FOUR
Joining the Guild
of Carnivores:
The Benefits of
Membership
A Guild of Carnivores shadows the big mammals of the
Pleistocene throughout Eurasia, living by tooth and
claw. Wild dogs and wolves choreograph the hunt.
The Late Pleistocene was the crowning age of big mammals and the “big hunt,” a final bash before glaciers retreated, and, like dinosaurs before them, giant warm-bloods toppled into the trash bin of extinction. That happened when the world’s warmth-loving plants and animals were spreading out to all continents and islands—except the big ice patch that is Antarctica. As yet there is no consensus as to the question of whether the extinct animals fell to human hunters overwhelmed with a bloodlust not seen in them before, or, boxed in by advancing forests, failed to slingshot into the future off the latest climatic gyration in an epoch marked by wild swings of global weather.
Whatever reasons finally account for it, the end of the Pleistocene some ten to twelve thousand years ago brought a massive dying of the largest terrestrial mammals—mastodons, wooly mammoths, rhinoceroses, giant deer, long-horned bison, aurochs, musk ox, horses and reindeer, short-faced bears, two-ton glyptodonts, giant Irish elk, hippopotamuses, and hulking vegetarian cave bears, among them. Some of them for hundreds of thousands of years had grazed their migratory routes deep into steppes and tundra, grooved mountain passes. Their constant companions in life and death were the carnivores—including giant spotted cave-skulking hyenas, smaller spotted hyenas, dirk cats, scimitar cats, jaguars, panthers, lions, tigers, cheetahs, raptors and huge scavenging condors, dire wolves, wild dogs, dholes, wolves, and Neanderthal (the big bipedal meat eater).
Excluding small cats and foxes, those animals formed the Guild of Carnivores, the collection of bone crushers, slicers and dicers, stalkers, slashers, bushwhackers, pouncers, scavengers, and long-distance pursuit artists who shadowed the herbivorous ungulates on their migrations. The big cats sat atop the Guild. Nothing could match them one on one, yet some of them, like the scimitar cat and dirk cat, were so highly specialized that they had virtually no ability to adapt to dramatically changing conditions. Nor could the hyenas, prime scavengers of Pleistocene encampments for thousands of years. who seemed unable to alter their behavior or temperament in response to exposure to several different species of early humans—although there is no evidence that any human group tried to befriend them.
This period in the world’s history is a tumultuous one filled with various animals unknown in the world and barely recognizable today, but which a few adaptations of nomenclature might bring into sharper focus. Following the emerging convention, I use hominin to refer specifically to members of the genus Homo who rank as our most direct ancestors but are not of our species. I also sometimes refer to members of all human species as naked bipeds, to keep them in their natural context as members of the Guild of Carnivores, as another species of hunters who nonetheless were different from other animals.
The naming system Swedish botanist Carl von Linné established in the eighteenth century to bring uniformity and order to the babel of names for the various representatives of life on Earth is essentially binomial. Each organism is assigned a species and genus name by which it is formally known, and subspecies and specific population designations are added when necessary to recognize unique genetic, ecological, or behavioral characteristics of a smaller group that might be on its way to forming a new species. Genus and species names lie at the base of a hierarchical chain of categories and subcategories. The system is old and clunky and favors splitters—people who want to name a new species every time they find individuals that are different in appearance from the description for that species, even if the description is of only one or two individuals. That is the way dog breeds are sometimes identified, but it is a narrow, restrictive view, especially when applied to wide-ranging species.
Despite its flaws, taxonomy provides uniform descriptions and definitions that allow people in different parts of the world to communicate in a meaningful fashion. Here are the taxonomies (only major categories provided) of three members of the Guild of Carnivores: cave hyena, scimitar cat, and Arabian wolf, the only generalist in the group and the only one of the three to survive the Late Pleistocene extinction, is categorized as follows:
Class: Mammalia
Order: Carnivora
Family: Hyaenidae
Genus: Crocuta
Species: C. crocuta
Subspecies: C. c. spelaea
The scimitar cat is:
Class: Mammalia
Order: Carnivora
Family: Felidae
Genus: Homotherium
Species: H. latidens
The Arabian wolf, a subspecies of the gray wolf, is:
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species: C. lupus
Subspecies: C. l. arabs
The order Carnivora has 23 families with 118 genera and 287 extant species. Families include Ursidae (bears), Mustelidae (weasels), Proccyonida (raccoons), Ailuropodas (pandas), Otariidae (eared seals), Odobenidae (walruses), Phocidae (earless seals), Felidae (cats), Viverridae (civets), Hyaenidae (hyenas), and Canidae. The Family Canidae is comprised of foxes (Vulpes); South American canids; the maned wolf and bush dog, which are not included with other South American canids; a group of ancient fox lineages made up of the island-gray fox, the gray fox, the bat-eared fox, and the raccoon dog; and the wolflike canids.
Carnivorans, as members of the order Carnivora are known in order to distinguish them from carnivores of different orders, like Homo sapiens, have binocular vision and specially adapted carnassials—the last premolar in the upper jaw and first molar in the lower jaw—like for shearing and tearing skin, muscle, tendons, and bone. They are possessed of a minimum of four and a maximum of five toes (the fifth toe on the forefeet of many is the vestigial dewclaw), relatively large, thick-boned heads with generally well-protected brains, and a pronounced preference for meat, although the giant panda is a bamboo-chewing vegetarian, who only occasionally indulges in eggs, insects, and fish; and raccoons and bears, except the polar, are omnivores. Cats and dogs walk on their toes; bears go heel first; and seals, sea lions, and walruses waddle with their flippers in lieu of walking. Carnivorans teach their off-spring how to find, kill, and protect the food they need to survive. The absence of such teaching has long been a major impediment to the success of captive breeding programs intended to reintroduce predators to the wild. Those animals did not have a clue how to survive because no one had taught them.
The catlike and doglike carnivorans are believed to have originated some forty to fifty million years ago in North America as miacids—ferret- to fox-sized animals with distinctive carnassials who gave way to larger five-toe hyenas (creodonts). Around thirty-five million years ago, Hesperocyon walked on its toes to found the Hesperocyoninae subfamily of Canidae and spawn two more subfamilies, Borophaginae and Caninae, both of which manifest a distinctive and crucial mutation in the lower carnassials that makes them versatile enough to rend flesh, tendon, and bone, or to grind fruits and vegetables. The Borophaginae remained in North America, crunching their way to extinction, while the Caninae went walking. From Hesperocyon also came the long-nosed Leptocyon, who gave rise to foxes five million years ago.1
The true dogs began with Eucyon five to seven million years ago, contemporaneous with the secession of South American canids, among which there were a few “hypercarnivores.” Weighing more than forty-five pounds, the hypercarnivores of the Pleistocene rose and fell in population and territory according to the rhythms of glaciations, which affected the amount of prey available to fuel their appetites. They were creatures of the cold because of the huge ungulates it supported. Eucyon headed for Asia and Europe and down into China, creating along the way the first Holarctic explosion of wolflike species.
Canis appeared in the form of C. ferox and C. lepophagus five to six million years ago in North America and almost immediately headed west and south, crossing Siberia and coming off the steppes, the same way Mongol horsemen would slingshot into Europe millions of years later. Canis burst upon Eurasia at the end of the Pliocene and, mixing, matching, radiating new species in all directions, joined the Guild of Carnivores.
Prehistory is a sand painting subject to constant revision by scholars, fossil hunters, and time itself. A great mystery revealed one day is forgotten the next. A paucity of physical evidence at many sites—for many periods, a shortage of sites—makes all species counts and time-lines provisional. In a recent revision, the start of the Pleistocene was moved from around 2 to 2.58 million years ago to include all the major glacial events of the last Ice Age. Because of the persistent and insistent cleaving of humans and human activities from natural events, geological time—in this case, the Pleistocene—covers natural events and all organic activities, except those related to humans, which are considered to have occurred in the Paleolithic, or Old Stone, Age. A major problem with this approach becomes apparent every time a natural disaster or an environmental catastrophe affects all life and reshapes the landscape. In an attempt to avoid confusion, I will use Pleistocene throughout, except in those cases that absolutely demand use of Paleolithic.
Many species of wolves rose and fell before the Pleistocene yielded to our own era, the Holocene, around twelve thousand years ago, most notoriously Canis dirus, the dire wolf, a hypercarnivore among hypercarnivores, and a whole group of previously unknown wolves recently unearthed in Alaska that liked to crunch bones of horses and bison. They are a powerful reminder that in many cases, we do not have a clue how much of the story of life we do not know, much less how its discovery would alter what is known. Other significant animals might still be buried in the permafrost or submerged with the ancient coast awaiting discovery and excavation, or lying among the unnamed fossil fragments in a long-forgotten collector’s neglected cabinet of natural history.
But based on what is known, for wolves—and for hominins—the last third or more of the Pleistocene was a time of diminishing species diversity and pulsating demographic expansions and contractions in response to the dance of glaciers. Nonetheless, the main canid species we know today slipped into place long before the Pleistocene ended: Canis adustus (side-striped jackal), C. aureus ( golden jackal), C. lupus (gray wolf), C.s latrans (coyote ), C. mesomeias (black-backed jackal), C.s simensis (Ethiopian wolf), and, perhaps, Canis rufus (red wolf). Scientists in India recently suggested that their indigenous wolves—C. lupus pallipes and C. chanco—played no role in the creation of the dog, despite a strong supposition among many paleontologists that both might have been involved, and also differed enough from the gray wolf to deserve separate species status.2
All the extant wolflike canids have 78 chromosomes and could be classed in the genus Canis, but two are not: the superpack-forming red dhole (Cuon alpinus) of India and Asia, and the patchwork-coated African wild dog (Lycaon pictus). They are assigned their own genera largely because of habit and peculiarities with their feet—they lack the vestigial fifth toe, and the middle two toes are fused, a characteristic that turns up in Canis lupus arabs as well. The dhole, like the wolf, was a member in good standing of the Pleistocene Guild of Carnivores, as was a larger wild dog, Xenocyon lycaonoides, also counted as Canis lycaonoides. For some time on both sides of a million years ago, X. lycaonoides and a smallish wolf, Canis mosbachensis, appear in tandem at caves in Spain, Italy, Greece, Germany, and the Levant. The reasons for this are unknown, since lycaonoides was primarily a courser of the steppes, not a scavenger of caves. The wild dog decamped for Africa around 700,000 or 800,000 years ago and is there still in the different, smaller form of Lycaon pictus, but until then it seemed to cap the size of prey that C. mosbachensis pursued at deer.3
Whether precursors of Canis lupus or evolutionary dead-ends, the various wolves that rose and vanished during the first million-plus years of the Pleistocene were, with the exception of North America’s dire wolf, not the biggest dogs in the Guild of Carnivores. Paleontologists call them mesopredators, or middlin’, down at the bottom of the list in terms of size and thus, according to modern scientific calculations, too small to bring down game much bigger than a deer. Certainly, that appears initially to be the situation with C. mosbachensis, who would have had to compete with larger carnivorans, including its running mate, Xenocyon lycaonoides, for bigger game. Often mentioned but not much discussed or described, C. mosbachensis is variously referred to as the size of a small wolf, like the Arabian wolf, or of a coyote or a medium-sized dog.
Once Xenocyon lycaonoides dropped into Africa, Canis mosbachensis turned to larger game and began to grow and evolve into lupus— or so runs one theory. Similar arguments are made today concerning deer-eating the coyote or coyote-wolf hybrid that is colonizing the Eastern United States. The gray wolf was different from the start, a leggy, wide-ranging, devoted carnivore with catholic tastes who nonetheless showed a pronounced preference for bison in England, reindeer in northern Europe, and reindeer and horse on the Mammoth Steppe—predilections it shared with hominin hunters. C. lupus had the added advantage of being highly adaptable through changes in shape and behavior to a range of ecosystems. Subspecies include the 24- to 50-pound Arabian wolf who tends to hunt singly or with one or two partners, when it is not scavenging dumps, and the 150-pound tundra wolf running in packs of twenty or more, pursuing the downsized megafauna that survived the Last Glacial Maximum.
What distinguished wolves, dholes, and wild dogs behaviorally—and still does—was their pronounced, we might even say exaggerated, ability, relative to other carnivorans, to run and attack together. Spotted hyenas also hunt in packs but they are not renowned for their social cohesion or toleration for long hunts. The wolflike canids were constantly aware of the position of everyone in the pack, spatially and temporally, as if they projected themselves into spatial comprehension. Pack-running hounds do get lost, usually because they surrender to their noses and eagerness in pursuit and as a result forget to recalibrate their positions. Once contact is lost, other demands take charge. But anyone who has hiked with a dog in the woods understands the phenomenon of the disappeared animal who suddenly materializes at their side and looks at them as if to say, “I knew where you were all along.” The ability of wolves to communicate over distance and act in concert amplified their strength, so that together they could drive all but the largest, most obstinate bears from their prey, or bring down animals considerably larger than themselves, like moose or giant elk. Wolves also adapted their hunting style to suit the prey a capacity that helped them through rough times.
Gray wolf