A Miocene Enigma
This copiously illustrated story of the pronghorn antelope (Antilocapra americana Ord 1818) is built upon two interlocking themes. The first explores the ongoing endeavor to describe and understand the species in terms of its origins, physical attributes, habits and habitats, and its interactions with other mammalian species, including humans. The second theme features highlights from the generations-long effort to arrive at a definitive image of the species, which began about 1650, and climaxed in the 20th century after high-speed camera lenses became common.
The two threads crossed in September of 1804 when the Corps of Discovery first encountered the pronghorn antelope at the eastern boundary of its native habitat, in today's southeastern South Dakota (Fig. 2). The Corps' journalists documented the men's learning curves as they figured out how to hunt them, wrestled with suitable names, observed their relations with Indians, analyzed their strengths and weaknesses, took their measurements from various perspectives, and above all learned to admire and respect them.
I. The First 15 Million Years
Pronghorn in flight–cheetah in pursuit
Sucking air through its flared nostrils and open mouth, the hairs of its mane erect in alarm, and those of its white rump patch erect to urgently signal the rest of the herd to follow, a pronghorn buck springs from his sagebrush lunch break less than a leap ahead of the carnivore whose ancestors may have helped to make the antelope what he is today, an "elegant speedster." For nearly two million years the pronghorn's principal predator was the North American cheetah (Miracinonyx inexpectatus),1 half again as large as the modern African cheetah. It might have been able to sprint faster than a pronghorn, but it was out of breath and out of the race at about the three-mile mark. Unknown natural forces extirpated the American cheetah some 12,000 years ago, leaving its prey the winner by a long shot.2
Birth of the species
Tracings of the faint lines of natural selection among mammals tell us but little of the points of origin of today's animals in time and place. Their evolutionary predecessors may have originated elsewhere on the globe. The North and South American land masses had been separated since they drifted westward from Europe and Africa, nearly 175 million years previously, and were not connected until the Isthmus of Panama rose up during the mountain-building era, approximately 2.5 million years ago. Thus for nearly 13 million years our pronghorn antelopes' ancestors were restricted to parts of western North America (Figs. 2, 3).
Estimated Range, 1900 BCE
At the close of the 19th century, zoologist C. Hart Merriam (1855-1942), the first chief of the federal agency that was to become the U.S. Fish and Wildlife Service, and one of the founders of the National Geographic Society, estimated the pronghorn's range as shown above. Note that whereas Lewis and Clark saw their first pronghorns near the eastern border of today's South Dakota, less than a hundred years later the surge of settlers across the plains had pushed the herds into the western quarter of the state.
Pronghorn Territory in 2000
A ghostly shape embraces the general region in which fossil evidence has suggested that pronghorn herds have thrived for millions of years on western savannas and high, relatively dry, wide-open prairies, but never in deep forests nor–as the words to "Home on the Range" fantasize it–"on mountaintops green" (see below, "Codicil: A more musical name"). On the basis of slim fossil evidence, paleobiologists have estimated that the ancestors of the animal we now call the pronghorn appeared on North America's grassy steppes and tree-dotted savannas during the middle Miocene, about 15 million years ago.3
By the time humans entered its native habitat some 20,000 to 15,000 years ago, the family Antilocapridae contained only two genera, one of which, the prong-horn, looked much as we see it now, and there may have been between 20 and 40 million of them alive at the peak of their population. Owing largely to human predation, which accelerated rapidly in the 19th century as new opportunities drew entrepreneurs and other settlers westward, the population fell steadily toward extinction until about 1910. Since then it has recovered nearly to the maximum number supportable by what remains of its favored habitat, reaching approximately one million head in the 1980s.
Perhaps 1.5 million years ago, Antilocapra americana was one of about a dozen species in the zoological family that only since 1866 has been known as Antilo-capridae (an-til-lo-cap-rid-dee, "goat-like antelopes"), but all except this one one have been extinct for thousands of years, and it has no close relatives living anywhere else on earth.
The pronghorn that we know owes its survival to its princi-pal defenses: speed and endurance. And as we shall see, nearly every aspect of its physical makeup contrib-utes in some way to those strengths. It can accelerate in seconds to a cruising speed of 40 or 45 miles per hour and keep it up flat-out for four miles or more, with bursts up to 55 or 60 mph in linked strides reaching nearly 30 feet each.
With its visual acuity equal to eight-power binoculars, it can detect motion three or four miles distant, but may suddenly stop in midflight from a perceived threat and trot back for a closer, stiff-necked, seemingly puzzled look at the supposed threat. At the time of its family's evolutionary peak its mem-bers all dreaded the claws and fangs of saber-toothed tigers, lions, hyenas, and especially American cheetahs. Today its enemies are hardly in the same league in terms of pursuit. The eager but low-gear coyote is an expert ambusher; the invincible grizzly bear culls out the sick and the lame; bobcats, ravens and golden eagles prey upon newborn fawns. But mature, healthy adults have little to fear from any predators but human hunters or–when deep midwinter snow drives them into populated neighborhoods–unattended domestic dogs.
Notwithstanding their peerless speed on the hoof and their awe-some endurance, pronghorns are relatively fragile beasts. A few members of a herd may live for 14 years, while most fawns are lucky to last one full year. The average life expectancy of adults is 4.5 years.4 Unfortunately, none of nature's wonder-workings have equipped the species to cope with humans' greed for land of their own, much less with the long fatal finger of a high-powered rifle. So, considering the pronghorn's relatively short lifespan, it may seem that nature has over-invested in this beautiful animal's survival.
First Impressions—before Lewis & Clark
For many years the known facts relating to this particular animal were few and often vague, but they added up to one question which, during the decade of the Lewis and Clark Expedition, was of fundamental importance in those climactic years of Enlightenment cosmography: Where to put it in a logical structure of an orderly universe, in the taxonomy–the epitome of scientific methodology–of four-legged, twin-hoofed and hollow-horned, cud-chewing grazers.
Tribes from central Siberia began migrating to North America via the Bering land-bridge perhaps 30,000 years ago, gradually spreading out across North and South America and, between periodic Ice Ages, traipsing the Old North Trail southward along the eastern margins of the Rocky Mountains to Mexico and beyond.5 En route they must have encountered many herds of this species or its predecessors, fed on their flesh, and made clothing of their hides. Numerous peoples also made up names for them.6
Antilocapra americana Ord, 1818
Built for speed
Crowned by lethal-looking bantamweight horns on a slightly oversized head and mounted on lithe runner's legs, this pronghorn antelope's deep-chested, stocky torso holds an exceptionally large heart and capacious lungs to power it through high-speed, long-distance flights from perceived threats.
—Bev Wigney is a naturalist-photographer whose galleries may be viewed at http://magickcanoe.com/
With the collapse of the Aztec Empire in 1519 under the onslaughts of Spanish invaders, sightings began to be written down in European languages laced with smatterings of Nahuatl, an Aztec tongue. Still, the references were vague and the descriptions confusing or even contradictory, mainly because the writers were neither naturalists nor philologists, but were preoccupied with more practical, political matters—expanding a Spanish empire in America, stealing gold and silver, consolidating political and military power, and leading heathens to Christianity. When they saw an animal that was new to them, the best they could do was make a guess, on the fly so to speak, that this or that one seemed to resemble some kind of a deer. Or goat, sheep, or gazelle, or an antelope—species that they had learned to identify back home, or had seen illustrated in books about apparently similar species. But what-ever the pronghorn seemed to resemble, its most obvious characteristic was its astonishing speed.
The first of the conquistadors to record anything that can be interpreted as a reference to the pronghorn was Pedro de Castañeda, who chronicled Coronado's search for Cibola, the mythical Seven Cities of Gold, in the early 1540s. Castañeda claimed to have been told by other Spaniards who had traveled through the region that flocks of "goats" had been seen "which ran so fast that they disappeared very quickly."7 Nearly 250 years later, two Franciscan missionaries traveling through what is now Utah, Colorado, Arizona and New Mexico saw some "wild sheep" that seemed to be larger and "much swifter" than domestic sheep.
Beginning in 1771, while Louisiana was still a French colony, Jean-Bernard Bossu (1720-1792) made an official sightseeing tour up the Mississippi River as far north as Fort Cahokia, opposite St. Louis. He reported being told that "wild goats and kids" which were "extremely agile and alert" could be seen near the source of the Missouri River, wherever that was. "The Frenchmen who have tasted them," he continued, "assure me that the young kids are as good [to eat] as Briançon sheep."8 Five years later, while the colonists on the opposite side of the continent were preoccupied with their War for Independence, the scribe for Lieutenant Don Jose Joaquin Moraga (1741-1785), in command of the contingent of Spanish soldiers and colonists who founded San Francisco and San Jose in California, wrote of seeing a
species of deer about the size of three-year-old sheep. They are similar in appearance to the deer, except that they have short horns and also short legs like the sheep. They live in the plains where they go in herds of 100, 200, or more. They run all together over the plains so fast that they seem to fly. . . . These animals are called berrendos.9
Berrendos, a Spanish word meaning "two-colored," is still the everyday Mexican name for Antilocapra americana.10
Little by little the glossary of facts, near-facts and vague impressions grew. Edward Umfreville (ca. 1755-1789), a British accountant with the Hudson's Bay Company during the 1780s, published his description of the new species in 1790 after having explored the farthest reaches of the Red and Saskatchewan Rivers. He admitted that he was "not sufficiently conversant in the science of Zoology to give this beautiful animal its proper name in the English language," and even allowed that "perhaps it has never yet been described in natural history."
The French people resident in these parts call it Cul Blanc ["white bottom"]. . . . A more beautiful creature is not to be found in this or perhaps any other country. Extreme delicacy of make, and similarity of proportion, are observable in all parts. No animal here is so swift of foot; not the fleetest horse or dog can approach it. They herd together in large droves, but sometimes three or four are found in a place. Its horns are not ossified like [those of] the other species, nor are they branched. . . . they resemble more the horns of the Goat than those of the Deer species. . . . The whole length may be about four feet and a half; the legs are white and slender; the rest of the body a light red.11
He also mentioned its Cree Indian name, "Apis-to-chik-o-shish," although he felt no compulsion to render it in English.12
By the end of the 18th century, naturalists in Europe and the eastern U.S. had heard a few rumors about it, and were eager to find the answers to some basic questions. How would you recognize it if you saw one? How would it act, and why? Where in the world would you be? What uses could be made of it? Above all, what would you call it? Lewis and Clark were to produce only a few answers, and many of those fell through the cracks of history for a considerable time when both Meriwether Lewis and Benjamin Smith Barton failed to produce the promised book on the expedition's scientific discoveries. Fortunately, however, the information about the pronghorn antelope that survived in Nicholas Biddle's paraphrase of the journals was enticing enough to earn the explorers some credit for a minor but durable role in the pronghorn's recorded history.13
Forgotten impressions—Lewis & Clark
On September 3, 1804, a clear and pleasantly cool day, the men of the Corps of Discovery caught their first glimpse of some pronghorn antelopes on the prairie a couple of miles up the Missouri from today's Springfield, South Dakota. Their French-Canadian engagés knew the elusive animals as Cabri, French for "goats," so that's what Clark called them. Otherwise, all he could say of them with certainty was that they were "wild and fleet"—timid and fast.
The two-toed, black front hooves of a mature pronghorn buck, with narrow rounded tips at the fore, are about 7.5 cm (3 in.) long. The rear hooves, similarly wider at the heel than at the fore tip, measure 5.5 cm (2.2 in). Lengths may vary by region. All four pairs of hooves are cushioned with cartilaginous padding to prevent bruising on hard or stoney ground. It can be very difficult to distinguish between deer and pronghorn prints, except that at a bounding run the latter's rear hooves push off on each stride, while the narrow tips of the front hooves bear most of the animal's weight upon landing.
On the fifth, some ten miles above the mouth of the Que Courre, or Rapid River (now the Niobrara, in northeast Nebraska), the captains evidently browsed their copy of Owen's Dictionary in search of some clues as to the animal's "real" identity. The search was fruitless; because of the date and place of its publication, Owen's was perforce limited to European species. The best that Lewis could do was to record that on that day they had spied some "wild goats or antelopes." Again the animals raced out of sight with supernatural speed. "We could not discover them sufficiently distinctly to discribe even their colour," he wrote. They were all left to contemplate the speedsters' hoofprints. Indeed, the hoof prints of a full-grown pronghorn are practically indistinguishable from those of a mature deer of any species, unless the ground is soft enough to show the deer's dewclaws–which the pronghorn lacks. Only a lifetime spent on stony soil would noticeably blunt the normally sharp leading edges of a pronghorn's hooves (Fig. 4).
Clark spent all day on the 14th walking on shore in search of the volcano that was rumored to be in the vicinity of the White River. Of course, he didn't find the phantom volcano, but he did come upon some more pronghorns, and somehow managed to shoot "a Buck goat." One wonders how he did it, considering the number of unsuccessful tries the Corps' other hunters made during the next few weeks. Did one of the engagés tip him off to the fact that he should rely on the animals' innate curiosity and wait for one of them to stop running and return within rifle range for a closer look at him?14
That evening he and Lewis together studied the pronghorn specimen and agreed upon some more or less accurate notes of its most prominent features. While Lewis turned his time and attention to another new species of interest, the jackrabbit (Lepus townsendii), it was up to Clark to write up the pronghorn. It was, he thought, "about the hight of the Grown Deer," although its body was shorter. Instead of antlers it had horns which were "not very hard," and they forked ⅔ of the way above the roots, with one prong flat and short, the other round, "Sharp arched," and growing immediately above its eyes. The animal's color was "light gray" with black behind its ears and on its neck. Its jaw was white round its neck, sides and rump. Its tail was "Short & white." The pronghorn specimen was "verry actively made," and had "only a pair of hoofs to each foot," which might be taken to mean that the captains noticed the absence of dewclaws. They obviously went beyond a superficial examination, for Clark noted that its brains were at the back of its skull. The nostrils were large. This animal was, Clark concluded, "more like the antelope or gazella of Africa than any other Species of Goat" (Fig. 5).
In his own journal for September 14, Lewis recorded the measurements of Clark's "wild goat": length (57"), height at withers (36"), height behind the withers (36"), girth of the breast (37"), girth of neck near shoulders (26"), girth near the head (19"). Lewis was especially struck by the eyes, which were "large and prominent." The pupils were "deep sea green, large percing and reather prominent, & at or near the root of the horn within one º inches."15 The green color Lewis saw was the tapetum, an iridescent membrane at the retina that enhances most wild animals' vision in dim light, and reflects the "eyeshine" that we notice in our nighttime photos–and in our automobile headlights.
Finally, Lewis recorded the weight of the specimen at 65 pounds, which is difficult to explain, considering that it was shot in mid-September when it should have been approaching its maximum weight for the year. The live weight of an average-sized mature male pronghorn (2 years of age or older) probably would have been between 102 and 134 pounds, averaging 118. Its hog-dressed or field-dressed carcass (i.e., minus the viscera, stomach and contents, diaphragm, and legs below the knees) would have weighed, on average, 84 pounds. Applying the currently accepted formula for reverse-calculating live weights from dressed weights, the captains' first pronghorn kill would have weighed only 88 pounds on the hoof.16 But Lewis gave no hint that it was "poor," so was his "65 pounds" just a rough and hasty estimate on his part, or did he misread the scale on their pocket steelyard (if they had one)? Or was it merely a slip of the pen?
On the 16th, "vast herds of Buffaloe deer Elk and Antilopes wer seen feeding in every direction as far as the eye of the observer could reach." By the 20th Clark had seen enough pronghorns to express his gut-level admiration in one angular but trenchant eight-word summary: "They are all keenly made, and is butifull." Nicholas Biddle noticed Clark's phrasing, but buffed it down to nothing but the facts. Those pronghorns were, he yawned, "delicately formed and very beautiful."
All of the foregoing details, and many more, were in the original manuscript journals. At Fort Mandan on May 5 of 1805, for example, Lewis observed two local, seasonal vulnerabilities which were among the principal causes of mortality in the pronghorn herds.
My dog caught a goat, which he overtook by superior fleetness, the goat it must be understood was with young and extreemly poor [undernourished]. a great number of these goats are devowered by the wolves and bear at this season when they are . . . passing the river from S. W. to N. E. they are very inactive and easily taken in the water, a man can out swim them with great ease; Indians take them in great numbers in the river at this season and in autumn when they repass to the S. W.
Beyond that, there were numerous ethnological tidbits, mainly about Indian clothing made of antelope skins, that Lewis or Barton might have included in the scientific summaries they never got around to writing.
But before Biddle's paraphrase was off the press in the winter of 1814 all of the original manuscript drafts of the captains' journals were laid away among family records and in the archives of the American Philosophical Society, where their contents were effectively hidden from sight for nearly 100 years. No one thought to look into them for more about the explorers' discoveries, nor even to wonder what else they might have said about the pronghorn beyond what Biddle had grasped. It was all yesterday's news.
In 1901 the American Philosophical Society, anticipating the centennial of the expedition, persuaded Dodd, Mead and Company, to publish the original journals under the editorship of the eminently qualified Reuben Gold Thwaites. With the appearance of the 8-volume edition in 1904-1905, the vast amount of "scientific" details–geographical, ethnological, and biological–that Biddle had purposely omitted from his paraphrase, was at last available to every student of the Lewis and Clark Expedition.
Clark's conclusion about the animal's correct name suggests that he and Lewis consulted the 8-volumes-in-4 encyclopedia they carried, familiarly known as Owen's Dictionary, for whatever help it might contain. The one-line entry for "antelope" referred the reader to the article on "gazella," which read:
GAZELLA, in zoology, the name of several species of goat: as, 1. the african gazella, called also antelope, . . . the horns of which are cylindric and half way arched. 2. The indian [referring to India, in Asia] gazella, or antelope, with very long, cylindric, and straight horns, annulated [ringed] at the base. 3. Another species of african gazella, with cylindric, arched, and perfectly annulated horns. This last is a small, but very beautiful species, and greatly resembles the common deer in shape: the horns, which arise from the middle of the forehead, are of a beautiful black colour, and annulated all the way from the base to the very tips.17
In 21st-century zoological taxonomy the goat, the gazelle and the pronghorn are classified in the order Bovidae and the Family Artiodactyla, which is to say that, first, they are ruminants having multi-chambered stomachs, and that they feed on vegetable matter; and second, that all have cloven or two-toed hoofs.
But whereas domestic and wild goats are in the genus Capra, and there are now 18 species of gazelle in 6 different genera, the pronghorn is the sole remaining species in the genus Antilocapra.
From the early 19th-century perspective, the crux of the problem was that the respective nomenclatures of European, Asian and African mammals were as yet unconsolidated. Therefore Lewis and Clark had no reason to suspect that their "goats" were unrelated to any existing African species of antelopes. Nor was it yet clear that African gazelles and gazellas did not have spiral horns, but there were nine species in two genera of spiral-horned African antelopes, including the bushbuck, the eland, and the kudu.18
II. Capturing the pronghorn's image
Again: If you had only seen an artist's impression of a pronghorn, how would you recognize it if you encountered a live one? From the middle of the 16th century to the end of the 19th, the publication of books containing engravings, often lovingly hand-tinted, of artists' impressions of mammals, plants, birds, insects and fish increased steadily, year by year. Surely the first European or American who caught sight of a pronghorn felt the impulse to try to capture its image with paint or pen. Almost certainly, however, it was the animal's two conspicuous qualities, shyness and swiftness, that doomed the best of intentions. Remember that after their second view of a herd, Lewis and Clark couldn't even recall the colors of those "goats" clearly enough to put them into words. The following seven images illustrate some of the stages in the emergence of the pronghorn's likeness by artists would could never see one in action, in the flesh.
The earliest drawings that seem to have been meant to represent the pronghorn appeared as embellishments or teasers on two sixteenth-century maps of southern Mexico, but the first published likeness appeared in 1569, when Father Bernardino Sahag˙n (1499-1590), a Franciscan missionary who documented the Spanish conquest of Mexico, published a drawing of the mysterious quadruped with a few words of explanation.19 Sahag˙n's description, however, was cryptic and his illustration was little more than a quick-draw cartoon, so neither could be taken very seriously.
Meanwhile, King Philip II of Spain ordered his court physician, Francisco Hernández de Toledo (1517-1587), to inventory all of the medicinal plants he could find in New Spain. Accompanied by his son Juan, and assisted by three Aztec painters who prepared illustrations, he spent seven years carrying out his assignment, with the assistance of various translators. A four-volume report on his results was published posthumously in 1615, in Spanish, translated by Brother Francisco Ximénez, but it contained only one illustration, a frontispiece. Thirty-six years later a new edition of Hernández's original Latin version appeared, supplemented by 800 illustrations, including 30 woodcuts of native fauna.20 One of those pictured a quadruped having two flat horns, each with a single anterior branch or prong. Owing perhaps to an unexplained disconnection between Hernández and his artists, the illustration didn't jibe with his description. In those days, though, when it came to discussions and illustrations of exotic species, readers were accustomed to making allowances. Indeed, it may have been this depiction of the Teuthlalmaçame (Fig. 6), or even mere rumors of it, that captured general interest as well as imagination, and led to more misunderstandings among naturalists in Europe and America.
A "tolerably good" figure
Certainly, Hernández's figure of the Teuthlalmaçame resembles a pronghorn antelope in that the horns appear to be flat, and there are white collars on the throat, and in the accompanying text he described it as being "a little larger than a medium sized goat." Furthermore, he wrote that it was "covered with gray hair, easily pulled out and, contrary to the drawing, "with sides and belly hoary white, hence the Spanish natives were accustomed to call them Berrendos," all of which would seem to identify it as what was later called a pronghorn. On the other hand, the tapered monochromatic body and the dewclaws in the figure do not belong to a pronghorn, any more than does the perky tail. In fact, the root of Teuthlalmaçame is the Nehuatl word Maçame or Mazama, which means deer. Hernández didn't mention its most obvious attribute, its blazing speed, which suggests that he himself never actually saw one in the flesh.
Nevertheless, Hernández's ambiguous drawing and confusing description, while defying logic and challenging interpretation, continued to draw attention for more than 150 years. William Dunbar (1750-1810) of Natchez, Mississippi, the Scot whom Jefferson engaged to lead an exploratory expedition up the Washita River, was aware of the confusion, either from Hernández's book or through his correspondence with Jefferson or Barton, or his occasional house guests from Philadelphia, William Bartram and Alexander Wilson. As Dunbar noted in his journal,
The great western prairies, besides the herds of wild cattle, (bison, commonly called buffaloe) are also stocked with vast numbers of wild goat (not resembling the domestic goat) extremely swift footed. As the description given of this goat is not perfect, it may from its swiftness prove to be the antelope; or it possibly may be a goat which has escaped, from the Spanish settlements of New Mexico.21
The eccentric Constantine Rafinesque contributed mightily to the confusion in 1817 by establishing Mazama as a genus and placing in it seven native American quadrupeds, including two that Lewis and Clark had discovered, which had already been named by George Ord (1781-1866)–the mountain goat and the pronghorn antelope.22 Rafinesque's taxonomy was sternly repudiated by Spencer Fullerton Baird (1823-1887) of the Smithsonian Institution in 1857.23 However, in 1850 the American botanist Asa Gray (1810-1888) reaffirmed his predecessor's decision concerning the mountain goat, and on those grounds a mountaineering club that was founded in Portland, Oregon, in 1894, took Mazama for its name, and a figure of a mountain goat for its logo.
By the beginning of the 21st century, the word Mazama had withdrawn to parts of Mexico and South America where it belonged, becoming the legitimate generic class containing four species of the rare brocket deer.
Sorting out the pieces
On March 5, 1806, the Philadelphia physician and naturalist Benjamin Smith Barton, friend of Thomas Jefferson and pre-expedition tutor of Meriwether Lewis, drew a bold line from Hernández's Teuthlalmaçame to Lewis and Clark's reported discovery of a new "wild goat or antelope." In his own magazine he posted a brief news item for his readers to contemplate, which was based on communications and specimens the captains had shipped to President Jefferson from Fort Mandan in April of 1805:
Among the animals which Captains Lewis and Clark met with . . . there is one which cannot fail to prove interesting to the naturalist. It is a species of the genus Antelope. The existence of this animal, as a native of North-America, has been hinted at by several writers, more than one hundred and fifty years ago, . . . and a tolerably good figure of the entire animal was published as early as the year 1651.24
Dr. Barton had been misled by Hernández's figure, and would not see the captains' specimens for another six months.
One month later Thomas Jefferson wrote to the Philadelphia naturalist William Bartram: "I dare say you know that Westward of the Misipi there is an animal of the Capra [goat] kind; & tho by some it is called a deer, & by some a goat that would not authorize us to call it the Cervicapra ["deer-goat"] of the East." The Osage Indians who had visited the President in Washington City the previous year had shown him a sample of leather cured from the animal's skin which, he exclaimed, was superior to anything of the kind he had ever seen. "It receives no injury from being wet," he said, adding confidently, "I count on special information as to this animal from Capt. Lewis, and that he will enrich us with other articles of zoology, in which he is more skilled than in botany."25
Jefferson was of course unaware that on that very same day, April 7, 1805, Lewis had shipped to him via the returning barge a large collection of plant and animal specimens of plants and animals, including, according to Biddle, who may have learned it from either Clark or Shannon, "a stuffed male and female antelope with their skeletons." The shipment arrived on Jefferson's doorstep in Washington City early in October, much the worse for the wear and tear of its long journey.26 The President, excited by the promises of Clark's invoice, but without taking enough time to open all of the boxes, hastened to tell Charles Willson Peale what had come from the far West, and prematurely expressed one reservation:
I have some doubts whether Capt Lewis has not mistaken the Roe for the Antelope, because I have received from him a pair of horns which I am confident are of the Roe (tho' I never before supposed that animal to be in America) and no Antelope horns came. These you know are hollow, annulated [ringed], single. Those of the Roe are horny, solid, & branching. I hope you will have the skeletons well examined to settle this point. You will recieve them in great disorder as they came here, having been unpacked in several places on the road, & unpacked again here before I returned, so that they have probably got mixed."27
Three days later Jefferson wrote to Peale apologetically, having found "the bony prominence to the cranium on which the horn is fixed," plus two pairs of horns that fit over them. He forwarded one pair to Peale. "These," he wrote, "sufficiently prove that the animal is of the Antilope family & of the Chamois branch of it. This is strengthened by the dressed skin which is softer, & stronger in it's texture than any Chamois I have seen."28
Peale responded thoughtfully, confident that the "several skins" would make a valuable addition to his Museum even though some parts of them were in bad condition "owing to the Moth & Dermest having made great havock."29
The Skeletons are much broken and I fear some of the bones are lost at the places where they have been opened. I can mend the broken bones but cannot make good the deficiency of lost bones, being mixed togather is of no great consequence, as every bone must find its fellow bone. Whether I can get an intire skeleton from this mass of bones, I cannot yet determine, it will be a work of time and the exercise of much patience.30
Time and patience, indeed. Five months later he informed Jefferson that "the Skins of the several Antilopes was so badly managed in the Skinning, and also so much eaten by Dermests, that it was with much difficulty I could mount one of them, but being so interesting an Animal, I conceived it was better to have one even in bad condition, than to let it be wanting in the Museum." Not having received a description of the animal from Lewis, he began one himself for possible publication in the Transactions of the American Philosophical Society. He included a copy of the 375-word rough draft in a letter to Jefferson, and enclosed a pen-and-ink drawing. However, upon Lewis's return to Philadelphia in June of 1807, Peale withdrew his description; his drawing has never been found.31
Early drawing from the mounted specimen
in Charles Willson Peale's Museum
by Charles Hamilton Smith, 1819
(Antilope furcifer, "Forked antelope")
The figure shown here accompanied a paper on the exotic quadruped that was read by the English antiquary, naturalist, artist and illustrator Charles Hamilton Smith (1776-1859) before the Linnæan Society of London in 1819. He himself had drawn its picture from the mounted specimen in Peale's Museum at Philadelphia—"the only one preserved of those which Messrs. Lewis and Clarke sent to the President . . . during their exploratory travels up the Missouri." Smith described it as a complete skin of an adult male, "stuffed with great skill, although in a very indifferent state of preservation." Peale had given him access to the glass case containing the specimen in order to record its dimensions. The average measurements of adult males arrived at by biologist Bart O'Gara in 1968 were: Length, nose to tail (straight line), 58 inches; height, shoulders to soles of hooves, 34 inches.32 Smith's and Lewis's corresponding measurements were nearly the same as O'Gara's.
Another drawing based on the specimen mounted in Peale's Museum, lithographed by Alexander Rider from a sketch by Titian Ramsay Peale, appeared in the 1830-34 edition of Doughty's Cabinet of Natural History and American Rural Sports. Like Smith, Rider portrayed the species in a lofty Alpine setting, notwithstanding Nicholas Biddle's numerous references to its presence on the plains.
The hairs of a pronghorn's mane are between 2.5 and 4 inches long. The mane extends from between the ears down the back of the neck to the withers, with the longest hairs toward the top. The mane in Hamilton's-Smith's drawing of Peale's mounted specimen suggests that this was one of the skins that was damaged in transit, and that what remained implied a topknot or plume rather than a continuous mane. No one had yet recorded any observations of pronghorns erecting their manes. Compare this detail in Smith-Hamilton's drawing with Titian R. Peale's watercolor (Fig. 9), Leseuer's drawing of Peale's mounting (Fig. 10), Audubon's treatment (Fig. 11), Caton's drawing from his photograph (Fig. 12), and Wallihan's photograph (Fig. 13).
Nicholas Biddle's legacy
One of Thomas Jefferson's greatest disappointments over the outcomes of the Expedition was the eight-year delay in the publication of the captains' journals. Worse yet was the indefinite postponement of a report on their scientific discoveries, the consequence of two other uncontrollable circumstances: Lewis's unexplained failure to produce a single word of the illustrated reports he had publicly announced in March and April of 1807; followed by Dr. Barton's declining health, which interrupted his promised collaboration with Biddle and, upon Barton's death in 1815 at age 49, doomed all hope of committing the captains' botanical and zoological notes to print. Whatever useful information Biddle omitted from his paraphrase of the captains' manuscripts would await Reuben Gold Thwaites's edition of the original journals in 1905, by which time it was no longer of much interest to scientists.
The "mere journals," as Jefferson referred to Biddle's two-volume History33 were the only sources of information about the captains' northwestern discoveries for almost 80 years while, as the president had predicted, the rest of their secrets remained for others to discover.
Meanwhile Patrick Gass's journal, published in 1807, was eagerly read, judging from the number of its U.S. reprints and European translations. It was, however, "the least minute and valuable" of all four of the journals then known, according to Paul Allen, who was the publishing editor of Biddle's paraphrase. The only informative entry among its manifold trivialities on pronghorns was a description of the way wolf packs took advantage of a herd's defensive reaction of bunching up and running in circles. Relays of wolf teams then spun the circle without letup until the weakest of the herd's members faltered, lagged, and were then brought down. Gass claimed (August 28, 1806) to have witnessed just such a scene during his time at the Falls of the Missouri on the return trip.
Nevertheless, Biddle's condensation of the captains' journals held some of the major keys to the earliest descriptions of the new animal. The assimilation of those keys was somewhat tedious because of the absence of an index, possibly owing to pressures of cost and urgency. Among the first qualified men to endure that tedium was George Ord, the Philadelphia naturalist who wrote the section on North American zoology for the 1815 edition of William Guthrie's popular New Geographical, Historical, and Commercial Grammar. For it Ord composed the first published scientific description of the pronghorn–drawn almost verbatim from Biddle's paraphrase–along with his initial choice of a binomial, Antilope americanus, which he revised in 1818 to Antilocapra americanus. Coincidentally, his new generic name was a contracted Latin translation of the captains' equivocation, "antelope or goat," which was their tentative first answer to the question.34
The men of the Corps of Discovery must have been electrified by their first sighting of the pronghorn antelope near the mouth of the Niobrara River at the northeast corner of today's state of Nebraska. In Biddle's words it read, matter-of-factly, "Between Pawnee Island and Goat creek [now Chouteau Creek] on the north is a cliff of blue earth, under which are several mineral springs, impregnated with salts; near this we observed a number of goats, from which the creek derives its name." He treated subsequent encounters with somewhat more sympathy. Here, in essence, are most of the points concerning the pronghorn antelope that Biddle chose from the captains' journals. It is important to bear in mind three facts: First, he had ample reasons to assume, as he penned his paraphrase, that Benjamin Smith Barton would write a summary of the expedition's scientific discoveries; second, Barton's abnegation of that responsibility owing to his ill health, was out of Biddle's control; and finally, for the next 80 years, readers of the Biddle-Allen version of the journals, published in 1814, could only assume that this was all the captains had to say about the subject.
• September 17, 1804 (Lewis): "Most wonderful fleetness; shy and timorous, . . . repose only on the ridges . . . acuteness of their sight . . delicate sensibility of their smell . . . more like the flight of birds than the movement of an earthly being . . . only male encircled the summit of the hill, as if to announce any danger to the females. . . . the smell [of Lewis's proximity] alarmed them. . . a speed equal to that of the most distinguished race-horse." • September 20, 1804 (Clark): "Female . . . smaller in size; . . . horns smaller and straighter . . . one sharp prong . . . no black about the neck . . . none of these goats have any beard." Biddle's transcription of Clark's summary, "keenly made, and is beautiful," lacked Clark's spontaneity in becoming "delicately formed and very beautiful." • October 17, 1804 (Clark, from an Arikara Chief through interpreter Joseph Gravelines): "large flocks . . . spend the summer . . . in the plains east of the Missouri, . . . at the present season are returning to the Black mountains, . . . subsist on leaves and shrubbery during the winter, . . . resume their migrations in the spring." • April 14, 1805. (Lewis): "a plant . . . similar to the camphor in smell and taste; . . . another plant of the same size, . . . of an agreeable smell and flavour, . . . a favourite food of the antelope, whose necks are often perfumed by rubbing against it." • July 25, 1805 (Lewis): "Collected in small herds . . . several females with their young, attended by one or two males . . . [some] males solitary or wander in parties of two . . . the plains, which the antelope invariably prefers to the woodlands."
Biddle selected several accounts of Indians' various ways of hunting the pronghorns.
• October 16, 1804. (Lewis): "great numbers of goats on the banks of the river, . . . saw large flocks of them in the water. . . . driven into the river by the Indians, who now lined the shore so as to prevent their escape, and were firing on them, . . . boys went into the river and killed them with sticks; . . . we counted 58 which they had killed." • November 5, 1804. (Clark): "camp of Mandans caught within two days 100 goats a short distance below us. . . . form a large strong pen or fold, from which a fence made of bushes gradually widens on each side. . . . surrounded by the hunters . . . driven toward this pen, . . . imperceptibly find themselves inclosed, . . . at the mercy of the hunters." • August 14, 1805. (Lewis): "chief game of the Shoshonees . . . is the antelope. . . . About 20 Indians, mounted on fine horses, and armed with bows and arrows, . . . descried a herd of ten antelopes. . . . separated into little squads of two or three, and formed a scattered circle round the herd for five or six miles, keeping at a wary distance, . . . a small party rode toward the herd. With wonderful dexterity the huntsman preserved his seat and the horse his footing, as he ran at full speed over the hills, down the steep ravines, and along the borders of the precipices. . . . lasted about 2 hours and considerable part of the chase in view from my tent. . . . about 1 A. M. the hunters returned had not killed a single Antelope, and their horses foaming with sweat."
As the Corps approached the mouth of the Yellowstone they became aware that they were entering a region where wild animals were numerous, and mutually fearless, and unanimously indifferent toward humans.
• April 22, 1805 (Lewis): "Large herds of deer, elk, buffalo, and antelopes in view of us." • April 26, 1805 (Lewis): "Elk, antelope, and buffalo suffered [Lewis] to approach them without alarm, and often followed him quietly for some distance."• April 30, 1805 (Lewis, April 29): "Antelopes are yet lean, . . females are with young. This fleet and quick-sighted animal is generally the victim of its curiosity. When they first see the hunters they run with great velocity; if he lies down on the ground and lifts up his arm, his hat, or his foot, the antelope returns on a light trot to look at the object, and sometimes goes and returns two or three times, till it approaches within reach of the rifle. . . . Sometimes leave their flock to go and look at the wolves, which crouch down, and, if the antelope be frightened at first, repeat the same maneuver, and sometimes relieve each other, till they decoy it from the party, when they seize it." • May 5, 1805. (Lewis): "The antelope are now lean and with young, so that they may readily be caught at this season, as they cross the river from S.W. to N.E." • June 26, 1805. (Lewis): "Females with their young, . . . generally two in number, and the males by themselves." • August 29, 1806 (Clark): "We observe that the greatest quantities of wild animals are usually found in the country lying between two nations at war."35
Biddle probably refrained from including Clark's short litany of facts simply because he expected they would be included in the volume of scientific data from the expedition that Barton had agreed to write. For the same reason, no doubt, he also omitted Clark's remark of September 5, the day he first saw a herd that ran away in such a blur that he couldn't recall their color, leaving only their hoofprints for his contemplation. Subsequently, Thwaites misplaced Lewis's measurements, as well as the flawed record of the weight of Clark's specimen.
Titian Ramsay Peale's watercolor
of an imaginary pronghorn family, 1820
Originally, the expedition that Major Stephen H. Long, of the U.S. Topographical Engineers, eventually led up the Missouri to the Rockies and back to the Mississippi via the Arkansas River, was part of a grand exploratory venture under the sponsorship of the federal government, which was to have made scientific investigations, recorded details of "the face of the land," and established a fort either at the Mandan villages or at the mouth of the Yellowstone. In practical terms, its outcomes were far less significant than those of the Lewis and Clark expedition. The fullest account of Long's journey, written by Captain John R. Bell, the official journalist for the party of soldiers, scientists and artists, was not published until 1957.36
Thomas Say was engaged as company's zoologist. The official botanist was the otherwise undistinguished physician William Baldwin (1779-1819). The principal artist was the landscape painter Samuel Seymour (1775-1823), who is believed to have produced 150 sketches during the expedition, from which he completed some 60 paintings, but only 17 of his works are known to exist today. Titian Ramsay Peale (1799-1885), Charles Willson Peale's youngest son, was hired as the party's "assistant naturalist." One hundred and six of the paintings and sketches he made during the expedition are in the collection of the American Philosophical Society, including four images of pronghorns.
Consistent with the naive pre-Darwinian tendency to find human social attributes in animal behavior, Titian Peale presented these four pronghorns as a family group in an unsentimental, undramatic pose. (Compare it with Wallihan's photograph, Fig. 12.) Titian borrowed the background from one of Seymour's landscapes, and based the two adults on sketches he had made earlier in the spring. He drew the two fawns from life on June 7, 1820, when a member of the party captured one and brought it to camp. (Fawns can easily be picked up during the first two days of their lives, when they tend to flatten their bodies against the ground and pretend to be invisible; by the end of their first week of life they can outrun any human.) Peale made his drawing and immediately set the little one loose. On June 17 Captain Bell, by no means a naturalist himself, recorded the event that may have prompted Titian to paint his watercolor on the 20th.
Lieut. Swift shot an antelope, or, as it is called by the french cabrie, this animal resembles the deer–is nearly of the same colour–but not quite as large–its horns are short, each having one short prang [sic]–the tail is shorter than that of the deer, the hair of it white and long, when running it is carried erect–their speed is about equal to the deer–but are not as wild, or they have more curiosity, for they will stand observing the hunter, until he arrives within rifle range. The meat when fat, is preferable to the deer.37
Titian's watercolor is a purely imaginary scene. In reality, females with fawns may herd up during the summer, but bucks do not keep company with them. Instead, they either spend the summer staking out their individual territories, or they roam in groups of from 5 to 20 bachelors. Also, horns on females, if present, are never longer than their ears. However, all of this mattered little, for Captain Bell's journal was not seen in print until 1957, and Titian's painting has never been published at all.
Leseuer's Drawing of Peale's Specimen, 1828
Antilocapra americana Ord,
from Godman's American Natural History (1828)
All that has been related concerning this animal which is worth repeating or remembering," declared John Godman, "was published in Lewis and Clarke's narrative, . . . and has since been confirmed by the observations of Dr. Richardson." Godman provided a 1,300-word description consisting largely of reworkings of Biddle's paraphrases of Lewis, and concluding with a paragraph by Sir John Richardson (1787-1865)—a physical description of a wild specimen he observed during the Northern Land Expedition with Sir John Franklin in 1827-28.38
Audubon's pronghorns, from nature, 1845
Preparing to flee
The buck at center is shown with a foreleg poised in the act of stamping. The narrative that accompanies Audubon's dramatic scene, written by the artist and edited into a lively homily by his associate, the Reverend John Bachman, reads in part:
Hurra for the prairies and the swift antelopes, as they fleet by the hunter like flashes or meteors, seen but for an instant, for quickly do they pass out of sight in the undulating ground, covered with tall rank grass. Observe now a flock of these beautiful animals; . . . they pause in their rapid course to gaze on the hunter, and stand with heads erect, their ears as well as eyes directed toward him, and make a loud noise by stamping with their forefeet on the hard earth; but suddenly they become aware that he is no friend of theirs, and away they bound like a flock of frightened sheep–but far more swiftly, even the kids running with extraordinary speed by the side of their parents–and now they turn around a steep hill and disappear, then perhaps come in view, and once more stand and gaze at the intruder.
Their walk is a slow and somewhat pompous gait, their trot elegant and graceful, and their gallop or 'run' light and inconceivably swift; they pass along, up or down hills, or along the level plain with the same apparent ease, while so rapidly do their legs perform their graceful movements in propelling their bodies over the ground, that like the spokes of a fast turning wheel we can hardly see them, but instead, observe a gauzy film-like appearance where they should be visible.39
The buck at center is stomping on the ground to alert his flock, who direct their gaze toward the object of their guardian's attention. Next he will urinate and then defecate in order to lighten his body for the exertion of running for his life.
Caton's drawing, traced from his photo, 1877
One of Caton's "prong-bucks"
John Dean Caton (1812-1895) was a prominent attorney and supreme court judge in the early years of Chicago's history, and a founder of what was to become the Western Electric Manufacturing Company, producing instruments for the new Western Union Telegraph Company. In addition, he was a serious naturalist with a particular interest in several species of deer, and in the Antilocapre americana that he preferred to call, informally, the "prongbuck."40
At his Ottawa, Illinois home Caton developed a private 200-acre park that he stocked with 50 or 60 deer, and a few pronghorns of various ages. Having no way at that point in time to know that many scientific details had deliberately been omitted from the only published account of the Lewis and Clark expedition, he nevertheless gave credit where credit was due:
It has been long known to hunters and trappers, but the scientific world is indebted to Lewis and Clark for the first accurate information concerning it; not from the description of it which they give, for they do not pretend to describe it, and only speak of a few of its peculiar habits; but rather from the specimen which they brought with them.
Caton studied his animals daily from season to season, with meticulous attention to minute details. In his book, The Antelope and Deer of America (1877), he contributed new details concerning the antelope's eyes, tail, horns, hairs of the coat, the palatability of the flesh as human food, and the possibility of domesticating the "prong-buck." He illustrated his book with his own drawings which, to insure accuracy, he made from photographs he took with his own camera.
Of the eye: He recognized that it is "very nearly the size of that of the elephant, and much larger than that of the horse or the ox. Those who examine only the dead subject would most likely be deceived in the largeness of the eye. The eye is black–intensely black–so that it is impossible to distinguish the pupil from the iris or its surroundings. No white part is ever visible, unless the eye is turned to one side; otherwise, all that is seen is one uniform brilliant black. But for all this, the expression is soft, gentle, and winning. In this respect it is the rival of the true antelope gazelle" Finally, Caton exulted, "How much the flashing of the great black eyes augments one's sense of admiration, the observer may himself be at a loss to determine." The eyelashes–Caton called them "eye-winkers"–were long, coarse and stiff, "more on the upper than the lower eyelid, but not very abundant on either."41
Of the tail: "It is very short, not more than three inches in length, and is covered with coarse hairs which are a little shorter on the under side than on the upper. . . . It is usually carried closely depressed for so short a member, and is never seen erected to a vertical position. When the animal is excited or animated the tail may be seen raised to a horizontal position or a little above it, but that is all the change in its position observed under any circumstances." The pronghorn's tail "is useless as a weapon for defense against the attacks of flies and mosquitoes," Caton observed, "from which, however, it does not suffer nearly so much as the deer, probably because of the odor with which it always surrounds itself." Undoubtedly he was referring to the pests Lewis referred to as eye-gnats, which are sometimes called pecker-gnats, from their attraction to genitals of large wild and domestic quadrupeds.
Of those horns: Caton's own microscopic analysis revealed the cellular transformations that take place during the conversion of epidermis into horn, a process that begins in the hairy skin covering boney projections that are part of the skull. The lower half of each horn, to a point a little above the prong, is laterally flattened and hollow, as is the prong. The upper half is solid, round, and pointed at its end; the ends are turned downward and inward toward each other. The growth of these horns has to be rapid, since they are shed soon after the rut is over; regrowth begins very soon thereafter. Caton classified the horn-buck as an intermediate link between those animals "which have persistent and those which have deciduous corneous appendages"–that is, antlers versus horns.
Of the pelage: "I have been unable," Caton wrote, "to detect the resemblance which the hairs of the Prong Buck bear to the wool of the sheep.
They are coarser than the hairs of any of the deer; they are hollow, with a larger internal cavity, are comparatively non-elastic, and exceedingly fragile. When bent short, they break down and never straighten again. . . . The internal cavity is filled with a light, spongy pith, and the whole is so fragile as to be readily crushed. . . . They have no more felting properties than dry brush-wood.42
Therefore, the Nez Perces' advice to Lewis on 30 May 1806 notwithstanding, the use of "goats hair" to stuff saddle pads would have been pointless.
Of the meat: In a letter that Lewis wrote to his mother from Fort Mandan on March 31, 1805, he remarked on the quality of the flesh of the "Cabri." As of that date, he wrote, he and his men found it "deliciously flaivored." By early May, however, when they were well into the game park west of the Yellowstone, the taste had declined in appeal. They all agreed that antelope meat was "less esteemed, and certainly . . . inferior" to the "fine veal and fat beef" of the buffalo. Fat was the keyword. Besides, Caton knew of no one who had tasted American antelope meat who did not find it "a delicate and choice morsel," but if it was used as a constant diet "one soon cloys of it and desires no more. After almost living upon it for two weeks, I quite forgot how much I enjoyed it at first."43 The men of the Corps of Discovery had similar reactions. But they didn't have to force themselves to like it, since the hunters rarely shot more than one or two pronghorns in a day. A mature male might yield between 50 and 60 pounds of meat–roughly half its live weight.44 But whereas they could eat well on a little more than two bison or five elk per week, they would have had to bag nearly 50 pronghorns a week to obtain an equal number of calories.45 Recent research at the University of Wyoming's Agricultural Experiment Station has determined that the musky odor of the skin is not imparted to the antelope's flesh, so it has no effect on the flavor of the meat.46
Domestication: As we have seen many times in these pages, the value of an animal was measured by naturalists of the Enlightenment in terms of its utility. That is not to say that sheer beauty, or size, or even ferocity, were not important. But animals that were new to science, or were relatively unfamiliar, were routinely examined for their possible utility as beasts of burden or as food for the table. Along that line, for example, Meriwether Lewis expressed his belief that bison hair might be used in the milling of cloth, and he was neither the first nor the last to think of it. However, only someone like John Caton, who had the opportunity to observe pronghorns day after day in captivity, was qualified to assess their potential cash value to the agricultural economy, and his experience made him skeptical. Early mortality among fawns doomed all hope of raising them to maturity for meat. They were first attacked by diarrhea, he wrote. "If they escape this, they live . . . one, two, or three months, growing slowly; but at the end of that time all the female kids, and almost all the male ones, become diseased, having scrofulous inflammation of the joints, get a cough, become lame and poor, and finally die after lingering some weeks."
The mature prong buck, he observed, was "very easily tamed, and soon lost all fear of the one who fed him, sought his society, and seemed to enjoy his company." It was easy, then, to yield to passionate anthropomorphism and persuade oneself that the captive animal "showed not only observation and intelligence, but even reflection."
Caton was aware that many wild animals in western America were already at risk of extinction, and that was reason enough to try to domesticate them by establishing private game parks and zoos. But Caton had seen the futility of such a plan with regard to the pronghorn. "I think there is little hope of their permanent domestication," he concluded, "by suddenly transferring them to the east of the Mississippi River, where they never roamed wild. We may keep them for a short time, but they will not prosper, and will soon sicken and die."
First published photo of pronghorns in the wild, 1894
This is one of four photos of pronghorns taken by Allen Wallihan (1859-1935) from a blind on the west slope of the Colorado Rockies in November of 1893. They were among the numerous images of various wild animals published by the photographer and his wife in Hoofs, Claws and Antlers of the Rocky Mountains, one of the first collections of photographs to document western American wildlife in the field. Wallihan described the experience of capturing this view: "A bunch came in and went down to the water about fifty yards below [me], but scared and came up and . . . right down to the place where you see them–forty-five feet. . . . You will notice the crooked horn one [fourth from left]. I saw that day every shape of horns that could possibly grow, I think."47
Wallihan titled his picture "Unconsciousness," but the combined effect of the location of pronghorns' eyes on the sides of their heads and the horizontal elongation of the pupils gives them a 320° range of vision, binocular directly in front of them, and monocular toward each side. Thus it is probable that all eight animals in the group were quite aware of the photographer's proximity but didn't spook and run because they had not yet seen any motion that they could interpret as threatening. Otherwise, at that distance their visual acuity might not have been sharp enough to enable them to recognize or interpret such anomalies such as two-legged animals hiding behind big black-and-brown things that made noises like tongue clicks. Strange.
In his introduction to the Wallihans' book, Theodore Roosevelt (1858-1919)–naturalist, explorer, hunter, author, soldier and 26th president of the United States (1901-1909)–lauded the collection as "absolutely unique" and "of the utmost value." Of the antelope photos he wrote, "The alertness of the queer prong-horned beasts is caught to perfection, as well as the difference in their attitudes when compared with the attitudes of deer."
The publisher explained a few of the technical challenges that affected the outcome, especially in terms of poses and lighting: "In the accumulation of the various subjects contained in this book, the photographer was rarely able to obtain the most favorable position or sunlight advantage. The extreme difficulty of securing first-class negatives under these conditions (many failures and few successes), is therefore apparent, and in some of the plates the work of reproduction by half-tone process necessarily shows a deficiency in the original, especially as the photographs have all been very much enlarged." The Wallihans' patience and determination as photographers was motivated by the growing evidence that civilization's westward march was about to kill off all the large wild animals in the American west. "Mr. Wallihan and I," wrote his wife Mary, "work together, happy in our effort of trying to preserve the game in photography for the world at large."
In the absence of collarbones, the pronghorn's scapulas, or shoulder blades, are linked to the skeleton by weight-bearing muscular attachments,48 allowing the pronghorn's shoulders to reach forward far enough to increase the length of stride by 20 percent over the maximum for species of similar size that have clavicles. Simultaneously, the flexible spine bends into a tight arch that places the hind feet ahead of the forefeet. When the power in that spring is released, the back muscles propel the body forward in bounding strides of up to 30 feet (Fig. 11). In place of the complex and bulky musculature that other grazers need in order to support heads and control necks that are long enough to reach food at their feet, and to keep the weights of necks and heads from interfering with the overall equilibrium that high-speed running requires, the pronghorn has a broad ligament connecting the back of the skull to the front end of the torso. This produces a running silhouette that shows little or no up-and-down movement and, combined with the instinctual grouping of a running herd into a tight ellipse, exhibits the fluent maneuverability of a flight of birds.
A cool head
Nature has equipped the pronghorn with a large trachea to supplement those large nostrils that Clark noticed, to supply to its lungs with the great amount of air required during a strenuous run. Its lungs are large enough to draw out the oxygen, and its broad chest holds a heart large enough to pump a high volume of oxygenated blood through an arterial network that includes a cooling system strategically placed in its oversized head to keep its brain from overheating. Its small four-chambered stomach allows the pronghorn to eat and run without spending much energy on digestion. Its system needs to be fueled almost constantly, and always selectively, so it browses on high-energy forbs, sagebrush, willow, and occasionally some cactus stems. But no grasses after the early spring green-up.49
The antelope's horns are built to save weight. They consist of a hollow sheath of keratin–the stuff of our fingernails–from 12 to 18 inches long, supported by a bony core that grows above the eye orbits. The horns' upper points are curved sharply down and inward, while a short hollow prong points forward several inches above the base. These flexible, fingernail-like horns are essentially sexual symbols that are useful chiefly during the rut, when their fragility is proven by damage (Fig. 9), and are shed soon after mating is over in early autumn. Regrowth begins almost immediately.
Although a majority of pronghorns belong to the subspecies Antilocapra americana americana, splitters have identified four other subspecies that differ from one another in minor details. Three occupy habitats south of the border: A. a. mexicana Merriam 1901, occupying central Mexico–see Fernando Hernández, above; A. a. peninsularis Nelson 1912, in Baja California; and A. a. sonoriensis Goldman, 1945, in southwest Sonora, Mexico. The fourth, A. a. oregona Bailey 1932, in Oregon, is said to be a far-western race. Other scientists, including the late Bart O'Gara, have maintained that the superficial distinctions among them, chiefly in minute differences in size, color, and conformation, make the validity of the division into subspecies a moot question.
"a gauzy, film-like appearance"
Antilocapra americana Ord
John Caton further observed, "I have seen them in my grounds make prodigious horizontal leaps across a ravine or depression in the ground from a standing position or a leisurely walk when there was no obstruction to impede their walking across it if they had chosen so to do."
These leaps seemed to require scarcely more effort than the walk. It was a horizontal bound so light and elastic that it seemed like a fleeting shadow, when the gentle walk would be instantly resumed with no more animation or excitement than if they had walked across the space. Still, . . . they are unable to make vertical leaps. I think it safe to say they cannot overleap an obstruction a yard in height."50
Around 1800 there were still perhaps 35 million pronghorns in western North America. In 1893, however, Elliott Coues commented that the species had "of late years become much restricted, both in numbers and in extent of distribution, and seems likely to share the fate of the buffalo."51 Naturalists such as John Caton had made a similar prediction 20 years earlier, so it was hardly a surprise that by 1920 the pronghorn population had dropped to less than 13,000 head, first as a result of the demand for meat in mining camps throughout their range, and then to the arrival of homesteaders, cattle ranchers and grain farmers with a similar need. Today wildlife biologists say there are more than a million pronghorns living in various parts of western North America. That surprising recovery is attributed to modern wildlife management policies firmly grounded in historical lessons, scientific understanding, landowner cooperation, and the dedication of sport hunters to whom flagging, for example, is outside the boundaries of "fair chase." Nevertheless, sport hunting remains an important management tool.
Hunting has accounted for the death of more than 100,000 pronghorn per year during recent years, but hunting is carefully controlled to maintain the health of herds. Sport hunting is needed to keep herds in balance with available habitat and prevent excessive crop damage.52
Thus by 21st-century standards pronghorns are classified as being of "least concern" on the IUCN Red List of Threatened Species.53 At least for now, those are encouraging words for the only living descendants of this ancient family of cloven-hoofed, horned mammals that began evolving millions of years ago.
"Antelope or goat"—or deer
From the pronghorn's first tentative identification in Mexico in the 16th century, until the end of the 20th, it was generally believed that the entire evolutionary history of the pronghorn took place in North America. However, in the early 1990s mitochondrial DNA sequencing reinforced the conviction held by increasing numbers of specialists, that antilocaprids, bovids, cervids and giraffids had all evolved separately from an unidentified common Eurasian ancestor that migrated across the Bering land bridge to North America during the early Miocene epoch. The radiation, or evolutionary diversification of the several genera and species of Antilocapridae is thought to have occurred over a period of 5 million years, between 23 and 28 million years ago, reaching the peak of their diversity some 7 million years ago.54 In the long run, from the time of Hernández until the second half of the 20th century, the long, persistent, and betimes uncertain fumbling over the genetic and morphologic identity of the pronghorn antelope may have been on track all along. Ultimately, the pronghorn may well prove to be part antelope, part goat, part deer, and part giraffe.55
Still, our "elegant speedster" remains perpetually poised to flee from the long-gone ghosts of predators such as the American cheetah, that may have made it what it has become–an enduring symbol of the romanticized American West's wide open spaces that are considerably less wide and free than they were before civilization took them over. While much more is known to us now than was apparent to Lewis and Clark and to the naturalists and zoologists of their time, it remains an enigmatic relic of the Miocene, bequeathed to us for safekeeping. And until Time calls "time!" on all pronghorn antelopes, William Clark's apostrophe will still describe it best: "They are all keenly made, and is beautiful"
Codicil: A musical name
". . . and the antelope play"
Beginning at about three weeks of age and continuing through their first summer of life, fawns spend much of their time in what appear to be boisterous, random frolics that were programmed into their genes millions of years ago. The secret forces of natural selection are merely preparing them for the vicissitudes of adult pronghorn survival. They practice changing directions on the run, making sharp turns or carving graceful arcs. They fidget, pivot, and hop from side to side. They experiment with stotting or pronking–springing into the air, all four legs at once–flexing their muscles to show imaginary pursuers that they are strong and healthy, and are not easy prospects for predators' appetites. They make the hairs of their manes stand stand erect, and flare their rump patches in mock alarm.56
Just as the colloquial "buffalo" trumps the official nickname "bison," and "elk" overrides the proper moniker "wapiti," "antelope" has stuck to the pronghorn in popular culture despite taxonomists' insistence that it is "not a real antelope." The upshot has been an onomastic compromise: Go ahead, call it a "pronghorn antelope" if you must, but remember, that by-name was contravened back in the 1870s in a popular song. In the refrain of Home on the Range, as well as the second line of the first stanza, and in the third line of the seldom sung fourth stanza–with its bold misstatements–"and the antelope flocks/That graze on the mountaintops green"–the animal is simply an antelope. "Pronghorn" may be anatomically correct, but it won't suit either the meter or the mood of that melody. There's something unpoetic and unmusical about it, whereas the light-footed, cantering lilt of "antelope" works both ways. So the pronghorn will be an antelope for as long as that song remains in the American memory as a catalog of our heartfelt sentiments about the Old West, concluding with those two shameless canards, "Where never is heard a discouraging word, and the skies are not cloudy all day."
As if a pronghorn could care, either way.
III. What Next?
As of January 30, 2012, it might have appeared that the two threads of this story had reached their mutual consummation. Google Images displayed 17,912 photos of pronghorns, and Flickr added nearly 8,000 more. WorldCat listed nearly 1,000 books about pronghorns both in and out of print, among them the comprehensive collection of essays and reports on the species' life-ways titled Pronghorn: Ecology and Management, by the late Bart W. O'Gara and his co-author Jim D. Yoakum, copiously illustrated by Edson Fichtler and Daniel P. Metz. It consists of 903 oversize pages, including a bibliography of books, articles, papers, theses and dissertations numbering more than 1,700 titles. It might seem that there is nothing more to be seen or said about Antilocapra americana Ord 1818. But the durable Miocene enigma is still alive and well, and it still challenges inquisitive minds.
1. The generic name Miracinonyx (mir-rah-see-nah-nix) refers to the cheetah's camouflage, black spots against a light-to-dark, yellowish tan background. The specific epithet inexpectatus (in-ex-spec=tah-toos), meaning "sudden surprise," underscores the benefit of its camouflage enables the cheetah to take its prey by surprise.
2. John A. Byers, Built for Speed: A Year in the Life of the Pronghorn, (Cambridge, Massachusetts: Harvard University Press, 2003), xvi, 16, 17. The genus Miracinonynx in the family Felidae–the cheetah, or cheetah-like cougar–was named by L. D. Martin in 1998. The Paleobiology Database, http://paleodb.org accessed 7 July 2010. Rothman's depiction, originally painted in acrylics, was commissioned by Sports Illustrated magazine in 1997. Source: Rothman Natural Science Illustration, Ridgefield, CT, http://www.michaelrothman.com
3. Bart O'Gara and Jim D. Yoakum, Pronghorn Ecology and management (Boulder: University Press of Colorado,2004), 27-39. As of the year of its publication, at 903 pages, this was the most comprehensive technical study of the pronghorn antelope ever undertaken.
4. John A. Byers, "Pronghorn," in George A. Feldhamer et al., Wild Mammals of North America, 2nd ed. (Baltimore: Johns Hopkins University Press, 2003), 998. Adrian Forsyth, Mammals of North America (Buffalo, New York: Firefly Books, 1999), 324-27. Mule deer (Odocoileus hemionus) live up to 10 years; white-tailed deer (Odocoileus virginianus) from 10 to 16.5 years; elk (Cervus elaphus) 14 to 26 years, moose (Alces alces) up to 27 years.
5. Stuart J. Feidel, "The Kennewick Follies: 'New' Theories about the Peopling of the Americas," Journal of Anthropological Research, Vol. 60, No. 1 (Spring 2004), 75-110. Use the search utility in the heading of this page to find other pages in which the "Old North Trail" is mentioned.
6. Three hundred and thirty-three different names from 219 Indian tribes–including two from the Aztec nation–are listed in Richard E. McCabe, Bart W. O'Gara and Henry M. Reeves, Prairie Ghost: Pronghorn and Human Interaction in Early America (Boulder: University Press of Colorado, 2004), 143-50. The authors did not translate of any of the Indian names into English, so it is impossible to tell how many duplicates there might have been, or whether any tribe called them anything like "pronghorn" or "antelope."
7. George Parker Winship, trans. and ed.,The Journey of Coronado, 1540-1542, from the City of Mexico to the Grand Canon of the Colorado and the Buffalo Plains of Texas, Kansas and Nebraska (New York: Allerton Book Company, 1922), 90.
8. Jean-Bernard Bossu, Travels in the Interior of North America, 1751-1762, translated and edited by Seymour Feiler (Norman: University of Oklahoma Press, 1962), 197. Briançon, which for centuries has been famous for its mutton and lamb, is a city in the RhÙne-Alpes region of southeastern France, not far from the Italian border.
9. George E. Dane and Francisco PalÛu, "The Founding of the Presidio and Mission of Our Father Saint Francis," California Historical Society Quarterly, Vol. XIV, No. 2 (June 1935), 104.
10. Diccionario Enciclopédico de la Lengua Castellana (Paris: Casa Editorial Garnier Hermanos, 1900), s.v. berrendo.
11. Edward Umfreyville, The Present State of Hudson's Bay (London: Charles Stalker, 1790), 165-166. Clearly embarrassed over his lack of knowledge about the species, Umfreyville launched into one of his frequent complaints about the Company: "Without a doubt, if the inland parts [of Canada] were explored by a person of ingenuity, many useful discoveries might be made in every branch of Nature's operations, as well relating to the vegetable as the animal world; but unfortunately those mercantile gentlemen who have hitherto been sent into the Terra incognita have been so very intent upon the pecuniary emolument, arising from the trade they are engaged in, as intirely to neglect every effort to obtain a knowledge of the country and its productions; though such an an undertaking would be attended with little or no expence, and would certainly redound much to their honour, if not to their profit."
12. McCabe et al. (see note 5) list 13 Cree names for it from east-central Saskatchewan and south-central Alberta, without translating any of them. Phonetically, the nearest of them to Umfreyville's expression is apeestat-choekoos, from Alberta.
13. Biddle's paraphrase, prepared for the press by Paul Allen, was titled History of the expedition under the command of Captains Lewis and Clark, to the sources of the Missouri: thence across the Rocky mountains and down the river Columbia to the Pacific ocean; performed during the years 1804-5-6; by order of the government of the United States (Philadelphia: Bradford and Inskeep, 1814).
14. In April of 1810, Clark explained to Nicholas Biddle the best method for hunting pronghorns: "[W]hen they first see the hunter they run for½ mile ahead in a straight line%mdash;if [the hunter] lies on his belly & lifts up his hat, arm, & foot, the animal curios to see the object, returns on a trot, & sometimes goes & comes 2 or 3 times, till within reach of the rifle." Biddle, anticipating the promised volume about natural history discoveries on the expedition, refrained from mentioning it in his paraphrase of the captains' journals. Needless to say, the strategy soon became well-known western folklore; eventually it was outlawed as unsportsmanlike. Jackson, Letters, 2:539.
15. Noah Webster, in his 1806 Compendious Dictionary of the English Language, defined sheepseye as "a loving sly look, an oblique view." The name antelope has been said to have a similar connotation; the French naturalist and zoologist Georges Cuvier (1769-1832) identified it as a corruption of Antholops, a word found in the writings of Eustathius, a famous orator of the early 4th century, in Cappadocia, a region of central Turkey. It is thought to have referred to the beautiful eyes of the Mid-Eastern gazelle. Georges Cuvier, The animal kingdom arranged in conformity with its organization, translated by H. M'Murtrie, 4 vols. (New York: Carvill, 1831), I:191n. From time to time throughout recorded history the doe-eyed "who, me?" look has been fashionable among young women.
16. George J. Mitchell, "Measurements, Weights, and Carcass Yields of Pronghorns in Alberta," The Journal of Wildlife Management, vol. 35, no. 1 (January 1971) 76-86. Bill Jensen, "Estimating Big Game Weights," North Dakota Outdoors, (September-October 2000), Table 2. O'Gara, ibid., p. 2. Weights vary slightly among the five subspecies, (A. a. americana Ord, 1815; A. a. mexicana Merriam, 1901; Nelson, 1912; A. a. oregona Bailey, 1932; and A. a. sonoriensis Goldman, 1945), and also vary seasonally.
17. A New and Complete Dictionary of Arts and Sciences Comprehending All the Branches of Useful Knowledge London: Printed for W. Owen, 1754), s.v. gazella.
18. David McDonald, ed., The Encyclopedia of Mammals (New York: Facts on File, 1984), 544, 548-51.
19. Bernardo de Sahag˙n, Earthly Things, Book 11 in C. E. Dibble and R. J. O. Anderson, eds., Florentine Codex (Salt Lake City: University of Utah Press, 1963). Gerald J. Rosenzweig, "Jean Louis Berlandier Papers, 1826-1851, and related papers to 1886," Smithsonian Institution Archives, Record Unit 7052, Series 7, Folder 30. http://siarchives.si.edu/findingaids/FARU7052.htm (accessed 2 July 2010).
20. Francisco Hernández, Nova Plantarum, Animalium et Mineralium Mexicanorum Historia (New plants, animals and minerals of Mexico), ed. Nardo Antonio de Recchi; 4 vols. (Rome: V. Mascardi for B. Deversini and Z. Masotti, 1651), 324-25. The supposed connections between the Teuthlalmaçame and the pronghorn antelope were first asserted in the widely circulated manuscripts of Jean Louis Berlandier (ca. 1805-1851), and published in The Mammals of North America, by Spencer Fullerton Baird (1823-1887), Vol. 2 (1857), p. 666; cited in the 1894 reprint of George Ord's North American Zoology, Appendix, p. 23 (Internet Archive, from the Marine Biological Laboratory, Woods Hole Oceanographic Institution; accessed 14 July 2010). The same were repeated in Edward Richard Alston et al., Biologia Centrali-Americana: Mammalia (1879-1882), 113.
21. Discoveries made in exploring the Missouri, Red River and Washita by Captains Lewis and Clark, Doctor Sibley, and William Dunbar, and compiled by Thomas Jefferson, Natchez, 1806; facsimile edition, ed. Doug Erickson, Jeremy Skinner, and Paul Merchant (Spokane, Washington: Arthur H. Clark, 2004), 255. Dunbar's comment got additional exposure in the surreptitious compilation of pirated gems titled New Travels Among the Indians of North America published in 1812 by the shadowy William Fisher. Stephen Dow Beckham, The Literature of the Lewis and Clark Expedition (Portland, Oregon: Lewis & Clark College, 2003), 125-126, 138-140.
22. C. S. Rafinesque, ed., "Descriptions of seven new genera of North American Quadrupeds," The American Monthly Magazine and Critical Review, Vol. 2, No. 1 (November 1817). American Periodicals Series Online, p. 40, accessed 12 July 2010. As a young man, George Ord worked in his father's rope-making business until he left it at age 25 to devote his life to science, principally ornithology. Ord was a close friend of ornithologist Alexander Wilson, as well as entomologist Thomas Say and illustrator Charles Alexandre Lesueur.
23. George Bird Grinnel, "Names of the White Goat," Forest and Stream, Vol. 33, No. 20 (December 5, 1889), 383-84. Grinnell quoted Baird's statement from the Pacific Railroad Reports of 1857, p. 663: "The generic name of Mazama, as established by Rafinesque, . . . has been quoted by some authors for the American antelope, as well as for the mountain goat and the smaller deer. An examination of his diagnosis will show very satisfactorily that the name cannot be used at all, on account of its embracing too many incongruous elements."
24. The Philadelphia Medical and Physical Journal, 5 March 1806, p. 194; American Periodicals Series Online, accessed March 3, 2010. Barton, who was then a professor of medicine at the University of Pennsylvania, founded this journal in 1804, and "collected and arranged" its contents for the next five years. Subjects ranged from medicine (especially yellow fever) to natural science, geography, agriculture, archaeology and, as in this instance, "Miscellaneous Facts and Observations."
25. Jefferson to William Bartram, April 7, 1805. Library of Congress, The Thomas Jefferson Papers, Series 1; General Correspondence, 1651-1827, Image 131. Jefferson's letter was in response to Bartram's offer of a horn of an African antelope for Jefferson's collection. Along with that item, Bartram mentioned that he also had the horn of a deer ("Cervi") from northeastern Canada that was "small & remarkable in being so flat & thin," which suggests it might actually have belonged to a pronghorn. William Bartram to Jefferson, "Zoological Notes," n.d., 1805. Library of Congress, The Thomas Jefferson Papers Series 1, General Correspondence.
26. The trip down the Missouri River to St. Louis took 45 days. At St. Louis it was transferred to another river barge and taken to New Orleans, then via brig USS Comet to Baltimore, then by wagon to Washington City, where it arrived on August 12, 1805, 18 weeks and one day after leaving Fort Mandan.
27. Jefferson to Charles Willson Peale, October 6, 1805. Jackson, Letters, 1:260-61. Jefferson was correct concerning the roe deer (Capreolus capreolus—"little goat; little goat"—also known as the chevreuil (French for roe deer), and its antlers. The roe is native to Western Europe. The cause of the confusion was that the captains had sent two sets of mule deer antlers, of which at least one apparently had been placed in the wrong box during one of the unpackings. There is no explanation of those several disturbances of the shipment, but the captains may have instructed Corporal Warfington to open them periodically to see that everything was dry and free from insects.
28. Jefferson to Charles Willson Peale, October 9, 1805. Jackson, Letters, 1:263. The chamois (Rupicapra rupicapra) belongs to the family Bovidae, in the goat-antelope subfamily, which is divided into two species and eight subspecies. They are not plains-dwellers like pronghorn, but are at home among the highest mountains in central and southern Europe, the Balkans, parts of Turkey, and the Caucusus. In the Swiss and Austrian Alps they are commonly known as gemse (beginning with a hard g as in "good," not a j, and ending with an unaccented short e—GEM-seh). They are mostly medium to dark brown in color, and often have white under-jaws and throats. Because properly cured skins will absorb and hold water, they were once popular for washing carriages, and later, automobiles. So-called "chamois" or "shammy" skins today are synthetic.
29. Peale's "Dermest" was either a member of the large (500 to 700 species worldwide) family of beetles called Dermestidae, commonly known as hide-, leather-, or carpet-beetles, or a similarly large family of Coleoptera called skin beetles (Trogidae), which are drawn to dried skin of dead animals. Gordon Gordh and David Headrick, A Dictionary of Entomology (Wallingford, Oxon, UK: CABI Publishing, 2001), 270, 945-46.
The long, well documented story of the shipment's 18-week, 4,000-mile transit by river barges, the sailing vessel Comet from New Orleans to Baltimore, thence overland to Washington City, is related by Paul Russell Cutright in Lewis & Clark: Pioneering Naturalists (Lincoln: University of Nebraska Press, 1969), 375-381.
30. Charles Willson Peale to Jefferson, November 3, 1805. Jackson, Letters, 1:268.
31. C. W. Peale to Jefferson, April 5, 1806, Jackson, Letters 1:302-03.
32. Bart W. O'Gara, "Physical Characteristics," in Pronghorn Ecology and Management, 129.
33. Jefferson to Alexander von Humboldt, December 6, 1813. Jackson, Letters, 2:596. In 1893 a new edition of Biddle's version appeared, with "copious critical commentary" by Elliott Coues (pronounced "cowz"), including numerous amplifications of Biddle's references to plants, animals, minerals, and geographical features. There was an index to Biddle's two volumes at long last, but only a very few additions from the original journals, which had recently come to light and were in the process of editing for publication by Reuben Gold Thwaites.
Worst of all was the failure to complete the validation of Clark's final map with trigonometric solutions to all of the celestial observations for longitude that Lewis acquired en route. After all, that was "the main object of the expedition." Jefferson to José Corrèa da Serra, Jackson, Letters, 2:618.
34. Nicholas Biddle was not a naturalist, and it is not known who might have recommended that he transcribe all of Lewis and Clark's tentative names as "antelope, with "antelope or goat" occurring only once in his paraphrase of the journals.
35. See Paul S. Martin and Christine R. Szuter, "War Zones and Game Sinks in Lewis and Clark's West," Conservation Biology, Vol. 13, No. 1 (February 1999), 36-45.
36. Harlin M. Fuller and LeRoy R. Hafen, eds., The Journal of Captain John R. Bell, Official Journalist for the Stephen H. Long Expedition to the Rocky Mountains, 1820 (Glendale, California: Arthur H. Clark, 1957), 127.
37. Kenneth Haltman, Looking Close and Seeing Far: Samuel Seymour, Titian Ramsay Peale, and the Art of the Long Expedition, 1820-1823 (University Park: Pennsylvania State University Press, 2008), 110.
38. John Davidson Godman, American Natural History, 3 vols (Philadelpphia: Carey, Le Carey, 1828), 2:321-25. William Guthrie, A New Geographical, Historical, and Commercial Grammar, and Present State of the Several Kingdoms of the World, 2 vols. (Philadelphia: Johnson & Warner, 1815), 2:166. Sir John Richardson (1787-1865), Fauna Boreali-Americana: or, The Zoology of the Northern Parts of British America, Containing Descriptions of the Objects of Natural History Collected on the Late Northern Land Expedition, Under Command of Captain Sir John Franklin, R.N. (London: J. Murray, 1829-37)
39. John James Audubon and The Rev. John Bachman, The Viviparous Quadrupeds of North America. 3 vols. (New York: F. G. Audubon, 1851), 2:195-96.
40. Abraham Dee Bartlett (1812-1897), who was superintendent of the London Zoo from 1859 to 1897, had suggested "prong-buck" as a common name in "Remarks upon the affinities of the prongbuck (Antilocapra americana)." . . . in Proceedings of the Zoˆlogical Society of London, 1865. Bartlett may have borrowed the name from Charles Hamilton Smith, who had identified, merely on the basis of a fragment of a skull he saw in a Philadelphia museum (Peale's?), a fourth species of American antelope, which was reportedly to be found in New Jersey, where descendants of Dutch colonists called it a spring-bok, which became spring-back in English. Smith also proposed a binomial—Antilope temamazama.
41. John Dean Caton, Antelope and Deer of America: a comprehensive scientific treatise upon the natural history; Including the characteristics, habits, affinities, and capacity for domestication of the Antilocapra and Cervidae of North America (New York: Hurd and Houghton, 1877), 24.
42. Ibid., 35.
43. Ibid., 41.
44. George J. Mitchell, "Measurements, Weights, and Carcass Yields of Pronghorns in Alberta," The Journal of Wildlife Management, Vol. 35, No. 1 (January 1971), 82. Allowance has been made for possible regional differences in whole weights.
45. McCabe, O'Gara and Reeves, p. 19; Table 1.
46. R. A. Field et al., "The Pronghorn Antelope Carcass," University of Wyoming Agricultural Experiment Station, B-565R (August 2003).
47. Allen Grant Wallihan and Mary Augusta Higgins Wallihan, Hoofs, Claws and Antlers of the Rocky Mountains (Denver: F.S. Thayer, 1894), Antelope, Plate 6, "Unconsciousness."
48. Bart O'Gara and Susan Kraft Ball, Chapter 7, "Skeletal and Visceral Anatomy," in O'Gara & Yoakum, Pronghorn Ecology and Management, 211-212.
49. David W. Kitchen, Social Behavior and Ecology of the Pronghorn, in Wildlife Monographs, No. 38 (August 1974), 3-96.
50. Caton, p. 57.
51. Ellliott Coues, ed., The History of the Lewis and Clark Expedition. 3 vols. (New York: Dover, ), 3:849-50, note 50.
52. O'Gara and Yoakum, 408.
53. M. Hoffmann, J Byers & J. Beckmann, Antilocapra americana, in the IUCN Red List of Threatened Species, at http://www.iucnredlist.org/apps/redlist/details/1677/0 Accessed June 3, 2010.
54. O'Gara and Yoakum, 110, 139, 273.
55. The real but hidden genetic connections between the pronghorn and the giraffe are described by Bart O'Gara and Christine Janis in "Scientific Classification," Chapter 1 of O'Gara and Yoakum, Pronghorn, 5.
56. O'Gara, Mammalian Species, No. 90, p. 6.
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