Ó 2012 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.biocon.2012.07.027
extends from the Great Lakes south to the Gulf of Mexico and from the eastern seaboard of the US, westward across the southern Great Plains to Colorado and New Mexico (Frey, 2004; Hall, 1981), and Kamler and Ballard (2002) concluded that V. v. fulva replaced the native V. v. regalis in the northern Great Plains. It also is thought to have been introduced and established in low elevation regions within various areas of western North America (e.g. Buskirk and Zielinski, 2003; Kamler and Ballard, 2002; Lewis et al., 1999), although specific records detailing the source population and the date, location, and numbers of animals introduced are generally lacking (but see Aubry, 1983, 1984; Whitlow and Hall, 1933). If these scenarios are true, then as suggested by Kamler and Ballard (2002), all current populations of the red fox in the contiguous US, with exception of relict populations occurring in the western mountains (V. v. macroura, V. v. cascadensis, V. v. necator) and Sacramento Valley (V. v. patwin), are not native to North America and, hence, present a threat to native species. This interpretation has practical management implications as the attitudes and approaches towards native and exotic species are usually diametrically opposed (e.g. Clout and Williams, 2009). Despite the important ramifications, the exotic status of V. v. fulva usually has been taken as fact without any reasonable attempt to validate the claim of introduction based on original sources of information (e.g. Kamler and Ballard, 2002). Most modern treatments cite relatively recent works, especially Churcher (1959), that are themselves based on earlier works that cite even earlier works. Some treatments go so far as to provide no citations at all (e.g. Hamilon and Whitaker, 1979; Presnall, 1958), which are
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subsequently cited as evidence by other works (e.g. Fields, 2007; Long, 2003). A notable exception is the recent work by Statham et al. (2012). They built upon a prior phylogeographic analysis of native red foxes in North America (Aubry et al., 2009) to explicitly test the origins of red foxes throughout the eastern contiguous US, which they considered to have been established since the mid-1800s (Statham et al., 2012). They found no evidence of European haplotypes anywhere in North America and concluded that ‘‘recently established’’ populations in the eastern US were primarily the result of a southward range expansion from eastern Canada and northeastern US, and were not derived from introductions from Europe (Statham et al., 2012). However, their sampling included only 18 individuals from populations considered recently established in the eastern US, and did not include key areas where colonial introductions are frequently said to have occurred (e.g. eastern Virginia, Maryland). In addition, the methods they employed were based on mtDNA and, hence, were only able to track matrilineal ancestry. To better understand the origins of the southeastern red fox, I compiled original information concerning the importation of red foxes to the American colonies, such as dates, source locations, release locations, and numbers of animals. In large part, this was made possible by the availability of electronic copies of rare old texts, such as through the Biodiversity Heritage Library, American Journeys, Internet Archive, Google Books, Proquest American Periodicals Services Historical database, and other library resources.
2. Evidence for importation The chain of evidence for all references to the importation of red foxes from Europe to colonial America appears to extend back to only two references. The first was a travel journal written by the Swedish naturalist Pehr Kalm describing his experiences in eastern North America from 1748 to 1751 (Egerton, 2006). In Philadelphia on 7 November 1748, Kalm (1772: pp. 221–222) wrote: ‘‘Mr. Bartram [i.e. American botanist John Bartram who lived in New Jersey near Philadelphia] and several others assured me, that, according to the unanimous testimony of the Indians, this kind of foxes never was in the country, before the Europeans settled in it. But of the manner of their coming over I have two different accounts: Mr. Bartram and several other people were told by the Indians, that these foxes came into America soon after the arrival of the Europeans, after an extraordinary cold winter, when all the sea to the northward was frozen: from hence they would infer, that they could perhaps get over to America upon the ice, from Greenland or the northern parts of Europe and Asia. But Mr. Evans [i.e. Lewis Evans, an engineer from vicinity of Philadelphia], and some some [sic] others, assured me, that the following account was still known by the people. A gentleman of fortune in New England, who had a great inclination for hunting, brought over a great number of foxes from Europe, and let them loose in his territories, that he might be able to indulge his passion for hunting.⁄This is said to have happened almost at the very beginning of New England’s being peopled with European inhabitants. These foxes were believed to have so multiplied, that all the red foxes in the country were their offspring.’’ [The asterisk is to a footnote added by John Reinhold Forster who translated Kalm’s work into English. Forster did not consider either of Kalm’s accounts satisfactory; he believed that the red fox immigrated to North America ‘‘in remote times’’ from eastern Asia and then spread across the continent.] The second source of information is from the first volume (1829) of the American Turf Register and Sporting Magazine, which primarily served as a venue to register the performance and pedi-
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grees of purebred horses. However, it also contained information on rural sports including articles on the natural history of American game species. The editor, John Stuart Skinner, solicited readers to submit letters describing anecdotes about wildlife. The most widely cited ‘‘records’’ of importations of the red fox are based on an editorial note and two letters to the editor in this volume. Skinner’s editorial note (Skinner, 1829: pp. 9) follows an article on the manner in which George Washington hunted foxes: ‘‘The red fox is supposed to have been imported from England, to the Eastern Shore of Maryland, by a Mr. Smith, and to have emigrated across the ice to Virginia, in the hard winter of 1779–80, when the Chesapeake was frozen over.’’ In response, one letter to the editor from Fredericksburg, Virginia (Skinner, 1829: pp. 74) stated: ‘‘This I do not doubt; but I think it probable they were brought over and turned out at other places, and at very early periods. In 1789, when quite a boy, I was at the death of the first red fox killed in Perry county, Pa. . . . Not a person present, or any one who saw it for some days had ever seen or heard of an animal of the kind. At last it was shown to a Mr. Lenarton, an old Jersey-man, who pronounced it an English fox. He said the red fox was imported into New York from England, by one of the first English governors, who was said to be a great sportsman, and turned out on Long island, where they remained for many years, but at last made their way, on the ice, to the main land, and spread over the country.’’ In another letter to the editor written from Richmond, Virginia (Skinner, 1829: pp. 197): ‘‘The tradition of this part of the country. . .differs from the conjecture which seems to prevail farther north, in this; that . . . he is supposed to have been brought from the continent—Germany, I think—and not from the island of Great Britain.’’ In an Outing magazine article on the origins of fox hunting in America, Hiss (1897: pp. 14–15) provided a greatly embellished version of the story told in Skinner’s (1829) editorial comment, which he described as a ‘‘tradition’’ and ‘‘old tale.’’ Cogent details of Hiss’ story are that in 1730 eight wealthy farmers from Talbot County, Maryland, instructed the captain of the tobacco schooner ‘‘Monocacy,’’ which was owned by one of them, to bring back eight pairs of red foxes on the next trip to Liverpool, England. The animals were released in Maryland on the eastern shore of the Chesapeake Bay and a festival celebrated the event. The foxes multiplied and expanded into Virginia across the ice during the hard winter of 1779–80. In his book on fox hunting in America Potts (1912: pp. 5–7), dismissed the two stories written as letters to the editor of American Turf Register and Sporting Magazine as interesting legends and he considered Hiss (1897) story as true. Potts (1912: pp. 7) stated that the story had been verified by Skinner. Thus, stories told by Skinner (1829) and Hiss (1897) are the same. Potts (1912:7) changed some of the details in Hiss (1897), calling the name of the schooner ‘‘Monaccasy,’’ and stating that the foxes were consigned to a Mr. Smith. These stories were frequently recounted in periodicals and natural history literature, sometimes word-for-word (although sometimes lacking citations) and sometimes with misconstrued information (e.g. Pennant, 1784; Doughty and Doughty, 1830; Hunter, 1876; Rhoads, 1903). Reports of introductions of red foxes in 1650 (e.g. Gilmore, 1946; Long, 2003) are apparently misconstrued based on reports of the first importation of fox hounds to North America by Robert Brooke, an event that has been interpreted as the beginnings of fox hunting in America (Hiss, 1897; Brooke, 1910). 3. Supporting evidence for native status I found no direct evidence or firsthand accounts of any importation of red foxes to colonial America. The total ‘‘evidence’’ consists
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of a vague second-hand report given to Kalm by Evans, the vague story told by Skinner, which might be the same as the one told by Kalm, a vague second-hand story by Lenarton given by another person in a letter to the editor, and a rumor of a different source population given in a letter to the editor. Hiss’ more detailed version of the one given by Skinner, was written ca. 250 years after the supposed event and likely represents literary license. Even the earliest story occurred a century or so after the alleged importation. A recurring theme in these stories is that the red fox expanded its range over ice during a hard winter, which might ultimately derive from the supposed comments by the Native Americans that were relayed to Kalm by Bartram. By applying evidentiary standards used for interpreting species occurrence records, these scant, second-hand reports are deemed highly questionable (Frey, 2006; McKelvey et al., 2008). In contrast, introductions of foxes to Alaskan islands and Australia are relatively well documented (Bailey, 1993; Long, 2003). Given the popularity of fox hunting in colonial America, importations might have been reported in local newspapers, at least if introduction occurred during the last few decades of the colonial period when there were more media outlets. I found no such articles in my searches of the Proquest American Periodicals Series database, which includes >1000 sources dating back to 1741. If such an importation occurred, the paucity of records suggests it would have been an exceedingly rare event, probably because foxes (including the gray fox [Urocyon cinereoargenteus]) that could be used for sport were already present in the eastern US. There is a common modern misperception that gray foxes were not considered suitable targets for equestrian fox hunts with hounds because they climb trees (e.g. Voigt, 1999; Peddicord, 2010). However, the early literature is replete with accounts extolling the virtues of gray foxes as a quarry for such hunts (e.g. Audubon and Bachman, 1846; Godman, 1831; Page, 1892). In Australia where foxes were entirely absent, the desire to establish a population for hunting would have been more keen. However, even in Australia most of the early attempts to introduce the red fox failed (Long, 2003). It seems unlikely that any rare release of imported red foxes into North America would have established a population, especially given that any such importation would have been for hunting purposes. Thus, I found no compelling evidence that red foxes from Europe were successfully introduced in colonial America. Morphological and genetic data support the interpretation that V. v. fulva is native to North America. If the red fox of the eastern US is the result of importations, then it would be referable to the subspecies native to Great Britain and other areas of Europe, V. v. crucigera, as it is in Australia (Ellerman and Morrison-Scott, 1951). However, Doughty and Doughty (1830) noted that none of the cabinets (i.e. Victorian natural history collections) in Philadelphia contained specimens of the European red fox, and all known specimens from the US at that time were referable to V. v. fulva (also recorded in the literature as V. fulvous, V. fulvus, V. fulva, and V. pennsylvanica). I used the Mammal Networked Information System (http://manisnet.org/) to search for specimens of red foxes in the eastern US; none were referred to V. v. crucigera (all were referred to V. v. fulva). All studies that have compared morphology of red foxes from eastern North America and Eurasia have found consistent differences, which have been recognized at either the species or subspecies level (Baird, 1857; Bangs, 1897; Churcher, 1959; Merriam, 1900; Paradiso, 1969). Molecular genetic data based on mtDNA have shown that red foxes from Eurasia are in a subclade different from populations in North America (Aubry et al., 2009). With that phylogeography as a framework, Statham et al. (2012) examined genetic signals of red foxes from throughout the eastern US and found no Eurasian haplotypes. Similarly, specimens of red foxes taken at low elevations in California that were hypothesized by Kamler and Ballard (2002) to represent
introduced V. v. fulva, had haplotypes within North American clades rather than in the Eurasian subclade (Sacks et al., 2010, 2011; Statham et al., 2012). Further, molecular data (Aubry et al., 2009; Statham et al., 2012) have not supported the views of Kamler and Ballard (2002) that ‘‘exotic’’ V. v. fulva spread like a wave across the US replacing native red foxes. Thus, Statham et al. (2012) concluded that ‘‘recently established’’ populations in the eastern US were the result of a natural range expansion from southern Canada.
4. Distribution of the red fox in the eastern US Today the red fox is thought to occur throughout the US east of the Rocky Mountains, with exception of coastal Florida, the southern Great Plains, and southern Texas (Cypher, 2003). Aubry et al. (2009) and Statham et al. (2012) each provided a map of the geographic range of red foxes that were considered to be unequivocally indigenous (i.e. no introductions have been proposed as influencing distribution) in North America prior to European settlement. Areas south of that range where red foxes are currently known to occur, were denoted ‘‘recently established populations’’ and an explicit goal of Statham et al. (2012) was to evaluate ancestry of those populations. In the US east of the Rocky Mountains, the only region considered within the unequivocal range of indigenous red foxes was northern New England (i.e. Maine and the northern portions of New York, Vermont, and New Hampshire); rationale for this interpretation was detailed in Statham et al. (2012), and was based primarily on information contained in Audubon and Bachman (1851); Churcher (1959); Newberry (1857), and Rhoads (1903). However, the pre-settlement geographic range of the red fox in the eastern US requires further evaluation. There is very little reliable information about wildlife in America prior to 1800. The early 1800s was the period during which most of the initial scientific exploration of the country occurred, and also was a period of proliferation of news periodicals and natural history publications. It is important to recognize that lack of a record is not evidence that a species was absent (Keay, 1901) or that the first documentation of a species in a region is not necessarily the first occurrence of that species in the area (Frey, 2009). For instance, the red fox was reported in the Great Plains during several of the first scientific expeditions to the region, including what is now North Dakota in 1805 by Meriwether Lewis and William Clark (http://lewisandclarkjournals.unl.edu/index.html), what is now Minnesota in 1805 by Zebulon Pike (Pike, 1810), and what is now Nebraska in 1819 by Edwin James (Genoways and Ratcliffe, 2008; James, 1823). Given that these observations preceded European settlement of those regions, there is no reason to presume that they represent anything other than native occurrence of the red fox on the Great Plains. There also are credible reports of red foxes in the American colonies. For instance, Thomas Morton and John Josselyn reported both red foxes and gray foxes in Massachusetts in the early 1600s at the beginning of European settlement (Josselyn, 1865a,b; Morton, 1883). Kalm (1772), who primarily visited Pennsylvania, stated that both the red fox and gray fox occurred in the colonies, but that the red fox was scarce. On basis of Kalm’s report, Pennant (1784) stated that red foxes decreased in abundance to the south and that none were found south of Pennsylvania. Further, red foxes, but rarely gray foxes, were viewed as an important depredator of livestock and poultry (e.g. Thompson, 1853; Warren, 1897), and hence bounties were instituted for their control. Bounties on red foxes were some of the earliest wildlife laws enacted, including 1635 in Massachusetts and 1641 in Rhode Island (Rabushka, 2008). In 1714 a bounty on wolves in New Jersey was expanded to include red foxes (Rhoads,
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1903). In Pennsylvania, a bounty on foxes was established in 1718, and records from Chester County, show bounties paid on more than 453 foxes during 11 years prior to 1747 (Barclay, 2002). Kalm (1772) stated that there were bounties for red fox in Pennsylvania and all other provinces. The idea that red foxes were introduced to the eastern US was fueled, in part, by studies of vertebrate remains at archeological sites in southern Pennsylvania that lacked evidence of red foxes (Gilmore, 1946; Miller, 1940). While the red fox may have been absent from some local faunas, it is now possible to evaluate late Quaternary mammal distributions across thousands of sites in the US using the Faunmap electronic database (http://www.ucmp. berkeley.edu/faunmap/about/index.html). Although Faunmap is a powerful tool for estimating historical mammal distributions, some records may not be verified. However, errors in this case are likely to be minimized because there are many skeletal characteristics that distinguish between the red fox and gray fox (Gilmore, 1946), and other North American members of the genus Vulpes (i.e. the arid-land foxes V. velox and V. macrotis) and Urocyon (i.e. U. littoralis) are unlikely to have been present east of the Great Plains during this epoch. Using Faunmap, I examined 2,861 sites from the late Holocene (450–4500 B.P.) for records of red foxes and gray foxes in order to reconstruct their distribution east of the Rocky Mountains in the US. Sites of this age appeared best represented in the Ozark-Quachita Interior Highlands and the Interior Low Plateaus and Central Lowlands; sites were sparse in the Appalachian Highlands and the Gulf and Atlantic Coastal Plain (except Florida; physiographic regions follow http://tapestry. usgs.gov/physiogr/physio.html). I found 112 records of red foxes and 178 records of gray foxes. Records of both species were dispersed across the eastern US and exhibited similar distribution patterns. Records of red foxes occurred from Maine to the Missouri Plateau (e.g. North Dakota and South Dakota) and as far south as southern Georgia, Arkansas, and Oklahoma. Records of red foxes were sparse in the Appalachian Plateau and Interior Low Plateaus regions; there were no records from Florida and the High Plains. Records of gray foxes were comparatively more dense throughout the eastern US, but were absent from New England (i.e. northeast of New York). The northernmost records of gray foxes were from southern Ontario, central Wisconsin, and southern Minnesota; to the west records were sparse or absent in most of the Great Plains and Osage Plains. Thus, historical records and other evidence support a preEuropean native occurrence of red foxes in the US east of the Rocky Mountains. Consistent with its boreal affinities, the evidence presented herein (see also Allen, 1876; Rhoads, 1903) suggests that the red fox was primarily associated with the Canadian and Transition life zones (sensu Merriam, 1898), which in the eastern US extends from Maine south along the Appalachian Mountains and westward from New York through the Great Lakes region and the Missouri Plateau of the Great Plains. The red fox was likely rare or absent in much of the Austral zone, especially in southern deciduous forests. However, given the species’ ecological plasticity and capacity for long distance movements (Lariviere and Pasitschniak-Arts, 1996), it also might have been sparsely distributed in some regions of the Austral zone where an open habitat structure was present, such as prairies, beaches, and perhaps some pine forests. However, based on repeated testimony of the appearance of red foxes in places where it had not previously been noticed, it appears that the red fox historically expanded its distribution and abundance within the Austral zone (e.g. Audubon and Bachman, 1851; Hahn, 1909; Rhoads, 1903; Schmidly, 2002). Several interrelated factors may account for the apparent increase in distribution and abundance of red foxes within the Austral zone of the eastern US. Prior to European settlement, various types of forests dominated most of the eastern US. Foster et al.
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(2002) demonstrated widespread changes to wildlife assemblages in response to changes in abundance of forests as a result of postsettlement land-use. For instance, red foxes were included in a group of open land species that experienced historical increases in abundance that peaked in the mid-1800s as land was cleared for settlements, timber, and farms, but have recently declined as forests have regenerated. In contrast, gray foxes were in a group of woodland species that experienced concomitant historical declines in abundance, but recent increases (Foster et al., 2002). Indeed, many early historical accounts of foxes noted apparent increases in abundance and distribution of red foxes, and concomitant decreases in abundance of gray fox, as areas were settled and cleared of forests in the Austral zone (e.g. Allen, 1876; Hahn, 1909; Newberry, 1857). These changes in abundance are consistent with the habitat selection, predator escape strategies, locomotion, and other adaptations of each species (Cypher, 2003; Feeney, 1999). The gray fox is a true omnivore that prefers plants and eats a high proportion of insects, in addition to small mammals and birds; in contrast, the red fox is primarily carnivorous with a diet dominated by mammals (Cypher, 2003; Hockman and Chapman, 1983; Trapp and Hallberg, 1975). The most important prey for red foxes in the eastern US are cottontails (Sylvilagus spp.), although diets vary by region, and may also contain large proportions of other mammals such as voles (Microtus spp.) and woodchucks (Marmota monax). Most of the important prey species for red foxes are associated with early successional habitats and their populations are thought to have increased with deforestation and settlement, peaking in the late 1800s and early 1900s (Litvaitis, 2001). This increase in abundance and distribution of favored prey would have preferentially benefited the red fox. Lastly, intraspecific competition is thought to have an important influence on distribution and abundance of canids. Historically, wolves (Canis spp.) occurred throughout all of the eastern US, but were extirpated from most of the region in the wake of settlement due to control efforts. Mesopredator release due to extirpation of wolves could have benefited red foxes (Elmhagen and Rushton, 2007; Ritchie and Johnson, 2009). However, following the loss of wolves, coyotes (C. latrans) subsequently expanded their range from the plains region east to the Atlantic seaboard, which may result in renewed suppression of red fox populations by coyotes (Cypher, 2003).
5. Conclusions The belief by some early naturalists that V. v. fulva was an exotic was influenced by observations of apparent increases in abundance and range expansions of the red fox into areas where the species originally had been scarce or absent in southeastern North America (e.g. Allen, 1876; Audubon and Bachman, 1851; Bailey, 1896; Baird, 1857; Gilmore, 1946; Hahn, 1909). More recently, apparent expansion of V. v. fulva into western North America likewise has been attributed to these foxes being ‘‘descendants from European stock and therefore . . . better adapted to human activities, urban areas, and human-induced changes to the environment’’ (Kamler and Ballard, 2002: pp. 375). My study, which is based on independent lines of evidence, provides broad support for the central conclusions of Statham et al. (2012) – that red foxes in the eastern US are not the descendants of importations from Europe. However, my results also suggest a need to reevaluate origins of populations of red foxes, particularly in the central US, given evidence that the species was historically present in that region. Further, native status of V. v. fulva in North America calls for a reinterpretation of mechanisms responsible for changes in its distribution and abundance. As with most game species, there is evidence of isolated translocations of red foxes within the US and there are reported instances of animals escaping from fur farms (e.g. Aubry, 1983, 1984;
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Kellogg, 1939). Thus, conservation issues related to introduced populations of V. v. fulva in some areas of the western US where red foxes did not occur historically will remain (e.g. Perrine et al., 2007; Sacks et al., 2010). In these regions, conservation and management decisions should be predicated on native status of the red fox in North America. Finally, as with the origins of V. v. fulva, the proliferation of inaccurate information can be averted by verifying the original sources of cited ‘‘facts.’’ The ready access to electronic copies of rare old text can provide new perspectives on old problems. References Ables, E.D., 1975. Ecology of the red fox in North America. In: Fox, M.W. (Ed.), The Wild Canids: their Systematics Behavioral Ecology and Evolution. Van Nostrand Reinhold, New York, 508p (pp. 216–236). Allen, J.A., 1876. The former range of some New England carnivore mammals. Am. Nat. 10, 708–715. Aubry, K.B., 1983. The Cascade Red Fox: Distribution, Morphology, Zoogeography and Ecology. Unpublished Ph.D. thesis, Univ. Washington. Aubry, K.B., 1984. The recent history and present distribution of the red fox in Washington. Northwest Sci. 58, 69–79. Aubry, K.B., Statham, M.J., Sacks, B.N., Perrine, J.D., Wisely, S.M., 2009. Phylogeography of the North American red fox: vicariance in pleistocene forest refugia. Mol. Ecol. 18, 2668–2686. Audubon, J.J., Bachman, J., 1846. The Viviparous Quadrupeds of North America, vol. I. J.J. Audubon, New York. Audubon, J.J., Bachman, J., 1851. The Viviparous Quadrupeds of North America, vol. II. V.G. Audubon, New York. Bailey, V., 1896. List of mammals of the District of Columbia. Proc. Biol. Soc. Washington 10, 93–101. Bailey, E.P., 1993. Introduction of foxes to Alaskan islands – history, effects on avifauna, and eradication. U.S. Fish Wildl. Serv. Res. Publ. 193, 1–54. Baird, S.F., 1857. Mammals, part 1. 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Comparative Osteology, Myology, and Locomotor Specializations of the Fore and Hind Limbs of the North American Foxes Vulpes Vulpes and Urocyon Cinereoargenteus. Unpublished Ph.D. Dissertation, Univ. Massachusetts, Amherst. Fields, S.E., 2007. Mammals of the north-central piedmont of South Carolina. Southeastern Nat. 6, 577–596. Foster, D.R., Motzkin, G., Bernardos, D., Cardoza, J., 2002. Wildlife dynamics in the changing New England landscape. J. Biogeogr. 29, 1337–1357. Frey, J.K., 2004. Taxonomy and distribution of the mammals of New Mexico: an annotated checklist. Occas. Papers Mus. Texas Technol. Univ. 240, 1–32. Frey, J.K., 2006. Inferring species distributions in the absence of occurrence records: an example considering wolverine (Gulo gulo) and Canada lynx (Lynx canadensis) in New Mexico. Biol. Conserv. 130, 16–24. Frey, J.K., 2009. Distinguishing range expansions from previously undocumented populations using background data from museum records. Divers. Distrib. 15, 183–187. 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