Antelopes, Gazelles, Cattle, Goats, Sheep, and Relatives
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© Copyright, Princeton University Press. No part of this book may be distributed, posted, or reproduced in any form by digital or mechanical means without prior written permission of the publisher. INTRODUCTION RECOGNITION The family Bovidae, which includes Antelopes, Cattle, Duikers, Gazelles, Goats, and Sheep, is the largest family within Artiodactyla and the most diverse family of ungulates, with more than 270 recent species. Their common characteristic is their unbranched, non-deciduous horns. Bovids are primarily Old World in their distribution, although a few species are found in North America. The name antelope is often used to describe many members of this family, but it is not a definable, taxonomically based term. Shape, size, and color: Bovids encompass an extremely wide size range, from the minuscule Royal Antelope and the Dik-diks, weighing as little as 2 kg and standing 25 to 35 cm at the shoulder, to the Asian Wild Water Buffalo, which weighs as much as 1,200 kg, and the Gaur, which measures up to 220 cm at the shoulder. Body shape varies from relatively small, slender-limbed, and thin-necked species such as the Gazelles to the massive, stocky wild cattle (fig. 1). The forequarters may be larger than the hind, or the reverse, as in smaller species inhabiting dense tropical forests (e.g., Duikers). There is also a great variety in body coloration, although most species are some shade of brown. It can consist of a solid shade, or a patterned pelage. Antelopes that rely on concealment to avoid predators are cryptically colored. The stripes and blotches seen on the hides of Bushbuck, Bongo, and Kudu also function as camouflage by helping to disrupt the animals’ outline. Sociable species that live in the open tend to have bold markings or a dark color which, along with conformation, help them to tell their own kind apart from all other species. The sexes may be differently colored, with males usually having the darker pelage (e.g., Sable Antelope). Frequently within species, there are also age-specific colorations. 5 4 3 1 2 Figure 1. Variety of size, weight, and body shape in Bovids: (1) Royal Antelope (Neotragus pygmaeus); (2) Kirk’s Dik-dik (Madoqua kirkii); (3) Nubian Ibex (Capra nubiana); (4) Sable Antelope (Hippotragus niger); (5) Gaur (Bos gaurus). Sizes compared to an adult human. Horns: One of the defining characteristics of the family Bovidae is the presence of unbranched horns (figs. 2 and 3). Horns are present in males of all Bovid species and in females of some genera, usually in large species. Horns in males are always thicker and more complex. The horns are permanently attached to the frontal bones of the skull, and are composed of a bone core covered with a keratin sheath which is never shed, with an air space separating these two layers. Horn size and shape vary greatly, often having a spiral, twisted, or fluted form. This unique horn structure is the only unambiguous morphological feature of Bovids that distinguishes them from other ruminants. Tetracerus is unique among wild Bovids in that males regularly bear four horns (two pairs); all other genera, with the exception of some domestic sheep, have only one pair. Horn growth in Bovids continues throughout the animal’s life, and horn size and number of rings (not the annulations seen in some species) may be used for age determination (figs. 5 to 7), although size and quality of horns may be altered by several factors, such as food supply. Horns in young animals may seem quite odd compared with the final stage reached in the adult, and can sometimes resemble those of a very different species. Antlers (fig. 4.1), which are characteristic of the family Cervidae (deer), are not true horns: they are made of bone 8 For general queries, contact [email protected] antelopes22reduced.indd 8 02-Nov-15 7:11:04 PM © Copyright, Princeton University Press. No part of this book may be distributed, posted, or reproduced in any form by digital or mechanical means without prior written permission of the publisher. INTRODUCTION 4 3 2 5 6 1 7 8 9 10 11 14 12 13 Figure 2. Variety of horn shapes and sizes in Bovids: (1) Gemsbok (Oryx gazella); (2) Addax (Addax nasomaculatus); (3) Transcaspian Urial (Ovis arkal); (4) Kirk’s Dik-dik (Madoqua kirkii); (5) Eastern White-bearded Wildebeest (Connochaetes albojubatus); (6) Common Eland (Taurotragus oryx); (7) Alpine Ibex (Capra ibex); (8) Northern Grant’s Gazelle (Nanger notata); (9) Mountain Goat (Oreamnos americanus); (10) Red Hartebeest (Alcelaphus caama); (11) Ellipsen Waterbuck (Kobus ellipsiprymnus); (12) Four-horned Antelope (Tetracerus quadricornis); (13) Red-flanked Duiker (Cephalophus rufilatus); (14) Cape Buffalo (Syncerus caffer). Not to the same scale. 9 For general queries, contact [email protected] antelopes22reduced.indd 9 02-Nov-15 7:11:07 PM © Copyright, Princeton University Press. No part of this book may be distributed, posted, or reproduced in any form by digital or mechanical means without prior written permission of the publisher. INTRODUCTION 1 2 Figure 3. Horns: Horns are composed of a bony core (1) covered with a sheath of keratin (2). They are never branched, although they do vary in shape and size. Neither the sheath nor the core is ever shed, and in many species, the horns never stop growing. Horn cores begin as small bony growths under the skin, over the skull. Found only in Bovids. Photo credits: Falln-Stock. 1 2 3 Figure 4. Antlers (1): Consist of bony outgrowths from the skull with no covering of keratin, usually large and branched, growing from an attachment point on the skull called a pedicel. While an antler is growing it is covered with highly vascular skin called velvet; once the antler has achieved its proper size, the velvet is lost and the antler’s bone dies. Antlers shed after each mating season. Found only in Cervids. Pronghorns (2): They differ from the horns of Bovids in two respects: they are branched, and, while the horns consist of a bony core and keratinous sheath, the sheaths are shed annually, while in Bovids, the sheaths are always permanent. Found only in the Pronghorn antelope. Ossicones (3): Giraffes have a pair of short, unbranched, permanent bony processes that are covered with skin and hair, called ossicones. They derive from ossified cartilage, as true horn do, but they remain covered in skin and fur, rather than horn, while antlers are derived from bone tissue. Photo credits: LaTaxidermia.com and Skulls Unlimited. without keratin, and are not permanent. Antlers will begin growing in the spring and will continue to grow until mating season, which is when they reach their full size. In late winter, the antlers are shed, and males may be without antlers for a few months until the cycle begins again. Antlers are branched and are only found on males (and female Caribou). Horns in the family Antilocapridae, with only a single extant species, the Pronghorn, are different from either true horns or antlers (fig. 4.2), but have characteristics of both: they are made of keratin growing on a bony core, like true horns, but they are shed annually; in addition, the horn sheath is branched in male Pronghorns, while true horns are always unbranched. Giraffes (family Giraffidae) have a pair of short, unbranched, permanent bony processes that are covered with skin and hair, called ossicones (fig. 4.3); they differ from horns in Bovids in that they do not project from the frontal bones, but lie over the sutures between the frontal and parietal bones. Rhinoceros horns differ from true horns because these horns have no core or sheath; they are made up of a multitude of epidermal cells and bundles of dermal papillae, extensions of the dermis, forming horny fibers similar to thick hair. A single horn is situated over the nasal bones; 10 For general queries, contact [email protected] antelopes22reduced.indd 10 02-Nov-15 7:11:09 PM © Copyright, Princeton University Press. No part of this book may be distributed, posted, or reproduced in any form by digital or mechanical means without prior written permission of the publisher. INTRODUCTION birth 6 months 12 months 18 months 2 years 3 years 6 years Figure 5. Horn growth in male Black-faced Impala (Aepyceros petersi): Age can be estimated by rings on horns. Note that horns of young animals may seem quite odd compared with the final stage reached in the adult, and can sometimes resemble those of a very different species. and in species that have two horns, the second horn lies over the frontal bones. Other ungulates may have tusks, which are elongated, continuously growing front teeth, that may protrude well beyond the mouth; they are most commonly canines, as with warthogs and pigs. Hooves (fig. 8): Keratinous hooves sheathe the terminal bone of each toe. Hooves are modified toenails, and are composed of two structures: the unguis and the subunguis, which connect the unguis to the pad of the digit. Unlike claws and nails, hooves are the principal point of contact between the legs and the ground; as a result, ungulates are said to have unguligrade limbs. The tough unguis encircles the tip of the digit as a cylinder, enclosing the subunguis within it. Since the unguis is harder than the subunguis, it does not wear down as quickly, resulting in a firm leading edge to the hoof. Bovids walk on their hooves and have paraxonic feet, in which two well-developed digits, the third and fourth digits, bear the weight of the body. The second and fifth digits are either absent or, more often, small, forming the so-called lateral hooves, or dewclaws.