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    Elephant

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    MAMMALIAN SPECIES wo. 122, pp. 1-8, 4 figs. Elephas maximus. By Jeheskel Shoshani and John F. Eisenberg Published 18 June 1982 by The American Society of Mammalogists Elephas Linnaeus, 1758 Blephas Linnaeus, 1758538. Type species Elephas maximus Lin- acus, by original designation Elephontus Geofftoy Seint-Hilaire and Cuvier, 1795:189 (not Cu vier and Geoffroy SaintHilaie). Type species F. maximus Linnaeus, by original designation, Pilgrimia Osborn, 1988: Type species Elephas falconeri Bush, by original designation; fos Sivalikig Osborn, 1924: Type species Loxodonta namadica Fa ‘coner and Cautley, by original designation (=Elephas na~ ‘madicus): fos Palaeolozodon Matsumoto, 1926:257. Type epecies "E. namad- ‘cus naumanni Makiyama, 1924," by original designation lephas namadieus, os Leith-Adamsia Matsumoto, 1927:214. Type species Leith-Adam- sa siwalikiensis Matsumoto, by original designation (= Ele has planifrons; fos Hesperotoxodion Osborn, 1981-21. Type species Elephas antiquas Falconer and Cautley, by orginal designation (~Blephas na madicus;fosi Platelephas Osborn, 194221358. Type species Elephas platy: ‘eephalus Osborn by original designation; fosi ‘Hypselephas Osborn, 1942:1340, Type species Elephas kysudri- ‘us Falconer and Cautley, by monotypy’ fossil Stegoloxodon Kretzoi, 1990:405, Type species Ste ‘donesicus Kretzoi, by original designation bens fossil Omoloxodon Deranivagala, 1955:25. Type species Elephas reckt Dietrich, by subsequent designation (Magli, 1973234, fossil loxodon in- Tephas cele- CONTEXT AND CONTENT. Superorder Pacnungulat, Order Proboscidea, Suborder Gomphatheriides, Family Ele: phantidae, Subfamily Elephantinae. The genus Blephas contains ten extine’ and ane living species (Coppens eta, 1978; Maglio, 1973), Elephas maximus Linnaeus, 1758 Asian Elephant Blephas mavimus Linnaeus, 1758:33. Type locality “Zeylonae now Sri Lanka Elephas asiaticus Blumenbach, 1797:124. Type locality “such: fen Asien, vorzuglich auf Cellan” (southern Asis, chiefly in Ceylon, Elophes indicus Cuvier, 1198:148-149. Type locality not given. Elephas gigas Perry, 18llcunnumbered. page. 259" and ‘un ‘aumbered plate 3 fin copy in Rare Book Reom at the Amer- jean Museum of Natural History). Type locality "Africa and ‘Asia (not'a nomen nudum because figure and mest of the text are based om the Asian elephant) Elephas sumairanus Temmincl, 1847:91, Type locality “Suma CONTEXT AND CONTENT. Contest noted in generic summary above. We follow Chasen (i940) und Uist only three sub- species: Em, indicus Cuvier, 1798, see above (signs Perry, sodactylus Horgton, keterodactylas Hodgson, bengalensis de Blainvlle, ddauatela Falconer and Cautley, mukna Falconer and Cau Tey, Mrsutus Lydekker, dathunencis, asuras, rubridens, bir= rmanicas, borncensis, and sondaicuy all of Deraniyagale are ‘synonyms E. m. maximus Linnaeus, 1758, see above (asiaticus Blumen- bbach, ceylonicus de Bisinville, eylanicus Lydekker,vilaliya and sinkaleyus both of Decaniyagala are synonym) Bm sumatranus Temuminck, 1847, see above. DIAGNOSIS. Elephat maximus has more lamellae and wid- x upper and lower thitd molars then entinet species of Elephas Otago, 1973) "The Asian elephant is smaller then the Aftican elephant (Loxodonta africana) and has the highest body point atthe head instead of shoulder, The head is compressed anteroposterioly, 4nd has two dorsal bulges and a concave forehead. The back iv ‘conver ot level instead of concave, The ears of E. maximus are Sruall and in mature individuale the doreal borders are folded Iaterally instead of medially. The tip ofthe tunk has one finger- like process instead of two. There ace up to 20 pairs of ribs and {34 caudal vertebrae instead of 21 and 33, respectively there are usually five naillike structures on each forefoot and four on each ‘ear foot instead of four and three, Only males generally carry tusks (instead of both sexes); cheekteets have up to 29 narrow, ‘compressed closed loops or lamellae (instead of up 10 15 lovenge- Shaped plates per tooth). Skin is smoother than in L- africana (Carrington, 1986; Deraniyagela, 1985; Frade, 1955; Laursen and. Bekoff. 1978; Sikes, 1970), GENERAL CHARACTERS. The distinctive trunk is an clongation of nose and upper lip combined: the nosuils are sti tip. The familar build is shown in Fig. 1 Large bulls weigh 5.400 keg and are 3.20 m atthe shoulder. Record height (Pils, 1941) i 3143 m, Females reach 2.54 m at the shoulder and weigh 4,160 ke, average weight is 2,720 kg. and average height is 2.28 m. Boyle (1929) found that twice the forefoot eirourference gave the shoulder height. The length ofthe body and head including teunk ig 55 to 6.5 m; the tall s 1.2 to 1.3 m long (Benedict, 1930) Elephants are smaller in more eastern populations, smallest in Borneo, Skin color is generally gray but becomes lighter in the ‘caste part of the species range. Elephants from Sri Lanke to Burma display white spots and blotches: those from Malaysia Ihave gray patches and Bm. sumaranas has small light spots on fears and trunk (Dereniyagala, 1955) DISTRIBUTION. Past and present distributions of B. ‘maximus are depicted in Fig. 2. Pleistocene fossils are known from'Borneo (Hooijer, 1964; Medway. 1977). Distribution Is patchy and mostly Lmited to forest-grassland ecotone (Olivier, {97ad), Estimated wild elephant population 1s 23,000 to 41,000; FicuRe 1 {imu maximus, at Yala National Park, Sei Lanka (photo by J Shoshan An adult solitary bull Asian elephant, Blephos max- FicURE.2, Disteibution ofthe Asian elephant, Blephas maximus, alter Olivier (9782-1978): B. m. maximus is confined to the {sland of Sri Lanks, F, m, sumatranu tothe island of Sumat and B. m indicus occupies the rest of the range crude densities within the habitat studied fall between 0.12 and {U0 animals per kin (Bisenberg, 1980), FOSSIL RECORD. Ten fossil species of Blephas were recognized; the earliest i from the middle Pliocene Ekora beds, Southeastern Turkana, Kenya, fozmed ebout 4.3 million years ago (Maglio, 1973). FORM. The skin is wrinkled and may be depigmented on the trunk, ears, head, or neck. Albino Asian elephants are re- suarded ae sacred (Mache, 1927, Hair in adults grows in speciic facts, 21 on the head and 10 elsewhere. Only three tracts are permanent, The longest baie is onthe lis, on the elbow and knee Joints, and on the end ofthe tall (Deraniyagala, 1953), Fetal hair “described by Rajagopal and Ayer (1954), Smith (1890) fed hairs and bristles: the former are much finer and lid in cross section; the later are coure, longer, and perforated by a foramen when viewed under the microscope. Hair of E ‘maximus has 2 lfar content of 4.1% and known amine acid Composition Gillespie, 1970) "The epidermis and dermis of the body average 18 mm thick (Smith, 1890) Shoshani et sl. (1980) found that skin on the dex sum was 30 mm thick. The approximately hexagonal “epidermal columns” or “studs” are similar to those of L- africana (Horst- ‘mann, 1966). Smith (1890) noted the absence af glands in the skin: Benedict and Lee (1938) also reported thatthe skin is gen- cally dey ‘The mammae_and temporal (musth) glands are the only known integumentary glands. Milk it 83.82% water. 1L82% al- Duminoids and sugar, 3.89% fat, and 0.7% ash ut mineral mat- ter. The specife gravity i 1.088 (Benedict, 1986). Peters etal (972) reported that elephant milk contains 1.94 to 3.0% protein, 0.63 to 629 fat, 4.0 10 8.36% carbohydrate, and 8.27 wo 17.3% solids. The temporal gland is composed of tubular alveolar glands ‘Aggregated inte lobules, It is located subeutancously, midway Detween the eye and the ear on each side ofthe head. The land's ‘orifice i about 20 mm long Femando etal. 1963). The skeleton constitutes about 15% ofthe body weight. The histology of Blephas bone was studied by Amprino and Godina (947) tnd compared to that of Mammuthae by Esra and Cook (1950). The long bones lack large marrow cavities and are filed ‘with spongy bone through which the marrow is disseminated, The epipbyses, particulary on long bones and vertebrae do aot fmly fuse until about 25 to 30 years of age. The vertebral formula of E, maximas is: C 7, 719-20, L 3-5, 8 3-5, CA 24-34. According te Temminck (1887), the Sumatran elephant has 20 pairs of ribs, ‘whereas the mainland form has 19 pairs. The first six pairs of ‘ibs contact the sternum, the rest do not. The long bones of each limb almost form a straight line perpendicular to the vertebral column. The scapula when laid on «fat surface, tilts toward the Infraspinous fossa, which is about twice as large as the supre spinous fossa The lium is almost vertical and expanded laterally ‘The acetabulum faces downward. The knee joint is at the level of, of below, the body contour. Radius and ulna are separate (rsay be fusca in old individuals) and held permanently crossed Ja xed position of pronation, Tibia and fibula are also separate: MAMMALIAN SPECIES 182 the latter articulates with the caleaneurm, Manus is larger than pes; both are short, broad, pentadactyl, and seemingly me nic. The manus is semi-digtigrade, the pes ie semi-plantigrade. ‘The carpal bones ofthe manus are serially arranged. The astraga- lus is lat and does not articulate with the cubuid. Each foot, including its reduced phalanges, rests on a pad of elastic tissue tnd is encased in a cylindrical integument. Body weight is sup ported on these ough, compressible, and shock-absorbent,Aibro- Fatty elastic connective tissues. Os centrale is present on acaph ‘id of young individuals. Os cordis, baculum, clavicle, astragalar foramen, and humeral entepicondiar foramen are absent. Third femoral vochanter is absent or greatly reduced (Beddard, 1902: Blair, 1710; Deranivagala, 1955; Flower and Lydekker, 1891; Frade, 1955; Le Gros Clark and Sonntag, 1925; Osborn, 1996, 1942; Shoshent eta, 1980) Extensive pneumatization occurs in the pariotals, nasals, premaxillae, maxilla and fronals (Fig. 3) Other features of the all include elevated external nates, wide premaxille-frontal ‘contact posterior to eaudal border of obit, thickened and laterally extended squamosal portions of mandibular fossa and zygoma, fos. The erbriform plate is hor mary. A secondary Scoustic mestus Is formed by enfolding af the squamosal bone. 4 condylar foramen is absent and the lacrimal foramen is either absent or rudimentary. Te internal carotid canal perforate tympanic bulla. An alisphenoid canal is present. Alargeinfraor- bital foramen (sometimes two or more) transmits the maxillary branch ofthe trigeminal nerve and blood vestle tothe proboscis. ‘The mandibular symphysis is spout-tike. The high ascending rami terminate in transverse condyles. The coronoid process is com- Dressed and the coronoid canal is present at its base (Beddard, 1902; Osborn, 1936, 1942) The dental formule of E. maximus is identical to that of L africana:'\ 1, ¢ 00, dp 3/3, m 33, otal 2. Deciduous tusks are laced by permanent second inelsars within 6 to 12 months iter bith (Deraniyagala, 1955). Permanent tusks are composed Sf dentine and grow continuously atthe rate of about lem per year (Colyer and Miles, 1957). Males grow incisors of two 1¥pes: 9) large tusks which mey protrude well beyond the lip. end b) tshes which extend barely heyond the mouth. The two conditions are parently under genetic control, A smoath conical cap of enarmel i present at the tip of the tusk until it wears off early in life Record tusk length i 8.02 m (Smith, 1930) and record single tusk weight is 39.0 kg ohn, 1928). Tn cross section, a tusk exhibits Concentric growth lines, or ‘lines of Owen, as well asa checkered pattern of diamond-shaped areas which become. progressively fmaller towards the center of the tusk (Fig. 3). This pattern is unique to Proboscidea (Miles and White, 1960) For practical field work, the cheekteeth are called molars I through VIL The plates of upper teeth diverge towards the chew- turduce while the plates of lower teeth converge. The check teeth are hypsodont and have multiple roots, ‘The lophodont finding surfaces are composed of closed enamel inope whose enters are filed with dentine and which are held together by ‘cementam (Fig. 4). The first cheektooth has a maximum of si Tops, and the number increases foreach successive toth. The ‘minimum and maximum reported numbers of laminae above and Below ares 46/46, 710/710, N1=14/12-14, 15e17/14-17, 17-211 17-20, and 20-2620-29. A sampling of lower cheekteth weighed fin a) 9.0, 125.0, and $68.0 for the first, second, and third Adectduous premolars, and 1,660.0, 3,685.4 and 5,159.0 for the frst, second, and third molars (Frade, 1985; Roth and Shoshani, ‘unpubl. The cheekteeth, unlike those of most mammels, move an- teriorlly a4 well ay occlusally as they emerge (Fig. 3). These teeth are the p2, dps, dps, Ml, M2, and MS (dpl is rarely pre=- ent). Except for the three deciduous premolars which are present from birth through the fist several years, only one tooth or parts ‘of two teeth are functional at one time. As each tooth wears away, ‘developing tooth from behind replaces it, until the last molar fs worn outs the elephant then dies of starvation. The fist tooth 6, 12,20 1969). Supernumerary and ‘leforined tusks and cheekteeth sometimes occur in wild and cap tive elephants (Colyer, 1936) The hyoid bone and associated musculature were illustrated by Gase (1967) and comparison ofthe stylohoidea among E. nau- rmanni, E. maximus and L. africana was given by Inuzuka etal 975). MAMMALIAN SPECIES 182 Froune 3 top middle, poste ‘Top left, frontal view of cranium of Elephas maximas sumatranus (Nat. Mus. Nat, Hist. No, 240999, Washington, D.C. view of same skull; tp tight, side view of same skal: lower left, N-M.N.H. No, 399199’ with bone cut away. (arrow shows direction of tooth replacement; inset shows crows seetion of tsk (mam scale) of Elephas}; lower middle, ventral view of 240930; lower right, skull 399499, cut down middle to show air cells within most bones (skull photos courtesy’ of Smitheonian Institution; tusk eros section tom Bucky Steele, photo by Kathleen Morehead). The scales (white insets) represent approximately 80 mm. ‘The trunk musculature consists of two major sets: 1) longi- tudinal and 2) rediting and transverse. The longitudinals are tostly superficial and are further subdivided into anterior, lat- ‘nd posterior. The deeper muscles are best seen as nmer= bus distinct fasciculin a crose section of the trunk (Harrison, 1847; Miall and Greenwood, 1878: Shindo ard Mori, 1955). Lewis (71 claimed thatthe trunk contain as many as 60,000 muscles, Hartson (18501) described a muscle extending from the back part ofthe bifurcation ofthe trachea to the forepart ofthe esoph- gus. Watson (18722) and Miall and Greenwood (1878) found no ‘ace of this muscle. Shoshani eta. (1980) found it in one ofthe three elephants they examined. Le Gros Clark and Sonntag (1925) ‘compared the musculature and other systems among the Tubu- Tidentata, Hyraccidea, Proboscidea, Perissodactyla, and Artio- dactya "The hear of the Asian elephant, like that of the Aftican species and Sirenia but unlike that of most other mammals, has {bifid apex and paired anterior venae cava. The tneurpid valve tmay have one or two small addtional eusps. There are two cor- ‘onary arteries. The heart is connected to the perieardivm at its base, and two fibrous cords pass from the dorsum of the peri cardium to the tendinous center ofthe diaphragm (Frade, 1955, Hill, 1938, Mall and Greenwood, 1878; Sikes, 1971; Watson, 1873a). According to Benedict (1936), hearts weigh about 0.570 of total body weight. Mill and Greenwood (1878) stated thet a striking peculiarity of the veina ies in the plexuses and fren anaa- tomoses which occur in various parts of the body. Valves were found in some but not al pleauses. Distinct and large valves were found by one of us (JS) in the temporal plexuses of one elephant blood of E. maximus to at of Homo sapiens, bod and the platelet count was in the high-nonnal human range” Nirman nd Nair 0911) analyzed E. maximus plasma in four groupe af Shimale: Protein made up 825 to 92.8 g/10 mi of plasma, the Highest reported value for ay mammal; lobuin content was also the highest among mammals maximum of 6.68 = 192 g/00 ml plasma), and the abuminiglobuln ratio was the lowest (0.33 8.12" 0.46 = 0.20, Hemoglobine of both Asian and Afican elephants havea higher oxygen effiny than normal human heme fdobin (Brown and White, 1980, Dene eta. 1580) demonst {hat the myoglobin of E: maximus is unique among ‘Amino acd sequencing of this protein revealed that po {Ei} doce not have the dstl histidine found in al other mammals sled; instead glotarine occupies this position. ‘At bith the brain weighs about 35% of the weight of the brain ofall grown reas The dil buy he a wel ‘onvolated cerebral hemiophere, resembling tha of porpoise or fiman: the hemispheres do not cover the eerebellam, however. {Te temporal labes are proportionally vey larg, bulging oat a= Ficune 4. Occlusal views of upper molars of Elephas maximus (left) and Loxodonta africana (eight. In both photos, anterior is at top. Scales represent 50 mun (photos couresy of Smithsonian Inatitaion) erally. The olfactory lobes are also large. The trigeminal and facial nerves are lage, The brain of «19 to l0-yearold Asian tlephant, which weighed 2 tons, weighed 4.5 kg (Beddard, 1902: Evans; 1910; Harrison, 1847; Mayer, 1847; Tennent, 1867). The cranial capacity of an adult E, maximus was 6,652 ce (Osborn, 1D, ‘The epiglots is thin and flexible. The lungs are simple and slighty lobulated, Thee venteal surface is attached to the de phragm and the lateral surfaces are attached to the pleuroperi- toneum, ao there is litle space for « pleural cavity (Hareson, 1850 Mia and Greenwood. 1878; Watson, 18720. Engle (1963) noted thatthe acini of the lung were small but extremely numer ‘ous, providing an estensive respiratory surface ‘The gape of the mouths relatively ama. The checks a ‘capable of distention. Two small openings leading to Jacobson’ ‘canals ate at the anterior end of the hard palate. The soft palate je'short and there is no uvula. The tongue i thick and rounded 4a its base, tapering and pointed in front. The tip i directed dlovenwvard and Ties mostly inthe groove formed by the lower lip Large circumvallate papllge, about two to four or more, are found atthe bate ofthe tongue. The salivary glands include the parotid, Submenilary, and sublingual and a gland that is situated at the internal angle of the mandible. A pharyngeal pouch is present and is supported in part by the hyoid apparatus and the associated ‘musculature. The esophagus enters the stomach near is middle, ‘loser to the eardiae than to the pyloric end. The stomach 1 imple, not ruminant, Externally, the stomach is smooth, elon fated, and nearly straights the cardiae end is much prolonged fand tapering, Internally, 4 numberof tansverse, neatly circular folds project inwards from the cardiac wal, These folds almost Asappear when the stomach is distended (Miall and Greenwood, Wit Shimizu eta, 1960). ‘The dundenum is separate and the jejunum and ileum are ‘composed of numerous Ips. The large intestine is shorter (about 12'm) than the smal intestine (about 18m) large ia diameter; the loops end in 8 very short, sraight rectum. Avnumber of aggre- tga glands were found inthe rectum, and a special ileo-eaecal land was described, The eaccum ls lage and saceulated. The proximal end of the colon i also saceulated. The inner dorsal tral of the eaecum displays @ median fold that might indicate an ‘original paired condition. The anus is protected externally by an ‘the base of tal (Miall and Greenwood, 1878: Mitchell, 1916). The liver is simple and divided unevenly into two lobes, the left being the smaller. Vilaseior (1968) reported on a new tcellular type inthe liver. Many workers (e.g, Benedict, 1936: Evans. 1910) hase reported on the absence of «gall biadder Harrison (1847) stated that a gall bladder is rudimentary. Mi and Greenwood (1878) noted thatthe ductus cholaedocus is sac ulated within the duodenum and also receives secretion from MAMMALIAN SPECIES 182 the pancreas, The pancreas, unlike that of most mammals, does hot come in contact withthe spleen ‘The male reproductive tystem includes intraahdominal testes which are almost globular and lorated near the kidneys. Four prostate glands, two on cach side, are small each gland ‘opens into the urethra by a single and separate duct. The two Cowper's glands are oval nd flatened; they open separately into tive urethra. The penis relatively lon, «glans penis fs present, land the prepuce ix well marked. The orifice of the urethra is Yishaped; the stem of the "Ys diected ventrally. The female reproductive system includes a parly bicomuate uterus. A uro- genital canal is twice as long a® the setual vagina, The litris hhas « prepuce, ie ong, and reaches beyond the anterior end of the vulval orice. The glans clitoris is terminal and semi-globula. ‘The vulval orifice extends down between the female's hindleg, similar to but aot as far forward as the position ofthe penis. This m has resulted in some incorrect sexing of individuals c rinaty system includes labulated kidneys. The urethra ‘empties into the urogenital sinus in females. Discrepancies exist, 4a to the number and distinetness ofthe bes ofthe kidney and ‘of the numberof calices and their mode of entry into the ureter Five seems to be the common number of lobes but to to nine lobes have been reported (all and Greenwood, 1878; Watson, 18725). Endocrine glands include the hypophyss, thyroid, part: thyroid, ovary, pancreas, and adrenal (Miall and Greenwood, 1878), The thyrold and perathyroid glands were studied by Fjuita and Kamiya (1963) Kladetaky (1952) described the hypophysie as elongated, pear-shaped sieuctute attached to the brain by narrow stalk ‘Watson (1874) noted « well-developed periosteal muscle in the eye, corresponding in position toa similar muscle in sheep land deer, The nitilating membrane moves transversely. No true Tnerimal apparatus is present (Harrcon, 1850). Miall and Green wood (1876) found the Harderian gland with its excretory duct "pening on the surface of the nictitating membrane FUNCTION. The generally thick skin provides protection against bites, buraps, and adverse weather. Its folds increste ‘rface area or heat dissipation, Because of small surface area to macs ratio, elephants, ke other large mammals, ean tolerate old better than excessive heat (Benedict, 1986). The color of the ‘kin may be masked by dir hecauve of dusting and wallowing ‘which may function in thermoregulation and provide protection ‘Senin insect bites. The skin is movable and contains many nerve Centers (pressure points) used by the mahout or oozie to control trained animals (Deranivagala, 1985) Although aweat glands are not documented in literature, Carringion (1958) noted that sweat was observed on the back immediately after removal of a saddle Wild and captive elephants requite at least one bath per day. Skin temperature does not depend on ambient temperature (Ben- edict and Lee, 1938); skin varies from 240°C (at 22°C ambient) to S2.7C tat between the mother and calves as well x among herd members, Newborn may consume 7.6 109.41 of milk per day. Calves have been raised auccessflly on diluted milk of domeatic cows, om human milk, and on other formulas (Evans, 1910; Ferrer, 1947; Reuther, 1969: Sikes, 197). The hindlimbs provide both support and propulsion, and the forelimbs function 4 supports. Walking gait ic about 6.5 kav, og is swifl, and a charge may reach over 48.2 km/h (Seo 1973) The tal protects the anal opening and may be used to drive away biting insects. When in danger, elephents often run holding thet tails up, which may signal other herd members of approach ing danger. Young elephants follow their mothers or older sisters Dy holding onto their tals. This behavior has been reinforced in felrous and other captive elephants “The short neck brings the head close tothe center of gravity: Independent movements of the head are limited but the elephant ‘ean move its eyes conciderably. The trunk is a multi-purpose prehensile organ. Its highly sensitive, innervated by the mail lary division ofthe trigeminal nerve and by th facial nerve. Pinch and grasp methods used by the trunk and laterality were de Seribed by Racine (1980), Many functions are attributed to the trunk, including feeding, watering, dusting, smelling, touching, ound production/eommonication, loading, defense and offense. land washing. The acute sense of smell uses both the trunk and Jacobson's organ. Elephants are crepuscular; eyesight is poor In bright light and probably best during twilight. Hearing is acute. ‘The large pinnae intercept sound waves and aid in thermaregs- MAMMALIAN SPECIES 182 Tation. An ability to distinguish low amplitude sounds was dem. ‘onstrated by Heffner and Hefiner (1980), Sikes (1971) suggested that eranial pneumatizaion somehow improves hearing. The tongue is tactile and aids in bringing fod into the mouth. Jaws ‘move montly backward and forward sight sideways movements tate also apparent, Up t 150 kg of food and 140 Taf water may be consumed in I day (Ishwaran, 1978; MeKay, 1973; Vancuy Tenberg, 1977 ‘Tisks serve to dig for water, salt, and rocks; to debark trees: as levers for maneuvering fallen trees and branches; for work Uy domestic animals; for display: for marking trees; ax weapon of Gefense and offense; as trunkerests; as protection for the trunk and perhaps as 2 status symbol, Elephants are known to be left ‘or right tusked. Ivory has been used in art and in implements for kes. Artists prefer ivory of the Afvican elephant over that ofthe ‘Asian; they claim itis denser and, therefore, anore suitable for carving (Carrington, 1958; Kung, 1916; Wylie, 1980, Elephants incessantly move thee trunk, ears, and til. The rest of the body is telatively quiet, except during "weaving’ (that when standing elephants =hit their weight back and forth; Bene= dict, 1936), Orhan and Stohl (1962) concluded thatthe low crea tinin level in the rine is indicative of low metabolism. The total basal metabolism of one Arian elephant ¢Jup') at 20°C ambient when lying and without digestive setivity, was 99,000 caoris!day, Jap weighed 3,672 kg and had a surface areuof 2.8m". Thecefore, hher basal heat production per kg was 15.3 ealoiesday, and 2,060 calories m=day'*. A large elephant produces the heat of about 30 men (Benedict, 1986). The heart beats more slowly than that of man: 28 beatsnin when standing and 35 beats/min when lying (Benediet, 1996). the 70% of inhaled sie throogh thelr trunks. Res- Diration rates are to 5 breaths/min in lying and calm elephants fnd 10 or more in standing and awake individuals Benedict, 1936), Breathing is periormed more wih the sid ofthe diaphragms than by expanding the nib eage (Beddard, 1902), The lack of 3 pleural cevity scems to be an adaptation tothe elephants mode ft drinking (Short, 1962), ‘Only about 14% of the dry matter is digested, as compared, with 50 to 708 in eatle, sheep, and horses, The passage of food Through the intestine takes 24 h. Chemical composition of feces is much like that of poor hay (Benedict, 1936) Palm leaves (Car- ota urens) supply more than adequate cobalt forthe production Of vitamin Bye by microbial syathesis (An 1980) ONTOGENY AND REPRODUCTION, Data on cout: ship, mating behavior, natality, growth, and morality in E. max {mas in the feld are scanty. Copulation is accomplished in the usual posteriori postion of quadruped animals. Precopulatory behavive involves wresting with intertwined trunks, neck biting and ettempted mounts. During eopilation, the female is passive tnd intromission is achieved theowgh independent movement of the penis. Inromision laste less than 8» and the total duration ‘ofthe mounts about 30s. The number of mounts per ejaculation by the males varies between 2 and 4, and the number of intror missions per ejaculation from 1.t to 3.5 (Eisenberg etal, 1971, Fertilization takes place in the uterine horn and implantation of the 2ygote follows. The developing embryo soon becames en closed in membranous sacs of maternal and foetal erigin« com> posing a non-deckduate and zonary placenta (Cooper etal. 1963; Evans, 1910) The anatomy af the foetus was studied by Hil (1938) and by Deraniyagala (1959) Gestation usually lasts fom 18 0 22 months with a minimum of 17 and maximum of 24 (Deranivagala, 1955; Flower, 1943), Pregnancy is not noticeable until nearly the end of gestation Pillay (1976) diagnosed early pregnancy in elephants, using male frogs and serum of the suspected female elephants. Near the end of estation, the mammae syell, the nipples distend and curve tuutwards, end a watery Buid may ooze from them (Anghi, 1982), Maberry (1962) reported that several hours pricr to parturition 4 sreat deal of thick, rubbery mucus was passed from the vaginal ‘opening. The period of labor may be short or continue for several hours and the act of parturition lasts for n short time (ap to 5 rin), The aterbieth is usually consumed by the cov. Birth can occur at any season of the year. Usually one ealf ic born, rarely twins (Hundley, 1927) or triplets (Mache, 1916), MeKay (1973) estimated that females in favorable habitats had mean intervals between births of 2.5 to 4 years whereas others had intervals of 5 to 8 years. Newborns weigh about 100 kg (60 {0 110 kg) and measure from 75 1090 em (Flower, 1943; Hundley, 1934). Calves are extremely hairy compared to adults, The newly bora ean stand on ther feet shortly after birth and ean follow the mother in her daly routines after a few days. The infant, often swith maternal help, applies ite mouth (not the trunk) tothe nipple find may suckle from its mother or from other nursing cows ‘Young nurse for several months and then begin to eat grass and azeen foliage. They may nurse occasionally for about 18 months. Parental supervision continues for several years (Crandall, 1904; Deraaivagala, 1955; Evans, 1910, extalmatufity ally occurs in males at years 14 to 15 but smetinies as eatly as year 9. Females usyaly fest give bieth in years 15 or 16 (Flower, 1943), Evans (1910) reported a female 9 ears 1 month old giving birth, and Crandall 1968) reported an Beyearold. Male elephants can reproduce at any dine, not aly when they are in musth. The only indication of estrus is the Uttering of low sounds by the female (Benedict, 1936: Evans, 1910), The Asian elephant is polyestrous; the estrus lasts 3 to 4 days in captivity (Asdell, 1968) The interval between estrous periods i 3 weeks, with « mean of 22 days and a range of 18 to 2Bo'days, The mean duration of estrus for 6 females is 4 days but the cow is receptive to mounting only on T day (Eisenberg et al 1971; Jainudeen etal, 1971. Longevity in elephants has been exaggerated (Sanderson, 1962), Deraniyagala (1985) believed that elephant in the wild may live as long as 120 years, and Dollman (1997) reported om an elephant 170 years old. Alleged and actusl ages of elephants in captivity were given by Flower (1948). The oldest documented ge of ¢ captive Asian elephant was that af “Jesse,” in the Te ‘nga Zoological Park, Sydney, Australia; her estimated age was 69 or 77 years (Plower, 1948; Patten, 1940) Mortality in Asian elephants can be duc to predation on young, diseate end parasites, accidents drought, stress, bunting. And poaching, old age, and fighting. Morality rate was roughly festimated by MeKay (1975) for a population in southern Sri Lan Ke Given a cohort of 0, mortality may be as high a= two to three Individuals per year during the fist 10 years of ite SCOLOGY. Formerly Asian elephants used a greater var ety of habitats, Now they use the single-monsoon. dry. thorn- Setub forest in South India and Sri Lanka and effectively forage in suitable habitats in Malaya which would support multistate, evergreen forests in uclima phase (Olivier, 197&x, 19786, 1978), ‘Asian elephants favor an ecotone with an inerdigitaion of grass: Tow woody plants. and forests. Continuously forested areas with few clearings do not support high densities of elephants. Flephants drink atleast once a day and ave never far from 1 permanent source of fresh water, Shade is essential during 2 major part of the day. Elephans radiate heat from their ears, flapping rate of the ears varies with wind velocity, ambient tem petature and cloud cover (MeKay, 1973). Elephants require so dium and other trace elements (Hubback, 1999; Morris, 1988). In Sri Lanka where elephant populations wecur quite close to the coast, use of salt licks isnot as prominent a3 In areas where toils have litle sodium (Seidensticker and MeNeeley. 1973) ‘Asian elephants may feed at any time, but two major Feeding peaks accur each 28 h. Approximately 12 to 90% of an adult's Sctivities involve moving toward food and feeding. Within feed {ng bout the rate of feeding ix slow nti the animal has Ioested 4 primary source of food generally grass). Rapid feeding follows 4nd gradually declines (Eisenberg, 1980), An average adult may ingest 7 Ke af food in 1h feed IB May, and, therefore, eat 190 Kg of vegetable matter (wet weight) per day (Vancuslenberg, 1977), An elephant may feed on more than 75 different species ‘of plasts, ut preferences are shown (Iahwaran, 1978). Seasonal migratory movements of Asian elephants (Tennent, 1867) have heen seriously disturbed by human agriculture (Oly. ier, 19780). Seasonal short-range moversente of 30 to 40 km sil occur in parts of south India and Sri Lanka (Eisenberg ad Lack: hart, 1972: McKay, 1973). Movement restricted within National Parks may show a eyeic tendency, influenced by weldry cycles. lephants generally do not feed more than a few days at one place. Adult males in Sri Lanka have home ranges of 10'to 17 kn herd of 25 females and young had wet season range of 25 kim* and a dry season range of 64 kt. There is no evidence of territoriality (MeKay, 1973). Areas with prolonged drought eles generally support fewer clephants, but during the wets son temporary aggregations may elevate densities to more than en ‘Conservation and management receive much attention as wild elephant populations are dwindling. Captive breeding pro rams are partof these efforts, Crandall (1968) pointed out sone {tthe problems of managing elephants in captivity, such as dif- Feulies in controlling mature bulls. Reproduction in captivity may be expedited by arifcial insemination, Inbreeding should juvenile mortality of inbred young wa higher than bred young (Ralls et a, 1979) ‘The following ref- ferences leo disuse health and management: Clark etal (1980), Evans (1910), Fersier (1947), Kane eal. (1976) Miller (1977, into et al, (1973), Pyakural etal (1976), Ratnesar (1966), and Schmidt (1978), BEHAVIOR. Bathing and vesting may occupy 12% of an sims tes moving entber 128, an fog the rn nen ipo) Yount snus tay’ Beda men sieyng for ‘air prorated evods whee ‘wm only bey (Ror, 1960) An ad lh ‘mane from predation except by humans Thrughout the Iter Fans of tel evolution, bumnoa have ad a deft infiuence Sn elephat population. Dring the Peocene, humans begs to Kae clephons and mammctes hunting lepance fr fod, ory and domestaton han persited (Wiliams, 1980; Wyle, toa, Yeung elephants aye led by tigers sd Hons ao, Styles (180 eprted on + fullgrwn tke led by gers: 2 Tprtetive formation tn meh cove lace themes ntl hd cide wih the ats between them ot “eward the cee, has probably evened ana recpnse fo pre dation by large predators against their young. ® ‘ery ew exponents have book conducted to measure memory ce Reach, 1550, It may take an nda lpn many alt once har mastered ates, wl “meni” {hat Sick" orang time Blond, 182 Macho ta 197). ‘Al rained lpi can recognize between 6 sn i worde tnd phrsses Lon, 197), An abit opvform ate hat require sicne lance Cairn cap bat soo bvercd In Seture, Behavior in whlch an elephant appeared to tink wat Aen by Pil! (940), Ts elephant oe ocaion would Setlower «pla nts unl dog sloping i the pt was Tous snd diven aay Renach and Alevot (958) concladed that elephants sce wll a lnc range tnd ener shapes of eject fr sme ies in sabrequent stodier devebanp tone scrim, Rentch SS) concladed that elephants fave exolnt sity to

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