Pronghorns

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).

Species History

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-riddee, “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.

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 pronghorn, 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.

The pronghorn that we know owes its survival to its principal 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 members 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 eagle 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 awesome 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]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 … Continue reading 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.

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.^[5]O’Gara and Yoakum, 110, 139, 273. 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.^[6]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 … Continue reading

Spanish Descriptions

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.^[7]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 … Continue reading 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.^[8]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: … Continue reading

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 whatever 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.”^[9]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: … Continue reading 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.”^[10]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 … Continue reading 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.^[11]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.

Berrendos, a Spanish word meaning “two-colored,” is still the everyday Mexican name for Antilocapra americana.^[12]Diccionario Enciclopédico de la Lengua Castellana (Paris: Casa Editorial Garnier Hermanos, 1900), s.v. berrendo.

British Descriptions

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.^[13]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 … Continue reading

He also mentioned its Cree Indian name, “Apis-to-chik-o-shish,” although he felt no compulsion to render it in English.^[14]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 … Continue reading

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? Meriwether Lewis and William Clark were to produce only a few answers, and many of those fell through the cracks of history for a considerable time when both 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.^[15]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 … Continue reading

Clark’s Description

On 3 September 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.

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.

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?^[16]In April of 1810, Clark explained to Nicholas Biddle the best method for hunting pronghorns: “[W]hen they first see the hunter they run for ½ mile ahead in a straight line—if [the hunter] … Continue reading

Owen’s Dictionary

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]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.

Lewis’s Description

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” (see above).

In his own journal for 14 September 1804, 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.”^[18]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 … Continue reading 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.^[19]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, … Continue reading 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.”

Jefferson’s Question

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.”^[20]Jefferson to William Bartram, 7 April 1805. Library of Congress, The Thomas Jefferson Papers, Series 1; General Correspondence, 1651-1827, Image 131. Jefferson’s letter was in response to … Continue reading

Jefferson was of course unaware that on that very same day, 7 April 1805, Lewis had shipped to him via the returning barge (called the ‘boat’ or ‘barge’ but never the ‘keelboat’) 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.^[21]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 … Continue reading 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.”^[22]Jefferson to Charles Willson Peale, 6 October 1805. Jackson, Letters, 1:260-61. Jefferson was correct concerning the roe deer (Capreolus capreolus—”little goat; little … Continue reading

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.”^[23]Jefferson to Charles Willson Peale, 9 October 1805. Jackson, Letters, 1:263. The chamois (Rupicapra rupicapra) belongs to the family Bovidae, in the goat-antelope subfamily, which is divided into two … Continue reading

Peale’s Answer

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.”^[24]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 … Continue reading

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.^[25]Charles Willson Peale to Jefferson, 3 November 1805. Jackson, Letters, 1:268.

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.^[26]C. W. Peale to Jefferson, 5 April 1806, Jackson, Letters 1:302-03.

Biddle’s Version

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 History^[27]Jefferson to Alexander von Humboldt, 6 December 1813. Jackson, Letters, 2:596. In 1893 a new edition of Biddle’s version appeared, with “copious critical commentary” by Elliott … Continue reading 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 (28 August 1806) to have witnessed just such a scene during his time at the Great 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.^[28]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 … Continue reading

Selected Encounters

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.

• 17 September 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.”
• 20 September 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.”
• 17 October 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.”
• 14 April 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.”
• 25 July 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.”

Indian Hunting Methods

Biddle selected several accounts of Indians’ various ways of hunting the pronghorns:

• 16 October 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.”
• 5 November 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.”
• 14 August 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.”

Near the Yellowstone

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.

• 22 April 1805 (Lewis): “Large herds of deer, elk, buffalo, and antelopes in view of us.”
• 26 April 1805 (Lewis): “Elk, antelope, and buffalo suffered [Lewis] to approach them without alarm, and often followed him quietly for some distance.”
• 30 April 1805 (Lewis, 29 April 1805): “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.”
• 5 May 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.”
• 26 June 1805 (Lewis): “Females with their young . . . generally two in number, and the males by themselves.”
• 29 August 1806 (Clark): “We observe that the greatest quantities of wild animals are usually found in the country lying between two nations at war.”^[29]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.

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 5 September 1804, 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.

Forgotten for 100 Years

All of the foregoing details, and many more, were in the original manuscript journals. At Fort Mandan on 5 May 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.

21st-century Taxonomy

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.^[30]David McDonald, ed., The Encyclopedia of Mammals (New York: Facts on File, 1984), 544, 548-51.

Characteristics

In the absence of collarbones, the pronghorn’s scapulas, or shoulder blades, are linked to the skeleton by weight-bearing muscular attachments,^[32]Bart O’Gara and Susan Kraft Ball, Chapter 7, “Skeletal and Visceral Anatomy,” in O’Gara & Yoakum, Pronghorn Ecology and Management, 211-212. 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. 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.

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.^[33]David W. Kitchen, Social Behavior and Ecology of the Pronghorn, in Wildlife Monographs, No. 38 (August 1974), 3-96.

Horns

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, and are shed soon after mating is over in early autumn. Regrowth begins almost immediately.

Four Subspecies

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.

Recovery

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.”^[34]Ellliott Coues, ed., The History of the Lewis and Clark Expedition. 3 vols. (New York: Dover, [1893]), 3:849-50, note 50. 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.^[35]O’Gara and Yoakum, 408.

Thus by 21st-century standards pronghorns are classified as being of “least concern” on the IUCN Red List of Threatened Species.^[36]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 3 June 2010. 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.

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”

The Playful “Antelope”

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.^[37]O’Gara, Mammalian Species, No. 90, p. 6.

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.

As of 30 January 2012, 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.