Open Access
25 January 2011 Rusa unicolor (Artiodactyla: Cervidae)
David M Leslie
Author Affiliations +

Rusa unicolor (Kerr, 1792), or sambar, is the largest Oriental deer. Seven subspecies occur in varied habitats and elevations from India and Sri Lanka throughout southeastern Asia. Body mass and antler length decrease from west to east. R. unicolor is considered ancestral relative to the form of its male-only antlers and social behavior. Populations are vulnerable because of overexploitation for subsistence and markets in meat and antlers. R. unicolor was elevated by the International Union for Conservation of Nature and Natural Resources from no status in 2006 to “Vulnerable” in 2008 because of >50% decline in many populations over the past 3 generations. It is well represented in zoos and private collections and is introduced in Australia, New Zealand, South Africa, and the United States.

Synonymies completed 15 September 2010

Cervus: Kerr, 1792:300. Part.

Rusa Hamilton-Smith, 1827a:104. Described as a subgenus of Cervus Linnaeus, 1758, to contain C. hippelaphus, C. unicolor, C. aristotelis, C. equinus, C. peronii, “Rusa of Malacca” (no binomial provided), and C. mariannus; type species Cervus unicolor Hamilton-Smith, 1827b, by subsequent designation (Kretzoi and Kretzoi 2000a:365, 2000b:574).

Stylocerus Hamilton-Smith, 1827b:319. Part.

Rusas Brookes, 1828:62. Incorrect subsequent spelling of Rusa Hamilton-Smith, 1827a.

Hippelaphus: Bonaparte, 1837:unnumbered page associated with fascicolo xv and xvi. Part; type species Cervus hippelaphus de Blainville, 1822, by absolute tautonomy (Palmer 1904:325); described as a subgenus of Cervus Linnaeus, 1758; preoccupied by Hippelaphus Goldfuss, 1820 and Hippelaphus Reichenbach, 1835.

Rusa: Hodgson, 1841a:219. First use as a genus.

Russa Gray, 1843:179. Incorrect subsequent spelling of Rusa Hamilton-Smith, 1827a.

Russa Müller and Schlegel, 1845:210. Incorrect subsequent spelling of Rusa Hamilton-Smith, 1827a.

Axis: Gray, 1843:180. Part, not Axis Hamilton-Smith, 1827b.

Hippelaphi Sundevall, 1846:177. Part; incorrect subsequent spelling of Hippelaphus Bonaparte, 1837; used as a subgenus of Cervus Linnaeus, 1758, to contain C. japonicus, C. duvaucelli, C. aristotelis, C. equinus, C. hippelaphus, C. moluccenis, C. peroni, C. kuhlii, C. philippinus, C. mariannus, C. lepidus, C. axis, C. pseudaxis, and C. nudipalpebra.

Cervulus: Gray, 1861:138. Part, not Cervulus de Blainville, 1816.

Rucervus: Gray, 1872:76. Part, not Rucervus Hodgson, 1838.

Melanaxis Heude, 1888a:8. Type species Cervus alfredi Sclater, 1870, by original designation.

Sambur Heude, 1888a:8. Type species Cervus aristotelis G. Cuvier, 1823, by original designation.

Roussa Heude, 1888a:8. Type species Cervus equinus Cuvier, 1823, by original designation.

Ussa Heude, 1888a:8; 1888b:22. Type species Ussa barandanus Heude, 1888a, by original designation.

Hippelaphus: Heude, 1896:49. First use as a genus.

Sambar Lydekker, 1915:91. Incorrect subsequent spelling of Sambur 144145146Heude, 1888a.

Context and Content. Order Artiodactyla, suborder Ruminantia, family Cervidae, subfamily Cervinae, tribe Cervini. Currently, 4 species of the Old World Rusa are recognized: unicolor (sambar), marianna (Philippine deer), timorensis (rusa), and alfredi (Prince Alfred's deer—Grubb 2005). The following key was developed from characteristics provided by Groves and Grubb (1987), Grubb and Groves (1983), Lydekker (1915), and Meijaard and Groves (2004), and information at (accessed 6 May 2009):

1. Small, short-legged deer; soft, fine, and spotted pelage on young and adults; shoulder height 64–71 cm; body length generally ≤ 128 cm; antler length of mature males about 24 cm; restricted to Panay and Negros islands, PhilippinesR. alfredi

Larger forms with coarse pelage and no spots on adults; shoulder height > 70 cm and body length generally > 130 cm; antler length of mature males often > 60 cm2

2. Male antlers generally thin and lacking rugosity; dorsal hairs of both sexes annulated; gregarious, occurring in herds; restricted to Java and islands of Indonesia, introduced to most of the latter by humansR. timorensis

Male antlers thicker and rugose; dorsal hairs of both sexes not annulated; not gregarious, typically occurring in mother–offspring social units with males solitary; ranging extensively from India through southeastern Asia or the Philippines3

3. Small deer; shoulder height about 70 cm; greatest length of skull 200–270 mm; mature males with antlers generally 30–45 cm in length; restricted to the PhilippinesR. marianna

Large deer, particularly western forms; shoulder height 130–142 cm; greatest length of skull 300–400 mm; mature males with antlers 70–120 cm in length (as small as 40–50 cm in Taiwan); wide ranging from India, Sri Lanka, and Nepal across southern China through southeastern Asia to the Pacific CoastR. unicolor

  • Rusa unicolor (Kerr, 1792)

  • Sambar

  • C[ervus]. Axis unicolor Kerr, 1792:300. Type localities “dry hilly forests of Ceylon [ =  Sri Lanka], Borneo, Celebes [ =  1 of 4 Greater Sundas Islands, Indonesia], and Java;” based on the “Middle-sized Axis” of Pennant (1781:106); restricted to “Ceylon” by Hamilton-Smith (1827b:310).

    C[ervus]. Axis major Kerr, 1792:300. Type localities “marshes of Borneo and Ceylon;” based on the “Great Axis” of Pennant (1781:106); restricted to “Ceylon” by Groves (2003:351).

    Cervus unicolor: Bechstein, 1799:112. Name combination.

    Cervus albicornis Bechstein, 1799:112. Type locality not given; based on the “Great Axis” of Pennant (1781).

    C[ervus]. Niger de Blainville, 1816:76. Type locality “l'Inde [ =  India].”

    C[ervus]. maximus de Blainville, 1822:264. Type localities “Ceylan ou Bornéo?”

    C[ervus]. hippelaphus de Blainville, 1822:265. Type localities “Archipel indien [ =  Indian Archipelago];” modified to “Bengal [ =  area of West Begal, India, and Bangladesh], Java, Sumatra, and other great islands of the Indian Archipelago” by Hamilton-Smith (1827a:108) and “Java? Bengal chiefly the Jungleberry district” by Hamilton-Smith (1827b:309); preoccupied by Cervus elaphus hippelaphus Erxleben, 1777:304 ( =  Cervus elaphus Linnaeus, 1758).

    cervus equinus G. Cuvier, 1823:45. Type locality “Sumatra.”

    cervus Aristotelis G. Cuvier, 1823:503. Type localities “Napaul [ =  Nepal], et vers l'Indus [ =  India];” restricted to “Bengal in the Prauss Jungles” by Hamilton-Smith (1827b:310).

    cervus Leschenauldii G. Cuvier, 1823:506. Type locality “côte de Coromandel [ =  southeastern coast of India].”

    C[ervus (Rusa)]. Hippelaphus: Hamilton-Smith, 1827b:309. Name combination.

    C[ervus (Rusa)]. Unicolor: Hamilton-Smith, 1827b:310. Name combination.

    C[ervus (Rusa)]. Aristotelis: Hamilton-Smith, 1827b:310. Name combination.

    C[ervus (Rusa)]. Equinus: Hamilton-Smith, 1827b:311. Name combination.

    Rusas hippelalphus?: Brookes, 1828:62. Name combination.

    C[ervus]. Malaccensis J. B. Fischer, 1829:451. Type locality “peninsula Malacca [ =  peninsular Malaysia]” (see “Nomenclatural Notes”).

    Cervus Jarâi Hodgson, 1831:321, pl. XXI. Type localities “sub-Himâlayan ranges, and Saul forest.”

    [Rusa] Jaraya Hodgson, 1841a:219. Name combination and incorrect subsequent spelling of Cervus jarai Hodgson, 1831.

    [Rusa] Nipalensis Hodgson, 1841a:219. Type locality “Nepal.”

    C[ervus]. heterocerus Hodgson, 1841b:unnumbered plate opposite page 722. Type locality “Nepal.”

    [Rusa] Nepalensis: Hodgson, 1841c:914. Corrected spelling of Rusa nipalensis Hodgson, 1841a.

    [Rusa] Heterocervus Hodgson, 1841c:914. Name combination and incorrect subsequent spelling of Cervus heterocerus Hodgson, 1841b.

    Russa Hippelaphus: Gray, 1843:179. Name combination.

    Rusa Aristotelis: Gray, 1843:179. Name combination.

    Rusa Equina: Gray, 1843:179. Name combination.

    Axis pennantii Gray, 1843:180. Type locality “India;” said to be Pennant's (1781:106) “Great Axis” of “Borneo or Ceylon.

    Cervus russa Müller and Schlegel, 1845:212, 217. Type locality “Java, is van daar naar Borneo.”

    C[ervus]. bengalensis Schinz, 1845:390. Type locality “Bengala? [ =  Bengal],” northwestern India.

    [Cervus (Hippelaphi)] Aristotelis: Sundevall, 1846:178. Name combination.

    C[ervus (Hippelaphi)]. equinus: Sundevall, 1846:178. Name combination.

    C[ervus (Hippelaphi)]. hippelaphus: Sundevall, 1846:178. Name combination.

    C[ervus]. Leschenaulti Sundevall, 1846:183. Incorrect subsequent spelling of Cervus leschenauldii G. Cuvier, 1823.

    Cervulus cambojensis Gray, 1861:138. Type locality “Camboja [ =  Cambodia].”

    Cervus [Rusa] swinhoii Sclater, 1862:151,152, pl. XVII. Type locality “Formosa [ =  Taiwan].”

    Rusa tarai Hodgson, 1863:ix. Incorrect subsequent spelling of Cervus jarai Hodgson, 1831.

    Rucervus cambojensis: Gray, 1872:76. Name combination.

    C[ervus]. heterocercus Jerdon, 1874:256. Incorrect subsequent spelling of Cervus heterocerus Hodgson, 1841b.

    C[ervus]. saumur Jerdon, 1874:256. Type locality not given; presented as synonym for Cervus aristotelis G. Cuvier, 1823, but attributed to Ogilby (1839:lxxii), who only used the local vernacular name “Saumer” in association with C. hippelaphus.

    C[ervus]. laschenaultii Jerdon, 1874:256. Incorrect subsequent spelling of Cervus leschenauldii G. Cuvier, 1823.

    Rusa Aristotelis, nigra: Fitzinger, 1875:284. Name combination and correction of gender concordance of Cervus niger de Blainville, 1816.

    Rusa Aristotelis, leschenaultia Fitzinger, 1875:286. Name combination and incorrect subsequent spelling of Cervus leschenauldii G. Cuvier, 1823.

    Rusa Aristotelis, unicolor: Fitzinger, 1875:287. Name combination.

    Rusa Aristotelis, heteroceros Fitzinger, 1875:289. Name combination and incorrect subsequent spelling of Cervus heterocerus Hodgson, 1941b.

    Rusa equina, malaccensis: Fitzinger, 1875:294. Name combination.

    Rusa equina, Pennantii: Fitzinger, 1875:296. Name combination.

    S[ambur]. curvicornis Heude, 1888c:41, 42. Type locality “Cochinchine [ =  17th century name for southern one-third of Vietnam];” restricted to “Tay-ninh” Province, Vietnam, by Braun et al. (2001:631).

    S[ambur]. longicornis Heude, 1888c:41, 42. Type locality “Cochinchine;” restricted to “Saigon,” Vietnam by Braun et al. (2001:632).

    S[ambur]. outreyanus Heude, 1888c:41, 42. Type locality “Cochinchine.”

    S[ambur]. planidens Heude, 1888c:41, 43. Type locality “Cochinchine.”

    S[ambur]. colombertinus Heude, 1888c:41, 43. Type locality “Cochinchine;” restricted to “Baria [ =  Bà Ria-Vung Tau]” Province, Vietnam, by Braun et al. (2001:632).

    S[ambur]. combalbertinus Heude, 1888c:41, 43. Type locality “Cochinchine;” restricted to “Baria [ =  Bà Ria-Vung Tau]” Province, Vietnam, by Braun et al. (2001:632).

    S[ambur]. heteroceros Heude, 1888c:41. Name combination and subsequent incorrect spelling of Cervus heterocerus Hodgson 1841b.

    S[ambur]. lemeanus Heude, 1888c:41, 44. Type locality “Cochinchine.”

    S[ambur]. errardianus Heude, 1888c:42, 45. Type locality “Cochinchine.”

    S[ambur]. joubertianus Heude, 1888c:42, 45. Type locality “Cambodia” based on lectotype selection by Braun et al. (2001:632).

    S[ambur]. latidens Heude, 1888c:42, 45. Type locality “Cochinchine.”

    S[ambur]. planiceps Heude, 1888c:45. Type locality “Cochinchine;” restricted to “Baria [ =  Bà Ria-Vung Tau]” Province, Vietnam, by Braun et al. (2001:632).

    S[ambur]. officialis Heude, 1888c:42, 46. Type locality “Cochinchine;” restricted to “Baria [ =  Bà Ria-Vung Tau]” Province, Vietnam, based on paratype selection by Braun et al. (2001:632).

    S[ambur]. simonianus Heude, 1888c:42, 46. Type locality “Cochinchine;” restricted to “Baria [ =  Bà Ria-Vung Tau]” Province, Vietnam, by Braun et al. (2001:632).

    S[ambur]. brachyrhinus Heude, 1888c:42, 46. Type locality “Cochinchine.”

    S[ambur]. lignarius Heude, 1888c:44. Type locality “Cochinchine.”

    S[ambur]. verutus Heude, 1888c:46. Type locality “Cochinchine.”

    Cervus brookei Hose, 1893b:206. Type locality “Mount Dulit, E. Sarawak,” Malaysia.

    Hippelaphus hamiltonianus Heude, 1896:49. Type locality “Sandakan [ =  Sahab, East Malaysia], nord de Bornéo [ =  North Borneo].”

    Rusa dejeani de Pousargues, 1896:12. Type locality “Setchuan [ =  Sichuan],” southwestern China.

    Cervus dejeani: Ward, 1896:22. Name combination.

    Russa equina: Jentink and Büttikofer, 1897:63. Name combination.

    Cervus (Rusa) Swinhoei Nitsche, 1898:32. Incorrect subsequent spelling of Cervus swinhoii Sclater, 1862.

    Cervus unicolor typicus Lydekker, 1898:146. Usage equivalent to Cervus unicolor unicolor and not intended as a new name.

    Cervus unicolor swinhoii: Lydekker, 1898:154. Name combination.

    Cervus unicolor dejeani: Lydekker, 1898:156. Name combination.

    Cervus spatulatus O. Thomas in Sclater, 1901:536. Type locality “central Borneo” (see “Nomenclatural Notes”).

    Rusa unicolor equinus J. A. Allen, 1906:464, 467. Name combination and incorrect gender concordance.

    Rusa brookei: Lyon, 1906:584. Name combination.

    Rusa unicolor: Pocock, 1910:946. First use of current name combination.

    Cervus unicolor equinus: Lydekker, 1915:78. Name combination.

    Cervus unicolor brookei: Lydekker, 1915:80. Name combination.

    Cervus unicolor swinhoei Lydekker, 1915:81. Incorrect subsequent spelling of Cervus swinhoii Sclater, 1862.

    Cervus unicolor oceana Chasen and Kloss, 1928:818. Type locality “Siberut Island, West Sumatra.”

    Rusa unicolor dejeani: G. M. Allen, 1930:15. Name combination.

    Rusa equina brookei: van Bemmel, 1949:210. Name combination.

    [Cervus (Rusa) unicolor] equinus: Haltenorth, 1963:19, 58. Name combination.

    Cervus (Russa) unicolor: Drozdz, 1973. Name combination.

    Rusa unicolor niger: van Bemmel, 1974:297. Name combination.

    Rusa equina brooki van Bemmel, 1974:297. Name combination and incorrect subsequent spelling of Cervus brookei Hose, 1893a.

    Rusa equina swinhoei van Bemmel, 1974:297. Name combination and incorrect subsequent spelling of Cervus swinhoii Sclater, 1862.

    Cervus unicolor dejeni Wang and Du, 1982:25. Incorrect subsequent spelling of Rusa dejeani de Pousargues, 1896.

    C[ervus]. unicolor deieni Wang and Du, 1982:29, fig. 2. Incorrect subsequent spelling of Rusa dejeani de Pousargues, 1896.

    Cervus unicolor hainana Xu, 1983:395. Type locality “Hainan Dao [ =  Island],” China.

    [Cervus (Rusa) unicolor] brookei: Groves and Grubb, 1987:42. Name combination.

    [Cervus (Rusa) unicolor] swinhoei: Groves and Grubb, 1987:42. Name combination and incorrect subsequent spelling of Cervus swinhoii Sclater, 1862.

    [Cervus (Rusa) unicolor] cambojensis: Groves and Grubb, 1987:42. Name combination.

    [Cervus (Rusa) unicolor] niger: Groves and Grubb, 1987:42. Name combination.

    Cervus unicolor nigar Varman and Sukumar, 1993:273. Incorrect subsequent spelling of Cervus niger de Blainville, 1816.

    Cervus unicornis Barman, Sarma, Das, and Patgiri, 1999:781. Incorrect subsequent spelling of Cervus unicolor Kerr, 1792.

    C[ervus]. u[nicolor]. malaccensis: Benirschke, 2002:unnumbered page. Name combination.

    [Rusa] albicornis: Grubb, 2005:670. Name combination (see “Nomenclatural Notes”).

    [Rusa] bengalensis: Grubb, 2005:670. Name combination.

    [Rusa] heterocerus: Grubb, 2005:670. Name combination.

    [Rusa] hippelaphus: Grubb, 2005:670. Name combination.

    [Rusa] jarai: Grubb, 2005:670. Name combination.

    [Rusa] leschenauldii: Grubb, 2005:670. Name combination.

    [Rusa] leschenaulti: Grubb, 2005:670. Name combination.

    [Rusa] major: Grubb, 2005:670. Name combination.

    [Rusa] maxima: Grubb, 2005:670. Name combination.

    [Rusa] nepalensis: Grubb, 2005:670. Name combination.

    [Rusa] nigra: Grubb, 2005:670. Name combination.

    [Rusa] pennantii: Grubb, 2005:670. Name combination.

    [Rusa] tarai: Grubb, 2005:670. Name combination.

    [Rusa] typica: Grubb, 2005:670. Name combination.

    [Rusa unicolor] brookei: Grubb, 2005:671. Name combination.

    [Rusa] hamiltoniana: Grubb, 2005:671. Name combination.

    [Rusa unicolor] cambojensis: Grubb, 2005:671. Name combination.

    [Rusa] brachyrhina: Grubb, 2005:671. Name combination.

    [Rusa] colombertina: Grubb, 2005:671. Name combination.

    [Rusa] combalbertina: Grubb, 2005:671. Name combination.

    [Rusa] curvicornis: Grubb, 2005:671. Name combination.

    [Rusa] errardiana: Grubb, 2005:671. Name combination.

    [Rusa] joubertiana: Grubb, 2005:671. Name combination.

    [Rusa] latidens: Grubb, 2005:671. Name combination.

    [Rusa] lemeana: Grubb, 2005:671. Name combination.

    [Rusa] lignaria: Grubb, 2005:671. Name combination.

    [Rusa] longicornis: Grubb, 2005:671. Name combination.

    [Rusa] officialis: Grubb, 2005:671. Name combination.

    [Rusa] outreyana: Grubb, 2005:671. Name combination.

    [Rusa] planiceps: Grubb, 2005:671. Name combination.

    [Rusa] planidens: Grubb, 2005:671. Name combination.

    [Rusa] simonina: Grubb, 2005:671. Name combination.

    [Rusa] veruta: Grubb, 2005:671. Name combination.

    [Rusa unicolor] dejeani: Grubb, 2005:671. Name combination.

    [Rusa unicolor] equina: Grubb, 2005:671. Name combination.

    [Rusa] malaccensis: Grubb, 2005:671. Name combination.

    [Rusa] oceana: Grubb, 2005:671. Name combination.

    [Rusa unicolor] hainana: Grubb, 2005:671. Name combination.

    [Rusa unicolor] swinhoii: Grubb, 2005:671. Name combination.

    R[usa]. u[nicolor]. equine Timmins, Steinmetz, Sagar Baral, et al., 2008:3. Incorrect subsequent spelling of Cervus equinus Cuvier, 1823.

    Context and Content. Context as for genus. Seven subspecies of Rusa unicolor are currently recognized (Groves and Grubb 1987; Grubb 2005; and information from Mammal Species of the World,, accessed 13 January 2009):

    R. u. brookei (Hose, 1893a:206). See above.

    R. u. cambojensis (Gray, 1861:138). See above.

    R. u. dejeani de Pousargues, 1896:12. See above.

    R. u. equina (G. Cuvier, 1823:45). See above.

    R. u. hainana (Xu, 1983:395). See above.

    R. u. swinhoii (Sclater, 1862:152). See above.

    R. u. unicolor (Kerr, 1792:300). See above.

    Nomenclatural Notes. Lydekker's (1898:145) contentions that “few groups of deer are more difficult to understand than the various kinds of sambar” and “very different views have been entertained as to whether the various modifications of the sambar type indicate distinct species, or merely races” are still valid today as morphologists (e.g., Meijaard and Groves 2004) and molecular systematists (e.g., Hernández Fernández and Vrba 2005; Randi et al. 2001) debate the monophyly of Rusa. The nomenclatural history of R. unicolor is complicated because of its wide distribution from India through southern China and southeastern Asia; its varied mass, color, and antler characteristics; similarity to and sympatry with other Asian cervids; and the rapid pace of collection, description, and publication by notable zoologists such as B. H. Hodgson (1831, 1841a, 1841b, 1841c, 1863) in Nepal and India and P. M. Heude (1888a, 1888b, 1888c, 1888d, 1896) in southeastern Asia in the 1800s. The masterful tome of Braun et al. (2001) on the extensive eastern Asian faunal collections of French Jesuit missionary, Father Pierre Marie Heude, served to clarify types and type localities of his many rusine deer. Lydekker (1915) recognized 13 subspecies of unicolor and included Philippine nigricans, nigellus, mariannus, and philippinus, and their various synonyms and name combinations, in his synonymies. Based on more recent taxonomic assessments of the rusine deer (Grubb and Groves 1983; Haltenorth 1963), Grubb (2005) did not include them as synonyms of R. unicolor. Rusine deer from the Philippines are currently considered by most to be R. marianna (Grubb 2005), albeit Francis (2009) still considered them to be R. unicolor. Additional phylogenetic analyses are still needed to clarify the phylogeny of rusine deer (Grubb and Groves 1983; Hernández Fernández and Vrba 2005).

    Van Bemmel (1949:211) gave “S. Müller & H. Schlegel, 1839–1844” as the authorities and dates for Russa, but I credited Gray (1843) with this incorrect spelling of Rusa Hamilton-Smith, 1827a. Confusion exists because of the different dates that the various volumes and parts of Verhandelingen over de Natuurlijke Geschiedenis der Nederlandsche Overzeesche Bezittingen, edited by C. J. Temminck, were complied and published (Husson and Holthuis 1955). Mammals were presented in a Zoologie volume with 12 numbered parts issued in 1839–1845, and Russa was mentioned on page 206 in Number 12, published 26 June 1845 (later than Gray 1843), based on Husson and Holthuis' (1955) evaluation of original wrappers in the Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands.

    International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature 1999:36) outlines protocols for gender determination of generic names formed from words that are not Latin or Greek. Further, Article 30.2.3 states that “if no gender was specified, the name takes the gender indicated by its combination with one or more adjectival species-group names of the originally included nominal species.” Because Latinized species names that Heude (1888c) associated with Sambur were masculine, or neuter, and I could find no specific statement by Heude regarding its gender (Article 30.2.2), his use of Sambur must be assumed to be masculine. Lydekker (1915) did not present both spellings, so his use of Sambar was considered a subsequent misspelling (A. L. Gardner, pers. comm.). Grubb (2005:671) intentionally changed the masculine gender of 11 of Heude's (1888c:42–46) species names to feminine, which was warranted to concord with his combination of Heude's (1888c) species names with Rusa, which he assumed to be feminine and rooted in Latin.

    Unlike Lydekker (1915:78) and Grubb (2005:671), I credited the synonym malaccensis to J. B. Fischer (1829) rather than F. Cuvier (1823). I could not find any reference to malaccensis after review of the ruminants in the classic Histoire Naturelle des Mammifères of É. Geoffroy Saint-Hilaire and F. Cuvier. This “publication” contains a very complicated collection of series (4), tomes (7), livraisons ( =  deliveries; 72), and plates (431), which were issued, numbered, renumbered, and dated from 1819 through 1842. Frédéric Cuvier authored some of the unnumbered pages of descriptive text associated with the plates that were apparently sold as separate livraisons; some plate-associated texts were simply dated with no author. From 1824 to 1842, livraisons were grouped and dated into 7 tomes, and finally, in 1842, all plates were numbered 1–431. Lydekker (1915:78) gave the abbreviated credit for malaccensis as “F. Cuvier, Hist. Nat. Mamm. vol. i, pl. x, 1824.” Similarly, J. B. Fischer (1829:451) provided “Fr. Cuv. et Geoffr. Mamm. fasc. 10.” In Tome Premier dated 1824, the “Biche de la Presqu'ile de Malac[c]a [ =  French for Malayan Peninsula]” was presented in Livraison X and individually dated September 1819. Nowhere on the plate or in the associated descriptive text (which is not attributed to Cuvier alone), or in various tables of contents and indexes, was the epithet malaccensis presented. In fact, in the 1842 retrospective “Table Générale et Méthodique,” this plate was numbered 358, attributed to Tome Deuxième, and aligned parenthetically with Cervus hippelaphus.

    Unlike Grubb (2005), Corbet and Hill (1992:256) included “Cervus spatulatus Thomas, 1901” as a synonym for Rusa unicolor. While Thomas (1901:284) “exhibited and made remarks on a peculiar Stag's frontlet and horns which [were] obtained from Mr. Charles Hose in Borneo,” he did not use the specific epithet Cervus spatulatus, at least in that published record. Oldfield Thomas was a prolific writer and published 1,090 papers from 1879 to 1929, many of them descriptions of new mammalian species from specimens sent to the British Museum (Hill 1990). Thomas withdrew proofs of only 2 of his many papers during that time period, and one of them included a never-published, but presumably more complete, description of Cervus spatulatus “with three illustrations intended for the Proceedings of the Zoological Society of London” (Hill 1990:31, 49). In the same year, condensed and paraphrased minutes of the Zoological Society of London, dated 18 June 1901 and authored by Secretary Philip L. Sclater were published in Zoologischer Anzeiger along with minutes from other European societies. In those secondarily published minutes, Sclater (1901:536) used the name Cervus spatulatus, provided a more detailed description of its “antlers” than appeared in Thomas (1901)—perhaps paraphrasing the withdrawn proofs, and gave the type locality as “Central Borneo,” albeit Thomas (1901:284) originally included “from a hitherto unvisited part of Borneo, the Pa Bauan country in the far interior.” Furthermore, Sclater (1901:536) elaborated well beyond Thomas (1901:284) by stating the antlers differed from “all other known deer in being highly complicated and many-branched, with the upper portion curved forward, and the brow-tines developed into broad horizontal paddle-like structures,” not at all like R. unicolor, the only large cervid that occurs on Borneo (see Cervus brookei Hose, 1983b). In describing Thomas' withdrawn paper, Hill (1990:31) said the antlers were “much deformed,” suggesting that caused Thomas to withdraw his proofs but not before Sclater perhaps had seen them and incorporated a description into his summary minutes for Zoologischer Anzeiger. Sclater and Thomas were close colleagues and had collaborated around the same time on their now classic, 4-volume The Book of Antelopes. Hill (1990:31) concluded that the name Cervus spatulatus “remains available in the literature, although undiscovered.” Under Article 50.1.1 of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature 1999), attributing Cervus spatulatus to Thomas is warranted if it was clearly attributed to Thomas in Sclater's publication and if Thomas previously satisfied 2 of 3 criteria of availability by offering a name (he did not), a description or indication (he did by the standards of the day), or a publication (he did—Thomas 1901; see Leslie and Sharma 2009:2–3 for a similar case involving the authority for Tetracerus).

    Hamilton-Smith (1827a) adapted rusa, the common Malayan and Bahasan (Indonesian) name for deer (also rusa-etam), as the generic epithet, and Kerr's (1792) unicolor is Latin for 1 color. The common name sambar is Hindi for a vegetable stew with a mix of spices called sambar powder, which gives the stew a yellowish brown color, similar to the pelage hue of R. unicolor in parts of India. Because of the wide distribution of R. unicolor, it has many common names. In German, it is often called Aristotle-Hirsch, in reference Aristotle's knowledge of the species, or Pferdehirsch, referencing its long legs (Benirschke 2002). The large preorbital gland, everted during rut, no doubt gave rise to the Chinese vernacular name, four-eyed deer (Swinhoe 1862). Whitehead (1993:511–512) provided a comprehensive listing of common names: for example, con hai (Annamese); menjangon (Bahasa Java); payoh (Bornean); connai (Burmese); hai-lu, shui-lu, twahé ( =  mountain horse; Chinese); jerao, jerrow (Himalayan); sabhar, sámar (Hindi); mila, kada maan (Tamil) kadaba, kadave, kadavay, kaddama (Kanarese); kullay marn, rusa etam (Malayalam); meru, samba (Marathi); jarai (female), jarao (male; Nepalese); gona, marrei (Sri Lanka); hei-lu, hei-lu-tsze (Sichuan); cheeang, tamil (Taiwan); and kwang, kwang-paa (Thailand). R. unicolor has been featured on at least 23 stamps issued by 10 countries from 1894 to 1978 (Whitehead 1993).


    All 4 species of Rusa (alfredi, marianna, timorensis, and unicolorGrubb 2005) are allopatric and vary greatly in mass, forming a morphocline (Groves and Grubb 1987). R. unicolor can be distinguished from other species of Cervini by their “robust, rugose antlers with a long [acutely angled—Pocock 1933] brow tine, very deep lacrimal pits, reduced auditory bullae, and dark eumelanic pelage” (Groves and Grubb 1987:42; Fig. 1). R. unicolor in the western part of its range is the largest Oriental deer (Pocock 1943a), and it can be distinguished from comparably sized Eurasian red deer and North American elk (Cervus elaphusClutton-Brock et al. 1982; Geist 1998; Peek 1982; Whitehead 1972, 1993) by its 3-tined antlers, near-uniform pelage without a pronounced rump patch, and a long, dark tail.

    Fig. 1

    Mature male Rusa unicolor in Ranthambhore National Park, Rajasthan, northern India; note the neck “ruff” and simple configuration of the antlers and the acutely angled left brow tine (right brow tine broken off). Photograph by Chris Brunskill ( used with permission.


    Along with taxonomic treatments, early accounts by sportsman-naturalists contain useful descriptive information on R. unicolor (Baker 1855, 1898; Brander 1923; Fletcher 1911; Gilbert 1888; Glasfurd 1896; Jerdon 1874; Peacock 1933; Stebbing 1911), which also is featured in more colorful accounts of hunting “man-eating” predators, notably from India (Sivaramakrishnan 2008; Smith 1904). Most contemporary scientific information on wild R. unicolor comes from India (e.g., Johnsingh 1983; Schaller 1967) and Sri Lanka (e.g., Eisenburg and Lockhart 1972; Kurt 1978), but considerable ecological, behavioral, and physiological insight comes from research on introduced populations (free-ranging and captive) in Australia (e.g., Slee 1984) and New Zealand (e.g., Semiadi et al. 1994b, 1995a, 1995b, 1996) and on a single island off the Florida Coast in the United States (Flynn et al. 1990; Lewis et al. 1990; Shea et al. 1990). Syntheses of that information are provided to supplement missing information when likely comparable to native populations of R. unicolor.


    Body mass and antler length of Rusa unicolor are highly variable and generally decrease from west to east across its distribution (Geist 1998; Pocock 1942a). Sexes of R. unicolor are distinguished by larger male mass (e.g., males in India 225–320 kg; females < 225 kg—Sankar and Acharya 2004), male-only antlers on short pedicles, and generally lighter color of females and young (Blanford 1888; Brander 1923; Jerdon 1874; Lydekker 1898, 1915; Fig. 2). Across the range of R. unicolor, head and body length is 162–246 cm, tail length is 25–30 cm, and shoulder height is 102–160 cm (Nowak 1999).

    Fig. 2

    Mature female and young-of-the year Rusa unicolor foraging on aquatic vegetation in Ranthambhore National Park, Rajasthan, northern India, January–February 2000. Photograph courtesy of James Warwick ( used with permission.


    Antlers of mature males are unique among cervids (Fig. 1), considered to exhibit an ancestral condition (Pocock 1942a), generally only 3-tined, rough, and corrugated as males age, often robust, and “consist[ing] of an anterior more or less straight brow tine, that comes off at an acute angle (Blanford 1888:543; Pocock 1933) to the main or posterior beam, which forks [typically] but once” (Allen 1940:1169; Jerdon 1874). The “anterolateral tine” tends to follow the main beam and is typically longer that the “posteromedial” tine (Groves 2003:351), but not always (Whitehead 1972). In exceptional cases, the brow tine can be about 50% of total length of the main beam (Brander 1923; Gilbert 1888); 10% of males can have a 4th tine on 1 antler (Ward 1896) and rarely the brow tine is bifurcate (Brander 1923). The space between antlers is V- or U-shaped, and tips of tines are often inturned (Brander 1923; Downes 1983b:figure 12; Lydekker 1898, 1915). Mean record antler length of R. unicolor is 109.8 cm, exceeded by only red deer–North American elk and caribou (Rangifer tarandus) among cervids (Clutton-Brock et al. 1980; Whitehead 1993).

    There is considerable variation in mass and color of the various subspecies of R. unicolor (Geist 1998). Early general descriptions of R. unicolor were provided by Allen (1940), Blanford (1888), Jerdon (1874), Lydekker (1898, 1915, 1916), and Pocock (1943a, 1943b): large robust deer; pelage coarse and shaggy, particularly on males in rut and winter; coat color basically uniform around the body with considerable variation in color ranging from yellowish brown to brown or almost “black or dark salty grey” (Blanford 1888:543); belly often as dark as the rest of the body, or darker, but sometimes “chestnut or whitish on the inner side of buttocks, and on the under parts” (Lydekker 1898, 1915:73); females and young generally lighter in color and young not spotted, but very faintly spotted in Malaysia (Lydekker 1898), with a dark dorsal line (Brander 1923); sometimes inconspicuous light to rusty brown rump patch but not as pronounced as in other cervids (Schaller 1967) or completely absent (Swinhoe 1862); throat and neck “with unkempt ruff” of long hairs, particularly developed on males and sometimes tipped in gray (Blanford 1888; Schaller 1967:135); ears large (17.8–20.3 cm—Jerdon 1874), broad, and obovate (Brander 1923), about one-half the length of the head (Lydekker 1915) and whitish on the back at the base (Schaller 1967) and inside, resembling “the frosted terminal leaves of a young teak tree, amongst which [they] are found” (Brander 1923:169); black-tipped tail, long (30.5–33.0 cm—Jerdon 1874) compared with other cervids, rather bushy, and whitish underneath; lack of nasal scent glands (Atkeson et al. 1988); pronounced eversible (Lydekker 1898) preorbital glands and associated deep lachrymal fossa, twice the size of those of red deer (Blanford 1888; Pocock 1910, 1943b); supraorbital glands lacking; metatarsal glands with tuft of hair but sparse (Pocock 1910); caudal glands with secretion granules of 20 nm (Evgenjeva 1991); and a “sore spot” on the throat, presumably a dermal gland associated with rut (Brander 1923; Geist 1998; Peacock 1933; Schaller 1967; Thom 1937; see “Function”).


    Rusa unicolor is the most widespread deer in Asia (Corbet and Hill 1992; Fig. 3) and occurs from southern Nepal (Chesemore 1970; Dinerstein 1979, 1980; Seidensticker 1976a, 1976b), India (Menon 2009; Sankar 2008; Schaller 1967), Sri Lanka (Eisenburg and Lockhart 1972), and Burma (U Tun Yin 1967) throughout southern China (MacKinnon 2008; Ohtaishi and Gao 1990) and southeastern Asia to the Pacific Coast and the islands of Borneo, Hainan, and Taiwan (Hsu and Agoramoorthy 1997; Timmins et al. 2008; Whitehead 1972, 1993). It occurs from sea level at various places in southeastern Asia to about 3,000 m in the Indian Himalayas (Green 1985) and Burma (U Tun Yin 1967) and to about 3,500 m in Taiwan (Whitehead 1972). Largely because of excessive harvest and habitat loss (Timmins et al. 2008), R. unicolor is now rare in Bangladesh (Basbar et al. 2001), Thailand (Ngampongsai 1987), Laos (Timmins and Evans 1996), and Vietnam (Khun and Kan 1991).

    Fig. 3

    Distribution of Rusa unicolor in India, Sri Lanka, southern China, and southeastern Asia adapted from Ohtaishi and Gao (1990), Timmins et al. (2008), and van Bemmel (1974). The general distributions of the 7 subspecies of R. unicolor are: 1, R. u. unicolor; 2, R. u. dejeani; 3, R. u. cambojensis; 4, R. u. hainana; 5, R. u. swinhoii; 6, R. u. equina; and 7, R. u. brookei.


    Intentionally introduced R. unicolor or deer-farm escapees have established populations in Victoria, New South Wales, French Island (Victoria), Western Australia, and the Northern Territory of Australia (Bentley 1957; Downes 1983a, 1983b; Moriarty 2004; Slee 1984; Whitehead 1993; Yamada et al. 2003). More than 70,000 individual R. unicolor have been released or escaped in Australia since the 19th century; such activities have resulted in 8 established populations (Moriarty 2004). Similarly, introduced populations occur in New Zealand (Douglas 1983; Forsyth and Duncan 2001; Fraser et al. 2000; Harris 1971; New Zealand Department of Conservation 2005; Nugent et al. 2001; Riney 1957; Veblen and Stewart 1982); California (Hopkins 2005; Presnall 1958), Florida (Davidson et al. 1987; Lewis et al. 1990), and Texas (Ables and Ramsey 1972; Mungall 2007; Richardson 1972) in the United States; and West Cape Province, South Africa (Lever 1985). Introduced populations are mainly of Indian or Sri Lankan origin (Lever 1985). An unknown number of the Philippine deer (R. marianna) were introduced along with R. unicolor in Victoria, Australia, in the 1860s, but the current free-ranging population there is considered typical of R. unicolor (Downes 1983b). Similarly, some Philippine deer from Java were introduced in New Zealand, and likely interbred with R. unicolor (Harris 1971); literature pertaining to these populations was used only if the investigators identified R. unicolor as the study species. R. unicolor failed to establish itself after introduction in Tasmania in the early 1890s, and the status of a private herd in Rio Tietê, Brazil, is unknown since the property was liquidated in 1986 (Whitehead 1993). Given current taxonomic distinctions, references to introductions of R. unicolor in Guam are incorrect; the Philippine deer was introduced there in 1770s (Conry 1988).


    Cervidae is a species-rich family of Eurasian origin (Gilbert et al. 2006) that radiated from tropical to temperate climates in the Pleistocene (Geist 1998; Groves 2007) and contains as many as 26 fossil genera and 80 fossil species from China alone (Dong 1993). Paleomerycines (e.g., Amphitragulus), procervulines (e.g., Procervulus), and cervulines (e.g., Eostyloceros) of the Miocene were likely precursors of species now included in the family Cervidae (Flerov 1952; Kurtén 1968; Miyamoto et al. 1990 cf. Gilbert et al. 2006), with fossil species of Cervavitus giving rise to tribe Cervini (Di Stefano and Petronio 2002; S. Mattioli, pers. comm.; Petronio et al. 2007). Despite a relatively recent radiation into South America during the Pliocene (Eisenberg 1987; Gilbert et al. 2006), Cervidae is mainly a family of the Northern Hemisphere (Geist 1998; Gentry 2000; Janis and Scott 1987; Webb 2000); the only cervid in Africa, Cervus elaphus barbarus (Barbary red deer), may have been introduced by humans from Europe along the southern Mediterranean coast as long as 8,000 years ago (V. Geist, in litt.). Despite availability of numerous cervid fossils, particularly antler parts, considerable speculation remains, much of it focused on the particularly active Plio-Pleistocene period and a transition of herbivorous species from forest to grasslands (Azzaroli et al. 1988; Di Stefano and Petronio 2002; Gilbert et al. 2006; Lister 1987, 1993).

    During the upper Pliocene, rusine deer were found in Europe (Di Stefano and Petronio 2002; Lydekker 1898; van Bemmel 1974), and early lower Pleistocene forms with 3-tine antlers, such as the Philis deer (Cervus philisi [ =  etuerarium or rhenanus]), have purported affinities with living Rusa (Kurtén 1968; Lister 1987). R. unicolor is among the most ancestral of living cervids, with characteristics little changed from the late Pliocene and paralleling other Chinese pliocervines (Petronio et al. 2007). It likely evolved in southern tropical areas (Flerov 1952), perhaps from the extinct Pleistocene forms such as Epirusa hilzheimeri (Di Stefano and Petronio 2002; Zhdanski 1925) or Eucladoceros (Geist 1998; Grubb 1990; Koizumi et al. 1993). Di Stefano and Petronio (2002) proposed that Rusa ( =  Cervus) elegans branched in the mid-Villafranchian, 2.0–2.5 million years ago, giving rise to the extinct R. hilzheimeri, which gave rise to R. unicolor and the high-elevation specialist Przewalskium albirostre of the Tibetan Plateau (Leslie 2009; Schaller 1998).

    Five rusine fossil species (elegans, microta, stehlini, unicolor, and yunnanensis) from the early Pleistocene have been found throughout China; R. timorensis apparently did not appear until the late Pleistocene (Dong 1993). A fossil Rusa, which was comparable but larger than extant R. unicolor, has been found in caves in northern Vietnam, dated from the middle Pleistocene 80,000–169,000 years ago (Bacon et al. 2004). In Sichuan, China, antlers of R. unicolor from the middle Pleistocene possibly were worked by humans prior to fossilization (Hooijer 1951). Bones of R. unicolor also occur in caves of Paleolithic origin in China (Huang et al. 1995; Si et al. 1993).



    The pelage of Rusa unicolor is generally shaggy and coarse, and individual hairs are “not distinctly banded with different coloured rings” (Lydekker 1898:145). Hairs on R. unicolor are thickest on the neck, back, and abdomen; thinner on dorsal side of the tail and legs; and thinnest around the preorbital glands, temples, and gaskins (lower thighs—Sokolov et al. 1987). Guard hairs of R. unicolor have a mean medullary width of 186 µm and mean cortical width of 29.0 µm (Sheng et al. 1993). Measurements (µm) of axial hairs vary depending on the location on the body; ventral neck hair: cortical thickness  =  15.0, medullary diameter  =  75.0, root diameter  =  147.0, hair diameter above the root  =  93.3; ventral tail hairs: 10.6, 75.0, 166.6, 107.0; lateral gaskin hairs: 10.0, 40.0, 86.7, 66.5; lateral thigh hairs: 10.0, 37.5, 80.0, 60.0; abdomen hairs: 10.0, 75.0, 100.0, 66.6; preorbital region hairs: 10.0, 22.2, 63.3, 50.0; and forehead hairs: 12.5, 55.0, 70.0, 43.3 (Sokolov et al. 1987).

    Related to the tropical origin of R. unicolor, development of the undercoat in young R. unicolor is modest or lacking; 2 of 4 captive 165-day-old (± 9.8 SE) R. unicolor did not have an undercoat, and the proportion of the undercoat relative to the total pelage weight on the other 2 young was only 0.7% (Semiadi et al. 1996). Pelage characteristics of those young R. unicolor, when present, were: depth of fiber undercoat, 26.8 mm; weight of fiber undercoat, 114 g/m2; length of fiber undercoat, 20.3 mm; length of guard hairs, 44.0 mm; diameter of undercoat fibers, 18.0 µm; and diameter of guard hairs, 277.0 µm (Semiadi et al. 1996). Shedding is said to occur in “large tufts … the old hair coming away in sections” (Brander 1923:169).

    Although numerous antler measurements have been published for R. unicolor (e.g., Downes 1983b; Lydekker 1898; Pocock 1943b; Whitehead 1993), skull measurements are less common. Nasal bones on the skull of R. unicolor “develop a plate at the posterior expansion, which tends to grow over the lachrymal vacuity” (Lydekker 1898:146, 1915); the vacuity is longer than in other cervids; and the skull has a robust appearance (Pocock 1943b; Fig. 4). The small auditory bullae rarely project below the basioccipital, and the preorbital “gland-pit” is large, usually exceeding the diameter of the orbit (54–64 mm in India—Pocock 1943b:27). Representative condylobasal lengths of skulls (mm, sexes combined) are 350–408 from India, 332–395 from Sumatra, 330–345 from Borneo, and 312–328 from Taiwan (Lyon 1906; Pocock 1942b, 1943b), illustrating the general decrease in size from west to east (Geist 1998). More recent representative mean skull measurements (mm ± SD) of R. unicolor include: greatest length of skull, 357.5 ± 35.0; condylobasal length, 340.5 ± 35.3; basal length, 318.6 ± 33.0; palate length, 212.0 ± 22.5; condylar breadth, 65.4 ± 7.1; rostrum length, 199.1 ± 26.7; nasal length, 104.7 ± 14.7; biorbital breadth, 139.4 ± 14.8; maximum breadth of nasals, 39.4 ± 7.4; interorbital breadth, 88.9 ± 13.5; and braincase breadth, 86.7 ± 7.5 (n  =  27–30, males and females combined—Meijaard and Groves 2004).

    Fig. 4

    Ventral, dorsal, and lateral views of skull and lateral view of mandible of male Rusa unicolor brookei (British Museum [Natural History], specimen with pedicels after antler shedding, collected in Sarawak, northwestern Borneo. Greatest length of skull is 354 mm.


    Unlike New World deer, new antlers of Old World deer such as R. unicolor begin to grow immediately after the previous antlers are shed (Geist 1998). During growth, antlers are covered in modified skin, or velvet, that nourishes the growing bone (Bubenik 1993); developing antlers and velvet of various cervids are prized in Asian medicine (Peacock 1933; Thom 1937) and grown commercially for that market (Luick 1983). Early anecdotal accounts suggested that R. unicolor did not replace its antlers annually (Baker 1898; Fletcher 1911; Gilbert 1888; Phythian-Adams 1951), but this is not the case and is probably related to the lack of a pronounced breeding season and observations of males in various stages of antler growth and ossification throughout much of the year (Brander 1923; Putman 1988; Schaller 1967; Thom 1937)—a pattern that persists even if males are transplanted to more temperate areas (Lao 1968).

    Unlike other cervids, particularly northern boreal species, antler bone of R. unicolor, as well as that of primitive species of Axis and Rucervus, has a very thick cortex suggesting rapid building processes of the osteon; a tubular cavity of 8 mm in diameter, apparently a blood sinus suggesting that the some bone remains alive after shedding of velvet; and concave seals in young males changing to flat seals in prime males (Acharjyo and Bubenik 1983). Males can be grouped into general age classes ≤ 6 years of age based on their antler characteristics (Downes 1983b). Length and mass of antlers increase with age but most notably in a male's 7th year; thereafter, length and mass change little (Downes 1983b), although Baker (1898) contended that maximum antler size (and mass) may not be achieved until a male is about 10 years old. Yearling males have a single spike antler, and 2- to 3-year-old males have a brow tine and main beam; both tend to have smooth antlers. By 4 years of age, males have “roughly corrugated” antlers with the typical 3 tines (Brander 1923:171; Downes 1983b). Yearling and adult females can be differentiated based on body mass, but no external characteristics are useful for aging adult females.

    The dental formula of adult R. unicolor is typical of cervids: i 0/3, c 1/1, p 3/3, m 3/3, total 34 (c1 incisiform). Upper canines may be absent; only 2 of 83 R. unicolor skulls had them in Sariska Tiger Reserve, India (K. Sankar, pers. comm.). Eruption of the 1st permanent teeth (m1–M1) begins at about 1 month, followed by m2–M2 at about 8 months and then i1–I1 at about 11 months; all permanent teeth are present by about 2.5 years (Slee 1984)—a pattern useful for aging (Humphries and Rowler 1976). Molars are notably hypsodont with “small accessory columns” (Blanford 1888:543); 2 or 3 cusps occur on the occlusal surfaces of the premolars and molars forming sharp lingual crests; and incisors are “hockey stick like” in shape (Shalini et al. 2004a:1102). Representative tooth measurements (mean ± SE, cm) of an adult R. unicolor are crown length: p1  =  0.60 ± 0.01, p2  =  1.36 ± 0.01, p3  =  1.30 ± 0.04, m1  =  1.66 ± 0.11, m2  =  1.83 ± 0.02, and m3  =  1.95 ± 0.04; and crown width: p1  =  0.56 ± 0.03, p2  =  0.77 ± 0.5, p3  =  0.73 ± 0.05, m1  =  1.06 ± 0.03, m2  =  1.32 ± 0.01, and m3  =  1.42 ± 0.02 (Shalini et al. 2004a).

    Tooth eruption-wear and cementum annuli of R. unicolor in Gir National Forest, India, were highly correlated (r2  =  0.974—Berwick 1974), but irregular layering of cementum was reported in R. unicolor from Ruhuna National Park, southern Sri Lanka, making interpretation of age more difficult (Ashby and Santiapillai 1986). Patterns of growth layers in dental cementum of the 1st mandibular molar of R. unicolor relative to age have been investigated in New Zealand and are comparable to those of red deer (Douglas 1970; Slee 1984). Flynn et al. (1990) noted that it was difficult to age R. unicolor from tooth eruption and wear alone; estimates could be off by 12 months.

    Anatomy of the 4-chambered stomach, particularly the rumenoreticulum, and poorly developed papillae on the roof of the rumen suggest that R. unicolor should be a grass-roughage feeder (Stafford 1995). Various attributes of each chamber are rumen: volume  =  9–20 l; reticulum: volume  =  0.2–1.0 l, wall thickness  =  1.5–4.0 mm; omasum: volume  =  0.2–1.1 l, mass  =  360–568 g, wall thickness  =  2.0–2.75 mm; and abomasum: volume  =  0.4–1.2 l, length  =  25–35 cm (9 male and 4 female R. unicolor, aged 1–5 years—Stafford 1995). Lengths (cm) of lower digestive organs of R. unicolor are: small intestine, male 480 and female 360; large intestine, male 450 and female 360; and cecum, male 45 and female 35 (Stafford 1995). Compared with other ruminants, those lengths are very short, and the near 50∶50 ratio of small to large intestinal length of R. unicolor is in contrast to that of the red deer, a true grass-roughage feeder, which has a ratio of about 75:25 (Dryden 2008; Stafford 1995). Combined, intestinal characteristics of R. unicolor suggest a species capable of efficiently digesting diets of grasses and browse, or an intermediate feeder. Morphology of the scapula (Sarma et al. 2003), atlas (C1 vertebra), and axis (C2—Shalini et al. 2004b) of R. unicolor also has been described.

    The reproductive tract of female R. unicolor generally is comparable to that of other cervids (Plotka 1999). R. unicolor has a bicornuate uterine horn with 3 mesometrially located caruncles, and the 3 associated cotyledons, in each horn; the 6 placental cotyledons are large (late gestation: 300–320 g and 5 by 9 cm); and the placenta implants on the 3 mesometrial caruncles of both horns (Benirschke 2002). A single corpus luteum was noted per pregnancy in 5 females from Perak, Malaysia (Khan and Khan 1968). The uterus of a near-term female R. unicolor from the San Diego Zoo was 130 cm long, and the placenta was epithelio-chorial without invasion into the maternal tissue (Benirschke 2002). The 7-cm umbilical cord in that gravid female was not spiraled, contained 4 blood vessels, and had a large, highly vascularized allantoic duct (Benirschke 2002).

    Mean (± SD) testicular and semen characteristics of 15 male R. unicolor from Taiwan include: single testicular volume, 126.9 ± 72.7 cm3; scrotal circumference, 23.5 ± 5.5 cm; semen volume, 1.4 ± 0.7 ml; spermatozoa concentration, 557 ± 344.8 × 106/ml; spermatozoa motility, 75.5% ± 9.3%; and normal spermatozoa morphology, ≥80% (Wu et al. 2002). Mean (µm ± SD) lengths of individual spermatozoa include: total length, 65.4 ± 1.7; head length, 9.3 ± 0.5; midpiece length, 12.9 ± 0.7; and tail length, 43.2 ± 1.8 (Wu et al. 2002). Fetal testes of R. unicolor have binucleated trophoblastic cells on the villi and histological characteristics that suggest a lack of active fetal gonadotropins (Benirschke 2002).


    Mean (± SD) hematological values of Rusa unicolor are: hematocrit, 37.7% ± 3.2%; red blood cells, 5.70 ± 0.22 × 106 cells/ml; hemoglobin, 13.0 ± 0.9 g/dl; mean corpuscular volume, 66.0 ± 3.2 µm3; white blood cells, 4.79 ± 1.18 × 103 cells/ml; eosinophils, 2.58% ± 2.75%; basophils, 1.43% ± 1.68%; and monocytes, 1.82% ± 0.76% (n  =  3 adults—Peinado et al. 1999a). Representative mean (± SD) blood chemistry and serum values of R. unicolor are: aspartate aminotransferase, 24.4 ± 4.1 IU/l; alanine aminotransferase, 28.7 ± 4.7 IU/l; creatinine phosphokinase, 205.0 ± 122.0 IU/l; lactic dehydrogenase, 452 ± 49 IU/l; gamma glutamyl transpeptidase, 38.1 ± 19.6 IU/l; alkaline phosphatase, 197 ± 74 IU/l; glucose, 8.50 ± 1.32 mmol/l; urea, 6.94 ± 2.77 mmol/l; uric acid, 29.1 ± 10.7 mmol/l; creatinine, 238 ± 23 mmol/l; cholesterol, 1.49 ± 0.42 mmol/l; triglycerides, 0.13 ± 0.05 mmol/l; protein, 7.45 ± 0.34 g/dl; albumin, 71.9% ± 5.7%; albumin–globulin ratio, 2.6 ± 0.7; and osmolality, 289 ± 6 mOsm/kg (n  =  3 adults—Peinado et al. 1999b).

    Unique among cervids, reproductively active male R. unicolor, females during late pregnancy and lactation, and even young-of-the-year can have a “sore spot” of 10–15 cm (Davar 1938; Geist 1998; Kurt 1978; Peacock 1933; Schaller 1967; U Tun Yin 1967) on their throats, ventrally halfway down the neck (Evans 1912). This spot is apparently glandular, centered in a whorl of hair, and often, but not always, exuding “whitish-looking oily or watery substance” from a blood-red spot (Davar 1938; Evans 1912; Morris 1938; Phythian-Adams 1951; Thom 1937:315). Thom (1937:313–317) summarized the colorful early speculation on the cause or function of the sore spot, which included disease (e.g., leprosy in Thailand), consumption of “wild olives,” ticks or some other parasite (Davar 1938; Evans 1912; Peacock 1933; Whitehead 1972), irritation from rubbing or moving through thick coarse grass, or a wound from an attack by a marten, likely Martes flavigula. The sore spot apparently does not occur (or has not been observed) on R. unicolor in Sri Lanka (Kurt 1978), among introduced populations on St. Vincent Island, Florida (Shea et al. 1990), and Australia (Downes 1983b), or under some captive conditions (Evans 1912; Mary and Balakrishnan 1984; Thom 1937; U Tun Yin 1967). Richardson (1972:58) noted that all sexes and ages of introduced R. unicolor in Texas had a “whirl of hair” on the throat that was “relatively bare,” but he did not notice any secretions or blood. Perhaps this “gland” does not manifest itself when densities are relatively high or group size is large, as in Sri Lanka or under confinement, suggesting a role in communication (scent dispersal—Mary and Balakrishnan 1984; Schaller 1967), particularly during breeding and at low densities (Geist 1998). The sore spot is associated with the breeding season in some places in India (Johnsingh 1980), but it does not bleed or secret any substance, even during breeding, in western or northern India (K. Sankar, pers. comm.).

    Muscle tissues of R. unicolor vary (mean ± SD) in their composition: longissimus dorsi, 74.40% ± 0.29% moisture, 2.93 ± 0.41 g fat/100 g dry weight, 21.99 ± 0.34 g fat/100 g wet weight, 101.30 ± 0.39 g cholesterol/100 g dry weight, and 4.73% ash, and biceps femoris, 74.52% ± 0.42% moisture, 4.06 ± 0.41 g fat/100 g dry weight, 21.83 ± 0.11 g fat/100 g wet weight, 79.52 ± 0.21 g cholesterol/100 g dry weight, and 2.35% ash (Dahlan and Norfarizan-Hanoon 2008). Detailed information on fatty acid profiles in muscles of R. unicolor was provided by Sinclair et al. (1982) and Dahlan and Norfarizan-Hanoon (2007).

    During captive feeding trials with <14-month-old R. unicolor on an ad libitum diet of chaffed alfalfa (Medicago sativa) hay and with a voluntary intake of 58.7 and 55.9 g/kg body weight−0.75/day in summer and winter, respectively, numbers of eating and ruminating bouts, and their duration, were about equally distributed during the day (0600–1800 h) and night (1800–0600 h) in both seasons (Semiadi et al. 1994a). Captive R. unicolor fed for 6.4 h in summer and 4.2 h in winter and ruminated for 9.2 h in summer and 9.1 h in winter (Semiadi et al. 1994a). Metabolizable energy for maintenance was 474 kJ/kg body weight−0.75/day; nitrogen retentions (h) as a percentage of intake on 26.6–47.3 and 33.7–44.4 g N/day were 15.1–31.7 and 15.3–29.4, respectively (Semiadi et al. 1998). Similar captive experiments showed that young R. unicolor responded to low-quality diets by lowering voluntary food intake, increasing chewing activity, and conserving nitrogen compared with young red deer (Howse et al. 1995; Semiadi et al. 1994a). The modulus of fineness (Poppi et al. 1980) of fecal particle size from digesta residue suggests that R. unicolor is an intermediate feeder (modulus of fineness  =  2.21); percentage of fecal particles passing through various sieve sizes are: 4-mm sieve, 0.33%; 2-mm sieve, 1.15%; 1-mm sieve, 5.36%; 0.25-mm sieve, 16.74%; and <0.125-mm sieve, 44.32% (Clauss et al. 2002). Additional insight on fecal particle size during captive feeding trials was provided by Semiadi et al. (1994a).

    During experiments with four 165-day-old (± 9.8 SE) R. unicolor in metabolic chambers at 5°C and 20°C without and with wind (6 km/h), respectively, mean heat production (kJ/kg body weight−0.75/day) was 615, 659, 460, and 490 (Semiadi et al. 1996). When ambient temperature was dropped from 20°C to 5°C, heat production increased 34% without wind and 44% with wind; lower critical temperature was 11.6°C without wind and 14.0°C with wind (Semiadi et al. 1996). Compared with young red deer (Semiadi et al. 1996), metabolic responses of young R. unicolor suggested that, given their tropical affinities, they need more shelter and food during cold weather, not unlike Crandall's (1964) observations that adults of R. unicolor seek shelter during cold weather at the New York Zoological Park.


    In the wild, female Rusa unicolor probably experience puberty at 18–24 months (Plotka 1999; Sheng and Ohtaishi 1993). Age of sexual maturity of 7 captive female R. unicolor in New Zealand was 7–19 months; mean (± SE) length of the luteal cycle was 17.2 ± 3 days; 6 of the 7 females were anestrus in November–February; and they displayed no seasonal patterns in prolactin secretion, suggesting little to no response to photoperiod (Asher et al. 1997). Mean birthing interval was 329 days ± 29.7 SD for 6 captive females in New Zealand (Semiadi et al. 1994b). A captive female in India reached sexual maturity at 18 months of age and gave birth at 26 months of age (Acharjyo and Misra 1971). Despite translocation of R. unicolor from its native tropical latitudes to temperate latitudes, lack of seasonal reproduction is demonstrated by births throughout the year (Asher et al. 1997; Duke of Bedford and Marshall 1942; Lao 1968; Zuckerman 1953). Gestation is about 8 months (Brand 1963; Hayssen et al. 1993), although some reports suggest that it can be longer (Plotka 1999; Sheng and Ohtaishi 1993). Among 525 birth records from 210 semidomesticated adult female R. unicolor in Taiwan, mean length of the estrous cycle was 18.2 days ± 0.5 SE (n  =  56), mean length of gestation was 258.6 days ± 0.3 SE (n  =  160), and mean birth interval was 369.9 days ± 2.3 SE (n  =  122—Chan et al. 2009).

    Estimates of productivity suggest that either females bred every other year, as was reported in Sri Lanka (Eisenburg and Lockhart 1972), mortality of young is high (Berwick 1974), low observability of offspring (Fig. 2) biases estimates of productivity (Shea et al. 1990), or some combination of all 3 (Schaller 1967). In Perak, Malaysia, only 9 of 23 females were pregnant when collected throughout the year (Khan and Khan 1968). The number of young-of-the-year per 100 females is typically <50:100, even where introduced: 11–44:100 (Johnsingh 1983), 16–43:100 (Berwick 1974), 17–24:100 (Varman and Sukumar 1993), 33.7:100 (Schaller 1967), 38.2:100 (Bagchi et al. 2008), and 55:100 (Berwick and Jordan 1971) in India; 50:100 in Nepal (Seidensticker 1976b); 43.3:100 in Thailand (Ngampongsai 1977); and 22.3:100 in Florida (Flynn et al. 1990). Female offspring may remain with their mothers as yearlings, but males leave their mothers after about 1 year (Lewis et al. 1990).

    Timing of the breeding and birthing seasons of R. unicolor has been discussed widely because of the variability in parturition dates and antler growth and shedding across the species' substantial latitudinal and longitudinal range (Baker 1898; Fletcher 1911; Lydekker 1898; Schaller 1967). To explain variable antler growth throughout the year, Comber (1904) suggested that 2 distinct breeding seasons occurred in India, but such variation more likely represents an opportunistic strategy, perhaps based on nutrient availability in varied locations across the species' range relative to antler growth, changing hormone levels, and photoperiod. In Chitwan National Park, Nepal, male R. unicolor can be observed in hard antler during any month of the year, reaching a low of 12–14% in July–August and a maximum of 81–92% in December–March; males without antlers occur in December and February–August (Mishra 1982 [not seen, cited in Putman 1988]). Similar patterns of variable antler development have been noted among captive males in New Zealand (Semiadi et al. 1994b).

    Twining in R. unicolor is uncommon, although Evans (1912:144) stated “sometimes two at birth.” Hayssen et al. (1993) give the average litter size of R. unicolor as 1.05. Only 2 (0.6%) of 320 births were twins among semidomesticated R. unicolor in Taiwan (Chan et al. 2009); only 1 (2.4%) of 41 births was twins at the New York Zoological Park (Crandall 1964); and only 1 (1.5%) of 66 births was twins among introduced R. unicolor in Florida (Flynn et al. 1990). Neonates in Australia are 5–6 kg at birth (Slee 1984). Measurements of 8 R. unicolor born in captivity throughout the year in New Zealand were: mass, 5.5–8.5 kg; body length, 36.0–43.1 cm; shoulder height, 44.4–55.0 cm; and body circumference, 44.2–54.3 cm (Semiadi et al. 1993); birth weights did not differ between sexes (Semiadi et al. 1994b). A near-term female fetus from the San Diego Zoo weighed 8.3 kg with a crown-to-rump length of 66 cm (Benirschke 2002). In captivity, neonates lick soil at 2–5 days, nibble on dead forage at 5–14 days, eat fresh forage at 13–23 days, browse lightly at 16–26 days, defecate without stimulation at 4–7 days, and begin to ruminate at 30–42 days (Semiadi et al. 1993).


    Population characteristics

    Despite the widespread distribution of Rusa unicolor in southern Asia and use of many different habitat types, it is no longer abundant throughout most of its native range, except in some protected areas (Sankar and Acharya 2004; Timmins et al. 2008). Because of its predominantly crepuscular to nocturnal behavior, small group size, and general shyness, it is difficult to census accurately (Eisenburg and Lockhart 1972; Schaller 1967). Observed densities of R. unicolor are generally low but vary depending on season and related grouping behavior, habitat conditions in native and introduced areas, competition, predation, and degree of protection. Representative densities are: 0.24–10.70 individuals/km2 in moist and dry deciduous tropical forests in India (Bagchi et al. 2003b; Balakrishnan and Easa 1986; Berwick and Jordan 1971; Biswas and Sankar 2002; Jathanna et al. 2003; Karanth and Sunquist 1992, 1995; Khan et al. 1996; Kurt 1978; Varman and Sukumar 1993); 0.70–1.17 individuals/km2 in lowland dry-zone scrub jungle in Sri Lanka (Eisenberg and Lockhart 1972); 2.0–11.5 individuals/km2 in riverine and Shorea forests and tall-grass habitats in Nepal (Seidensticker 1976b); 1.9–4.2 individuals/km2 in dry tropical forests in Thailand (Srikosamatara 1993); 0.62–1.42 individuals/km2 in lowland rain forest in Sumatra, Indonesia (O'Brien et al. 2003); and 1.76–6.01 individuals/km2 in feral populations in Florida (Flynn et al. 1990). Relative abundance indices of R. unicolor in Bukit Barisan Selatan National Park, Sumatra, Indonesia, are 5.6 times higher in areas of low human density (0–9 villages/area) than areas of high human density (16–30 villages/area—O'Brien et al. 2003).

    Longevity records in captive subspecies include 6 years, 7 months for a male R. u. brookei, 7 years, 11 months for a male R. u. swinhoii, 26 years, 5 months for a female R. u. equina, and 28 years, 5 months for a female R. u. unicolor (Manville 1957; Weigl 2005). Based on life-table analyses, a typical wild R. unicolor dies before about 12 years of age in Gir National Forest, India (Berwick 1974). Mean life expectancy of wild R. unicolor in Ruhuna National Park, southern Sri Lanka, was about 10 years, with no difference between sexes; maximum life expectancy was 24 years; and somewhat surprisingly, mortality of individuals < 2 years old was only about 6%/year (Ashby and Santiapillai 1986). Maximum life span of exotic R. unicolor in New Zealand is estimated at 12 years for males and 17 years for females (Forsyth and Duncan 2001).

    Age structures from populations in India suggest relatively low productivity and male-biased mortality: 43–45% adult females, 5–11% yearling females, 16–19% adult males, 11% yearling males, 19–20% young-of-the-year (n  =  242 [Karanth and Sunquist 1992], n  =  674 [Karanth and Sunquist 1995]); 51.4%, 9.8%, 15%, 3.5%, 20.4% (n  =  363—Schaller 1967); and 58%, 7%, 21%, 2%, 12% (n  =  1,242—Varman and Sukumar 1993). In Gir Lion Sanctuary, India, average age structure in 1987–1989 was 57.6% adult females, 31.4% adult males, 3.7% yearlings, and 7.1% young-of-the-year, suggesting very low recruitment (Khan et al. 1995). Schaller (1967) contended that about 50% of young R. unicolor in central India die before reaching 1 year of age; similarly, Berwick (1974) observed high mortality of young in Gir National Forest in western India.

    Adult sex ratios of R. unicolor favor females, sometimes remarkably: 6 males ∶ 100 females (Ngampongsai 1977); 16–50:100 (Johnsingh 1983); 26–53:100 (Berwick 1974; Berwick and Jordan 1971); 27:100 (Varman and Sukumar 1993); 28:100 (Mohammad Ali 1982); 29:100 (Schaller 1967); 54:100 (Khan et al. 1995); and 83:100 (Bagchi et al. 2008). Comparison of sex ratios from kills by Indian tigers (Panthera tigris tigris—120 males ∶ 100 females) and from direct observations (30∶100) in Kanha National Park, central India, suggests differential predation on young and adult males (Schaller 1967). Where major predators are rare or lacking, male numbers seem to be higher; for example, Eisenburg and Lockhart (1972) noted an usually high male-dominated sex ratio of 123 males ∶ 100 females in Sri Lanka, and Flynn et al. (1990) noted 73 males ∶ 100 females in Florida where R. unicolor was introduced.

    A typical adult male R. unicolor is solitary throughout much of the year, and its low observability may bias estimates of sex ratios. Nevertheless, high rates of mortality are common for male ungulates in general, and disparate sex ratios of R. unicolor could be the result of rutting mortality among mature males. Combat between mature males in rut has been described as antler-to-antler “pushing matches,” but “often severe wounds are inflicted by the [pronounced and lethally angled] brow antler;” Fletcher (1911:346) observed a likely lethal blow by the victorious male to the abdomen of the fleeing vanquished, which he shot. Phythian-Adams (1951:7) noted that mature males “fight desperately” and found 2 dead males with their antlers interlocked. Although mature males give the “impression of harmony” in captivity, deaths have been reported from rutting injuries (Semiadi et al. 1994b:82). Such mortality is difficult to quantify in the wild, but for related North American elk, rutting mortality may be more common than once thought (Leslie and Jenkins 1985).

    Space use

    Although Rusa unicolor is remarkably flexible in its habitat affinities, it prefers areas relatively free from human disturbance (Kushwaha et al. 2004; O'Brien et al. 2003). It mainly prefers forested landscapes (Sankar and Acharya 2004). In India, it occurs “wherever there are hilly ranges covered with jungle” and has an affinity for the tall-grass ecotone between dense forest and open grasslands (Fletcher 1911:333; Lydekker 1916; Ngampongsai 1977) where it finds food and protective cover. In Nepal, R. unicolor uses dense climax sal forests, where it may have a competitive advantage (Dinerstein 1979, 1980). In Thailand, R. unicolor tends to preferentially bed where forest canopy cover is >90% with north and east aspects (Brodie and Brockelman 2010). R. unicolor is nonmigratory over much of it range, but in mountainous areas, it may leave higher elevations during winter (Green 1987). R. unicolor is rather sedentary, although movements become more extensive during rut (Schaller 1967).

    Size of home range of deer varies depending on habitat quality and conditions (Putman 1988), and given the vast geographical distribution of R. unicolor, its home-range size is probably quite variable, although few studies have been conducted in its native habitat. In Sariska Tiger Reserve, India, mean annual home ranges (ha) were 1,500 for males and only 300 for females (Sankar 1994 [not seen, cited in Sankar and Acharya 2004]). Where R. unicolor has been introduced in coastal Texas, annual home ranges (ha) from direct observations of known individuals were larger for males (69–124, n  =  4) than females (38–51, n  =  2); smallest seasonal home ranges occurred in winter (4 for females and 10–86 for males—Richardson 1972). On St. Vincent Island, Florida, mean annual home ranges (ha) of radiocollared individuals were 201.2 for females and 406.6 for males; seasonally, home ranges of females (summer, 36.8; autumn, 45.4; winter, 83.0; spring, 59.8; n  =  4) were smaller than those of males (summer, 81.2; autumn, 23.7; winter, 106.2; spring, 189.3; n  =  3), except in autumn (Shea et al. 1990). Home-range size in New Zealand is comparable to that in Florida (Lo 1985 [not seen, cited in Shea et al. 1990]). Reflecting a generally sedentary nature of R. unicolor, introduced populations in New Zealand dispersed only 0.64 km/year from the time of release, the lowest level of 9 exotic translocated ungulates (maximum 8.64 km/year for chamois [Rupicapra rupicapra]—Caughley 1963).


    Rusa unicolor is a herbivorous ruminant with great variation in dietary selection depending on forage availability (Geist 1998; Schaller 1967). Lewis et al. (1990) provided a cogent review of the food habits of R. unicolor and noted that its ability to consume a wide variety of grass and browse (Fig. 5), while meeting nutritional needs, is correlated with its wide geographical range (Timmins et al. 2008) and success when translocated to alien continents (Fraser et al. 2000; Harris 1971; Kelton and Shipworth 1987; Lewis et al. 1990). Although the size of R. unicolor suggests that it could favor grasses and herbs, it is an intermediate feeder and consumes a great variety of shrubs and trees (Khan et al. 1994; Schaller 1967; Srivastava et al. 1996)—not unlike the larger North American elk in temperate coastal forest in the Pacific Northwest (Leslie et al. 1984).

    Fig. 5

    Mature female Rusa unicolor browsing from her hind legs, a common position of the species but relatively uncommon in other cervids, Kheoladeo National Park, Bharatpur, Rajasthan, northern India, February 2006; note the relatively long tail. Males also assume this position to mark branches as high 3 m at “stomping grounds,” used repeatedly during rut. Photograph courtesy of Lee Dalton ( used with permission.


    Dietary selection of R. unicolor varies considerably depending on season, location, habitat variety and its effect on forage availability and quality, competitive interactions, and human activities (Kushwaha et al. 2004), whether in native habitat (Bagchi et al. 2003a; Johnsingh and Sankar 1991; Khan 1994; Padmalal et al. 2003; Schaller 1967; Shukla and Khare 1998) or introduced locations (Kelton and Skipworth 1987; Shea et al. 1990; Stafford 1997). In India, R. unicolor consumes a greater variety of plants than any other ungulate (Schaller 1967), often uses cultivated areas, and is not deterred by fences as high as 2 m (Baker 1898; Brander 1923).

    Rusa unicolor is 1 of 3 primary consumers of the fleshy fruits of Choerospondias axillaris, a large tropical canopy tree in Thailand, but it deposits most of its seeds under the female canopy trees where regeneration is low (Brodie et al. 2009). Low crude protein and low plant cell-wall contents of R. unicolor feces in November–December suggest that fruits were important dietary constituents in the Indian Himalayas (Green 1987). Although R. unicolor ate fleshy and dry seeds in Nepal, it was not considered an important seed disperser (Dinerstein 1989).

    Limited information exists on the nutritional ecology of R. unicolor, but as with most dietary generalists, it is presumed that nutritional needs are met by varying dietary selection throughout the year. Using fecal nitrogen and other fecal constituents (e.g., Leslie et al. 2008; Leslie and Starkey 1985), Green (1987) demonstrated that R. unicolor in the Indian Himalayas obtained the highest dietary quality, relative to crude protein, in spring and summer and that high ash content in spring, summer, and autumn suggested ingestion of soil, perhaps as a buffer to presumed high levels of ingested volatile oils common in woody species. In Horton Plains National Park, Sri Lanka, nitrogen in feces of R. unicolor suggested highest nutrient availability in May–June (Padmalal et al. 2003). Similar to North American elk in temperate rain forests (Leslie et al. 1984), R. unicolor in the Indian Himalayas consumed significant qualities of ferns (about 40% of the diet) in winter, as well as bamboo and woody browse (Green 1987:figure 10). Despite deficiencies of copper and selenium in soils in New Zealand, introduced populations of R. unicolor obtain sufficient amounts of dietary selenium (>850 nmol/kg in 22 liver samples) and vitamin B12 (>220 nmol/kg), but often insufficient amounts of copper (<100 µmol/kg—Stafford 1997).

    Rusa unicolor regularly drinks water (Hose 1893a; Peacock 1933; Thom 1937; Whitehead 1972) and usually is not far from free water (Brander 1923; Sankar and Acharya 2004; Fig. 6). In Sri Lanka, temporary aggregations of R. unicolor occur at dusk around regularly used water sources (Eisenburg and Lockhart 1972). During feeding trials, forage intake by captive R. unicolor in India decreased from about 400 bites/day, with ad libitum water, to <200 bites/day after 3 days of water deprivation (Berwick 1974). Related Javan rusa (R. timorensis russa) in captivity in Australia tolerated saline water of 1,000–6,000 mg/kg of total dissolved salts with little effect on food intake, food digestibility, and nitrogen balance, but when salinity reached 8,500 mg/kg, they showed signs of stress such as rapid breathing, head shaking, and swelling of the preorbital gland (Yape Kii and Dryden 2005). R. unicolor occurs near estuaries and ocean coasts (Geist 1998; Whitehead 1972), and it may have a similar tolerance of salty water. In Florida, however, R. unicolor prefers freshwater habitats and avoids saltwater habitats (Flynn et al. 1990), which may reflect preferences for forage growing in such areas rather than intolerance of salt water.

    Fig. 6

    Mature male Rusa unicolor wallowing in the muddy wetland in Ranthambhore National Park, Rajasthan, northern India; note the acutely angled brow tine, approaching 50% of the main beam. Photograph by Chris Brunskill ( used with permission.


    Mineral licks are used regularly by R. unicolor (Brander 1923; Matsubayashi et al. 2007a; Schaller 1967), particularly at night (Matsubayashi et al. 2007b; Peacock 1933). In tropical rain forests of Borneo, lactating female R. unicolor use natural mineral licks more frequently during the wet season than the dry season, and both sexes visit such licks more often at night (85% of visits) than during daylight (15% of visits—Matsubayashi et al. 2007b). Water associated with mineral licks in Borneo provides substantially more macronutrients (mean ± SD, µg/ml ) than control water from local ponds and streams: calcium, 83.4 ± 50.0 versus 13.8 ± 8.5; magnesium, 21.4 ± 9.8 versus 2.7 ± 1.0; potassium, 14.4 ± 12.6 versus 1.6 ± 0.6; and sodium, 801.8 ± 1,173.5 versus 6.9 ± 2.4 (Matsubayashi et al. 2007a).

    Diseases and parasites

    No infectious diseases or disease agents have been reported to cause substantial population declines of Rusa unicolor (Presidente 1978, 1984a, 1984b; Slee 1984), but foot-and-mouth disease in Sri Lanka (Brooksby 1973) and India (37 of 104 individuals affected—Barman et al. 1999), sarcocystotic cysts with associated pathology of a bluetonguelike disease in India (Acharjyo and Rao 1988; Gangadharan et al. 1992), erythrocyte sickling in Borneo (Dunn 1964; Undritz et al. 1960), and mucosal disease virus, malignant catarrhal fever, and, rarely, infectious bovine rhinotracheitis in Australia (Presidente 1984a; Slee 1984; Slee and Presidente 1981) have been reported. Diseases, or disease-related deaths, have been reported in captive individuals: chronic (erosive) arthritis, tuberculosis, and arteriosclerosis (Fox 1939); foot-and-mouth disease (Kar et al. 1983; Presidente 1984a; Sarma et al. 1983); malignant catarrhal fever (Semiadi et al. 1994b); and bovine tubercule bacilli (Datta 1954). Intestinal rupture and resulting acute peritonitis, perhaps from a parasitic infection, was reported in a wild pregnant female R. unicolor in India (Bhattacharjee 1986).

    By the mid-1980s, 18 endoparasites (9 nematode species, 6 flukes, 2 protozoans, and 1 tapeworm) and 40 ectoparasites (35 tick species, 2 sucking lice, 2 keds, and 1 flea) had been reported in native, feral, and captive populations of R. unicolor (Presidente 1984a). Parasites of free-ranging R. unicolor in native habitat include the nematodes Bunostomum, Haemonchus, Oesophagostonmum, Strongyloides, and Trichuris in India (Bhat and Manickam 1998; Hiregoudar 1976) and Ashwortius sidemi, Rinadia andreevae, and Spiculopteragia houdemeri in Vietnam (Drozdz 1965, 1973), and trematodes Calicophoron microbothrioides in Malaysia (Lee et al. 1987), and Fischoederius elongates, Homologaster poloniae, Paramphistomum explanatum (Patnaik and Acharjyo 1970; Rao and Acharjyo 1969), and Gastrothylax crumenifer in India (Agrawal and Ahluwalia 1980). Feral and captive populations of R. unicolor also harbor a variety of internal parasites including nematodes Gongylonema pulchrum (Chakrborty 1994) and cestode cysts Coenurus gaigeri (Varma et al. 1994) in India; Spiculopteragia asymmetrica in New Zealand (Andrews 1973) and Australia (Presidente 1984a); trematodes Ceylonocotyle streptocoelium in Australia (Keith and Keith 1969) and Paramphistomum explanatum in India (Rao and Acharjyo 1984); and protozoans Theileria aristotelis (Levine 1971) and Toxoplasma gondii in India (Ippen et al. 1981).

    External parasites of native and feral R. unicolor include ticks Haemaphysalis ramachandrai and H. davisi in India and Nepal (Dhanda et al. 1970; Hoogstraal et al. 1970); H. anomala and H. papuana in southeastern Asia (Hoogstraal et al. 1965, 1967); H. davisi in Malaysia (Hoogstraal and El Kammah 1971); H. mjoebergi in Borneo and Sumatra (Hoogstraal and Wassef 1982); and Dermacentor variabilis in Florida (Davidson et al. 1987). Captive R. unicolor in eastern India, and no doubt wild populations elsewhere, are affected by the hematophagous flies (Tabanus rubidus, T. striatus, Stomoxys calcitrans, Haematobia irritans exigua, and Musca crassriostris) that can transmit trypanosomiasis and other diseases (Veer et al. 2002). When flying parasites are most active (e.g., humid, rainy seasons), larger than normal groups of R. unicolor are seen often in open areas and near or submerged in water (Prater 1980).

    Interspecific interactions

    Because of the extensive native range of Rusa unicolor in southern Asia (Fig. 3), it can be sympatric with many other large herbivores (e.g., Schaller 1967), with which competitive interactions for food may exist (Sankar et al. 2007). In India, for example, R. unicolor can be sympatric with axis deer or chital (Axis axis), hog deer (A. porcinus), barasingha (Rucervus duvaucelii), red muntjac (Muntiacus muntjakKushwaha et al. 2004), blackbuck (Antilope cervicapra), chinkara (Gazella bennettii), chowsingha (Tetracerus quadricornisLeslie and Sharma 2009), nilgai (Boselaphus tragocamelusLeslie 2008), gaur (Bos gaurus), water buffalo (Bubalus bubalisBerwick 1974; Jathanna et al. 2003; Karanth and Sunquist 1992), Asian elephants (Elephas maximusShoshani and Eisenberg 1982), and Indian rhinoceroses (Rhinoceros unicornisLaurie et al. 1983). Significant dietary overlap of R. unicolor and livestock can occur locally (e.g., Shukla and Khare 1998; Srivastava et al. 1996). In extreme eastern Asia and Taiwan, R. unicolor can be sympatric with the smaller sika deer (Cervus nipponFeldhammer 1980; MacKinnon 2008). In southern China, the distributional range of R. unicolor could overlap with red deer, forest musk deer (Moschus berezovskii), tufted deer (Elaphodus cephalophus), red muntjac, Reeves' muntjac (M. reevesi), Chinese water deer (Hydropotes inermisMacKinnon 2008), and white-lipped deer (P. albirostreLeslie 2009), but habitat affinities and decimated populations likely limit direct interactions in most areas (Timmins et al. 2008).

    Typically, some degree of habitat (Bagchi et al. 2003a, 2003b; Berwick 1974; Dinerstein 1979; Kushwaha et al. 2004) and dietary (Berwick 1974; Khan 1994; Shukla and Khare 1998) differentiation occurs among R. unicolor and other ungulates. In Ranthambhore National Park, India, R. unicolor and chital form a “cervid guild” that prefer AnogeissusGrewia forests, and nilgai and chinkara form a “bovid guild” and select AcaciaButea habitats during summer and winter; unlike the cervids, nilgai were tolerant of livestock grazing and associated degradation of grass cover (Bagchi et al. 2003a, 2003b). In contrast, high dietary overlap of 90–93% was noted among R. unicolor, chital, and nilgai in the semiarid Sariska Tiger Reserve, northwestern India (Sankar et al. 2007). In the Himalayas of northern India, R. unicolor is sympatric with Himalayan musk deer (Moschus chrysogaster), serow (Capricornis sumatraensis), and goral (Nemorhaedus goralMead 1989); competitive interactions are avoided by inverse relationships of habitat use and diet, but diets of R. unicolor and serow were most similar in autumn (Green 1985, 1987). In Nepal, R. unicolor uses riverine forests (80% of its time) with muntjac (78%) and chital (75%) but not hog deer (<5%—Mishra 1982 [not seen, cited in Putman 1988]). On St. Vincent Island, Florida, R. unicolor and white-tailed deer (Odocoileus virginianusSmith 1991) partition habitat and food resources; in particular, R. unicolor uses freshwater habitats and associated aquatic vegetation (11.1–56.2% aquatic vegetation in seasonal diets; Figs. 2 and 6) to a much greater extent than white-tailed deer (1.5–4.4%—Shea et al. 1990).

    Cattle egrets (Bubulcus ibis; Fig. 7) and Indian, or rufous, tree-pies (Dendrocitta vagabunda) associate with and forage on R. unicolor, respectively. Bharucha (1987) observed a female R. unicolor standing awkwardly with a rear leg raised up and away from her body to accommodate an Indian tree-pie that was apparently removing parasites from her groin area.

    Fig. 7

    Male Rusa unicolor and cattle egret (Bubulcus ibis) foraging together in wetland in Ranthambhore National Park, Rajasthan, northern India. Photograph by Chris Brunskill ( used with permission.


    The endangered Indian tiger preys extensively on R. unicolor (Mazák 1981; Ramesh et al. 2009; Schaller 1967; Wang and Macdonald 2009) and, anecdotally, is said to mimic the call of R. unicolor to deceive it while hunting (Whitehead 1972). In Nagarahole, southern India, tigers prey preferentially on male R. unicolor (43 kills: adult males, 40.0%; adult females, 30.6%; yearling males, 5.7%; yearling females, 5.7%; 122 scats: young-of-the-year, 18.0%) and take more young individuals (58.6%) than prime (34.5%) or old (6.9%) individuals (Karanth and Sunquist 1995). R. unicolor constitutes 30.5% of prey items in tiger scats and 36.8% of tiger kills in Bandipur Tiger Reserve, southern India (Johnsingh 1983), and 24.9% and 28.6% in Nagarahole National Park, southern India (Karanth and Sunquist 1995, 2000). Frequency of occurrence of R. unicolor in tiger scats is 59.8% in Mudumalai Tiger Reserve, southern India (Ramesh et al. 2009); 51.4% in Sariska Tiger Reserve, northwestern India (Sankar and Johnsingh 2002); 36.9% in Ranthambhore National Park, northern India (Bagchi et al. 2003b); 14.6% in Pench National Park, central India (Biswas and Sankar 2002); and 27.6% in Nagarjunasagar Srisailam Tiger Reserve, south-central India (Reedy et al. 2004). Seidensticker (1976a) considered R. unicolor a primary prey species of tigers in Royal Chitwan National Park, Nepal, as did Wang and Macdonald (2009) for tigers in Jigme Singye Wangchuck National Park, central Bhutan. The Sumatran tiger (P. t. sumatrae) in Bukit Barisan Selatan National Park, Sumatra, Indonesia, also preys on R. unicolor (O'Brien et al. 2003).

    Rusa unicolor, primarily fawns, constitutes 14% of prey items in scats and 5% of the kills of the leopard (Panthera pardus) in Bandipur Tiger Reserve (Johnsingh 1983) and 13.5% and 9.6% in Nagarahole National Park, southern India (Karanth and Sunquist 1995, 2000). In southern India, frequency of occurrence of R. unicolor in leopard scats is 29.0% in Mudumalai Tiger Reserve (Ramesh et al. 2009), 9% and 11.7% on the Mundanthurai Plateau and in the Mudumalai Wildlife Sanctuary, respectively (Ramakrishnan et al. 1999), and 5.9–7.8% in Sigur and Thalamalai reserve forests (Arivazhagan et al. 2007). Scats of leopards in Wolong Reserve, Sichuan, China, have only 0.5–1.6% frequency of occurrence of R. unicolor (Johnson et al. 1993). In Sri Lanka, only 1 of 29 identified prey items of leopards was R. unicolor (Eisenberg and Lockhart 1972). R. unicolor comprised 14.8% of 142 kills by highly endangered Asiatic lions (Panthera leo persicaHaas et al. 2005) in Gir Forest, India (Berwick 1974; Chellam and Johnsingh 1993).

    Frequency of R. unicolor in scats of the group-hunting dhole (Cuon alpinusCohen 1978; Earle 1987) was 14.7% in Mudumalai Wildlife Sanctuary in southern India (Cohen et al. 1978), and elsewhere in India, comprised 9–15% of the dhole diet (Arivazhagan et al. 2007). In Bandipur Tiger Reserve, fawns of R. unicolor (31.4%) were about 3 times as frequent in dhole scats as adults (10.6%—Johnsingh 1983). In Nagarahole National Park, southern India, 10.2% of prey items in dhole scats and 3.0% of prey kills were R. unicolor (Karanth and Sunquist 1995, 2000). Composite samples of scats from large carnivores, primarily dhole and leopards, in Huai Kha Khaeng Wildlife Sanctuary, Thailand, contained only 1.2–6.4% R. unicolor (Rabinowitz and Walker 1991). In Khao Yai National Park, Thailand, R. unicolor was the most common prey found in dhole scats (Austin 2002). The highest reported frequency of occurrence of R. unicolor in dhole scats was 60.9% in in Jigme Singye Wangchuck National Park, central Bhutan (Wang and Macdonald 2009).

    In India, R. unicolor has not been reported as prey of the endangered Indian wolf (Canis lupus pallipes) or the striped hyenas (Hyaena hyaenaArivazhagan et al. 2007), as have other species (Leslie 2008). Green (1985) found small amounts of hair of R. unicolor in scats of red fox (Vulpes vulpes) in Kedernath Sanctuary in the Indian Himalayas, believed to be from fawns or scavenged animals.


    Rusa unicolor adapts readily to captivity and is well represented in zoos, private ranches, and research facilities throughout the world (Crandall 1964; Semiadi et al. 1993, 3141994b, 1995a, 1995b, 1998; Weigl 2005). R. unicolor is reared in captivity in New Zealand (e.g., Semiadi et al. 1993), Malaysia (Dahlan and Norfarizan-Hanoon 2007, 2008), and Bangladesh (Basbar et al. 2001). Some of these captive populations involve research and production of meat and other by-products, and others are focused on restoration of wild populations (Basbar et al. 2001). Some described R. unicolor as very alert and nervous in captivity (Semiadi et al. 1994b), but Crandall (1964) suggested otherwise.

    In New Zealand, 6 of 8 neonatal R. unicolor were reared successfully on commercially available sheep milk replacer, containing 38.5% lactose, 27% milk fat, 4.6% nitrogen, and 23.6 kJ/g dry matter, and a liquid vitamin replacement; 2 neonates died because of bloat and severe diarrhea (Semiadi et al. 1993). Milk intake peaked early at week 3 when neonates consumed <400 g dry matter/day and declined thereafter until self weaning at about 10 weeks (Semaidi et al. 1993). They consumed 312 g dry matter/day and gained 241 g/day ± 99.6 SE from birth through their 1st week of age, 387 ± 46.9 g/day in weeks 1–4, and 322 ± 34.0 g/day thereafter (n  =  8—Semiadi et al. 1993). Neonatal mortality from adult aggression, presumably by unrelated individuals, has been noted in captivity (Semiadi et al. 1994b).

    To minimize births in captivity, chemical contraceptives have been successfully applied: melengestrol acetate as a feed additive (Raphael et al. 2003) and injections of porcine zonae pellucidae vaccine, albeit the later resulted in health problems to neonates if their mother was inoculated while pregnant (Kirkpatrick et al. 1996). Because of conservation concern and possible need for captive breeding, semen cryopreservation has been investigated for R. u. swinhoii in Taiwan (Cheng et al. 2004). R. unicolor can be tranquilized for handling with a variety of drugs: 0.07–0.14 mg succinylcholine chloride/kg of body weight, but doses ≥ 16.0 mg/kg resulted in mortality (Lentz et al. 1986); a combination of 1 mg ketamine/kg of body weight and 0.75 mg xylazine/kg of body weight, which can be reversed by yohimbine (Ibrahim 1998); and oral doses of diazepam (Thomas et al. 1967), although capture myopathy has been reported (Presidente 1978).

    Rusa unicolorCervus elaphus and R. unicolorR. timorensis hybrids have been reported (Idris and Moin 2009; Muir et al. 1997; New Zealand Department of Conservation 2005; Slee 1984; van Mourik and Schurig 1985). R. unicolorR. timorensis hybrids at 10 months old attain the same weight as pure R. timorensis at 20–24 months old, and male hybrids are fertile with pure R. timorensis females, as are female hybrids with pure R. unicolor males (Slee 1984).


    Grouping behavior

    Kurt (1978:233) described the social and reproductive systems of Rusa unicolor as varying, depending on habitat, from “non-seasonal, alternatingly territorial” in stable rain forest to “seasonal, synchronized territorial” in varied deciduous forest and “solitary, aggregational” in stable grass jungles. Unlike most comparably sized cervids characterized by large groups (Geist 1998), R. unicolor typically occurs in small groups, most often a single female that dominates the group, her young-of-the-year, and perhaps her female yearling; mature males > 6 years old are typically solitary, with young males grouping together, close to females, or as satellites to solitary mature males (Eisenberg and Lockhart 1972; Khan et al. 1995; Schaller 1967). In some areas (e.g., Sri Lanka—Kurt 1978), R. unicolor occurs with regularity in groups of 30–40, usually related to more abundant patches of forage and water availability (Geist 1998).

    Maximum group size of R. unicolor in Gir National Forest, western India, was 5 individuals (Berwick 1974), and in Gir Lion Sanctuary, there were no seasonal differences in group size (Khan et al. 1995). Group sizes were 1–10 individuals in Nagarahole, southern India: 52% of the groups, 1 individual; 44%, 2 or 3 individuals; and 4%, 4–10 individuals (Karanth and Sunquist 1992). Group sizes were 1–16 individuals in Bandipur Tiger Reserve, southeastern India: 39% of the groups, 1 individual; 27%, 2 individuals; 21%, 3 or 4 individuals; 7%, 5 or 6 individuals; 3%, 7–9 individuals; 3%, 10–16 individuals (Johnsingh 1983:figure 6). In Mudumalai Wildlife Sanctuary, southern India, group size averaged 3.1 individuals but varied seasonally: maximum dry-season group size was 19 individuals and maximum wet-season group size during 2 years was 44 and 50 individuals (Varman and Sukumar 1993). Mean summer and winter group sizes in Ranthambhore Tiger Reserve, India, were 4.2 and 3.4 individuals, respectively (Bagchi et al. 2008). In Sri Lanka, group size was 2–8 individuals, and 60% of 230 individuals were solitary (Eisenberg and Lockhart 1972). Group size is often largest near water holes (Johnsingh 1980, 1983).

    Reproductive behavior

    Downes (1983b:36) described the mating system of Rusa unicolor as “polygamous male dominance … in dispersed facultative meeting-territories.” Reproductive behavior of R. unicolor is “primitive with unique elements” (Geist 1998:76); males do not establish harems (Brander 1923; Schaller 1967). At the beginning of the breeding season, male and female R. unicolor “suddenly [begin] to wander widely, even at mid-day, becoming highly conspicuous in sharp contrast to their usual elusiveness,” and appear “nervous, as if looking for something” (Schaller 1967:144).

    The following description of reproductive behavior was synthesized from Brander (1923), Downes (1983b), Fletcher (1911), Geist (1998), Prater (1980), Schaller (1967), and Thom (1937). During rut, mature males establish nonexclusive breeding territories to attract females and from which they challenge competitors of comparable rank; younger males often occupy peripheries of such territories. Mature males in rut have swollen necks, strong odor, and everted preorbital glands and appear aggressive, often thrashing vegetation. They are usually coated with mud from regular wallowing in wet spots (Fig. 6), accentuating their generally dark pelage, and frequently rub their muddy necks on tree trunks and vegetation while patrolling their territories for receptive females and competing males. Females are said to wallow but less frequently than males (Peacock 1933). Males paw and stomp the ground, and thereby create areas as large as 3–13 m in diameter, devoid of vegetation; “stomping grounds” may occur in dense forest or in the open atop hills. Often such areas are below an overhanging branch 2.3–3.3 m off the ground; male “preach” at these sites by standing erect on their hind legs and rubbing their scent-soaked preorbital glands and antlers on the branch (a behavior not observed among introduced R. unicolor in Florida). Males copiously spray themselves, even their faces, with urine from their mobile penis, the structure of which is unique among cervids.

    Aggressive behavior between competing males includes head-up and head-down displays, pawing and thrashing, and head-to-head pushing matches, with tail cocked up and mane and back hairs erected, until the weaker gives up. Geist (1998:76) noted that, unlike any other deer species but goat-like, male R. unicolor will “rise on their hind legs and clash downward into one another.” Females also rise on their hind legs and hit each other on the head with their forelegs, resulting in a “noise [that] resounds through the jungle;” the same behavior is used against predators (Brander 1923:177).

    Sexual behavior of R. unicolor has not been described in detail, but accounts by Brander (1923), Geist (1998), and Schaller (1967) provide some insight. Females actively seek or court adult males, moving widely among breeding territories; courtship is based on pair-bonding without serious vocal advertisement; males do not clasp females during mounting, front legs hang loosely (Fig. 8); and intromission is a “copulatory jump” (Geist 1998:76). Schaller (1967) observed mature males sniffing and licking females' vulvae. Another male trotted after a female in a low-stretch display with his neck parallel to the ground and preorbital glands everted. Satellite males assist dominant males by “warding off lesser rivals” and may breed if >1 receptive female enters the dominant male's territory (Geist 1998:77).

    Fig. 8

    Mature male Rusa unicolor mounting a female in Ranthambhore National Park, Rajasthan, northern India, January–February 2000. Photograph courtesy of James Warwick ( used with permission.


    No published observations of parturition of wild R. unicolor were found. Presumably, females separate themselves from other individuals, seek secluded places to give birth, and hide their neonates. Neonates may rest alone, hidden, for much of their first 3 months of life, with their mother returning at regular intervals (Eisenberg and Lockhart 1972; Shea et al. 1990).


    Brander (1923) noted that eyesight of Rusa unicolor was only moderately developed, but Peacock (1933) contended that all senses were highly developed. Along with highly developed scent-marking routines and persistent acrid odor, particularly among males in rut (see “Reproductive behavior”), foot stomping by female R. unicolor is used to alert (and summon) neonates and conspecifics of threats (Brander 1923; Mason 1994). Despite their size, R. unicolor can move quietly (Stebbing 1911) and with stealth through dense forest; mature males, when moving rapidly “enclose their necks and shoulders” with their antlers by “carrying their heads thrust out before them” (Brander 1923:182). Ever alert and a “high-stepper” stomping its forelegs when alarmed (Stebbing 1911:59), R. unicolor makes “sharp, short ‘pooks,’” “tits,” or “honks” ( =  “sharp, high notes”) when disturbed (Brander 1923:180; Lydekker 1916; Mason 1994:23; Peacock 1933:126; Thom 1937). Brander (1923:180–181) described 2 other call types: a “loud metallic bellow” of rutting males and a “death cry” consisting of “a prolonged hoarse scream.”

    Miscellaneous behavior

    Rusa unicolor is largely crepuscular to nocturnal depending on location (Brander 1923; Fletcher 1911; Peacock 1933), but it can be active throughout the day in areas with minimal human disturbance (Johnsingh 1983; O'Brien et al. 2003) and in introduced locations such as Texas (Richardson 1972). Anecdotally, the Malayan form may be more nocturnal than Indian forms (Evans 1912), but Peacock (1933:125) described R. unicolor in Burma as “very nocturnal … ordinarily seek[ing] heavy cover with the first light at dawn and not leave[ing] the same till dusk.” U Tun Yin (1967) contended that nocturnal habits of R. unicolor in Burma were related to continual human harassment.

    Much like the moose (Alces alcesFranzmann 1981), R. unicolor is 1 of the few deer that will readily face wild predators (e.g., leopards and dholes) and hunting dogs defensively (Brander 1923; Geist 1998; Johnsingh 1980; U Tun Yin 1967). Females defend their young in a low-head posture, “barking loudly” and stomping front legs, with tail and ears erect and mane hairs flared; if several females are present, they stand rump-to-rump, facing outward toward the threat (Geist 1998:75)—reminiscent of muskoxen (Ovibos moschatusLent 1988) and the wild yak (Bos mutusLeslie and Schaller 2009). If the threat is minimal (e.g., a single dhole), they may not react at all (Divyabhanusinh 1988). Brander (1923) recounted an incident when a female R. unicolor aggressively defended its young from a leopard who successfully carried the offspring 3 m up into a tree; when a hunter arrived on the scene, the leopard dropped the young, which ran off with its mother. Females can be “savage” while protecting their neonates in captivity (Crandall 1964:569). When perceiving or facing a threat, R. unicolor will not typically run off but often stands motionless, its dark pelage blending into the surrounding vegetation; alternately, it may creep off in a “semicrouch trot” with its neck held horizontally (Schaller 1967).

    Rusa unicolor readily swims with its body fully submerged and only its head above the water (Prater 1980; Fig. 9), often to avoid insects and to forage (Richardson 1972; Shea et al. 1990; Shukla and Khare 1998). When ambient temperatures approach freezing, R. unicolor may lie in water that is warmer than the air (Brander 1923; Prater 1980).

    Fig. 9

    Mature male Rusa unicolor swimming through wetland in Ranthambhore National Park, Rajasthan, northern India, January–February 2000. Photograph courtesy of James Warwick ( used with permission.



    Diploid number (2n) of Rusa unicolor varies, apparently among subspecies, from 56 in New Zealand specimens originally from Sri Lanka and India (Muir et al. 1997) to 58 in India (Chandra et al. 1967) and 62 in southwestern China (Wang and Du 1982) and Malaysia (Idris and Moin 2009); fundamental number (FN) is 70; there are 44–64 acrocentric autosomes and 2–14 meta- and submetacentric autosomes (Bonnet-Garnier et al. 2003:table 1; Groves and Grubb 1987:table 2). The X chromosome is acrocentric, and the Y chromosome is acro- or submetacentric (Bonnet-Garnier et al. 2003; Chandra et al. 1967; Groves and Grubb 1987). Seven Robertsonian translocations have been identified in R. unicolor (Bonnet-Garnier et al. 2003).

    Chromosomal evidence suggests that Rusa is sister to a clade consisting of Przewalskium and Rucervus (Groves and Grubb 1987). R. unicolor forms a clade closest to R. timorensis from Java and the Timor islands (e.g., Emerson and Tate 1993), followed by the critically endangered Rucervus eldi from India, and their speciation was proposed by Bonnet-Garnier et al. (2003) to have occurred from monobrachial centric fusions (Baker and Brickham 1986). Ongoing interest in the molecular systematics of Cervidae (e.g., Cronin 1991; Di Stefano and Petronio 2002; Liu et al. 2003; Miyamoto et al. 1990; Pitra et al. 2004; Randi et al. 2001) led Groves (2006:21) to note that conspicuous external features (e.g., antler configurations of males, rump patch) that have led to taxonomic affiliations are more likely convergent, caused by shared “climatic-related lifestyle factors” rather than phylogeny. Although some evidence suggests that rusine deer split from other cervids about 5 million years ago, the validity of Rusa as a monophyletic genus is still debated by morphologists (Meijaard and Groves 2004) and molecular systematists (Hernández-Fernández and Vrba 2005; Randi et al. 2001).

    Characteristics of the mitochondrial 16S rRNA gene have been used forensically to differentiate R. unicolor from sympatric species such as nilgai, chital, and blackbuck (Guha and Kashyap 2005). Electrophoretic analysis of mitochondrial DNA fragments also have been used to differentiate R. unicolor from other ungulates (Cronin et al. 1991). Five exclusive monomorphic random amplified polymorphic DNA markers of 150–520 base pairs have been identified for R. unicolor from Malaysia (El-Jaafari et al. 2008).


    Conservation challenges for Rusa unicolor are daunting because it is so widespread geographically and affected by numerous local customs, national laws, and even civil unrest (e.g., Sri Lanka and Myanmar—Timmins and Evans 1996; Timmins et al. 2008). R. unicolor was elevated by the International Union for Conservation of Nature and Natural Resources from no status in 2006 to “Vulnerable” in 2008 because of >50% decline over the past 3 generations in many populations, with probable local extinctions, notably in Vietnam, Laos, Thailand, Cambodia, Myanmar, Malaysia, Bangladesh, Borneo, and Sumatra (Timmins et al. 2008). Populations of R. unicolor in Taiwan, India, and Nepal are more stable, mostly in protected or remote areas (Hsu and Agoramoorthy 1997; Timmins et al. 2008). R. unicolor is considered “Lower Risk, Schedule III” in India with an estimated population size of >100,000 countrywide (Sankar 2008), although nowhere is it abundant (Sankar and Acharya 2004). Even a century ago, game wardens and sportsman expressed concern that Indian and Burmese wildlife, including R. unicolor, was being overexploited for subsistence and sport and to minimize depredation of agricultural areas (Glasfurd 1903; Peacock 1933; Stebbing 1911; Wang et al. 2006). Well ahead of their time, Glasfurd (1903) and Stebbing (1911) commented that fire suppression caused forest encroachment into grasslands and diminished important habitat of R. unicolor and other large herbivores.

    Rusa unicolor is a preferred meat throughout southeastern Asia (Duckworth et al. 1999; Gardner 1993), although some historic local taboos have existed; Evans (1918:194) noted that among the Sakai of Malaysia “women and children may not eat, cook, or touch deer's flesh, or go near the body of a dead deer.” Nevertheless and despite strict laws against hunting in India and elsewhere, exploitation for subsistence and marketing of meat and antlers are the most important ongoing problems facing R. unicolor throughout southeastern Asia (Datta et al. 2008; Duckworth et al. 1999; Khan and Khan 1968; Sammaiah et al. 2008; Steinmetz et al. 2006; Timmins et al. 2008). Timmins et al. (2008) noted that expanding urban wealth and increasing demand for exotic meat and “medicines,” rather than rural poverty, are having the greatest pernicious impacts on declining populations of R. unicolor and other fauna in southeastern Asia. Other conservation challenges recently summarized by Timmins et al. (2008) include fragmentation and loss of forested habitats through indiscriminate logging for wood products or conversion to agriculture (Johnsingh 1983; Kumara et al. 2004; Kushwaha et al. 2004), associated loss of or access to specialized habitats such as salt licks (Matsubayashi et al. 2007a, 2007b), mining and energy development, urban expansion, and roads and associated human traffic (Griffiths and van Schaik 1993). Improved education and partnerships with local communities and conservation initiatives that work together to identify critical wildlife issues are useful to enhance local buy-in and forward-thinking conservation action (Steinmetz et al. 2006).

    Few comprehensive ecological studies of R. unicolor have been conducted within its native range, and much of what is known comes from demographic assessments of R. unicolor as prey for highly endangered and charismatic species such as the Indian tiger or the Asiatic leopard. Studies of introduced populations provide some insight (e.g., Downes 1983b; Lewis et al. 1990), but clearly, more basic information on status, ecology, and behavior of native populations is needed to assure adequate conservation of southeastern Asia's largest and most ecologically generalized cervid. Without such knowledge, R. unicolor may indeed “be, like the Hog Deer and Eld's Deer are already, almost absent from South-east Asia” (Timmins et al. 2008:19).


    I thank C. Carr, Oklahoma State University, for her assistance with assembling literature for this review; A. L. Gardner, United States Geological Survey, and C. P. Groves, Australian National University, for their many constructive comments on the synonymies; S. Mattioli, Z. Roehrs, K. Sankar, and G. B. Schaller for their helpful reviews; A. Albert, B. Horner, and the entire staff of Interlibrary Loan Services, Oklahoma State University, D. Wingreen-Mason, L. Overstreet, and C. Shaw, Cullman Collection, Smithsonian Libraries, and M. Boothe, Department of Interior Library, Washington, D.C., for providing copies of many seminal pages from rare literature; K. Anderson and L. Tomsett of the British Museum (Natural History) for assistance with preparation of the skull images, and S. Hallgren and C. Zhou, Oklahoma State University, for help with French and Chinese translations, respectively. The Oklahoma Cooperative Fish and Wildlife Research Unit (Oklahoma State University, Oklahoma Department of Wildlife Conservation, United States Geological Survey, United States Fish and Wildlife Service, and Wildlife Management Institute cooperating) provided technical support during the preparation of this monograph.



    E. D Ables and C. W Ramsey 1972. Indian mammals on Texas rangelands. Journal of the Bombay Natural History Society 71:18–25. Google Scholar


    L. N Acharjyo and A. B Bubenik 1983. The structural peculiarities of antler bone in genera Axis, Rusa, and Rucervus. Pp. 195–209 in Antler development in Cervidae ( R. D Brown ed.). Caesar Kleberg Wildlife Research Institute, Kingsville, Texas. Google Scholar


    L. N Acharjyo and R Misra 1971. Age at sexual maturity of three species of wild animals in captivity. Journal of the Bombay Natural History Society 68:446. Google Scholar


    L. N Acharjyo and A. T Rao 1988. Sarcocystosis in some Indian wild ruminants. Indian Veterinary Journal 65:169–170. Google Scholar


    R. D Agrawal and S. S Ahluwalia 1980. A note on the occurrence of Gastrothylax crumenifer in sambar (Cervus unicolor). Indian Veterinary Journal 57:436. Google Scholar


    G. M Allen 1930. Pigs and deer from the Asiatic expeditions. American Museum Novitates 430:1–19. Google Scholar


    G. M Allen 1940. The mammals of China and Mongolia. Natural history of central Asia. Vol. XI. Part 2 American Museum of Natural History, New York. Google Scholar


    J. A Allen 1906. Mammals from the Island of Hainan, China. Bulletin of the American Museum of Natural History 22:463–490. Google Scholar


    J. R. H Andrews 1973. A host–parasite checklist of helminths of wild ruminants in New Zealand. New Zealand Veterinary Journal 21:43–47. Google Scholar


    C Arivazhagan R Arumugam and K Thiyagesan 2007. Food habits of leopard (Panthera pardus fusca), dhole (Cuon alpinus) and striped hyena (Hyaena hyaena) in tropical dry thorn forest of southern India. Journal of the Bombay Natural History Society 104:178–187. Google Scholar


    K. R Ashby and C Santiapillai 1986. The life expectancy of wild artiodactyl herbivores, water buffalo (Bubalus bubalis), sambar (Cervus unicolor), spotted deer (Axis axis), and wild pig (Sus scrofa) in Ruhuna National Park, Sri Lanka, and the consequences for management. Tigerpaper 13(2):1–7. Google Scholar


    G. W Asher P. D Muir G Semiadi K. T O'Neill I. C Scott and T. N Barry 1997. Seasonal patterns of luteal cyclicity in young red deer (Cervus elaphus) and sambar deer (Cervus unicolor). Reproduction, Fertility and Development 9:587–596. Google Scholar


    T. D Atkeson V. F Nettles R. L Marchington and W. V Branan 1988. Nasal glands in the Cervidae. Journal of Mammalogy 69:153–156. Google Scholar


    S. C Austin 2002. Ecology of sympatric carnivores in Khao Yai National Park, Thailand. Ph.D. dissertation, Texas A&M University, College Station. Google Scholar


    A Azzaroli C De Giuli G Ficcarelli and D Torre 1988. Late Pliocene to early mid-Pleistocene mammals in Eurasia: faunal succession and dispersal events. Palaeogeography, Palaeoclimatology and Palaeoecology 66:77–100. Google Scholar


    A-M Bacon et al. 2004. The Pleistocene Ma U'Oi cave, northern Vietnam: palaeontology, sedimentology, and palaeoenvironments. Geobios 37:305–314. Google Scholar


    S Bagchi S. P Goyal and K Sankar 2003a. Niche relationships of an ungulate assemblage in a dry tropical forest. Journal of Mammalogy 84:981–988. Google Scholar


    S Bagchi S. P Goyal and K Sankar 2003b. Prey abundance and prey selection by tigers (Panthera tigris) in semi-arid, dry deciduous forest in western India. Journal of Zoology (London) 260:285–290. Google Scholar


    S Bagchi S. P Goyal and K Sankar 2008. Social organization and population structure of ungulates in a dry tropical forest in western India (Mammalia, Artiodactyla). Mammalia 72:44–49. Google Scholar


    R. J Baker and J. W Bickham 1986. Speciation by monobrachial centric fusions. Proceedings of the National Academy of Sciences 83:8245–8248. Google Scholar


    S. W Baker 1855. Eight years' wandering in Ceylon. Longman, Brown, Green, and Longmans, London, United Kingdom. Google Scholar


    S. W Baker 1898. Wild beasts and their ways: reminiscences of Europe, Asia, Africa, and America. Macmillan and Company, Limited, New York. . Google Scholar


    M Balakrishnan and P. S Easa 1986. Habitat preferences of the larger mammals in the Parambikulam Wildlife Sanctuary, Kerala, India. Biological Conservation 37:191–200. Google Scholar


    N. N Barman D. K Sarma S Das and G. P Patgiri 1999. Foot-and-mouth disease in wild and semi-domesticated animals of the north-eastern states of India. Indian Journal of Animal Sciences 69:781–783. Google Scholar


    M. A Basbar F Begum and K. K Mondal 2001. Sambar deer breeding in captivity in Dulahazra Safary (Managed National Reserve), Bangladesh. Tigerpaper 28:21–23. Google Scholar


    J. M Bechstein 1799. Thomas Pennant's Allgemeine uebersicht der Vierfüssigen Thiere. Aus dem Englischen übersetzt und mit Anmerkungen und Zusätzen Versehen. Erster Band. Verlage des Industrie, Comptoir's, Weimar, Germany. Google Scholar


    K Benirschke 2002. Indian sambar deer. Comparative Placentation Project, University of California–San Diego, San Diego,, accessed 8 June 2009. Google Scholar


    A Bentley 1957. A brief account of the deer in Australia. Journal of Wildlife Management 21:221–225. Google Scholar


    S. H Berwick 1974. The community of wild ruminants in the Gir Forest ecosystem, India. Ph.D. dissertation, Yale University, New Haven, Connecticut. Google Scholar


    S. H Berwick and P. A Jordan 1971. First report of the Yale–Bombay Natural History Society studies of wild ungulates at the Gir Forest, Gujarat, India. Journal of the Bombay Natural History Society 68:412–423. Google Scholar


    E. K Bharucha 1987. An observation on the relationship between a sambar and a tree-pie. Journal of the Bombay Natural History Society 84:675. Google Scholar


    M. N Bhat and R Manickam 1998. Intensity of nematode infection and corpoculturing in sambar (Cervus unicolor). Indian Journal of Animal Sciences 68:643–644. Google Scholar


    K. K Bhattacharjee 1986. Intestinal rupture in a free-ranging sambar deer (Cervus unicolor) in India. Journal of Wildlife Diseases 22:136. Google Scholar


    S Biswas and K Sankar 2002. Prey abundance and food habit of tigers (Panthera tigris tigris) in Pench National Park, Madhya Pradesh, India. Journal of Zoology (London) 256:411–420. Google Scholar


    W. T Blanford 1888. The fauna of British India, including Ceylon and Burma. Taylor and Francis, London, United Kingdom. Google Scholar


    C. L. J. L Bonaparte 1837 Iconografia della fauna Italica per le quattro classi degli animali vertebrati. Fascicolo XV–XVI. Salviucci, Rome, Italy. [Dated 1836, but published in 1837 according to Salvadori 1888.]. Google Scholar


    A Bonnet-Garnier F Claro S Thévenon M Gautier and H Hayes 2003. Identification by R-banding and FISH of chromosome arms involved in Robertsonian translocations in several deer species. Chromosome Research 11:649–663. Google Scholar


    D. J Brand 1963. Records of mammals bred in the National Zoological Gardens of South Africa during the period 1908 to 1960. Proceedings of the Zoological Society of London 140:617–659. Google Scholar


    A. A. D Brander 1923. Wild animals in central India. Edward Arnold & Company, London, United Kingdom. Google Scholar


    A Braun C. P Groves P Grubb Q-S Yang and L Xia 2001. Catalogue of the Musée Heude collection of mammal skulls. Acta Zootaxonomica Sinica 26:608–660. Google Scholar


    J. F Brodie and W. Y Brockelman 2010. Bed site selection of red muntjac (Muntiacus muntjac) and sambar (Rusa unicolor) in a tropical seasonal forest. Ecological Research 24:1251–1256. Google Scholar


    J. F Brodie O. E Helmy W. Y Brockelman and J. L Maron 2009. Functional differences within a guild of tropical mammalian frugivores. Ecology 90:688–698. Google Scholar


    J Brookes 1828. A catalogue of the anatomical and zoological museum of Joshua Brookes, esq. F.R.S., F.L.S. &c. George Robins, London, United Kingdom. Google Scholar


    J. B Brooksby 1973. Observations on foot-and-mouth disease in Sri Lanka. Ceylon Veterinary Journal 21:40–45. Google Scholar


    G. A Bubenik 1993. Morphological differences in the antler velvet of Cervidae. Pp. 56–64 in Deer in China: biology and management ( N Ohtaishiand H-L Sheng eds.). Elsevier Science Publishers, Amsterdam, The Netherlands. Google Scholar


    G Caughley 1963. Dispersal rates of several ungulates introduced into New Zealand. Nature 200:280–281. Google Scholar


    A Chakrborty 1994. Occurrence and pathology of Gongylonema in captive wild herbivores. Veterinary Parasitology 52:163–167. Google Scholar


    J. P-W Chan H-Y Tsai C-F Chen K-C Tung and C-C Chang 2009. The reproductive performance of female Formosan sambar deer (Cervus unicolor swinhoei) in semi-domesticated herds. Theriogenology 71:1156–1161. Google Scholar


    H. S Chandra D. A Hungerford J Wagner and R. L Snyder 1967. Chromosomes of five artiodactyl mammals. Chromosoma 21:211–220. Google Scholar


    F. N Chasen and C. B Kloss 1928. Spolia Mentawiensis—mammals. Proceedings of the General Meetings for Scientific Business of the Zoological Society of London 1927:797–840 [Dated 1927, but Part IV published in January 1928.]. Google Scholar


    R Chellam and A. J. T Johnsingh 1993. Management of Asiatic lions in Gir Forest, India. Symposia of the Zoological Society of London 65:409–424. Google Scholar


    F-P Cheng et al. 2004. The effect of different extenders on post-thaw sperm survival, acrosomal integrity and longevity in cryopreserved semen of Formosan sika deer and Formosan sambar deer. Theriogenology 61:1605–1616. Google Scholar


    D. L Chesemore 1970. Notes on the mammals of southern Nepal. Journal of Mammalogy 51:162–166. Google Scholar


    M Clauss M Lechner-Doll and W. J Streich 2002. Faecal particle size distribution in captive wild ruminants: an approach to the browser/grazer dichotomy from the other end. Oecologia 131:343–349. Google Scholar


    T. H Clutton-Brock S. D Albon and P. H Harvey 1980. Antlers, body size, and breeding group size in the Cervidae. Nature 285:565–567. Google Scholar


    T. H Clutton-Brock F. E Guinness and S. D Albon 1982. Red deer: behavior and ecology of two sexes. University of Chicago Press, Chicago, Illinois. Google Scholar


    J. A Cohen 1978. Cuon alpinus. Mammalian Species 100:1–3. Google Scholar


    J. A Cohen M. W Fox A. J. T Johnsingh and B. D Barnett 1978. Food habits of the dhole in south India. Journal of Wildlife Management 42:933–936. Google Scholar


    E Comber 1904. The breeding season of big game. Journal of the Bombay Natural History Society 16:176–179. Google Scholar


    P. J Conry 1988. Management of feral and exotic game species on Guam. Transactions of the Western Section of The Wildlife Society 24:26–30. Google Scholar


    G. B Corbet and J. E Hill 1992. The mammals of the Indomalayan region: a systematic review. Oxford University Press, Oxford, United Kingdom. Google Scholar


    L. S Crandall 1964. The management of wild mammals in captivity. University of Chicago, Chicago, Illinois. Google Scholar


    M. A Cronin 1991. Mitochondrial-DNA phylogeny of deer (Cervidae). Journal of Mammalogy 72:553–566. Google Scholar


    M. A Cronin D. A Palmisciano E. R Vyse and D. G Cameron 1991. Mitochondrial DNA in wildlife forensic science: species identification of tissues. Wildlife Society Bulletin 19:94–105. Google Scholar


    G Cuvier 1823. Recherches sur les ossemens fossils, où l'on rétablit les charactères de plusieurs animaux dont les revolutions du glore ont detruit les espéces. Tome quatrieme. Troisiéme partie. Sur les ossemens fossils de ruminans. Chez G. Dufour et E. D'Ocagne, Libraries, Paris, France. Google Scholar


    I Dahlan and N. A Norfarizan-Hanoon 2007. Fatty acid profiles and cholesterol composition of venison from farmed deer. Journal of Animal and Veterinary Advances 6:650–657. Google Scholar


    I Dahlan and N. A Norfarizan-Hanoon 2008. Chemical composition, palatability and physical characteristics of venison from farmed deer. Animal Science Journal 79:498–503. Google Scholar


    A Datta M. O Anand and R Naniwadekar 2008. Empty forests: large carnivore and prey abundance in Namdapha National Park, north-east India. Biological Conservation 141:1429–1435. Google Scholar


    S Datta 1954. Preliminary report on incidence and prevalence of tuberculosis in cattle and domestic animals with special reference to localization and type of bacilli. Indian Journal of Tuberculosis 1:89–92. Google Scholar


    S Davar 1938. Cause of sore neck in sambar. Journal of the Bombay Natural History Society 40:118–122. Google Scholar


    W. R Davidson J. L Blue L. B Flynn S. M Shea R. L Marchinton and J. A Lewis 1987. Parasites, diseases and health status of sympatric populations of sambar deer and white-tailed deer in Florida. Journal of Wildlife Diseases 23:267–272. Google Scholar


    H. M. D de Blainville 1816. Sur plusieurs espèces d'animaux mammifères, de l'ordre des ruminans. Bulletin des Sciences par la Société Philomatique de Paris 1816:73–82. Google Scholar


    H. M. D de Blainville 1822. Sur les caractères distinctifs des espèces de cerfs. Journal de Physique, de Chime, d'Histoire Naturelle et des Arts, avec des Planches en Taille-douce 94:254–285. Google Scholar


    E de Pousargues 1896. Sur la faune mammalogique de Setchuan et sur une espèce asiatique du genre Zapus. Bulletin du Muséum d'Histoire Naturelle 2:11–16. Google Scholar


    V Dhanda H Hoogstrool and H. R Bhat 1970. Haemaphysalis (Kaiseriana) ramachandrai sp. n. (Ixodoidea, Ixodidae), a parasite of man and domestic and wild mammals in northern India and Nepal. Journal of Parasitology 56:823–831. Google Scholar


    E Dinerstein 1979. An ecological survey of the Royal Karnali-Bardia Wildlife Reserve, Nepal. Part II. Habitat/animal interactions. Biological Conservation 18:265–300. Google Scholar


    E Dinerstein 1980. An ecological survey of the Royal Karnali-Bardia Wildlife Reserve, Nepal. Part III. Ungulate populations. Biological Conservation 18:5–38. Google Scholar


    E Dinerstein 1989. The foliage-as-fruit hypothesis and the feeding behavior of South Asian ungulates. Biotropica 21:214–218. Google Scholar


    G Di Stefano and C Petronio 2002. Systematics and evolution of the Eurasian Plio-Pleistocene tribe Cervini (Artiodactyla, Mammalia). Geologica Romana 36:311–334. Google Scholar


    Divyabhanusinh. 1988. Interaction between sambar (Cervus unicolor) and Indian wild dog (Cuon alpinus) in Sariska National Park. Journal of the Bombay Natural History Society 85:410–411. Google Scholar


    W Dong 1993. The fossil record of deer in China. Pp. 95–102 in Deer in China: biology and management ( N Ohtaishiand H-L Sheng eds.). Elsevier Science Publishers, Amsterdam, The Netherlands. Google Scholar


    M. J. W Douglas 1970. Dental cement layers as criteria of age for deer in New Zealand with emphasis on red deer, Cervus elaphus. New Zealand Journal of Science 13:352–358. Google Scholar


    M. J. W Douglas 1983. Status and future management of the Manawatu sambar deer herd. FRI Bulletin 30:1–15. Google Scholar


    M Downes 1983a. The Sambar Consultancy. Second report, October, 1979. Australian Deer 4(6):4–15. Google Scholar


    M Downes 1983b. The forest deer project 1982. Australian Deer Research Foundation Limited, Melbourne, Australia. Google Scholar


    J Drozdz 1965. Studies on helminths and helminthiasis in Cervidae. I. Revision of the subfamily Ostertagiinae Sarwar, 1956 and an attempt to explain the phylogenesis of its representatives. Acta Parasitologica Polonica 13:445–481. Google Scholar


    J Drozdz 1973. Materials contributing to the knowledge of the helminth fauna of Cervus (Russa) unicolor Kerr and Muntjacus muntjac Zimm of Vietnam including two new nematode species: Oesophagostomum labiatum sp. n. and Trichocephalus muntjaci sp. n. Acta Parasitologica Polonica 21:465–474. Google Scholar


    G. M Dryden 2008. Animal nutrition science. CABI Press, Wallingford, United Kingdom. Google Scholar


    J. W Duckworth R. E Salter and K Khounboline (comps) 1999. Wildlife in Lao PDR: 1999 status report. World Conservation Union, Wildlife Conservation Society, and Centre for Protected Areas and Watershed Management, Vientiane, Laos. Google Scholar


    Duke of Bedford. F. H. A Marshall 1942. On the incidence of the breeding season in mammals after transference to a new latitude. Proceedings of the Royal Society, B. Biological Sciences 130:396–399. Google Scholar


    F. M Duncan 1937. On the dates of publications of the Society's ‘Proceedings,’ 1859–1926. With an appendix containing the dates of publication of ‘Proceedings,’ 1830–1858, complied by the late F. H. Waterhouse, and of the ‘Transactions,’ by the late Henry Peavot, originally published in P.Z.S. 1893, 1913. Proceedings of the Zoological Society of London, A. General and Experimental 107:71–84 . Google Scholar


    F. L Dunn 1964. Erythrocyte sickling in the barking deer of Borneo. Journal of Mammalogy 45:492–493. Google Scholar


    M Earle 1987. A flexible body mass in social carnivores. American Naturalist 129:755–760. Google Scholar


    J. F Eisenberg 1987. The evolutionary history of the Cervidae with special reference to the South American radiation. Pp. 60–64 in Biology and management of the Cervidae ( C. M Wemmer ed.). Smithsonian Institution Press, Washington, D.C. Google Scholar


    J. F Eisenburg and M Lockhart 1972. An ecological reconnaissance of Wilpattu National Park, Ceylon. Smithsonian Contribution to Zoology 101:1–118. Google Scholar


    H. A. A El-Jaafari J. M Panandam I Idris and S. S Sriaj 2008. RAPD analysis of three deer species in Malaysia. Asian-Australasian Journal of Animal Sciences 21:1233–1237 . Google Scholar


    B. C Emerson and M. L Tate 1993. Genetic analysis of evolutionary relationships among deer (subfamily Cervinae). Journal of Heredity 84:266–273. Google Scholar


    J. C. P Erxleben 1777. Systema regni animalis per classes, ordines, genera, species, varietates cum synonymia et historia animalium. Classis I. Mammalia. Impensis Weygandianis, Lipsiae, Germany. Google Scholar


    G. P Evans 1912. Big-game shooting in Upper Burma. Longmans, Green, and Company, London, United Kingdom. Google Scholar


    I. H. N Evans 1918. Some Sakai beliefs and customs. Journal of the Royal Anthropological Institute of Great Britain and Ireland 48:179–197. Google Scholar


    T Evgenjeva 1991. Integumentary glands of sambar deer (Cervus unicolor Kerr) in culture. Proceedings of the International Symposium Ongulés/Ungulates 91:459–460 . Google Scholar


    G. A Feldhammer 1980. Cervus nippon. Mammalian Species 128:1–7. Google Scholar


    J. B Fischer 1829. Addenda, emendanda et index ad synopsis mammalium. Sumtibus J. G. Chottae, Stuttgardtiae, Germany. Google Scholar


    L. J Fitzinger 1875. Kritische Untersuchungen uber die Arten der naturlichen Familie der Airsche (Cervi). Sitzungsberichte de Kaiserlichen Akademie der Wissenschaften, Wein 70:239–333 [Dated 1874, but published in 1875.]. Google Scholar


    K. K Flerov 1952. Musk deer and deer. Fauna of USSR. Mammals. Vol. 1. No. 2. National Science Foundation and Smithsonian Institution, Washington, D.C. [Israel Program for Scientific Translation.]. Google Scholar


    F. W. F Fletcher 1911. Sport on the Nilgiris and in Wynaad. Macmillan and Company, Limited, London, United Kingdom.  Google Scholar


    L. B Flynn S. M Shea J. C Lewis and R. L Marchinton 1990. Part III. Population statistics, health and habitat use. Pp. 63–107 in Biology of sambar deer on St. Vincent National Wildlife Refuge, Florida. Bulletin of Tall Timbers Research Station 25:1–107. Google Scholar


    D. M Forsyth and R. P Duncan 2001. Propagule size and the relative success of exotic ungulate and bird introductions to New Zealand. American Naturalist 157:583–595. Google Scholar


    H Fox 1939. Chronic arthritis in wild mammals. Being a description of lesions found in the collections of several museums and from a pathological service. Transactions of the American Philosophical Society, New Series 31:73–148. Google Scholar


    C. M Francis 2009. A guide to the mammals of southeast Asia. Princeton University Press, Princeton, New Jersey. Google Scholar


    A. W Franzmann 1981. Alces alces. Mammalian Species 154:1–7. Google Scholar


    K. W Fraser J. M Cone and E. J Whitford 2000. A revision of the established ranges and new populations of 11 introduced ungulate species in New Zealand. Journal of the Royal Society of New Zealand 30:419–437. Google Scholar


    B Gangadharan K. V Balsala M. G Nair and A Rajan 1992. Sarcocystosis in sambar deer (Cervus unicolor). Indian Journal of Animal Sciences 62:127–128. Google Scholar


    P. M Gardner 1993. Dimensions of subsistence foraging in South India. Ethnology 32:109–144. Google Scholar


    V Geist 1998. Deer of the world: their evolution, behavior, and ecology. Stackpole Books, Mechanicsburg, Pennsylvania.  Google Scholar


    A. W Gentry 2000. The ruminant radiation. Pp. 11–25 in Antelopes, deer, and relatives: fossil record, behavioral ecology, systematics, and conservation ( E. S Vrbaand G. B Schaller eds.). Yale University Press, New Haven, Connecticut. Google Scholar


    É Geoffroy Saint-Hilaire and F Cuvier 1819. Unnumbered page associated with pl. 358, vol. i, livr. 10 in Historie naturelle des mammifères, avec figures originales, coloriées, dessinées d'après des animaux vivants; publiée sous l'autorité de l'Administration du Muséum d'Histoire Naturelle. Tome cinquième. Chez A. Belin, Libraire-Éditeur, Paris, France. Google Scholar


    C Gilbert A Ropiquet and A Hassanin 2006. Mitochondrial and nuclear phylogenies of Cervidae (Mammalia, Ruminantia): systematics, morphology, and biogeography. Molecular Phylogenetics and Evolution 40:101–117. Google Scholar


    R Gilbert 1888. Notes on sambhur and sambhur stalking. Journal of the Bombay Natural History Society 3:224–232. Google Scholar


    A. I. R Glasfurd 1896. Rifle and romance in the Indian jungle: a record of thirteen years. John Lane: The Bodley Head, London, United Kingdom. Google Scholar


    A. I. R Glasfurd 1903. Leaves from an Indian jungle gathered during thirteen years of a jungle life in the central provinces, the Deccan, and Berar. Times Press, Bombay, India. Google Scholar


    G. A Goldfuss 1820. Handbuch der Zoologie. Johann Leonhard Schrag, Nürnberg, Germany. Google Scholar


    J. E Gray 1843. List of the specimens of Mammalia in the collection of the British Museum. George Woodfall and Son, London, United Kingdom.  Google Scholar


    J. E Gray 1861. List of Mammalia, tortoises and crocodiles collected by M. Mouhot in Camboja. Proceedings of the Zoological Society of London 1861:135–140. Google Scholar


    J. E Gray 1872. Catalogue of ruminant Mammalia (Pecora, Linnaeus) in the British Museum. Taylor and Francis, London, United Kingdom. . Google Scholar


    M. J. B Green 1985. Aspects of the ecology of the Himalayan musk deer. Ph.D. dissertation, University of Cambridge, Cambridge, United Kingdom. Google Scholar


    M. J. B Green 1987. Ecological separation in Himalayan ungulates. Journal of Zoology (London), Series B 1:693–719 . Google Scholar


    M Griffiths and C. P van Schaik 1993. The impact of human traffic on the abundance and activity periods of Sumatran rain forest wildlife. Conservation Biology 7:623–626. Google Scholar


    C. P Groves 2003. Taxonomy of ungulates of the Indian subcontinent. Journal of the Bombay Natural History Society 100:314–362. Google Scholar


    C. P Groves 2006. The genus Cervus in eastern Eurasia. European Journal of Wildlife Research 52:14–22. Google Scholar


    C. P Groves 2007. Family Cervidae. Pp. 249–256 in The evolution of artiodactyls ( D. R Protheroand S. E Foss eds.). Johns Hopkins University Press, Baltimore, Maryland. Google Scholar


    C. P Groves and P Grubb 1987. Relationships of living deer. Pp. 21–59 in Biology and management of the Cervidae ( C. M Wemmer ed.). Smithsonian Institution Press, Washington, D.C. Google Scholar


    P Grubb 1990. Cervidae of Southeast Asia. Pp. 169–179 in Horns, pronghorns, and antlers ( G. A Bubenik A. B Bubenik eds.). Spring Verlag, New York. Google Scholar


    P Grubb 2005. Order Artiodactyla. Pp. 637–722 in Mammal species of the world: a taxonomic and geographic reference ( D. E Wilsonand D. M Reeder eds.). 3rd ed. Johns Hopkins University Press, Baltimore, Maryland. Google Scholar


    P Grubb and C. P Groves 1983. Notes on the taxonomy of the deer (Mammalia, Cervidae) of the Philippines. Zoologischer Anzeiger, Jena 210:119–144. Google Scholar


    S Guha and V. K Kashyap 2005. Development of novel heminested PCR assays based on mitochondrial 16s rRNA gene for identification of seven pecora species. BMC Genetics 6:42–49. Google Scholar


    S. K Haas V Hayssen and P. R Krausman 2005. Panthera leo. Mammalian Species 762:1–11. Google Scholar


    T Haltenorth 1963. Handbuch der Zoologie Band VII: Mammalia. 1. Teil: Die Klassifikation der Säugetiere. 18. Ordnung Paarhufer Artiodactyla Owen, 1848. Walter de Gruyter & Co., München, Germany. Google Scholar


    C Hamilton-Smith 1827a. The class Mammalia. Supplement to the order Ruminantia. Pp. 105–165 in The animal kingdom, arranged in conformity with its organization, by the Baron Cuvier, with additional descriptions of all the species hitherto named, and of many not before noticed ( E Griffith C Hamilton-Smithand E Pidgeon eds.). Vol. IV. G. B. Whittaker, London, United Kingdom. Google Scholar


    C Hamilton-Smith 1827b. Synopsis of the species of the class Mammalia, as arranged with reference to their organization. Order VII.—Ruminantia. Pecora, Lin. Pp. 303–321 in The animal kingdom, arranged in conformity with its organization, by the Baron Cuvier, with additional descriptions of all the species hitherto named, and of many not before noticed ( E Griffith C Hamilton-Smithand E Pidgeon eds.). Vol. V. G. B. Whittaker, London, United Kingdom. Google Scholar


    L. H Harris 1971. Notes on the introduction and history of sambar deer in New Zealand. New Zealand Wildlife 35:33–42. Google Scholar


    V Hayssen A van Tienhoven and A van Tienhoven 1993. Asdell's patterns of mammalian reproduction: a compendium of species-specific data. Comstock University Press, Ithaca, New York.  Google Scholar


    M Hernández Fernández and E. S Vrba 2005. A complete estimate of the phylogenetic relationships in Ruminantia: a dated species-level supertree of the extant ruminants. Biological Review 80:269–302. Google Scholar


    P. M Heude 1888a. Etude sur les ruminants de l'Asia orientale. Cerfs des Philipines et de l'Indo-Chine. Mémoires Concernant l'Histoire Naturelle de l'Empire Chinois par des Péres de la Compagnie de Jesus 2:1–10. Google Scholar


    P. M Heude 1888b. Etude sur les ruminants de l'Asia orientale. Essai d'un catalogue des cerfs des Philippines. Mémoires Concernant l'Histoire Naturelle de l'Empire Chinois par des Péres de la Compagnie de Jesus 2:19–41. Google Scholar


    P. M Heude 1888c. Etude sur les ruminants de l'Asia orientale. Notes sur le type Sambur, tel qu'il est en Cochinchine. Mémoires Concernant l'Histoire Naturelle de l'Empire Chinois par des Péres de la Compagnie de Jesus 2:41–46. Google Scholar


    P. M Heude 1888d. Etude sur les ruminants de l'Asia orientale. Axis et Melanaxis. Mémoires Concernant l'Histoire Naturelle de l'Empire Chinois par des Péres de la Compagnie de Jesus 2:47–51. Google Scholar


    P. M Heude 1896. Aperçu sommaire du genre Hippelaphus, groupe de la famille des cervidés proper aux Iles Malaises. Mémoires Concernant l'Histoire Naturelle de l'Empire Chinois par des Péres de la Compagnie de Jesus 3:47–52. Google Scholar


    J. E Hill 1990. A memoir and bibliography of Michael Rogers Oldfield Thomas, F.R.S. Bulletin of the British Museum (Natural History), Historical Series 18:25–113. Google Scholar


    L. S Hiregoudar 1976. Some parasites of wild ruminants in Gir Forest of India. Indian Veterinary Journal 53:237. Google Scholar


    B. H Hodgson 1831. Contributions in natural history. Gleanings in Science 3:320–324. Google Scholar


    B. H. Hodgson 1838. [readings from “A classified catalogue of Nepalese Mammalia”]. Annals and Magazine of Natural History, Series 1 1:152-154. Google Scholar


    B. H Hodgson 1841a. Classified catalogue of mammals of Nepal, corrected to end of 1841, first presented 1832. Calcutta Journal of Natural History, and Miscellany of the Arts and Sciences in India 2:212–223. Google Scholar


    B. H Hodgson 1841b. Note on Cervus Elaphus (?) of the Sâl Forest of Nepâl. Hodie, C. Affinis, nob. Journal of Asiatic Society of Bengal 10:721–724 . Google Scholar


    B. H Hodgson 1841c. Classified catalogue of mammals of Nepal (corrected to end of 1841, first presented 1832). Journal of Asiatic Society of Bengal 10:907–916. Google Scholar


    B. H Hodgson 1863. Account of the collection. Pp. iii–xii in Catalogue of the specimens and drawings of mammals, birds, reptiles and fishes of Nepal and Tibet presented by B. H. Hodgson Esq., to the British Museum ( J. E Grayand G. R Gray eds.). Taylor and Francis, London, United Kingdom. Google Scholar


    H Hoogstraal V Dhanda and H. R Bhat 1970. Haemaphysalis (Kaiseriana) davisi sp. n. (Ixodoidea, Ixodidae), a parasite of domestic and wild mammals in northeartern India, Sikkim, and Burma. Journal of Parasitology 56:588–595. Google Scholar


    H Hoogstraal and K. M El Kammah 1971. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. (H.) traubi Kohls, redescription of male, description of female, and new artiodactyl host and Malayan distribution records. Journal of Parasitology 57:426–431. Google Scholar


    H Hoogstraal G. M Kohls and H Trapido 1967. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. (Kaiseriana) anomala Warburton: redescription, hosts, and distribution. Journal of Parasitology 53:196–201. Google Scholar


    H Hoogstraal H Trapido and G. M Kohls 1965. Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. (Kaiseriana) papuana nadchatrami spp. n. and redescription of H. (K.) semermis Neumann. Journal of Parasitology 51:433–451. Google Scholar


    H Hoogstraal and H. Y Wassef 1982. Haemaphysalis (Garnhamphysalis) mjoebergi: identity, structural variation and biosystematic implications, deer hosts, and distribution in Borneo and Sumatra (Ixodoidea, Ixodidae). Journal of Parasitology 68:138–144. Google Scholar


    D. A Hooijer 1951. On the special evidence of early man in the middle Pleistocene of Southwest China. Southwestern Journal of Anthropology 7:77–81. Google Scholar


    R. A Hopkins 2005. Sambar deer Cervus unicolor. California Wildlife Habitat Relationships System, California Department of Fish and Game., accessed 20 May 2009. Google Scholar


    C Hose 1893a. A descriptive account of the mammals of Borneo. Edward Abbott, London, United Kingdom. Google Scholar


    C Hose 1893b. Description of a new deer from Mount Dulit, eastern Sarawak. Annals and Magazine of Natural History, Series 6 12:206 . Google Scholar


    A. J Howse G Semiadi K. J Stafford T. N Barry and P. D Muir 1995. Digestion and chewing behaviour of young sambar and red deer consuming low quality roughage. Journal of Agricultural Science, Cambridge 125:399–405. Google Scholar


    M. J Hsu and G Agoramoorthy 1997. Wildlife conservation in Taiwan. Conservation Biology 11:834–836. Google Scholar


    W Huang X Si Y Hou S Miller-Antonio and L. A Schepartz 1995. Excavations at Panxian Dadong, Guizhou Province, southern China. Current Anthropology 36:844–846. Google Scholar


    I Humphries and D Rowler 1976. Data analysis: sambar deer. Australian Deer 1(4):3–7. Google Scholar


    A. M Husson and L. B Holthuis 1955. The dates of publication of “Verhandelinge over de Natuurlijke Geschiedenis de Nederlansche Overzeesche Bezittingen” edited by C. J. Temminck. Zoologische Mededelingen 34:17–24. Google Scholar


    M. A Ibrahim 1998. Extirpation of an eyeball in a sambar. Animal Keepers' Forum 25:249–250. Google Scholar


    I Idris and S Moin 2009. Somatic chromosomes of the Borean sambar deer and rusa deer interspecific hybrids. American Journal of Applied Sciences 6:862–868. Google Scholar


    International Commission on Zoological Nomenclature. 1999. International code of zoological nomenclature. 4th ed International Trust for Zoological Nomenclature, London, United Kingdom. Google Scholar


    R Ippen V Kozojed and J Jíra 1981. Toxoplasmosis in zoo animals. Folia Parasitologica 28:109–115. Google Scholar


    C. M Janis and K. M Scott 1987. The interrelationships of higher ruminant families with special emphasis on the members of the Cervoidea. American Museum Novitates 2893:1–85 . Google Scholar


    D Jathanna K. U Karanth and A. J. T Johnsingh 2003. Estimation of large herbivore densities in the tropical forests of southern India. Journal of Zoology (London) 261:285–290. Google Scholar


    F. A Jentink and J Büttikofer 1897. Zoological results of the Dutch scientific expedition to central Borneo. The mammals. Notes from the Leyden Museum 19:26–66. Google Scholar


    T. C Jerdon 1874. The mammals of India; a natural history of all the animals known to inhabit continental India. John Wheldon, London, United Kingdom. Google Scholar


    A. J. T Johnsingh 1980. Sambar, a magnificent deer. Australian Deer 5(6):3–6. Google Scholar


    A. J. T Johnsingh 1983. Large mammalian prey–predators in Bandipur. Journal of the Bombay Natural History Society 80:1–57. Google Scholar


    A. J. T Johnsingh and K Sankar 1991. Food plants of chital, sambar and cattle on Mundanthurai Plateau, Tamil Nadu, south India. Mammalia 55:57–66. Google Scholar


    K. G Johnson W Wang D. G Reid and J Hu 1993. Food habits of Asiatic leopards (Panthera pardus fusea) in Wolong Reserve, Sichuan, China. Journal of Mammalogy 74:646–650. Google Scholar


    B. C Kar N Hota and L. N Acharjyo 1983. Occurrence of foot-and-mouth disease among some wild ungulates in captivity. Indian Veterinary Journal 60:237–239. Google Scholar


    K. U Karanth and M. E Sunquist 1992. Population structure, density and biomass of large herbivores in the tropical forests of Nagarahole, India. Journal of Tropical Ecology 8:21–35. Google Scholar


    K. U Karanth and M. E Sunquist 1995. Prey selection by tiger, leopard and dhole in tropical forests. Journal of Animal Ecology 64:439–450. Google Scholar


    K. U Karanth and M. E Sunquist 2000. Behavioral correlates of predation by tiger (Panthera tigris), leopard (Panthera pardus) and dhole (Cuon alpinus) in Nagarahole, India. Journal of Zoology (London) 250:255–265. Google Scholar


    R. K Keith and K Keith 1969. Ceylonocotyle streptocoelium in feral ruminants in the Northern Territory of Australia. Australian Veterinary Journal 45:594. Google Scholar


    S. D Kelton and J. P Skipworth 1987. Food of sambar deer (Cervus unicolor) in a Manawatu (New Zealand) flax swamp. New Zealand Journal Ecology 10:149–154. Google Scholar


    R Kerr 1792. The animal kingdom, or zoological system, of the celebrated Sir Charles Linnaeus; Class I. Mammalia: containing a complete systematic description, arrangement, and nomenclature, of all the known species and varieties of the Mammalia, or animals which give suck to their young; being a translation of that part of the Systema Naturae, as lately published, with great improvements, by Professor Gmelin of Geottingen. J. Murray and R. Faulder, London, United Kingdom. Google Scholar


    J. A Khan 1994. Food habits of ungulates in dry tropical forests of Gir Lion Sanctuary, Gujarat, India. Acta Theriologica 39:185–193. Google Scholar


    J. A Khan R Chellan and A. J. T Johnsingh 1995. Group size and age–sex composition of three major ungulate species in Gir Lion Sanctuary, Gujarat, India. Journal of the Bombay Natural History Society 92:295–302. Google Scholar


    J. A Khan R Chellan W. A Rodgers and A. J. T Johnsingh 1996. Ungulate densities and biomass in the tropical dry deciduous forests of Gir, Gujarat, India. Journal of Tropical Ecology 12:149–162. Google Scholar


    J. A Khan W. A Rodgers A. J. T Johnsingh and P. K Mathur 1994. Tree and shrub mortality and debarking by sambar Cervus unicolor (Kerr) in Gir after a drought in Gujarat, India. Biological Conservation 68:149–154. Google Scholar


    M Khan and M Khan 1968. Deer biological data. Malayan Nature Journal 21:159–164. Google Scholar


    D. Z Khun and L. S Kan 1991. Populations of some mammals in the sclerophylous evergreen tropic forests near Konhkanyng (South Vietnam). Zoologicheskii Zhurmal 70:114–118. Google Scholar


    J. J Kirkpatrick P. P Calle P Kalk I. K. M Liu and J. W Turner Jr 1996. Immunocontraception of captive exotic species. II. Formosan sika deer (Cervus nippon taiouanus), axis deer (Cervus axis), Himalayan tahr (Hemitragus jemlahicus), Roosevelt elk (Cervus elaphus roosevelti), Reeves' muntjac (Muntiacus reevesi), and sambar deer (Cervus unicolor). Journal of Zoo and Wildlife Medicine 27:482–495. Google Scholar


    T Koizumi N Ohtaishi K Kaji Y Yu and K Tokida 1993. Conservation of white-lipped deer in China. Pp. 309–318 in Deer in China: biology and management ( N Ohtaishiand H-L Sheng eds.). Elsevier Science Publishers, Amsterdam, The Netherlands. Google Scholar


    M Kretzoi and M Kretzoi 2000a. Index generum et subgenerum Mammalium. Fossilium Catalogus I: Animalia. Pars 137, Section 1. Backhuys Publishers, Leiden, The Netherlands. Google Scholar


    M Kretzoi and M Kretzoi 2000b. Index generum et subgenerum mammalium. Fossilium Catalogus I: Animalia. Pars 137, Section 2. Backhuys Publishers, Leiden, The Netherlands. Google Scholar


    H. N Kumara M Ananda Kumar A. K Sharma H. S Sushma M Singh and M Singh 2004. Diversity and management of wild mammals in tea gardens in the rainforest regions of the Western Ghats, India: a case study from a tea estate in the Anaimalai Hills. Current Science 87:1282–1287. Google Scholar


    F Kurt 1978. Socio-ecological organization and aspects of management of South Asian deer. Pp. 219–239 in Threatened deer: proceedings of a working meeting of the Deer Specialist Group of the Survival Service Commission. International Union for the Conservation of Nature and Natural Resources, Gland, Switzerland. Google Scholar


    F Kurtén 1968. Pleistocene mammals of Europe. Weidenfeld and Nicolson, London, United Kingdom. Google Scholar


    S. P. S Kushwaha A Khan B Habib A Quadri and A Singh 2004. Evaluation of sambar and muntjac habitats using geostatistical modelling. Current Science 86:1390–1400. Google Scholar


    D Lao 1968. Beitrag zur Geweihentwicklung und Fortpflanzungsbiologie de Hirsche. Zeitscrift für Säugetierkunde 33:193–214. Google Scholar


    W. A Laurie E. M Lang and C. P Groves 1983. Rhinoceros unicornis.. Mammalian Species 211:1–6. Google Scholar


    C. C Lee A. R Sheikh-Omar and A Jones 1987. Calicophoron microbothrioides (Price and McIntosh, 1944) (Paramphistomidae: Paramphistominae) in Malaysian sambar deer (Cervus unicolor). New Zealand Veterinary Journal 35:190–191. Google Scholar


    P. C Lent 1988. Ovibos moschatus. Mammalian Species 302:1–9. Google Scholar


    M Lentz R. L Marchington L. B Flynn S. M Shea and P. J Stuart 1986. The immobilization of sambar deer with succinylcholine chloride. Australian Deer 11(1):3–9. Google Scholar


    D. M Leslie Jr 2008. Boselaphus tragocamelus (Artiodactyla: Bovidae). Mammalian Species 813:1–17. Google Scholar


    D. M Leslie Jr 2009. Przewalskium albirostre (Artiodactyla: Cervidae). Mammalian Species 42(849):7–18. Google Scholar


    D. M Leslie Jr R. T Bowyer and J. A Jenks 2008. Facts from feces: nitrogen still measures up as a nutritional index for mammalian herbivores. Journal of Wildlife Management 72:1420–1433. Google Scholar


    D. M Leslie Jr and K. J Jenkins 1985. Rutting mortality of male Roosevelt elk. Journal of Mammalogy 66:163–164. Google Scholar


    D. M Leslie Jr and G. B Schaller 2009. Bos grunniens and Bos mutus (Artiodactyla: Bovidae). Mammalian Species 836:1–17. Google Scholar


    D. M Leslie Jr and K Sharma 2009. Tetracerus quadricornis (Artiodactyla: Bovidae). Mammalian Species 843:1–11. Google Scholar


    D. M Leslie Jr and E. E Starkey 1985. Fecal indices to dietary quality of cervids in old-growth forests. Journal of Wildlife Management 49:142–146. Google Scholar


    D. M Leslie Jr E. E Starkey and M Vavra 1984. Elk and deer diets in old-growth forests in western Washington. Journal of Wildlife Management 48:762–775. Google Scholar


    C Lever 1985. Naturalized mammals of the world. Longman Group Limited, London, United Kingdom. Google Scholar


    N. D Levine 1971. Taxonomy of the piroplasms. Transactions of the American Microscopical Society 90:2–33. Google Scholar


    J. C Lewis L. B Flynn R. L Marchinton S. M Shea and E. M Marchinton 1990. Part I. Introduction, study area, and literature review. Pp. 1–12 in Biology of sambar deer on St. Vincent National Wildlife Refuge, Florida. Bulletin of Tall Timbers Research Station 25:1–107. Google Scholar


    C Linnaeus 1758. Systema naturae per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Holmiae, Impensis Direct. Laurentii Salvii, Stockholm, Sweden. Google Scholar


    A. M Lister 1987. Diversity and evolution of antler form in Quaternary deer. Pp. 81–98 in Biology and management of the Cervidae ( C. M Wemmer ed.). Smithsonian Institution Press, Washington, D.C.. Google Scholar


    A. M Lister 1993. The stratigraphical significance of deer species in the Cromer Forest-bed Formation. Journal of Quaternary Science 8:95–108. Google Scholar


    X-H Liu Y-Q Wang Z-Q Liu and K-Y Zhou 2003. Phylogenetic relationship of Cervinae based on sequence of mitochondrial cytochrome b gene. Zoological Research 24:27–33. Google Scholar


    P Lo 1985. Movement, home ranges and habitat utilisation by sambar deer (Cervus unicolor) in Santoft State Forest, Manawatu. New Zealand Forest Service, Wellington, New Zealand. (not seen, cited in Shea et al. 1990). Google Scholar


    J. R Luick 1983. The velvet antler industry. Pp. 329–337 in Antler development in Cervidae ( R. D Brown ed.). Caesar Kleberg Wildlife Research Institute, Kingsville, Texas. Google Scholar


    R Lydekker 1898. The deer of all lands: a history of the family Cervidæ living and extinct. Rowland Ward, Limited, London, United Kingdom. Google Scholar


    R Lydekker 1915. Catalogue of the ungulate mammals in the British Museum (Natural History). Vol. IV. Artiodactyla, families Cervidæ (deer), Tragulidæ (chevrotains), Camelidæ (camels and llamas), Suidæ (pigs and peccaries), and Hippopotamidæ (hippopotamuses). Trustees of the British Museum, London, United Kingdom.  Google Scholar


    R Lydekker 1916. Wild life of the world: a descriptive survey of the geographical distribution of animals. Frederick Warne and Company, London, United Kingdom. Google Scholar


    M Lyon 1906. Mammals of Banka, Mendanan, and Billiton Islands, between Sumatra and Borneo. Proceedings of the United States National Museum 31(1498):575–612. [Dated 1907, but published 18 December 1906.]. Google Scholar


    J MacKinnon 2008. Order Artiodactyla. Pp. 451–480 in A guide to the mammals of China ( A. T Smithand Y Xie eds.). Princeton University Press, Princeton, New Jersey. Google Scholar


    R. H Manville 1957. Longevity of captive mammals. Journal of Mammalogy 38:279–280. Google Scholar


    E Mary and M Balakrishnan 1984. A study on olfactory signals in sambar deer, Cervus unicolor. Proceedings of the Indian Academy of Sciences 93:71–76. Google Scholar


    E Mason 1994. Auditory and visual communications of sambar. Australian Deer 19(2):22–26. Google Scholar


    H Matsubayashi P Lagan N Majalap J Tangah J. R. A Sukor and K Kitayama 2007a. Importance of natural licks for the mammals in Bornean inland tropical rain forests. Ecological Research 22:742–748. Google Scholar


    H Matsubayashi P Lagan J. R. A Sukor and K Kitayama 2007b. Seasonal and daily use of natural licks by sambar deer (Cervus unicolor) in a Bornean tropical rain forest. Tropics 17:81–86. Google Scholar


    V Mazák 1981. Panthera tigris. Mammalian Species 152:1–8. Google Scholar


    J. I Mead 1989. Nemorhaedus goral. Mammalian Species 335:1–5. Google Scholar


    E Meijaard and C. P Groves 2004. Morphometrical relationship between South-east Asian deer (Cervidae, tribe Cervini): evolutionary and biogeographic implications. Journal of Zoology (London) 263:179–196. Google Scholar


    V Menon 2009. Mammals of India. Princeton University Press, Princeton, New Jersey. Google Scholar


    H. R Mishra 1982. The ecology and behaviour of chital (Axis axis) in the Royal Chitwan National Park, Nepal. Ph.D. dissertation, University of Edinburgh, Edinburgh, United Kingdom (not seen, cited in Putman 1988). Google Scholar


    M. M Miyamoto F Kraus and O. A Ryder 1990. Phylogeny and evolution of antlered deer determined from mitochondrial DNA sequences. Proceedings of the National Academy of Sciences 87:6127–6131. Google Scholar


    S Mohammad Ali 1982. Some observations on the group size of the sambar Cervus unicolor Kerr, with reference to their biomass at Hazaribagh National Park, Bihar. Proceedings of the Symposium of the Ecology of Animal Populations, Zoological Survey of India 4:105–130. Google Scholar


    A Moriarty 2004. The liberation, distribution, abundance and management of wild deer in Australia. Wildlife Research 31:291–299. Google Scholar


    R. C Morris 1938. ‘Stomping grounds’ and ‘sore neck’ in sambar. Journal of the Bombay Natural History 40:560–561. Google Scholar


    P. D Muir G Semiadi G. W Asher T. E Broad M. L Tate and T. N Barry 1997. Sambar deer (Cervus unicolor) × red deer (C. elaphus) interspecies hybrids. Journal of Heredity 88:366–372. Google Scholar


    S. S Müller and H Schlegel 1845. Over de Herten van den Indischen Archipel. Verhandeligen over de Natuurlijke Geschiedenis der Nederlandsche Overzeesche Bezittingen door de leden der Natuurkundige Commissie voor Nederlansch-Indie ( C. J Temminck ed.). Zoologie, part 1. S. en J. Luchtmans en C. C. van der Hoek, Leiden, The Netherlands. [Dated 1844, but published 26 June 1845 according to Husson and Holthuis 1955.]. Google Scholar


    E. C Mungall 2007. Exotic animal field guide: nonnative hoofed mammals in the United States. Texas A&M University Press, College Station. Google Scholar


    New Zealand Department of Conservation. 2005. Public consultation of sambar deer management in the Horowhenua, Manawatu, Rangitikei and Wanganui area. Wanganui Conservancy, Wanganui, New Zealand. Google Scholar


    C Ngampongsai 1977. Habitat relations of the sambar (Cervus unicolor) in Khao-Yai National Park, Thailand. Ph.D. dissertation, Michigan State University, East Lansing. Google Scholar


    C Ngampongsai 1987. Habitat use by the sambar (Cervus unicolor) in Thailand: a case study for Khao-Yai National Park. Pp. 289–298 in Biology and management of the Cervidae ( C. M Wemmer ed.). Smithsonian Institution Press, Washington, D.C. Google Scholar


    H Nitsche 1898. Studien über Hirsche (Gattung Cervus im Weitesten Sinne). Heft 1. Untersuchungen über Mehrstangige Geweihe und die Morphologie der Huftheirhömer im Allgemeinen. Verlag von Wilhelm Engelmann, Leipzig, Germany.  Google Scholar


    R. M Nowak 1999. Walker's mammals of the world. 6th ed. Johns Hopkins University Press, Baltimore, Maryland.  Google Scholar


    G Nugent K. W Fraser G. W Asher and K. G Tustin 2001. Advances in New Zealand mammalogy 1990–2000: deer. Journal of the Royal Society of New Zealand 31:263–298. Google Scholar


    T. G O'Brien M. F Kinnaird and H. T Wibisono 2003. Crouching tigers, hidden prey: Sumatran tiger and prey populations in a tropical forest landscape. Animal Conservation 6:131–139. Google Scholar


    W Ogilby 1839. Memoir on the mammalogy of the Himalayas. Pp. lvi–lxxiv in Illustrations of the botany and other branches of the natural history of the Himalayan Mountains and of the flora of Cashmere ( J. F Royle ed.). W. H. Allen, London, United Kingdom.  Google Scholar


    N Ohtaishi and Y Gao 1990. A review of the distribution of all species of deer (Tragulidae, Moschidae and Cervidae) in China. Mammal Review 20:125–144. Google Scholar


    U. K. G. K Padmalal S Takatsuki and P Jayasekara 2003. Food habits of sambar Cervus unicolor at the Horton Plains National Park, Sri Lanka. Ecological Research 18:775–782. Google Scholar


    P. S Pallas 1766. Miscellanea zoological quibus novae imprimis atque obscurae animalium species describuntur et observationibus iconibusque illustrantur. Hague comitun. Petrum van Cleef, The Hague, The Netherlands.  Google Scholar


    T. S Palmer 1904. Index generum mammalium: a list of the genera and families of mammals. North American Fauna 23:1–984. Google Scholar


    M. M Patnaik and L. N Acharjyo 1970. Notes on the helminth parasites of vertebrates in Baranga Zoo (Orissa). Indian Veterinary Journal 47:723–730. Google Scholar


    E. H Peacock 1933. A game-book for Burma & adjoining territories: the types, distribution and habits of large and small game, together with notes on game preservation, photography, tracking, still-hunting and the care and measurement of trophies. H. F. & G. Witherby, Edinburgh, United Kingdom.  Google Scholar


    J. M Peek 1982. Elk (Cervus elaphus). Pp. 851–861 in Wild mammals of North America: biology, management, and economics. Johns Hopkins University Press, Baltimore, Maryland.  Google Scholar


    V. I Peinado J. F Celdrán and J Palomeque 1999a. Basic hematological values in some wild ruminants in captivity. Comparative Biochemistry and Physiology, A. Comparative Physiology 124:199–203. Google Scholar


    V. I Peinado J. F Celdrán and J Palomeque 1999b. Blood chemistry values in some wild ruminants in captivity. Comparative Haematology International 9:175–181. Google Scholar


    T Pennant 1781. History of quadrupeds. Vol. 1. Hoofed quadrupeds. B. White, London, United Kingdom.  Google Scholar


    C Petronio T Krakhmalnaya L Bellucci and G Di Stefano 2007. Remarks on some Eurasian pliocervines: characteristics, evolution, and relationships with the tribe Cervini. Geobio 40:113–130 . Google Scholar


    E. G Phythian-Adams 1951. Jungles memories. Part IX—antelope and deer. Journal of the Bombay Natural History Society 50:1–12. Google Scholar


    C Pitra J Fickel E Meijaard and P. C Groves 2004. Evolution and phylogeny of Old World deer. Molecular Genetics and Evolution 33:880–895. Google Scholar


    E. D Plotka 1999. Deer. Pp. 842–857 in Encyclopedia of reproduction ( E Knobiland J. D Neill eds.). Vol. 1. Academic Press, San Diego, California.  Google Scholar


    R. I Pocock 1910. On the specialised cutaneous glands of ruminants. Proceedings of the Zoological Society of London 78:840–986. Google Scholar


    R. I Pocock 1933. The homologies between the branches of the antlers of the Cervidae based on the theory of dichotomous growth. Proceedings of Zoological Society of London 1933:377–406. Google Scholar


    R. I Pocock 1942a. The skull-characters of some forms of sambar (Rusa) occurring to the east of the Bay of Bengal.—Part I. Annals and Magazine of Natural History, Series 9 11:516–525. Google Scholar


    R. I Pocock 1942b. The larger deer of British India. Journal of the Bombay Natural History Society 43:298–317. Google Scholar


    R. I Pocock 1943a. The larger deer of British India. Part II. Journal of the Bombay Natural History Society 43:553–572. Google Scholar


    R. I Pocock 1943b. The larger deer of British India. Part III—The sambar (Rusa). Journal of the Bombay Natural History Society 44:27–37. Google Scholar


    D. P Poppi B. W Norton D. J Minson and R. E Hendricksen 1980. The validity of the critical size theory for particles leaving the rumen. Journal of Agricultural Science 94:275–280. Google Scholar


    S. H Prater 1980. The book of Indian animals. Bombay Natural History Society, Bombay, India.  Google Scholar


    P. J. A Presidente 1978. Diseases and parasites of captive rusa and fallow deer in Victoria. Australian Deer 3(1):23–38. Google Scholar


    P. J. A Presidente 1984a. Ectoparasites, endoparasites and some diseases reported from sambar deer throughout its native range and in Australia and New Zealand. Deer Refresher Course for Veterinarians, University of Sydney, Proceedings 72:543–557. Google Scholar


    P. J. A Presidente 1984b. Parasites of farmed and free-ranging deer in Australia. Deer Refresher Course for Veterinarians, University of Sydney, Proceedings 72:623–643. Google Scholar


    C Presnall 1958. The present status of exotic mammals in the United States. Journal of Wildlife Management 22:45–50. Google Scholar


    R Putman 1988. The natural history of deer. Comstock Publishing Associates, Ithaca, New York.  Google Scholar


    A. R Rabinowitz and S. R Walker 1991. Carnivore community in a dry tropical forest mosaic in Huai Kha Khaeng Wildlife Sanctuary, Thailand. Journal of Tropical Ecology 7:37–47. Google Scholar


    U Ramakrishnan R. G Coss and N. W Pelkey 1999. Tiger decline caused by the reduction of large ungulate prey: evidence from a study of leopard diets in southern India. Biological Conservation 89:113–120. Google Scholar


    T Ramesh V Snehalatha K Sankar and Q Qureshi 2009. Food habits and prey selection of tiger and leopard in Mudumalai Tiger Reserve, Tamil Nadu, India. Journal of Scientific Transactions in Environment and Technovation 2:170–181. Google Scholar


    E Randi N Mucci F Claro-Hergueta A Bonnet and E. J. P Douzery 2001. A mitochondrial DNA control region phylogeny of the Cervinae: speciation in Cervus and implications for conservation. Animal Conservation 4:1–11. Google Scholar


    A. T Rao and L. N Acharjyo 1969. Pathological lesions in livers of two Indian sambars (Cervus unicolor niger) infected with Paramphistomum explanatum (Creplin, 1847); Nasmark, 1937 Gigantocotyle explanatum. Indian Veterinary Journal 46:916–918. Google Scholar


    A. T Rao and L. N Acharjyo 1984. Diagnosis and classification of common diseases of captive animals at Nandan in Orissa (India). Indian Journal of Animal Health 33:147–152. Google Scholar


    B. L Raphael P Kalk P Thomas P. P Calle J. G Doherty and R. A Cook 2003. Use of melengestrol acetate in feed for contraception in herds of captive ungulates. Zoo Biology 22:455–463. Google Scholar


    H. S Reedy C Sprinivasulu and K. T Rao 2004. Prey selection by the Indian tiger (Panthera tigris tigris) in Nagarjunasagar Srisailam Tiger Reserve, India. Mammalian Biology 69:384–391. Google Scholar


    A. B Reichenbach 1835. Bildergallerie der Thierwelt order Abbilungen des Interessantesten aus dem Thierreiche mit ausführliher Beschreibung. 2nd ed. Heft 7. E. Pönicke & Sohn, Leipzig, Germany (not seen, cited in Kretzoi and Kretzoi 2000). Google Scholar


    W. A Richardson II 1972. A natural history survey of the sambar deer (Cervus unicolor) on the Powderhorn Ranch, Calhoun County, Texas. M.S. thesis, Texas A&M University, College Station. Google Scholar


    T Riney 1957. Sambar (Cervus unicolor) in Sand Hill Country. Proceedings of the New Zealand Ecological Society 5:26–27. Google Scholar


    T Salvadori 1888. Le date della pubblicazoine della “Iconografia della fauna Italica” del Bonaparte ed indice delle specie illustrate in detta opera. Bollettinu dei Musei di Zoologia ed Anatomia Comparata della Reale Università di Torino 39:1–25. Google Scholar


    C Sammaiah E Narayana C Samatha and C Sravanthi 2008. Depletion of wildlife in Eturnagaram Wildlife Sanctuary, Warangal, Andhra Pradesh. Pp. 251–255 in Wildlife Biodiversity Conservation ( M. V Reddy ed.). Daya Publishing House, New Delhi, India.  Google Scholar


    K Sankar 1994. The ecology of three large sympatric herbivores (chital, sambar and nilgai) with special reference for reserve management in Sariska Tiger Reserve, Rajasthan. Ph.D.dissertation, University of Rajasthan, Jaipur, India (not seen, cited in Sankar and Acharya 2004). Google Scholar


    K Sankar 2008. Ungulate conservation in India. Pp. 37–50 in Wildlife biodiversity conservation ( M. V Reddy ed.). Daya Publishing House, New Delhi, India.  Google Scholar


    K Sankar and B Acharya 2004. Sambar (Cervus unicolor Kerr, 1792). ENVIS Bulletin (Wildlife and Protected Areas): Ungulates of India 7., accessed 10 May 2009. Google Scholar


    K Sankar and A. J. T Johnsingh 2002. Food habits tiger (Panthera tigris) and leopard (Panthera pardus) in Sariska Tiger Reserve, Rajasthan, India, as shown by scat analysis. Mammalia 66:285–289. Google Scholar


    K Sankar A. J. T Johnsingh and R Mathur 2007. Food habits of three major ungulate species in a semi-arid zone of Rajasthan, India. Cheetal 44:18–39. Google Scholar


    G Sarma S. K Das and P. K Dutta 1983. Outbreak of foot-and-mouth disease in deer in the Assam State Zoo. Veterinary Record 113:420–421. Google Scholar


    K Sarma A Kalita S Suri and M. M. S Zama 2003. Comparative anatomical studies on the scapula of Bakarwali goat and sambar deer. Indian Journal of Animal Health 43:15–18. Google Scholar


    G. B Schaller 1967. The deer and the tiger: a study of wildlife in India. University of Chicago Press, Chicago, Illinois.  Google Scholar


    G. B Schaller 1998. Wildlife of the Tibetan steppe. University of Chicago Press, Chicago, Illinois.  Google Scholar


    H. R Schinz 1845. Systematisches Verzeichniss aller bis jetzt bekannten Säugethiere oder Synopsis Mammalium nach dem Cuvier'schen System. Vol. 2. Jent and Gassmann, Solothurn, Germany.  Google Scholar


    P. L Sclater 1862. Note on the Formosa deer. Proceedings of the Zoological Society of London 1863:150–152. Google Scholar


    P. L Sclater 1870. Report on additions to the Society's menagerie during the month of May, and description of Cervus alfredi. Proceedings of the Zoological Society of London 1870:380-381. Google Scholar


    P. L Sclater 1901. Mittheilungen aus Museen, Instituten, etc. 2. Zoological Society of London. Zoologischer Anzeiger 24:535–536. Google Scholar


    J Seidensticker 1976a. On the ecological separation between tigers and leopards. Biotropica 8:225–234. Google Scholar


    J Seidensticker 1976b. Ungulate populations in Chitwan Valley, Nepal. Biological Conservation 10:183–210. Google Scholar


    G Semiadi T. N Barry and P. D Muir 1993. Growth, milk intake, and behaviour of artificially reared sambar deer (Cervus unicolor) and red deer (Cervus elaphus). Journal of Agricultural Science, Cambridge 121:273–281. Google Scholar


    G Semiadi T. N Barry and P. D Muir 1995a. Comparison of seasonal patterns of growth, voluntary feed intake and plasma hormone concentrations in young sambar deer (Cervus unicolor) and red deer (Cervus elaphus). Journal of Agricultural Science, Cambridge 125:109–124. Google Scholar


    G Semiadi T. N Barry P. D Muir and J Hodgson 1995b. Dietary preferences of sambar (Cervus unicolor) and red deer (Cervus elaphus) offered browse, forage legume and grass species. Journal of Agricultural Science, Cambridge 125:99–107. Google Scholar


    G Semiadi T. N Barry K. J Stafford P. D Muir and C. S. W Reid 1994a. Comparison of digestive and chewing efficiency and time spent eating and ruminating in sambar deer (Cervus unicolor) and red deer (Cervus elaphus). Journal of Agricultural Science, Cambridge 123:89–97. Google Scholar


    G Semiadi C. W Holmes T. N Barry and P. D Muir 1996. Effects of cold conditions on heat production by young sambar (Cervus unicolor) and red deer (Cervus elaphus). Journal of Agricultural Science, Cambridge 126:221–226. Google Scholar


    G Semiadi C. W Holmes T. N Barry and P. D Muir 1998. The efficiency of utilization of energy and nitrogen in young sambar (Cervus unicolor) and red deer (Cervus elaphus). Journal of Agricultural Science, Cambridge 130:193–198. Google Scholar


    G Semiadi P. D Muir and T. N Barry 1994b. General biology of sambar deer (Cervus unicolor) in captivity. New Zealand Journal of Agricultural Research 37:79–85. Google Scholar


    A Shalini Kalita K Sarma and M. M. S Zama 2004a. Comparative dental anatomy of Bakarwali goat, local sheep of Jammu region and sambar deer. Indian Journal of Animal Sciences 74:1102–1104. Google Scholar


    K Shalini Sarma A Kalita and M. M. S Zama 2004b. Comparative anatomy of the atlas and axis of Bakarwali goat, local sheep, and sambar deer. Indian Veterinary Journal 81:71–73. Google Scholar


    S. M Shea L. B Flynn R. L Marchinton and J. C Lewis 1990. Part II. Social behavior, movement ecology, and food habitats. Pp. 13–62 in Biology of sambar deer on St. Vincent National Wildlife Refuge, Florida. Bulletin of Tall Timbers Research Station 25:1–107. Google Scholar


    A Sheng E Zhang Q Chen and B Ni 1993. A comparative study on morphology of deer hair. Pp. 73–84 in Deer in China: biology and management ( N Ohtaishiand H-L Sheng eds.). Elsevier Science Publishers, Amsterdam, The Netherlands.  Google Scholar


    H Sheng and N Ohtaishi 1993. The status of deer in China. Pp. 1–11 in Deer in China: biology and management 9 N Ohtaishiand H-L Sheng eds.). Elsevier Science Publishers, Amsterdam, The Netherlands.  Google Scholar


    J Shoshani and J. F Eisenberg 1982. Elephas maximus. Mammalian Species 182:1–8. Google Scholar


    R Shukla and P. K Khare 1998. Food habits of wild ungulates and their competition with livestock in Pench Wildlife Reserve, central India. Journal of the Bombay Natural History Society 95:418–421. Google Scholar


    X Si J Liu H Zhang and C Yuan 1993. Preliminary report on the excavation of Panxian Dadong, a Paleolithic cave-site in Guizhou Province. Acta Anthropologica Sinica 12:113–119. Google Scholar


    A. J Sinclair W. J Slattery and K O'Dea 1982. The analysis of polyunsaturated fatty acids in meat by capillary gas–liquid chromatography. Journal of the Science of Food and Agriculture 33:771–776 . Google Scholar


    M Sivaramakrishnan 2008. True chronicles: the jungle narratives of Jim Corbett and Kenneth Anderson—from big game hunting to conservation of wildlife and biodiversity. Pp. 37–50 in Wildlife biodiversity conservation ( M. V Reddy ed.). Daya Publishing House, New Delhi, India.  Google Scholar


    K. J Slee 1984. The sambar deer in Victoria. Deer Refresher Course for Veterinarians, University of Sydney, Proceedings 72:559–572. Google Scholar


    K. J Slee and P. J. A Presidente 1981. Biological and pathological features of sambar in Victoria. Part I. Haematology, biochemistry, and serology. Australian Deer 6(4):7–14. Google Scholar


    A. M Smith 1904. Sport and adventure in the Indian jungle. Hurst and Blackett, Limited, London, United Kingdom.  Google Scholar


    W. P Smith 1991. Odocoileus virginianus. Mammalian Species 388:1–13. Google Scholar


    V. E Sokolov T. P Evgen'eva and M Balakrishnan 1987. Architecture of axial hairs in the sambar (Cervus unicolor) by scanning electron microscopy. Doklady Akademii nauk Soiuza Sovetskikh Sotsialisticheskikh Respublik 6:1490–1494. Google Scholar


    S Srikosamatara 1993. Density and biomass of large herbivores and other mammals in a dry tropical forest, western Thailand. Journal of Tropical Ecology 9:33–42. Google Scholar


    K. K Srivastava A. K Bhardwaj S George and V. J Zacharias 1996. Micro-histological studies on the food habits of sambar, gaur and cattle in Periyar Tiger Reserve in winter. Indian Forester 122:933–936. Google Scholar


    K. J Stafford 1995. The stomach of the sambar deer (Cervus unicolor unicolor). Anatomia, Histologia, Embryologia 24:241–249. Google Scholar


    K. J Stafford 1997. The diet and trace element status of sambar deer (Cervus unicolor) in Manawatu district, New Zealand. New Zealand Journal of Zoology 24:267–271. Google Scholar


    E. P Stebbing 1911. Jungle by-ways in India: leaves from the note-book of a sportsman and a naturalist. John Lane: The Bodley Head, London, United Kingdom.  Google Scholar


    R Steinmetz W Chutipong and N Seuaturien 2006. Collaborating to conserve large mammals in Southeast Asia. Conservation Biology 20:1391–1401. Google Scholar


    C. J Sundevall 1846. Methodisk öfversigt af Idislande djuren, Linnés Pecora. Kongliga Vetenskapsakademiens Handlinger, för år 1844, New Series 32:174–185. [Dated 1844, but published in 1846 according to Corbet and Hill 1992.]. Google Scholar


    R Swinhoe 1862. On the mammals of the island of Formosa (China). Proceedings of the Zoological Society of London 1862:347–365. Google Scholar


    W. S Thom 1937. The Malayan or Burmese sambar. Journal of the Bombay Natural History Society 39:309–319. Google Scholar


    J. W Thomas R. M Robinson and R. G Marburger 1967. Use of diazepam in the capture and handling of cervids. Journal of Wildlife Management 31:686–692. Google Scholar


    O Thomas 1901. Exhibition of, and remarks upon, a peculiar stag's frontlet and horns from Borneo. Proceedings of the Zoological Society of London 1901(2):284. Google Scholar


    R. J Timmins and T. D Evans 1996. Wildlife and habitat survey of the Nakai-Nam Theun National Biodiversity Conservation Area. Wildlife Conservation Society, New York, and Centre for Protected Areas and Watershed Management, Vientiane, Lao People's Democratic Republic.  Google Scholar


    R. J Timmins et al. 2008. Rusa unicolor. 2008 IUCN Red list of threatened species. International Union for Conservation of Nature and Natural Resources., accessed 15 September 2010. Google Scholar


    E Undritz K Betke and H Lehmann 1960. Sickling phenomenon in deer. Nature 187:333–334. Google Scholar


    U Tun Yin. 1967. Wild animals of Burma. Rangoon Gazette, Limited, Rangoon, Burma.  Google Scholar


    A. C. V van Bemmel 1949. Revision of rusine deer in the Indo-Australian Archipelago. Treubia 20:191–262. Google Scholar


    A. C. V van Bemmel 1974. The concept of superspecies applied to Eurasiatic Cervidae. Zeitschrift für Säugetierkunde 38:295–302. Google Scholar


    S van Mourik and V Schurig 1985. Hybridization between sambar (Cervus (Rusa) unicolor) and rusa (Cervus (Rusa) timorensis) deer. Zoologischer Anzeiger 214:177–184. Google Scholar


    T. K Varma B. M Arora and H. C Malviya 1994. A note on Coenurus gaigeri (Hall, 1916) recovered from the thigh muscles of a sambhar (Cervus unicolor). Indian Veterinary Journal 71:618. Google Scholar


    K. S Varman and R Sukumar 1993. Ecology of sambar in Mudumalai Sanctuary, southern India. Pp. 273–284 in Deer in China: biology and management ( N Ohtaishiand H-L Sheng eds.). Elsevier Science Publishers, Amsterdam, The Netherlands.  Google Scholar


    T. T Veblen and G. H Stewart 1982. The effects of introduced mammals on New Zealand forests. Annals of the Association of American Geographers 72:372–397. Google Scholar


    V Veer B. D Parashar and S Prakash 2002. Tabanid and muscoid haematophagous flies, vectors of trypanosoiasis or surra disease in wild animals and livestock in Nandankanan Biological Park, Bhubaneswar. Current Science 82:500–503. Google Scholar


    S. W Wang P. D Curtis and J. P Lassoie 2006. Farmer perceptions of crop damage by wildlife in Jgme Singye Wangchuck National Park, Bhutan. Wildlife Society Bulletin 34:359–365. Google Scholar


    S. W Wang and D. W Macdonald 2009. Feeding habits and niche partitioning in a predator guild composed of tiger, leopards and dholes in a temperate ecosystem in central Bhutan. Journal of Zoology (London) 277:275–283. Google Scholar


    Z Wang and R Du 1982. Evolution of karyotype of the genus Cervus. Acta Genetica Sinica 9:24–31. Google Scholar


    R Ward 1896. Rowland Ward's records of big game. Rowland Ward and Company, Limited, London, United Kingdom.  Google Scholar


    S. D Webb 2000. Evolutionary history of New World Cervidae. Pp. 38–64 in Antelopes, deer, and relatives: fossil record, behavioral ecology, systematics, and conservation ( E. S Vrbaand G. B Schaller eds.). Yale University Press, New Haven, Connecticut.  Google Scholar


    R Weigl 2005. Longevity of mammals in captivity; from the living collections of the world. Kleine Senckenberg-Reihe 48:1–214. Google Scholar


    G. K Whitehead 1972. Deer of the world. Viking Press, New York.  Google Scholar


    G. K Whitehead 1993. The Whitehead encyclopedia of deer. Swan Hill Press, Shrewsbury, United Kingdom.  Google Scholar


    J Wu G Dong H Fong C Ge P Cai and F Zeng 2002. Evaluation of reproductive characteristics of farmed Formosan sika deer and Formosan sambar deer. Taiwan Veterinary Journal 28:204–210. Google Scholar


    L Xu 1983. Cervus unicolor hainana.. Pp. 395–398 in Birds and mammals of Hainan Dao ( L. H Xu Z. H Liuand S. M Yu eds.). Science Press, Beijing, China.  Google Scholar


    K Yamada J Elith M McCarthy and A Zerger 2003. Eliciting and integrating expert knowledge for wildlife habitat modelling. Ecological Modelling 165:251–264. Google Scholar


    W Yape Kii and G Mcl Dryden 2005. Effect of drinking saline water on food and water intake, food digestibility, and nitrogen and mineral balance of rusa deer (Cervus timorensis russa). Animal Science 81:99–105. Google Scholar


    O Zhdanski 1925. Fossile Hirsche Chinas. Palaeontologia Sinica 3:1–90. Google Scholar


    E. A. W von Zimmerman 1780. Geographische Geschichte des Menschen und der allgemein verbreiteten vierfüssigen Thiere: nebst einer heiher gehörigen Zoologischen Weltcharte Enthält ein vollständiges Verzeichniss aller bekannten Quadrupeden. Vol. II. Weygand, Leipzig, Germany.  Google Scholar


    S Zuckerman 1953. The breeding seasons of mammals in captivity. Proceedings of the Zoological Society of London 122:827–905. Google Scholar


    [1] Edited by Associate editors of this account were Jamie Harris and Pamela Owen. Synonymies were reviewed by Alfred L. Gardner and Colin P. Groves. Editor was Meredith J. Hamilton.

    American Society of Mammalogists
    David M Leslie "Rusa unicolor (Artiodactyla: Cervidae)," Mammalian Species 43(1), 1-30, (25 January 2011).
    Published: 25 January 2011
    exotic species
    Southeast Asia
    ungulate ecology
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