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Audubon's red wolf. Audubon described the red-colored wolves of Texas as nothing more than a color phase.

For us to fully understand the problems inherent in deeming the red wolf a unique species, we need to look at this animal from an historical perspective.

This gets complicated very quickly, for the records we do have of wolves in the American South are not as good as one might think.

European man killed off the wolves– and anything that looked like a wolf– as soon as he arrived.

Most of the historical records make mention that the wolf of what became the United States came in several color.  Yellowish, reddish, black, gray, and white individuals were all recorded, but the black individuals were always mentioned. In some areas, like Florida, it appears that black individuals comprised nearly the entire population.

William Bartram, a famed botanist from the Philadelphia area, was the first person to describe these black wolves in Florida. He would give them the name Lupus niger, which later authorities would move into Canis niger, which is an archaic name given to the supposed red wolf species.

Bartram traveled into Florida during the 1770’s, and he later compiled his notes and stories into a book called Travels through North and South Carolina, Georgia, East and West Florida, the Cherokee Country, the Extensive Territories of the Muscogulges or Creek Confederacy, and the Country of the Chactaws. Containing an Account of the Soil and Natural Productions of Those Regions; Together with Observations on the Manners of the Indians. Usually referred to as “Travels” or “Batram’s Travels,” the book includes detailed historical accounts of fauna and flora of the a huge swathe of the Southeast. It was published in 1791,  and his descriptions of the land of Florida made his American readers quite covetous of this territory, which was Spanish territory at the time.

Bartram first describes the wolves feeding on the carcass of a horse:

Observing a company of wolves (lupus niger) under a few trees, about a quarter of a mile from shore, we rode up towards them; they observing our approach, sat on their hinder parts until we came nearly within shot of them, when they trotted off towards the forests, but stopped again and looked at us, at about two hundred yards distance: we then whooped, and made a feint to pursue them; when they separated from each other, some stretching off into the plains, and others seeking covert in the groves on shore. When we got to the trees, we observed they had been feeding on, the carcase of a horse. The wolves of Florida are larger than a dog, and are perfectly black, except the females, which have a white spot on the breast; but they are not so large as the wolves of Canada and Pennsylvania, which are of a yellowish brown colour (pg. 197).

Bartram would later go on to describe the black wolves as coming in other besides black:

I have been credibly informed that the wolves here are frequently seen pied, black and white, and of other mixed colours. They assemble in companies in the night time, howl and bark altogether, especially in cold winter nights, which is terrifying to the wandering bewildered traveller.

The idea of there ever being black and white wolves may sound a bit dubious, but keep in mind that there are coyotes with Irish spotting. These spots likely came from crossing with dogs, and this description of black and white wolves further strengthens the case that black wolves were developed through crossing with domestic dogs.

It also argues against these animals being red wolves as we know them now. The red wolves that we know them today. Wolves and dogs are much more closely related than dogs are to coyotes, and the black coloration that originated in dogs is far more common in wolves than it is in coyotes. And although we do see coyotes with dog coloration, these animals are not that common. However, black wolves and wolves with dewclaws on the hind legs have been reported– which are definitely derived from wolves mating with dogs.

Coyotes have much stronger reproductive barriers against breeding with dogs than wolves do.  It’s easy to find stories of male wolves coming in on bitches in heat. In the old days, a bitch in heat was great bait to bring in the male wolves to the gun. Sometimes the hunters would wait until the two had mated, then they would come in and either club the wolf to death or chop it up with a hatchet while the two were tied.

Coyotes and dogs don’t readily breed with each other. If they did, the coyote population would have absorbed far more dog genes than it currently has.

However, Native American dogs and wolves were likely exchanging genes all the time, and it more strongly suggests that Bartram’s black wolf was part of Canis lupus, rather than a coyote or close relative of the coyote.

Further evidence for this close relationship between Florida wolves and native dogs comes from a dog that Bartram observed guarding a band of Seminole ponies:

One occurrence remarkable here, was a troop of horse under the controul and care of a single black dog, which seemed to differ in no respect from the wolf of Florida, except his being able to bark as the common dog. He was very careful and industrious in keeping them together; and if any one strolled from the rest at too great a distance, the dog would spring up, head the horse, and bring him back to the company. The proprietor of these horses is an Indian in Talahasochte, about ten miles distance from this place, who, out of humour and experiment, trained his dog up from a puppy to this business: he follows his master’s horses only, keeping them in a separate company where they range; and when he is or wants to see his master, in the evening he returns to town, but never stays at home a night (pg. 220-221).

The Seminoles had learned from the Spanish how to use dogs as livestock guardians. Raising the dogs with the stock, they ensured that the dog bonded with the animals. The Spanish used large mountain dogs and mastiffs, but the Seminoles used their native dogs, which may or may not have had some wolf ancestry. This black, wolf-like dog suggests very strongly for the hybrid origin of melanism in wolves, and that the Florida black wolf was actually part of Canis lupus, along with the dog.

The Pennsylvania physician and naturalist Benjamin Smith Barton would also describe similar dogs. In an article entiled “Some Account of Native American or Indian Dogs” that appeared in the The Philadelphia Medical and Physical Journal in 1804. He aligns this particular dog with the black wolf, as did Bartram, but he points out some differences:

The dogs which are now in use among the Creeks, Chikkasah, and other southern tribes, are of different kinds. As far as I have been able to collect information concerning them, they, in general, bear a very strong family resemblance to the wolf. One kind is very similar to the Canis Lycaon, or black wolf. It is not, however, always black, but of different colours, commonly of a bay colour, and about one third less [in size] than the wild black wolf. It carries its ears almost erect, and has the same wild and sly look that the wolf has (pg. 12).

Barton never saw these dogs, but he got his information from travelers in the region. His source for this information was Bartram, whose previous description of the Florida black dog  matches this description of the Creek dogs in every way but size. Thus, both black color and spotting could have worked its way into the Southern wolf population without dramatically affecting phenotype.

Canis lycaon became a very common name to describe this black wolf, which Barton described as being quite large and common in the region where the Creeks lived in his Notes on the Animals of North America

Canis lycaon is a name sometimes used to refer to the supposed “Eastern wolf” today, but for much of its use, it referred to black North American wolves. Black European wolves were called Canis lupus niger,  but later texts, in supposed deference to Bartram, red wolves would be called Canis niger.

Nowhere do we find mention of these wolves being particularly different from other wolves. The only thing they have that is unusual is that they are very often black in color.

As I noted earlier, this black color likely originated in dogs. Black wolves living in North America today derive their black coloration from crossbreeding with domestic dogs. In Western North American wolves, this black coloration was introduced about 12,000 years ago. However, in these Eastern wolf populations there would have likely been a large number of Native American dogs running about, exchanging genes with wolves all the time.

It is theorized that melanism is associated with greater immune function, so there would have been a selection pressure for the black color in warm, humid places like Florida and much of the Eastern United States. This could explain why so many wolves in the Eastern US and the South were black. Warm and humid places are great environments for various pathogens to live, so a stronger a immune system that is associated with the same gene that gives the black coloration could be a major asset.

Okay, now that we understand that the Southeastern US was home to large black wolves that somehow got connected to so-called red wolves of East Texas, when does someone first mention the red wolf?

Well, the first text to include a mention of a red wolf in North America is John James Audubon and John Bachman’s The Quadrupeds of North America, Volume 2 (1851). This book was published after Audubon’s death, but it is from this description that we get the idea that the coyote-like animals that once roamed East Texas and Louisiana were “red wolves.” The text describes the red wolf as a color phase of wolf , not a unique species. The section is called the “Red Texan Wolf”:

This variety is by no means the only one found in Texas, where Wolves, black, white and gray, are to be met with from time to time. We do not think, however, that this Red Wolf is an inhabitant of the more northerly prairies, or even of the lower Mississippi bottoms, and have, therefore, called him the Red Texan Wolf.

The habits of this variety are nearly similar to those of the black and the white Wolf, which we have already described, differing somewhat, owing to local causes, but showing the same sneaking, cowardly, yet ferocious disposition.

***

In all species of quadrupeds that are widely diffused over our continent, it has often appeared to us that toward the north they are more subject to become white—toward the east or Atlantic side gray—to the south black—and toward the west red. The gray squirrel, (S. migrulorius), of the Northern and Eastern States presents many varieties of red as we proceed westwardly towards Ohio. In the south, the fox squirrel in the maritime districts is black as well as gray, but not red. On proceeding westwardly, however, through Georgia and Alabama,a great many are found of a rufous colour. In Louisiana, there are in the southern parts two species permanently black as well as the foxsquirrel, which in about half the specimens are found black, and the remainder reddish. The same may be said in regard to the Wolves. In the north there is a tendency towards white—hence great numbers are of that colour. Along the Atlantic coast, in the Middle and Northern States, the majority are gray. To the south, in Florida, the prevailing colour is black, and in Texas and the southwest the colour is generally reddish. It is difficult to account, on any principles of science, for this remarkable peculiarity, which forms a subject of curious speculation.

This variety of Wolf is traced from the northern parts of the State of Arkansas, southerly through Texas into Mexico; we are not informed of its southern limits….

The Wolves present so many shades of colour that we have not ventured to regard this as a distinct species; more especially as it breeds with those of other colours, gangs of Wolves being seen, in which this variety is mixed up with both the gray and black (pg 241-243.).

Audubon and Bachman didn’t think that the red wolf was a unique species. It is clearly described as a color variant that popped up in wolves living in East Texas. There is no description of this animal having many unique features or even a remote suggestion of it being a separate species. That notion is entirely rejected in the text– which is probably why they call it the “Red Texan Wolf” instead of the “Texas red wolf.”

Audubon and Bachman would have also been aware that the term “red wolf” referred to another species.

It is first mentioned in Georges Cuvier Le règne animal distribué d’après son organisation [The Animal Kingdom] (1817). In the text Cuvier refers to a very different animal than the one that Audubon and Bachman would have met in Texas:

Le Loup Rouge D’amerique

D’un beau roux cannelle, une courte crinière noire tout le long de l’épine. Des marais de l’Amérique méridionale (pg. 180).

Translation:

“A beautiful brown cinnamon, a short black mane along the spine. Marshes of South America.”

Cuvier gave this animal the scientific name of Canis jubatus, and his description points to the maned wolf of South America (Chrysocyon brachyurus). It actually is red in color, but it does have black legs and a black mane. It’s not found in the marshes of South America. Instead, it is a creature of the grasslands and scrub forest.

Both Audubon and Bachman would have been familiar with Cuvier’s work. Audubon was a native French-speaker from Haiti, and Bachman was a well-read naturalist of his day.

Further, we know Audubon knew of Cuvier’s work from an unusual story.  Audubon shot an unusual red-crowned kinglet at his father-in-law’s farm on the Schuylkill River in Pennsylvania in 1812. He thought was unique enough to be declared a new species, which he called Cuvier’s kinglet (Regulus cuvieri).

I have named this pretty and rare species after Baron Cuvier, not merely by way of acknowledgment for the kind attentions which I have received at the hands of that deservedly celebrated naturalist, but more as a homage due by every student of nature to one at present unrivalled in the knowledge of General Zoology.

–John James Audubon, Ornithological Biography (1831) (pg. 288).

So it’s very likely that Audubon, who was an admirer of Cuvier, wanted to make sure that the red wolves of Texas were clearly delineated from Cuvier’s Le Loup Rouge D’amerique.

So it’s very likely that the red wolf was actually just a color phase of wolf that was common in Texas, Louisiana, and Arkansas.

And nothing more.

Wolves of all colors were heavily persecuted in the region.

In Bruce Hampton’s The Great American Wolf there is a photo of a red wolf that was captured in the Arkansas in the early 1900’s. Its jaws are bound shut with wire, and it is very likely that the dogs are about to be set on it. Wolves were so hated  that they weren’t even accorded human dispatch. Tortuous methods were very commonly used, including this macabre spectacle in which dogs would tear a wolf to pieces. With its jaw bound, it would be unable to defend itself.

However, the photo of this wolf reveals that it looks nothing like the animals currently called red wolves. The photo shows an animal with broad jaws and relatively small ears, which are being slightly pulled back against the skull. The look on the animal’s eyes is nothing but pure terror.

This photo seems to agree with Audubon and Bachman. The red wolf was nothing more than a color phase of the wolf subspecies native to the region.

And that wolf subspecies was eventually killed off until just a few remained.

But they had already started mating with coyotes as soon as the widespread persecution of their species began, and those that remained soon got absorbed into the coyote population.

In the 1970’s, animals that were deemed red wolves based upon morphology were trapped in the region, and they were bred in captivity. The Endangered Species Act was passed in 1973, and this was to be one of the first major attempts at saving a species.

The wolves were bred in captivity for several decades.

And they were “reintroduced” to Eastern North Carolina in 1987 at the Alligator River National Wildlife Refuge.

The  animals did fine, but then coyotes started moving in. And the red wolves started mating with coyotes.

To keep the animals “pure,” the US Fish and Wildlife Service started trapping coyotes, and that’s what they’ve been doing ever since.

This project is deemed a major success of the ESA. So-called red wolves are roaming in several reserves in the Southeast now, but the main population is in Eastern North Carolina.

In 199w, Robert Wayne released the first extensive mitochondrial DNA studies on wolves in North America. These analyses revealed that the most red wolves, both historical and living, had coyote mtDNA. This suggested that the red wolf was a hybrid species.

This finding was not well-received. The ESA is the “Endangered Species Act, ” not the “Endangered Hybrids Act.”

Ron Nowak threw a fit. He contended that red wolves were actually primitive wolf-like canids that were ancestral to both wolves and coyotes.  He proposed several ancient species for the potential ancestors of the red wolf, ending with Canis mosbachensis. Canis mosbachensis might be the ancestor of all wolves in both the Old World and the New, so the argument might be a bit moot. Other proposed ancestors of the red wolf include Canis edwardii, which was red wolf-sized that lived from the Pliocene to the Irvingtonian stage of the Pleistocene. It may be the ancestor of all wolves and coyotes.

But it’s not the red wolf that we know today.

After Wayne’s study came out, there were some other genetic studies that looked a limited parts of the y-chromosome or limited microsatellite analyses.

Wayne stated that the red wolf should be conserved. Lots of wolves have coyote mtDNA, but these animals have all been listed as subspecies of Canis lupus, not a unique species. So he suggested that the red wolf be considered an endangered subspecies. (The Eastern wolf as Canis lycaon canard hadn’t really developed yet.)

It was only last year, when the genome-wide assessment was released that it became obvious that the red wolves that live today aren’t a unique species after all. They really can’t be called a hybrid species. Instead, it might be more accurate to call them coyotes with wolf ancestry– which is not a rarity in North America.

So the historical record shows that the red wolf as we know it now never existed in the wild until after wolves had been persecuted in East Texas and Louisiana.

The wolves of the South were very likely Canis lupus subspecies, judging from their description from contemporaries. These southern wolves, which were very often black– not red–are now extinct.

Some might say that the coyote-derived impostors that have been introduced into the former range of the Southeastern wolf are merely fulfilling the same niche as the large black wolves that once roamed there.

They might be.

But they are still impostors.

The Southeast of the late eighteenth and early nineteenth century no longer exists.

We can only revisit it in the words of Audubon and Bartram.

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This depiction of a cross fox comes from The Viviparous Quadrupeds of North America by John Bachman and John James Audubon.

The cross fox is a color phase of the red fox, which is found almost exclusively in the North American populations. In fact, I’ve never heard of a European cross fox, but if they exist, I’ve not heard of them.

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