carnivore

—carnivoral /kahr niv"euhr euhl/, adj.

/kahr"neuh vawr', -vohr'/, n.

1. an animal that eats flesh.

2. a flesh-eating mammal of the order Carnivora, comprising the dogs, cats, bears, seals, and weasels.

3. an insectivorous plant.

[1850-55; < L carnivorus CARNIVOROUS]

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Any meat-eating animal, but especially any member of the order Carnivora, consisting of 12 families of primarily predatory mammals: Canidae (e.g., dogs), Ursidae (bears), Procyonidae (raccoons), Mustelidae (weasels), Mephitidae (skunks), Viverridae (civets), Herpestidae (mongooses), Hyaenidae (hyenas), Felidae (cats), Otariidae and Phocidae (seals), and Odobenidae (the walrus).

Though most carnivores eat only meat, some rely heavily on vegetation (e.g., the panda). Most have a complex tooth structure and a lower jaw that can move only vertically but can exert great power. The earliest carnivores, which probably evolved from an insectivorous ancestor, appeared during the Paleocene Epoch (about 65–55 million years ago). Carnivores are highly intelligent.

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▪ mammal order

Introduction

 any member of the mammalian (mammal) order Carnivora (literally, “flesh devourers” in Latin), comprising more than 270 species. In a more general sense, a carnivore is any animal (or plant; see carnivorous plant) that eats other animals, as opposed to a herbivore, which eats plants. Although the species classified in this order are basically meat eaters, a substantial number of them, especially among bears and members of the raccoon family, also feed extensively on vegetation and are thus actually omnivorous.

 The order Carnivora includes 12 families, 9 of which live on land: Canidae (canine) (dogs (dog) and related species), Felidae (feline) (cats), Ursidae ( bears), Procyonidae (procyonid) ( raccoons and related species), Mustelidae (mustelid) ( weasels, badgers, otters, and related species), Mephitidae ( skunks and stink badgers), Herpestidae ( mongooses), Viverridae (viverrid) ( civets, genets, and related species), and Hyaenidae ( hyenas). There are three aquatic families: Otariidae ( sea lions and fur seals), Phocidae (true, or earless, seals), and Odobenidae (the walrus). These aquatic families are referred to as pinnipeds.

Importance of Carnivora

  Two carnivores are probably the animals most familiar to people: the domestic dog and cat (cat, domestic), which are both derived from wild members of this order. On the other hand, various bears, felines, canines, and hyenas are among the few animals that occasionally attack humans. These large, dangerous carnivores are often the objects of hunters, who kill them for display as trophies. Most luxurious natural furs (fur) ( ermine, mink, sable, and otter, among others) come from members of Carnivora, as do many of the animals that attract the largest crowds at circuses and zoos. Producers of livestock worldwide are concerned about possible depredations upon their herds and flocks by this group of mammals.

  Being meat eaters, carnivores are at the top of the food chain and form the highest trophic level within ecosystems. As such, they are basic to maintaining the “balance of nature” within those systems. In areas of human settlement, this precarious balance has frequently been upset by the extermination of many carnivores formerly considered undesirable because of their predatory habits. Now, however, carnivores are recognized to be necessary elements in natural systems; they improve the stability of prey populations by keeping them within the carrying capacity of the food supply. As a result, the surviving animals are better fed and less subject to disease. Many of these predators dig dens and provide burrows in which other forms of wildlife can take refuge. Digging also results in the mixing of soils and the reduction of water runoff during rains. The carnivores best known for their burrow building are badgers and skunks, but bears, canines, and felines regularly engage in this behaviour as well.

 Carnivore numbers are limited by food, larger predators, or disease. When human influence removes larger predators, many of the smaller carnivores become extremely abundant, creating an ideal environment for the spread of infection. The disease of most concern to humans is rabies, which is transmitted in saliva via bites. Rabies is most common in the red fox, striped skunk, and raccoon, but it also occurs in African hunting dogs and can infect practically all carnivores. Billions of dollars are spent annually throughout the world to manage and control the incidence of this disease. In some countries, abundance of vector species, especially red foxes, is controlled by culling or by dropping vaccine-laden bait from the air. In other countries, programs of “capture-vaccinate-release” are in place to reduce the vulnerability of individual animals. Other infectious diseases carried by carnivores and of concern to humans include canine distemper, parvovirus (canine parvovirus disease), toxoplasmosis, and leptospirosis.

Behaviour

  Carnivores rank high on the scale of intelligence among mammals. The brain is large in relation to the body, an indication of their superior mental powers. For this reason, these animals are among the easiest to train for entertainment purposes, as pets, or as hunting companions. The highly developed sense of smell among dogs, for instance, supplements the sharper vision of man. Dogs are the carnivores most commonly trained for hunting, but the cheetah, caracal, and ferret have also been used to some extent. In China the otter is trained to drive fish (commercial fishing) under a large net, which is then dropped and pulled in. Dependent for survival upon their ability to prey upon living animals in a variety of situations, carnivores have evolved a relatively high degree of learning ability (see animal learning).

      Carnivorous mammals tend to establish territories, though omnivorous carnivores, such as the black bear, striped skunk, and raccoon, are less apt to do so. Territories (territorial behaviour) are often exclusive, defended by the residents against other animals of their own kind. Such areas may sometimes be marked by secretions produced by anal or other scent glands and by deposition of feces in prominent locations.

 There is a wide range of social patterns among carnivores. Many (bears, various foxes, genets, most cats, and most mustelids) are solitary except during the breeding season. Some remain paired throughout the year (black-backed jackal and lesser panda (panda)) or occasionally roam in pairs ( gray fox, crab-eating fox, and kinkajou). Other carnivores, such as the wolf, African hunting dog, dhole, and coati, normally hunt in packs or bands. Various pinnipeds form sedentary colonies during the breeding season, sea otters congregate during a somewhat larger part of the year, and meerkats are permanently colonial.

  Mating systems vary among families, ranging from monogamy in the wolf and polygyny in most bears and mustelids to harems in elephant seals. Copulation is vigorous and frequent in many species, including the lion, and many species possess reproductive peculiarities as adaptations to their environments. Induced ovulation, for instance, allows females to release egg cells during or shortly after copulation. Delayed implantation of the fertilized egg in the wall of the uterus is another phenomenon that allows births to occur when resources are abundant. This phenomenon is most prominent in species living in highly seasonal environments. Delayed implantation is most extreme in the pinnipeds and bears but is absent from canines.

Form and function

      The smallest living member of Carnivora is the least weasel (Mustela nivalis), which weighs only 25 grams (0.9 ounce). The largest terrestrial form is the Kodiak bear (Ursus arctos middendorffi), an Alaskan grizzly bear that is even larger than the polar bear (Ursus maritimus). The largest aquatic form is the elephant seal (Mirounga leonina), which may weigh 3,700 kg (8,150 pounds). Most carnivores weigh between 4 and 8 kg (9 and 18 pounds).

 The vast majority of species are terrestrial, but the pinnipeds are highly adapted to life in the water. Some nonpinnipeds, such as the sea otter, are almost fully aquatic, while others, such as the river otter and polar bear, are semiaquatic, spending most of their lives in or near water. Aquatic and semiaquatic forms have developed specializations such as streamlined bodies and webbed feet.

  Carnivores, like other mammals, possess a number of different kinds of teeth: incisors in front, followed by canines, premolars, and molars in the rear. Most carnivores have carnassial, or shearing, teeth that function in slicing meat and cutting tough sinews. The carnassials are usually formed by the fourth upper premolar and the first lower molar, working one against the other with a scissorlike action. Cats, hyenas, and weasels, all highly carnivorous, have well-developed carnassials. Bears and procyonids (except the olingo), which tend to be omnivorous, and seals, which eat fish or marine invertebrates, have little or no modification of these teeth for shearing. The teeth behind the carnassials tend to be lost or reduced in size in highly carnivorous species. Most members of the order have six prominent incisors on both the upper and lower jaw, two canines on each jaw, six to eight premolars, and four molars above and four to six molars below. Incisors are adapted for nipping off flesh. The outermost incisors are usually larger than the inner ones. The strong canines are usually large, pointed, and adapted to aid in the stabbing of prey. The premolars always have sharply pointed cusps, and in some forms (e.g., seals) all the cheek teeth (premolars and molars) have this shape. Except for the carnassials, molars tend to be flat teeth utilized for crushing. Terrestrial carnivores that depend largely on meat tend to have fewer teeth (30–34), the flat molars having been lost. Omnivorous carnivores, such as raccoons and bears, have more teeth (40–42). Pinnipeds have fewer teeth than terrestrial carnivores. In addition, pinnipeds exhibit little stability in number of teeth; for example, a walrus may have from 18 to 24 teeth.

      Several features of the skeleton are characteristic of the order Carnivora. Articulating surfaces (condyles) on the lower jaw form a half-cylindrical hinge that allows the jaw to move only in a vertical plane and with considerable strength. The clavicles (clavicle) (collarbones) are either reduced or absent entirely and, if present, are usually embedded in muscles without articulation with other bones. This allows for a greater flexibility in the shoulder area and prevents breakage of the clavicles when the animal springs on its prey.

      The brain is large in relation to the weight of the body, and it contains complex convolutions characteristic of highly intelligent animals. The stomach is simple as opposed to multichambered, and a blind pouch ( cecum) attached to the intestine is usually reduced or absent. Since animal tissues are in general simpler to digest than plant tissues, the carnivore's dependence on a diet with a high proportion of meat has led to less-complex compartmentalization of the stomach and a decrease in the length and folding (and therefore surface area) of the intestine. The teats are located on the abdomen along two primitive lines (milk ridges), a characteristic of mammals that lie down when nursing.

      Many carnivores have a well-developed penis bone, or baculum. It appears that this structure plays a role in helping to increase the success of copulation and fertilization of eggs in species where numerous males mate with a single female. Cats have a vestigial baculum or none at all, but the baculum of the walrus can measure up to 54 cm (21 inches).

Distribution and abundance

 Carnivores are found worldwide, although Australia has no native terrestrial members except for the dingo, which was introduced by aboriginal man. Terrestrial forms are naturally absent from most oceanic islands (island), though the coastlines are usually visited by seals. However, people have taken their pets, as well as a number of wild species, to most islands. For example, a large population of red foxes now inhabits Australia, having been introduced there by foxhunters. Introduction of carnivores to new environments has at times devastated native fauna. In New Zealand, stoats, ferrets and weasels were introduced to control rabbits, which had also been introduced. As a result, native bird populations were decimated by the carnivores. Birds were also a casualty of mongooses introduced to Hawaii and Fiji, where populations of introduced rodents and snakes had to be controlled. In Europe, American minks released from fur farms contributed to the decline of the native European mink.

 Because carnivores are large and depend on meat, there must be fewer carnivores in the environment than the prey animals they feed upon. The maintenance of territories limits the number of predators to the ecosystem's carrying capacity of prey. In general, carnivores have a population density of approximately 1 per 2.5 square km (1 per square mile). By comparison, omnivorous mammals average about 8 per square km (20 per square mile), and herbivorous rodents attain densities of up to 40,000 per square km (100,000 per square mile) at peak population. Relatively low population density makes carnivores vulnerable to fluctuations of prey density, habitat disturbance, infectious disease, and predation by man. The mobility and adaptability of some carnivores has enabled them to shift ecological roles and survive changes brought about by human activities. For example, the red fox, coyote, raccoon, and striped skunk can all be found in urban and suburban areas of North America. In Europe, the red fox lives in most large cities. Most other species do not fare nearly as well. The gray, or timber, wolf and brown bear once lived across much of the Northern Hemisphere, but their ranges have shrunk following habitat destruction, reduction of prey abundance, and persecution as competitors with man. In Africa and southern Asia the same can be said for lions and tigers. Numerous cats and bears and some seals have become rare and are threatened with extinction.

Classification

Distinguishing taxonomic features

      There is great diversity in Carnivora, especially among the highly specialized pinnipeds. Thus, the characteristics used to separate Carnivora from other mammalian orders and to define the subdivisions of Carnivora are primarily structural. Of great importance are certain features of the skull (such as jaw articulation), feet (number of toes, lack of opposability of the hind toe, type of claws, and fusion of certain bones), and teeth (both the overall tooth pattern and the shape of individual teeth). Dentition is especially important in determining the relationships of fossil forms. Also useful in the taxonomy of modern carnivores are the convolutions around the lateral, or Sylvian, fissure of the brain, the relative weights of the adrenal and thyroid glands, the type of uterus and placenta, and the position of the nipples.

Critical appraisal

      The taxonomy of the major categories of major groups placed in the Carnivora has been in a state of flux for more than a century, and these categories do not seem to be stabilizing, even today. Most mammalogists at present regard the seals and terrestrial carnivores as belonging to different orders, the Pinnipedia and Carnivora. There are, in reality, only a few features common to the seals and their terrestrial relatives because of the extensive and numerous adaptations the aquatic forms have undergone to make them efficient carnivores of the sea. Mammalogists who have studied seals intensively now realize that there is no anatomical structure unmodified by the extensive aquatic adaptations; every organ and tissue examined has been found to be different in some way from its counterpart in terrestrial forms. Other mammalogists, tending toward conservative taxonomy, think the relationship of the terrestrial and aquatic carnivores can be best expressed by retaining them in two suborders, the Fissipedia (“split-footed”) and Pinnipedia (“feather-footed”), of the single order Carnivora. This more conservative taxonomy is followed in this article.

      Of the living families recognized in the Carnivora, two have separated from their lines most recently and are most easily associated with other existing families: the Odobenidae (walrus) with the Otariidae (eared seals) and the Hyaenidae (hyenas) with the Viverridae (civets). Moreover, a new family, the Mephitidae (skunks and stink badgers), has been proposed as an offshoot from the Mustelidae (weasels). It appears that skunks do indeed possess enough differentiation in features and genetics to warrant the new grouping. Taxonomy of several species of carnivore remains uncertain. Among those, two of the most problematic species are the lesser, or red, panda (panda) (Ailurus fulgens) and the giant panda (panda, giant) (Ailuropoda melanoleuca). Both species have been classified equally often in the Ursidae (bears) or the Procyonidae (raccoons). However, the latest classification includes them under Ursidae. Another lesser-known species, the fossa (Cryptoprocta ferox), is regarded as a viverrid but retains characteristics of cats as well. It has been alternatively placed in Herpestidae, Viverridae, and even Felidae.

      The arrangement of the nine terrestrial families into two distinct superfamilies, Canoidea and Feloidea (or Aeluroidea), appears to be a natural arrangement dating back to the works of W.H. Flower and H. Winge in the late 1800s. In Canoidea, as revealed by studies in comparative anatomy and the fossil record, the families Canidae, Ursidae, and Procyonidae seem to be most closely related. Also placed in the Canoidea is the family Mustelidae, although some of the more primitive members show resemblances to the primitive viverrids as well as to the canids. In the Feloidea, the families Viverridae and Hyaenidae seem most closely related, the Felidae being the most aberrant. Those families that contain rather diverse lines have been divided into subfamilies, the number of subfamilies in each family indicating the amount of evolutionary divergence that has occurred. The groups that have probably been distinct the greatest length of time have the most subfamilies, Viverridae with six and Mustelidae with five.

      As a result of such complicated taxonomic appraisal, the formal classification of Carnivora is in some ways an artificial system set up for the sake of convenience. Ideally, the system reflects real evolutionary relationships, but these must be inferred from a scanty fossil record and from comparisons of modern species. Since there are differences of opinion among specialists as to which taxonomic characteristics should be given priority, there are certain to be alternate classifications, the acceptability of which depends on new information continually being discovered. Undoubtedly, advanced genetic fingerprinting and DNA analyses will allow a more objective classification of species within the order.

Howard James Stains Serge Lariviere Order Carnivora (carnivores)

 274 species found worldwide but introduced to Australia.

      Suborder Fissipedia

 

      Family Mustelidae (mustelid) ( weasels and related species)

 54 species in 21 genera belonging to 5 subfamilies, found worldwide except for Australia.

      Family Herpestidae (mongoose) ( mongooses)

 37 species in 18 genera belonging to 2 subfamilies, found in the Old World.

      Family Felidae (feline) ( felines, or cats)

 37 species in 18 genera belonging to 3 subfamilies, found worldwide except for Australia.

      Family Canidae (canine) ( canines)

 35 species in 13 genera, found worldwide but introduced to Australia.

      Family Viverridae (viverrid) ( civets and related species)

 35 species in 20 genera belonging to 6 subfamilies, found in the Old World.

      Family Procyonidae (procyonid) ( raccoons and related species)

 18 species in 6 genera belonging to 2 subfamilies, found in the New World.

      Family Mephitidae ( skunks and stink badgers)

 11 species in 4 genera, found in the New World and Southeast Asia.

      Family Ursidae (bear) ( bears)

 9 species in 6 genera belonging to 2 subfamilies, found in North America, South America, Europe, and Asia.

      Family Hyaenidae ( hyenas and the aardwolf)

 4 species in 4 genera belonging to 3 subfamilies, found in Africa and southern Asia.

      Suborder Pinnipedia (pinniped) ( pinnipeds)

 34 species in 18 genera belonging to 3 families, found primarily in marine waters.

      Family Phocidae (true, or earless, seals)

 19 species in 10 genera.

      Family Otariidae (eared seals)

 14 species in 7 genera.

      Family Odobenidae ( walrus)

 1 Arctic species.

Additional Reading

For general information on the biology of the carnivores, see E.P. Walker et al., Mammals of the World, 3 vol. (1964), in which each genus is described and illustrated, along with a brief summation of its biology. The taxonomy of carnivores is discussed in G.G. Simpson, “Principles of Classification and a Classification of Mammals,” Bull. Am. Mus. Nat. Hist., vol. 85 (1945), a classic work on classification, followed by most recent mammalogists; and H.J. Stains, “Carnivores and Pinnipeds,” in S. Anderson and J.K. Jones, Jr. (eds.), Recent Mammals of the World: A Synopsis of Families (1967). F.E. Beddard, Mammalia (1902, reprinted 1958), is a definitive early work on mammalian anatomy. The paleontology of the Carnivora is summarized in two works by A.S. Romer: Vertebrate Paleontology, 3rd ed. (1966), fundamental to an understanding of fossil forms, and Notes and Comments on Vertebrate Paleontology (1968), containing additional information not found in the general text. Books devoted to particular subgroups of the carnivores include A. Denis, Cats of the World (1964), an excellent summary of the status of all members of the Felidae; C.J. Harris, Otters (1968), a fine summary of the status of the otters around the world; H.E. Hinton and A.M.S. Dunn, Mongooses: Their Natural History and Behaviour (1967), which contains much interesting information that is difficult to find elsewhere, except in scattered literature; R.J. Harrison et al. (eds.), The Behavior and Physiology of Pinnipeds (1968), an excellent summation of knowledge on these aquatic carnivores; and V.B. Scheffer, Seals, Sea Lions, and Walruses: A Review of the Pinnipedia (1958), a major work on the taxonomy of the Pinnipedia.Howard James Stains

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