Equus asinus

Abstract

EquusLinnaeus, 1758:73. Type species [Equus] caballusLinnaeus, 1758:73 (domestic), by original designation. CaballusRafinesque, 1815:55. Substitute name for [Equus] caballusLinnaeus, 1758. AsinusGray, 1825:244. Type species Equus vulgaris (= [Equus] asinusLinnaeus, 1758:73) by subsequent designation. Not AsinusBrisson, 1762:70 which is not Linnaean. Not AsinusFrisch, 1775, which is not Linnaean. HippotigrisSmith, 1841:321. Type species [Equus] zebraLinnaeus, 1758:74, by original designation. ZebraAllen, 1909:163. Type species Equus burchelli grantide Winton, 1896:319, by original designation. In legend to figures 7, 8d, and 8e (but not figures 3–6 and 9–19, where Equus is employed). Evidently a lapsus. DolichohippusHeller, 1912:1. Type species Equus grevyiOustalet, 1882:12, by original designation and monotypy. MicrohippusReichenau, 1915:152. Type species Equus przewalskiiPolyakov, 1881:1, by monotypy. Not MicrohippusMatschie, 1922:68, which was used in a subgeneric sense. Not MicrohippusPocock, 1948:764, which is preoccupied. QuaggaShortridge, 1934:397. Type species [Equus] quaggaGmelin, 1788:213, by monotypy. OnagerQuinn, 1957:10. Type species [Equus] onagerBoddaert, 1784:160, by original designation. Not OnagerBrisson, 1762: 72, which is not Linnaean. Not OnagerZimmerman, 1780: 80, which was used in a subgeneric sense. HemippusDietrich, 1959:14. Type species Equus hemippusI. Geoffroy Saint-Hilaire, 1855a:1214, 1217, by monotypy. HemionusDietrich, 1959:14. Type species Equus hemionusPallas, 1775:394, by original designation. Not HemionusCuvier, 1821:555, a nomen nudum (Trumler 1961). Not HemionusStehlin and Graziosi, 1935, which was used as a subgenus. HydruntinusRadulesco and Samson, 1962:174. Type species Equus hydruntinus Regalia, 1904. Mesolithic to Neolithic subfossil. Order Perissodactyla, suborder Hippomorpha, family Equidae, subfamily Equinae. In general, names used as subgenera or based on fossil specimens are not included in the synonymy above. Equus has 1 recently extinct (E. quagga) and 7 extant species (Bennet 1980; Groves 1986; Grubb 1993; Schlawe 1986). The following key to the species is modified from Ansell (1971) and Corbet (1978) separates E. kiang from E. heminous (Bennett 1980; Corbet 1978; Groves and Mazak 1967). Chestnuts (epidermal callosities) on inner surfaces of fore- and hind legs; whole of tail long haired and growing from base; mane long and coarse, but in domestic horses, long and flowing -------------------------------------------------------------- E. caballus Chestnuts confined to forelegs; tail tufted, long hair confined to distal half ----------------------------------------------------------------------------- 2 Body striping limited to a dorsal line, shoulder stripe, or leg stripes; muzzle pale ---------------------------------------------------------------- 3 Body either wholly striped or with striping at least on head and neck; muzzle dark ----------------------------------------------------------- 4 Ears very long, >200 mm in adults ---------------------------- E. asinus Ears shorter, usually <150 mm in adults -------------------------------- 5 Size up to ca. 150 cm at withers; head and ears large in proportion to body; body markings with numerous vertical stripes behind shoulders, ending on haunches in a triangle pattern ---------------------------------------------------------------- E. grevyi Smaller size; pattern of striping not as above -------------------------- 6 Dorsal pelage dark reddish brown in summer, dark brown in winter; white ventral pelage with a prominent dorsal extension behind shoulder ----------------------------------------- E. kiang Body striping limited to transverse markings on legs, little or no dorsal line --------------------------------------------------- E. hemionus Stripes on croup forming gridiron pattern; dewlap present on throat ------------------------------------------------------------------------ E. zebra Stripes on croup not forming gridiron pattern; no dewlap -------------------------------------------------- 7 Striping whitish on dark ground, usually restricted to head, neck, and foreparts ------------------------------------------- E. quagga Striping dark on a light ground, over whole of body, usually at least partly on legs ------------------------------------------- E. burchellii Equus asinusLinnaeus, 1758:73. Type locality “in oreinte,” restricted by Linnaeus (1746:12) to “Magnatum;” Magnes or Magnesia, eastern Thessaly (Lewis and Short 1907), now in western Turkey, near Manias (Bridgewater and Kurtz 1963), ca. 38.4°N, 27.3°E. Linnaeus based his name on the domesticated ass of Europe, and did not designate a holotype. Asinus vulgarisGray, 1825:244–245. Type locality “Inhabits Tartary in the wild state,” Tatar, or Turkish region of eastern Europe and western Asia. Asinus domesticusSmith, 1845:314. Type locality “Syria.” Asinus ferusI. Geoffroy Saint-Hilaire, 1855b:1221. Type locality “d'Abyssinie.” Asinus africanusFitzinger, 1858:667. Type locality “Nubia,” vide Allen 1939:563; a nomen nudum (Groves 1986). E[quus]. taeniopusHeuglin, 1861a:15. Type locality “Die Wildesel heissen auf Arabisch: Hamár el Wadi. Mass.: Atgibaraha. Dan.: Debu-Kollo. Som.: Damer-debadeh;” considered “indeterminable” by Harper (1940:201). Type locality “near Berbera, Somalia,” vide Groves (1986). Type may be based on hybrids of domestic and wild asses (Groves 1986; Ziccardi 1970). Considered a nomen dubium by Groves (1986). Asinus taeniopus: Heuglin, 1861b:1 (description of figure 1). Name combination. Asinus africanusHeuglin and Fitzinger, 1867:588. Type locality “im südlichen Nubien und im gazen nordöstlichen Sennaar, swischen Sauakin und der Provinz Berber kis an den Nil,” restricted to “Ain Saba (etwa 16,0° N/38,0° O [= W]), Tigre, Eritrea (Schlawe 1980:104). Equus asinus somaliensisNoack, 1884a:101. Type locality “Somalikuste,” restricted to “Berbera (etwa 10,0° N/45,0° O [= W]),” northwest Somalia (Schlawe 1980:106). Asinus taeniopus somaliensis: Noack, 1884b:374. Name combination. Equus asinus atlanticusThomas, 1884:45. Type locality “l'oued Sequen,” Algeria, Pleistocene; according to Boule (1900), not an ass, but a zebra, E. burchelli mauritanicus. Equus asinus somalicusSclater, 1885:540, 542. Type locality “Somalica.” Equus nubianus somalicus: Peel, 1900:300. Name combination. Type locality “Guban, the great maritime plain between the [Red] sea and the Gulis range of mountains … between Hargaisa and Berbera,” Somaliland. E[quus]. (A[sinus].) a[sinus]. taeniopus: de Beaux, 1928:1. Name combination. Equus asinus atlanticusWerth, 1930:349, figure 3. Type locality “Enfouss (Annex d'Aflou, El Richa) in Algerien,” ca. 300 km SSW Algiers; 34.08°N, 2.03°E. Based on fresco and rock art; renaming of atlanticusThomas, 1884, thus a homonym and unavailable. Asinus asinus dianaeDollman, 1935:132. Type locality “Hafta Wadi 17° 43′ N., 37° 36′ E., Red Sea Province, south of Tokar, near Eritrean boundary,” Anglo-Egyptian Sudan. Equus asinus africanus: Allen, 1939:563. Name combination. E[quus]. a[sinus]. aethiopicusDenman, 1957:116–117. Type locality “in the Danakil country,” Ethiopia; ca. 13°N, 41°W; a nomen nudum. Asinus africanus arabicusTrumler, 1961:116. Type locality “Inneren Arabiens,” Saudi Arabia; a nomen nudum. Equus asinus hippagrusSchomber, 1963:121. Type locality “Sahara;” perhaps a homonym of Equus hippagrus Smith, 1836, used for a feral horse (Groves 1986). Equus asinus africanus saharienisDupuy, 1966:44. Type locality “Sahara Central … sud du massif du Hoggar;” unavailable because epithet is a quadrinomial. Equus asinus palestinaeDucos, 1968:28, 30. Type locality “Tel-Gat,” Jatt. Israel, 32.24°N, 35.02°W (The Times 1967). According to Ducos (1986) a nomen dubium. Equus melkeinsisBagtache et al., 1984:610. Type locality “Allobroges, Algerie,” Late Pleistocene. Equus africanus mureybetiDucos, 1986:241. Type locality “Muraibit … in Syria on the right bank of the Euphrates north of Meskene,” = Mesken; 36.02°N, 38.04°E (The Times 1967); renaming of palestinaeDucos, 1968. Context as above. Four subspecies of E. asinus are recognized. E. a. africanus may have been recently exterminated; wild populations of E. a. asinus disappeared within historic time, and E. a. melkiensis disappeared in early Holocene. E. a. asinusLinnaeus, 1758:73, see above (domesticus C. H. Smith, mureybeiti Ducos, palestinae Ducos, and vulgaris Gray are synonyms). E. a. africanus (Heuglin and Fitzinger, 1867:588), see above (dianae Dollman is a synonym). E. a. melkeinsisBagtache et al., 1984:610, see above. E. a. somaliensisNoack, 1884a:101, see above (nubianus Peel and somalicus Sclater are synonyms). Externally, E. hemionus (together with kiang and onager) and E. asinus (subgenus Asinus; Fig. 1) can be distinguished from E. caballus by possession of chestnuts only on forelimbs, rather than on both fore- and hind limbs (Groves and Mazak 1967). In addition, hairs forming mane of E. asinus and E. hemionus are shorter and softer than in E. caballus; tail of E. caballus is stiffer, with longer hairs beginning at base, rather than terminally tufted tail as in E. asinus or E. hemionus. E. asinus and E. hemionus lack striping found on E. burchelli, E. grevyi, E. quagga, and E. zebra. E. kiang has a wedge of white behind shoulders that reaches toward rump, and is covered in winter with longer hairs than other Equus species (Groves 1974). E. asinus can be distinguished from E. hemionus by presence of transverse leg stripes, which are much less common in E. hemionus (Groves and Mazak 1967). Adult male Equus asinus, Ajo, Arizona. Used with permission of the photographer J. R. Morgant Skulls of E. asinus (Fig. 2) and E. hemionus are similar to each other, but contrast with those of 6 other species in the genus. External auditory meatus (meatus acousticus) of E. asinus projects beyond the squama temporalis, and is therefore easily visible from above, whereas in E. caballus and zebras, it is little, if at all visible, being much shorter. Mastoid bone is visible from above in E. caballus and zebras but not in asses (Bennett 1980). Other distinguishing cranial features of more primitive Asinus are a narrower and squarer lambdoidal crest than in caballines; a greater basicranial–basifacial flexion in Asinus than in caballines and zebras; an occipital plane less vertical than in caballus so that a horse skull can be balanced on its occiput, whereas Asinus skulls are unstable; and when skulls of Asinus are viewed from rear, frontal bones cannot be seen, because facial region declines steeply, and top of cranium obscures them (Bennett 1980). In caballines and zebras, frontals are visible beyond line of juncture with line formed by connecting the postorbital bars. In addition, skull of E. caballus is elongated with a rounded occiput, a long molar row, and a broad incisor region. E. kiang, E. grevyi, E. burchelli, and E. quagga have a high, convex and the of the frontal bones to (Groves and Mazak 1967). postorbital of skull in E. asinus is elongated to E. hemionus (Groves and Mazak 1967). and of cranium and of of an male Equus asinus of skull is mm (1986) and of of and of bones for of and Equus and 1986). E. asinus can be distinguished from E. hemionus by long, of the no can be used to designate between the 2 species because subspecies of are 3 species kiang, and in which of The of E. hemionus of the is of the The of E. asinus only of the of the the is of the (Groves 1974). of the and are more in E. hemionus than in E. asinus, and on of E. asinus (Groves is usually in caballines and of the between and of the is and either in horses, in and in (Groves and Mazak 1967); or in zebras and in horses, and in from in the to in the (Groves and of E. asinus at shoulder is cm Ears are are and with no in between and (Groves 1974). hairs are restricted to distal of tail or is dorsal reaches from to tail and is it white either a the shoulder or dark transverse on legs and a from in less to no at all in wild asses have 1 or 2 shoulder stripes and transverse on legs (Groves 1986). wild asses by a shoulder stripe, and legs, with white on or of in all is a each of and inner and much of of Equus asinus is the species in the genus. and body and other of are (Groves and E. a. africanus cm at the whereas E. a. cm at subspecies in of wild ass is more whereas wild ass is more in but in Eritrea ass a. 1986). The of E. asinus from of the to the in eastern to in Somalia and to (Fig. 1986). historic E. asinus has a range the of the it was from in the et and of wild in of the E. asinus in and of the (Groves and in 1986). may in and and in the in but not in or (Groves 1986; 1986). range of Equus E. a. E. a. E. a. E. a. The wild ass in an and In E. asinus the western of the Red Sea in the of at Groves 1974). of E. a. africanus in and E. a. in the Danakil and the region of of the only E. asinus in the wild was E. a. in populations of E. asinus in and are in the western and western E. a. asinus in 1986). of in are from of to up to the present and from which E. asinus is are from in the early (E. and the and Red Sea by the of the in the may have included of and asses (Bennett or and asses may be in or Pleistocene. asses in of and E. asinus Neolithic E. may have been in to and by Equus asinus has vertical that are for (Groves 1974). has chestnuts on the to be of the and (Bennett The is of or to white in the but being more in and more in winter (Groves is are but with from than to than are used to and or the whole on the to a that early in (Groves 1974). Size and of are greater than that of and other early of E. is an to the that is found in the and asses and have molar than caballines (Bennett 1980; and are pattern on is for that and have the The in greater of which is when on 1980). Equus asinus in with et E. asinus to and by et have a for than other in of and to when in 2 feral E. asinus at on a was 2 or ca. et at was 2 or 5 and did not The of was of to whereas and The and of male domestic is by may to and is in the of the to and 5 considered to be of E. asinus africanus on was which was of that when the et in to when was half the when a E. asinus a body when was not of and in E. asinus and on its for E. asinus is to its but only if is et between and of 7 E. asinus was with a range of and when the wild asses to of between and to The between and body between and was of to not was whereas at an of was was ca. in when E. asinus populations in the of have no and in have a in and summer, but are and at all of the The was 1974). in and In a western of the of the no et of E. asinus is are at of but not are et and The of is 4 but up to and all are by 2 of et In a of feral wild asses at was not The of in a from for feral wild asses in western et an of for wild asses from 3 in eastern et to for wild asses in and and to 1 or 2 et 1974). on at 5 of but not 1974). its E. asinus with which are used as (Groves 1986). in the E. asinus can range km from The present range of E. asinus of Somalia and In the feral E. asinus has as the of the and and western asses an with the (E. in the and (Groves 1986; are by the on ground, and the on and (Groves 1986). Equus asinus or on species the of a in the of and and The of and is for E. asinus and are feral E. asinus have on The near but a to in has in with E. asinus and In E. asinus in of and with and E. asinus in populations with a than in populations and In the western and in wild asses not have and and from body has more in populations than in populations of E. asinus of large and In of the was in of in of and in of usually of a male with a or are all male or all adults have 1 or more and up to large of of and the and of with the to but up in the of feral E. asinus in the feral and wild populations has domesticated but the on wild populations is (Groves 1986). populations are and can in from can long to Groves 1974). male E. asinus are and for The is by the presence and of the at the in a with ears The male in an male to within of his to the and the following for up to 1 but from have been et are large with large are not at the In where the is and are may a male to his of in large et for to to 1974). may the the when and the because of a of and to and of in in and and are over of both are over is in large can the for a when being from the to if the male of the was but when the male to the et In only feral and can 1974). less to be and more at other from the than up to 2 et in a with hind legs and tail at a 1974). The male from behind and the to et The at the male in a and from in a 1974). The male the for or the the and a in a and is the male and the a from The male at the and on both the to its or from the male 1974). are by of the the and at the other between the or or with the have been in the and the with at least at The of the within 1 of the not other and by and the a the of its within 1 of its 1 to 2 of the of its within 1 of its of its is from 1 the of its within of its 1974). Equus asinus that from et The common of are a or and an the head is at a with the ears from 1 to and are in large E. asinus by for ca. to a for up to its ears and by a or it its head to the are with from to E. asinus is and the and of with the and The on of with a at of was and and was 1974). Equus asinus and The is the and is used when in and by from and are are used to to other The is an can be frontal and in of toward a the ears a and a usually by a to an in the toward the a a with 1 and with 2 the of or or of the 1974). of in which the head is with the muzzle toward the the is and the and are and are a 1974). is by and 1974). of in the and and are used in in the of or other them 1974). with other and with are Equus asinus is the of the domestic or feral domesticated feral that have been with wild E. asinus The domestic male E. asinus is with the E. caballus to the usually The large head and long ears of a E. asinus, whereas its body is that of E. The E. asinus is less with the male E. caballus to the usually The head and ears of the E. whereas its body E. asinus Equus asinus has = of E. asinus of and of et range of in of and and is a is an of 4 are in the that as in which the of all the and the of The is The at the but a prominent is present in the of each long and de The of the is for a of E. asinus from the line of and de of of E. asinus with those of E. caballus a and are in the to 2 in the E. asinus to a in the region is Four 2 and 1 are to E. asinus The species can be 3 based on and has a of than the 1 is to the and is above et that E. asinus has only 1 with in contrast to in the horse and are wild and zebras are more to each other than are to 1974). 6 based on the 3 zebra species grevyi, and usually a whereas asinus and hemionus and caballus E. a from the other 2 and 1986). The for the of and has E. a. as an species and may be the species on E. asinus was as of by the on in of and as an The of E. a. is in the of the The is and it is a between the and The Eritrean and the to populations of E. is on the which the wild ass by them with no and by at which the wild ass not Based on of the wild ass in either the de or based on in or may have in (Groves et but was based on the that the and wild asses be as on (Groves 1986). that of domestic asses of of the which as a in are in the early and wild from early domestic asses is the of wild ass is to be from Syria to ca. whereas the domestic ass in ca. and in de or in and by ca. 1986). is by ca. than the of the horse and and with horse to the north and may have ass The that the of wild asses in or the and domestic ass and north and Europe can only be to and the Sea to and it that the ass was Europe from by the the wild was The of the ass with other may be as as its The to be to the domestic was the the being ca. in 1986). the in the domestic ass was the common but wild for and is for 1986). when domestic from the domestic asses with them to the in is now and by the of the to be used in being by and the to be in are to ca. 1986). populations of feral E. asinus in the western and and The of asses the by was that of de in may have and other that by to in (Bennett little more than a in to the at (Bennett and a in present near El et to feral populations the of the in the no large feral populations because to be to and In the feral populations of E. asinus have and with and for and and and and and The and of the and of wild and on The is from and has feral E. asinus in have been whereas have been and up for 1980). similar has in E. asinus was to that for and the formed large feral E. asinus and of feral E. asinus from and from the E. asinus has a in and is to et The name of the wild ass is from 2 Equus and asinus ass, which is from an from are to on and the skull The of the