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Annales Zoologici Fennici 35: 149-162

Annales Zoologici Fennici 35: 149-162

Ann. Zool. Fennici 35: 149–162 ISSN 0003-455X Helsinki 11 December 1998 © Finnish Zoological and Botanical Publishing Board 1998

Ammotragus lervia: a review on systematics, biology, ecology and distribution

Jorge Cassinello

Cassinello, J., Estación Experimental de Zonas Aridas, CSIC, C/General Segura 1, ESP-04001 Almería, ; and Departamento de Ecología Evolutiva, Museo Nacional de Ciencias Naturales, CSIC, C/José Gutiérrez Abascal 2, ESP-28006 Madrid, Spain 1) Received 28 April 1998, accepted 9 August 1998

A revision on the current knowledge of the Ammotragus is provided. There is only one , A. lervia, which is considered an ancestor of both and . Six sub- species originally distributed in the North of Africa, but also introduced elsewhere, have been described. Particularly the study of the introduced wild ranging American populations, and recent research carried out on a captive population in Spain have expanded our knowl- edge on the species’ social behaviour, reproduction, female fitness components, behav- ioural ecology, feeding habits and ecology. Native and introduced populations of arruis are facing different problems; the former ones are generally threatened by human pression, and the latter ones pose a serious risk to native and plants.

1. Short foreword , or on American wild populations (e.g., Haas 1959, Simpson 1980, my own work). Herewith a Ammotragus lervia is an African retain- review on the literature available and suggestion ing some primitive and unique characteristics for future research are provided. which makes it particularly interesting for re- Common names for the species are: aoudad, search. It is also considered as a vulnerable spe- audad, udad, uaddan, ouaddan, aroui, arui, arrui, cies by IUCN (1996). The species was poorly Barbary , Barbarian, africain, mou- known until its introduction to the flon de Barbaris, muflón, muflón del Atlas, mouflon of America in the late 1930s. This gave a begin- à manchettes, ruffed mouflon, Saharan mouflon, ning to ecological and management studies (e.g., Mähnenspringer, Mähnenschaf, muflone berbere, Ogren 1965, Simpson 1980). Paradoxically, apart Tassilin, Fechstal, Naded, Naddan, kebach, Kabsh from few works mainly dealing with its distribu- Mai, Al Nakar, larrouy and bearded , among tion, research on this species has rarely been car- others (see e.g., Ogren 1965, Gray & Simpson ried out in its natural, African, environment (e.g., 1980a, Shackleton 1997). Depending on the region, Le Houérou 1992, Loggers et al. 1992, but see people and culture, we find a great diversity of com- Clark 1964). Therefore, most of the knowledge mon names for Ammotragus, some of them mak- we have about Ammotragus comes out from re- ing an evident reference to its relationship with the search on populations in European and American genus Ovis (see below). Barbary sheep is the Eng-

1) Address for correspondence 150 Cassinello • ANN. ZOOL. FENNICI Vol. 35 lish common denomination of the species; but, in 3. Phylogeny: comparative morphology order to prevent any taxonomic misinterpretation, and biochemistry I have preferred to use the Arabic name “arrui” in this review (see Valdez & Bunch 1980, and be- Ammotragus still represents a challenge for evo- low). lutionists, since due to its particular morphology and behaviour, its phylogeny is a complicated and controversial subject. 2. Both (genus Capra) and sheep (genus Ovis) share a series of characteristics with arruis The genus Ammotragus Blyth, 1840, with only (see Table 1). Some authors even suggest that the one species, A. lervia Pallas, 1777, has a com- latter should be included in one of these genera monly accepted taxonomic status: (e.g., Corbet 1978, see below). ‘Ammotragus’ Class: Mammalia means ‘sand ’, probably in reference to its color, Order: Artiodactyla which is pale, tawny brown or rufous, grading to a Suborder: Ruminantia whitish underside with dark brown areas around Infraorder: Pecora the head and forequarters (Valdez & Bunch 1980); Family: variability in color tones, however, is notable among Subfamily: Tribe: Caprini the . The general aspect resembles a ro- bust goat, with a relatively long head, short and However, the denomination of both genus and spe- stocky legs, and a long, naked tail underneath. As a cies has changed several times until reaching an unique feature, a mane extends from under the accord (Gray & Simpson 1980a, Gray 1985): throat down the front of the neck to the brisket, Genus: Ammotragus Blyth, 1840 bifurcating then and continuing down the forelegs Aegoceros Heughlin, 1861 in mature , where it is named chaps. Arruis Species: Antilope lervia Pallas, 1777 do not have a goatee, and like some sheep species Ovis Afzelius, 1815 Ovis ornata I. Geoffroy Saint-Hilaire, 1827 (Geist 1971a, Delibes 1986), they have a haired Ammotragus lervia Thomas, 1902 chin and a short erect fringe on the back. True horns are elliptical and similar to those of mouflon (Ovis Moreover, other taxonomic designations, which musimon). Unlike sheep, arruis lack preorbital, appeared between 1840 and 1902, may have con- interdigital and flank , but they have sub- tributed to creating a certain confusion in the tax- caudal glands. Their behaviour resembles that of onomy of the species (Gray & Simpson 1980a): sheep (Geist 1971a), although an ancestral pattern Ovis (Ammotragus) tragelaphus Blyth, 1840. does predominate (Katz 1949, Haas 1959). There- Ammotragus tragelaphus Gray, 1850. fore, morphologically arruis show intermediate Musimon tragelaphus Gervais, 1855. characteristics of sheep and goats (Geist 1971a, Aries tragelaphus Lataste, 1887. Ovis (Ammotragus) lervia Lydekker, 1898. Schaffer & Reed 1972). Ovis lervia Anderson and de Winton, 1902. The chromosome number of the arrui (2n = 58) is identical to that of Ovis vignei (Nadler et al. Finally, six subspecies have been described, mainly 1974, Bunch et al. 1977). Besides, the seroprotein according to their distribution (see the section about (Schmitt 1963) and immunoglobulin (Curtain & the distribution of the species below): Fudenberg 1973) analyses show a close relation- Ammotragus lervia lervia Pallas, 1777. ship to Ovis. However, the sequence of amino- Ammotragus lervia ornata I. Geoffroy Saint-Hilaire, 1827. acids of several hemoglobin chains (Manwell & Ammotragus lervia sahariensis Rothschild, 1913. Ammotragus lervia blainei Rothschild, 1913. Baker 1975) shows a closer relation to Capra hir- Ammotragus lervia angusi Rothschild, 1921. cus, and some unique characteristics. On the other Ammotragus lervia fassini Lepri, 1930. hand, immuno-diffusion studies by Hight and Nad- ANN. ZOOL. FENNICI Vol. 35 • Ammotragus lervia: a review 151 ler (1976) paradoxically establish a closer rela- and geographic distribution of the species, the cline tionship between Ovis and Capra than between is named argali or ammon cline in reference to the any of them and Ammotragus. Altai or Siberian argali, Ovis ammon ammon, which Some authors (Ansell 1971, Corbet 1978) place is placed in one end of the cline, while the arrui is the arrui in the genus Capra, due to morphological in the other end. To determine in which direction similarities and because it may interbreed with goats sheep evolved, Geist (1971a) firstly compared all and produce live hybrids (Petzsch 1957 in Geist the species to ancestral forms to establish which is 1971a). But Geist (1971a) discounted the ability of the evolutionary older form, and secondly he re- Capra to hybridize with Ammotragus as an indica- lated the cline to the history of Pleistocene glaci- tor of a closer relationship, and hypothesized that ations. reproductive barriers between Ovis and Ammo- Paleontologists suggest that the caprids evolved tragus arose due to their sympatric distribution in from the Tribe Rupicaprinae (Thenius & Hofer during the Pleistocene, whereas arruis 1960), which has four or six genera, according to and goats were not sympatric. various authors (Schaller 1977, Eisenberg 1981, Geist (1971a, 1985) hypothesized that the arrui Lovari & Perco 1982, MacDonald 1985). Two char- is ancestral to the Paleartic sheep lineage. He re- acteristic genera, the North American mountain lated all the Eurasian mountain sheep by means of goat, Oreamnos; and the , Capricornis; both a cline based on the morphological characteristics share a great number of characteristics with the most

Table 1. Anatomical, chromosomal and blood protein differences between sheep, arruis and goats (from Valdez & Bunch 1980). ————————————————————————————————————————————————— Phenotypic character Ovis Ammotragus Capra ————————————————————————————————————————————————— Preorbital Present Absent Absent Foot (interdigital) glands Present Absent Absent Inguinal glands Present Absent Absent Subcaudal glands Absent Present Present Chin beard Absent Absent Present in males; Horns In males usually in Supracervical usually scimitar-like spiral but sometimes or twisted and pointing bent backwards over the up or bent backwards neck (supracervical) over the neck, with the tips pointing inwards and up Tail length Short (≤ 6.0 inches) Long (≥ 6.0 inches) Long (≥ 6.0 inches) No. chromosomes (2n) 52 to 58 58 60 Transferrins (Tf) I, A, G, B, C, D, Identical to D of A and B differ in M, E and F sheep mobility from A and B of sheep Hemoglobins (Hb) Normal adult Amino acid Normal adult hemoglobin A, B and sequencing has hemoglobin A, B, D and D; with the exception indicated several E; amino acid sequence of mouflon all wild differences in the of either α or β chains sheep have Hb B Hb B-chains from differ from sheep that of sheep and goats ————————————————————————————————————————————————— 152 Cassinello • ANN. ZOOL. FENNICI Vol. 35 primitive caprid, the thar, Hemitragus, which, on cies of Ovis, Lydekker (1912) commented that the other hand, clearly resembles Ammotragus. these ‘sheep were akin to the modern arrui of Therefore, this author concludes that the latter must North Africa’. Osborn (1910) mentioned a ‘wild be the most primitive species in the evolutionary sheep (Ovis paleotragus) very similar to the ex- argali cline (Geist 1971a). As for the second issue, isting Barbary sheep’ when writing of the Pleisto- according to the so-called ‘Theory of Dispersion’ cene in . On the other hand, fossil (Geist 1971b, 1985), Ammotragus must also be the remains of Ammotragus have been found in , beginning of the cline. Firstly, it can be seen that in a number of deposits which ranged in antiquity the argali cline goes from North Africa, across Iran from about 85 000 to 2 000 years B.P. (McBurney into Turkestan, then into the Pamir Mountains, and 1967, Cremaschi & Di Lernia 1996), as well as in ends up in the Altai Mountains. This cline prob- Quaternary deposits in Cyrenaican Libya (Bate ably could not have existed during Pleistocene gla- 1955). Vaufrey (1955) noted the species among ciations, for all the mountains inhabited by the ar- the Pleistocene fauna in the Maghreb, and Aram- galis today, were then probably under ice (Frenzel bourg et al. (1934) recorded it at Beni Segoual, in 1968). This implies that the sheep found today in . central Asia are of rather recent evolutionary ori- This species was intensively exploited by Epi- gin, which surged from the unglaciated terrain into palaeolithic hunter-gatherers at the beginning of the mountain ranges after glacial withdrawal (Geist Holocene (Di Lernia & Cremaschi 1996), and a 1971a). After all this reasoning, Geist (1971a) con- potential or incipient domestication might have oc- cludes that Ammotragus should be placed in the curred at that time (Saxon 1976, Smith 1992). Cur- origin of the argali cline, and discusses carefully rent surveys and excavations in the Libyan Sahara all the morphological changes that have gradually are providing new evidence about this issue (Cre- taken place from arruis to . maschi & Di Lernia 1996, Di Lernia 1998). Ammotragus could also be an ancestor form of the genus Capra, at least of the round-horned goats (Geist 1971a). Disregarding its beard and mane, it is 4.2. Anatomy and morphology very similar to the young tur male (Capra cylindri- cornis) or the blue sheep (). Ammotragus Bourdelle (1924) carried out a detailed study on combines a number of goatlike characteristics and the anatomical and osteological characteristics of can be linked with intermediate forms not only to Ammotragus. The cranial morphology was ex- the argalis but also to the (Capra ibex plained by Schaffer and Reed (1972), the verte- sibirica). bral formulae established by Lydekker (1913), and There is also a recent study on the phylogenetic a first dental description made by Bourdelle relationships of nine genera of the Caprinae (Ca- (1924). Body measures were taken from individu- pra, Ammotragus, Hemitragus, Pseudois, Ovis, als in New (McClellan 1955, Ogren 1965), , Oreamnos, Nemorhaedus and Capri- Africa (Clark 1964), and Spain (Cassinello 1997a). cornis) based on the behaviours involved in court- The mean values obtained in a Spanish captive ship-mating and social status establishments (Kurt population show a strong sexual dimorphism in & Hartl 1994). The results obtained are in good sexually mature individuals: mean body weight, agreement with biochemical-genetic data avail- 82.07 ± 6.29 kg (males, N = 20) vs. 41.34 ± 1.92 able. kg (females, N = 42); mean body length, 146.76 ± 4.66 cm (males, N = 21) vs. 128.14 ± 1.82 cm (females, N = 44) (Cassinello 1997a). The study 4. Descriptive studies of length and shape, as well as the analysis of their rings and annuli have originated a series 4.1. Paleontology of works trying to find a relationship with either body weight (Gray & Simpson 1979), or with the In relation to fossil sheep remains recovered from age of the individual (Gray & Simpson 1985). Sex superficial deposits in at the end of XIX and age classes can be identified by face and horn century, and which were referred to as several spe- morphology (Gray & Simpson 1980b), although ANN. ZOOL. FENNICI Vol. 35 • Ammotragus lervia: a review 153 a new and more reliable identification key has re- success in captive Saharan arrui: longevity, fe- cently been published (Cassinello 1997b). Finally, cundity, offspring one-month survival rate and the a general description has already been provided age at first birth. Longevity accounts for 69.9% above, when comparing Ammotragus with other of the variance of reproductive success, fecun- caprids (see Section 3). dity for 54.2%, offspring one-month survival rate for 29.8%, and the age at first birth for 10.4% (see Brown 1988). A detailed study of these com- 4.3. Biochemistry and genetics ponents leads to the following conclusions: (a) longevity is higher in those individuals in better The hemoglobin polypeptide chains were studied physical condition, (b) fecundity is related to age in detail (see e.g., Huisman et al. 1958, Huisman and social rank, (c) heavier offspring at birth have 1974, Manwell & Baker 1975, 1977). Serum bio- a higher probability of surviving during their first chemical and hematological parameters (Schmitt month of life, and (d) the age at first birth is de- 1963, Tumbleson et al. 1970, Brady & Ullrey layed by high levels of population density, in- 1975), and immunoglobulins (Curtain & Fudenberg breeding coefficients, and birth weights. On the 1973) were also analysed. other hand, high-ranking females are character- The species’ karyotype was published by Heck ized by shorter inter-birth intervals and give birth et al. (1968), Schmitt and Ulbrich (1968), and to a higher proportion of twins. Nadler et al. (1974); Buckland and Evans (1978a, 1978b) analysed in detail its characteristics. Other studies in the field of genetics were carried out by 5. Factors affecting phenotypic traits Schreiber and Prosi (1988), and Schreiber and Ma- tern (1989). Following Cassinello (1997a), as adults, single- ton females become larger than females who had a littermate, also, when reaching sexual maturity, 4.4. Physiology females raised by older mothers are heavier. At birth, singletons are heavier than twins, but males Ogren (1965) studied the reproductive tracts and are only heavier than females when their mother sperm structure. Hormonal studies were carried holds a high social rank. Furthermore, high-rank- out by Hamon and Heap (1990). In addition, the ing mothers tend to produce heavier calves than effects caused by some drugs have been reported low ranking ones (see Cassinello & Gomendio by Nouvel et al. (1969), and Hampy (1978, in Gray 1996). High inbreeding coefficients yield lighter & Simpson 1980a). Robbins and Robbins (1979) calves. Finally, birth weights increase with ma- carried out a comparison of birth weight for vari- ternal age (Cassinello 1997a). ous ungulates species in relation to the maternal reproductive effort. 6. Behavioural ecology

4.5. Reproduction Pioneer studies on behavioural patterns in arrui, the so-called ethograms, were carried out by Katz Fourteen-months-old males and nine-months-old (1949), and Haas (1959). Succeedingly, a series females can be regarded as sexually mature (Cas- of works on the characteristic primitive agonistic sinello 1997a). Gestation and reproductive cycle behaviour of males, and on reproductive behav- was studied by several authors (e.g., Brown 1936, iour (arruis are polygynous and precocial mam- Zuckerman 1953, Ogren 1965, Cassinello & Ala- mals) have been published (e.g., Ogren 1965, dos 1996). The mean gestation period is 5.5 months Schaffer 1968, Hamdy & Schmidt 1972 in Gray and mating season peak occurs from September & Simpson 1980a, Schaffer & Reed 1972, Habibi to November, so that breeding season tends to be 1987); as well as others on more general aspects focused in spring. Cassinello and Alados (1996) of their behaviour (Solbert 1980, Gray & Simpson analysed four components of female reproductive 1982a). A more detailed study carried out by Cas- 154 Cassinello • ANN. ZOOL. FENNICI Vol. 35 sinello (1995) on a captive Saharan arrui popula- 7. Ecology tion in Spain shows that the species is character- ised by an absolute , which The habitat selection was analysed in Africa is also near-linear and fairly stable. Females may (Rothschild 1921 in Gray & Simpson 1980a), change their hierarchical position under some cir- America (Ogren 1965, Dickinson & Simpson cumstances: mating and parturition may cause a 1980, Johnston 1980), and Europe (R. C. Bigalke, rank increase, and offspring weaning may lead to unpubl.). Arruis tend to inhabit rocky and pre- a lower status. Thus, it seems plausible that fe- cipitous areas, from the sea level up to the extent male rank variation is related to proximal factors of snow-free areas at about 3 900 metres in their affecting social behaviour, because a female may African endemic range. Johnston (1980) estab- then challenge her hierarchical status, and acquire lished the following habitat preference according a higher social position. It is also postulated that to the season: woodlands during summer, grass- females may present a variable social rank, re- lands during autumn and winter, and protective lated to age, at the beginning of their reproduc- rocky slopes during spring, the main breeding sea- tive life, till they reach a fairly steady social sta- son (see above). Research on population para- tus, in accordance with their reproductive experi- metres were done in wild populations from ence and inherited family status. (Gray & Simpson 1983) and (R. C. According to a recent work (Cassinello 1996), Bigalke, unpubl.),the proportion of adults and ju- high ranking females allocate their resources pref- veniles being quite similar in all instances: ap- erably towards their sons, according to Trivers proximately 30% juveniles, 20% adult males and and Willard’s (1973) hypothesis. Cassinello and 50% adult females. Concerning group dynamics, Gomendio (1996) also showed that litter size and Gray and Simpson (1982b) verified that group sex ratio at birth are strongly influenced by the leadership is ascribed to females when adults of parity and maternal dominance rank at the time both sexes are present, while group composition of conception in arrui. When females give birth and size vary depending on the season. Six group for the first time they always produce a single off- types can be distinguished in the wild: solitary, spring, while multiparous females produce both nursery (females, calves and juveniles), mixed, singletons and twins. As maternal rank increases, all male, all female and all juveniles (Gray & females tend to produce the following sequence: Simpson 1982b). The knowledge of the feeding F-FF > M > MF > MM, which differs from Wil- habits of the species is circumscribed to wild liams’ (1979) prediction in that FF are produced American (e.g., Ogren 1965, Bird & Upham 1980, by females of lower rank that M. This may be so, Krysl et al. 1980, Simpson et al. 1980), and Span- because of the strong sexual dimorphism of the ish populations (Luengo & Piñero 1987, Piñero species in which the differential costs of sons and & Luengo 1992, R. C. Bigalke, unpubl.); but arruis daughters may be greater than in William’s model, are not selective at all and their diet may com- and in which dominant females have much to gain prise shrubs, succulent forbs, forbs, creepers, from producing exceptionally good males. Despite dwarf shrubs and grasses, depending on season lower levels of investment in females, single fe- availability. Concerning territories size and popu- males are more likely to survive than single males lation movements, Dickinson and Simpson and twins. (1980), and Simpson et al. (1978) provided some Behavioural conflict between mother and off- information, thus, the mean home range size ranges spring was investigated in the same captive popu- from 259 to 3 367 ha.; arruis’ dispersal is particu- lation of Saharan arruis by Cassinello (1997c). larly accentuated in summer. There are also stud- Mothers that conceive in the following mating sea- ies on potential interspecific competition son accelerate the weaning process of their cur- (Simpson et al. 1978) and parasites (e.g., Allen et rent offspring; furthermore, high-ranking females al. 1956, Allen 1960, Yeruham et al. 1996), al- wean their male calves earlier. Behavioural con- though arruis seem to be remarkably disease and flict occurs during the resume of mother’s sexual parasite-free both in captivity and in the wild (e.g., activity; however, no conflict appears to happen Ogren 1965, J. Ortiz pers. comm., see a review in during weaning. Pence 1980). Finally, a novel study on potential ANN. ZOOL. FENNICI Vol. 35 • Ammotragus lervia: a review 155 energetic costs due to heat loss through the horn United States of America (Gray 1985); and, surface reached the conclusion that metabolic according to Traweek (in Jones & Jones 1992), costs of possessing large horns in cold climates only in Texas there are currently more than may impose constraints on morphology and sexual 5 000 individuals. This could also be the sub- selection (Picard et al. 1994). species introduced in the Sierra Espuña moun- tains of Murcia, in Spain (Gray 1985). — Ammotragus lervia ornata. This subspecies is 8. Distribution native to various desert areas in (see Gray 1985), but it may already be extinct (Hei- Apart from its native distribution in Africa, the nemann 1972, Amer 1997). It is named Egyp- information on the arrui distribution in non-na- tian arrui. tive lands, as a consequence of human introduc- — Ammotragus lervia sahariensis. It has a very tions, has been collected mainly from the avail- large geographic distribution that, according able bibliography and some few personal com- to Gray (1985), includes parts of southern Mo- munications; but I have not intended to make an rocco and the Western Sahara (see also Aula- exhaustive listing of its presence in zoological gar- gnier & Thévenot 1997), the Sahara of south- dens, and private preserves. Undoubtedly, ern Algeria (but see de Smet 1997b), southern and because of the high reproductive rate and , southeastern Libya (see also Shack- adaptability of the species (see e.g., Ogren 1965, leton & de Smet 1997), , (see also Simpson et al. 1978), it may be present in a great Lamarche 1997a), , (see also number of zoos and game preserves, particularly Lamarche 1997b), and the . in temperate zones (south of Europe and USA). It This is also the subspecies found in (Me- follows the distribution of wild-ranging popula- konlaou & Daboulaye 1997, but see Alados et tions as well as the occurrence in zoos mentioned al. 1988), today probably restricted to the sand- by the more relevant authors. stone massifs in Ennedi (Mekonlaou & Da- boulaye 1997). Named Saharan arrui, it has been introduced into the Estación Experimen- 8.1. Endemic populations: Africa tal de Zonas Aridas (EEZA) in Almería, Spain (Cano & Vericad 1983, Alados et al. 1988, The arrui is endemic to North Africa (see Brentjes Alados & Vericad 1993). 1980); and, in theory, it might be found in practi- — Ammotragus lervia blainei. It used to be rela- cally any rugged terrain or mountain chain through- tively widespread from western Sudan to the out the whole northern part of the continent (see Red Sea coast, but currently it might be re- Gray 1985, Le Houérou 1992, Loggers et al. 1992, stricted to the Red Sea hills of eastern Sudan Shackleton 1997). There are six subspecies (Allen (Nimir 1997). It could also be found in the Enne- 1939), whose distribution is as follows: di and Uweinat mountains in Chad (Alados et — Ammotragus lervia lervia. Named Atlas arrui, al. 1988); but according to Mekonlaou and Da- it can be found in the mountains of boulaye (1997), that is the subspecies A. l. saha- (Aulagnier & Thévenot 1997) and Tunisia (de riensis. In 1923, A. l. blainei was introduced Smet 1997a), the northern part of Algeria (de into the Sabaloka reserve on the Sixth Cataract Smet 1997b), and in the regions of Aïr and the of the (Gray 1985). It has also been re- Tibesti massif (see Aulagnier & Thévenot ported in Libya (see Shackleton & de Smet 1997). More than 30 years ago, it was ap- 1997). It is named Kordofan arrui. proaching extinction in Algeria and Tunisia — Ammotragus lervia angusi. Named Aïr arrui, (Schomber & Kock 1960). Presumably, this it inhabits the Aïr and Asben, in Niger (Magin is the subspecies that was imported to Euro- & Newby 1997), west to the Adrar des Iforhas pean zoological gardens at the end of last cen- of Algeria (but see de Smet 1997b), and east tury, and from there to American zoos about to Tibesti in Chad (Alados et al. 1988, but see 1900 (Ogren 1965). This subspecies also form Mekonlaou & Daboulaye 1997). the basis of free-raging populations in the — Ammotragus lervia fassini. It can be found in 156 Cassinello • ANN. ZOOL. FENNICI Vol. 35

Libya (Shackleton & de Smet 1997) and in tions was successful (see Gray 1985). However, the extreme southern part of Tunisia (Gray the species is common in German zoos. 1985, de Smet 1997a). There are no current population estimates. Likewise the Saharan arrui, this subspecies was introduced into the 8.2.1.2. Italy EEZA (Cano & Vericad 1983), but the whole population was moved into the Barcelona According to Zammarano (in Gray 1985), the spe- in the late 1980s. It has been denominated Lib- cies was introduced into some game preserves yan arrui. The color of its fur is more pale than during the first half of this century. I do not know that of the Saharan arrui and it is character- whether the introductions were successful. ized by a more tranquil and even-tempered mood (EEZA staff communication). 8.2.1.3. Spain Currently, two main issues urge to be revised: first a more accurate definition of the subspecies, Definitely, this is the European country where the as from the distribution areas it may be inferred arrui has been more successfully introduced, due in various possible hybridation zones (the regions part to the mild climate and the geographic pro- of Aïr and the Tibesti massif, with three subspe- file which, particularly in the south east of the cies supposedly inhabiting, A. l. lervia, A. l. angusi country, resembles the native African lands. The and A. l. sahariensis); also, it remains unclear current distribution of the Spanish arrui popula- whether the populations found in the Western Sa- tions is difficult to assess accurately, as there are hara and Mauritania correspond with A. l. saha- more and more private game preserves which are riensis, which type locality is sited in the Alge- introducing the species, since it is easy to adapt to rian Sahara (Gray 1985). A second issue would most of the environments and habitats, being prac- obviously be to carry out reliable surveys of the tically a generalist (see e.g., Ogren 1965, Simpson distribution of the species in Africa, although they et al. 1978, Barrett & Beasom 1980, Dickinson & may sadly be confronted with political and eco- Simpson 1980, Johnston 1980, Luengo & Piñero nomic interests. Small groups scattered on large 1987, R. C. Bigalke, unpubl.). However, those in- territories are the typical pattern of the distribu- troductions carried out by public institutions into tion of the species in the wild (Savino di Lernia, natural parks or reserves can be easily enumer- accounting anonymous Tuareg informants), so ated, such as Sierra Espuña in Murcia and the Par- that intense surveys would need to be done to test que Nacional de la Caldera de Taburiente, in La their presence. Palma, (see below). The arrui can be found in the following Spanish regions: 8.2. Introduced populations — Murcia: thirty-six arruis from the Frankfurt Zoo (Germany), and the Ain Sebad Zoologi- Unequivocally, arrui introductions in Europe and cal Park, in Casablanca (Morocco), were in- America have always been related to game inter- troduced in the Sierra Espuña Natural Park in ests. Captive populations in zoological gardens, 1970 (Ortuño & De la Peña 1979). The popu- on the other hand, might help to preserve the most lation expanded rapidly, reaching up to 2 000 threatened subspecies. individuals in 1991 which also inhabited the surrounding mountains, such as Sierra de las Cabras and Sierra del Burete (ARMAN-Mur- 8.2.1. Europe cia, unpubl.). Unfortunately, in 1992 a mange epidemic drastically disminished the arrui pop- 8.2.1.1. Germany ulation (S. Eguia & P. Jimenez, unpubl.), and the most optimistic estimations established that The arrui was introduced in two locations in the scarcely 200 individuals survived (R. Sánchez north, near Lopshorn in Lippe in 1883, and later pers. comm.). As a preventive measure, 63 ar- in the Teutoburger Wald; none of these introduc- ruis were kept in an enclosure, inside the Natu- ANN. ZOOL. FENNICI Vol. 35 • Ammotragus lervia: a review 157

ral Park, in 1995 (DGMN-Murcia, unpubl.). San Simeon was disbanded and consequently Since then, the population has recovered again, 85 arruis began a wild existence. This popula- and it is supposedly expanding once more and tion expanded to over 500 within 10 years (see reaching nearby mountains (M. A. Sánchez also Gray 1985). pers. comm.). The arruis of Sierra Espuña prob- — : according to Ogren (1965), in ably belong to the subspecies A. l. lervia (see 1950, 57 arruis from private ranchs were re- Gray 1985). leased at Old Mills Canyon and on a state game — Canary Islands: 16 individuals from Sierra Es- refuge near the Canadian River Gorge. Thus, puña were introduced close to the Caldera de the Canadian River arrui population stems Taburiente National Park in La Palma in June from these animals. Apart from this popula- 1972 (Luengo & Piñero 1987, Piñero & Luen- tion, the Picacho herd is also known to be the go 1992). They expanded successfully since source of other arrui populations in central and then on, inhabiting nowadays the most remote southern New Mexico. Finally, 21 individu- and mountainous regions. The population in- als were planted in Canyon Largo by the San creased without an effective management Juan County Wildlife Federation in 1956 (O- (anon. comm.). Gray (1985), erroneously, gren 1965). Furthermore, periodic escapes of quoted this introduction as a failed one. arruis from the McKnight game encloure have — : there are some populations of arrui resulted in a another wild breeding popula- in private game preserves in Cádiz (Gray tion in the Guadalupe Mountains National Park 1985), Málaga and (anon. comm.). I (Dickinson & Simpson 1980). As a whole, do not know the origin of these private intro- there may be about 5 000 individuals in the ductions and, in consequence, which subspe- state (B. Morrison, pers. comm. quoted by cies they are composed of. Libyan arruis can Gray 1985). be found at a zoological garden in Córdoba. — Texas: in December 1957, 31 arruis were re- Finally, there is a captive population of the leased in (DeArment 1971 subspecies A. l. sahariensis at the EEZA, in in Gray 1985). According to Gray (1985), Almería (e.g., Alados et al. 1988), originated another 13 individuals were liberated in Bris- from two individuals which were shipped from coe County in February 1958. This author also the western Sahara in 1975. comments the dispersal of this species through- — Castile-La Mancha: according to Gray (1985), out the state. The arrui population at Palo Duro there are some private herds of arruis in the seemed to be as high as about 2500 animals in provinces of Ciudad Real and Toledo. the late 1970s (Simpson et al. 1978). A series — Catalonia: at least there is a population of Liby- of releases and escapes from private enclo- an arruis at the Barcelona zoo (see above). sures have resulted in some small free-rang- ing populations of the species since 1960’s (Decker 1978, Simpson & Krysl 1981). Free- 8.2.2. America ranging herds now exist on caprock along much of the eastern edge of Llano Estacado, 8.2.2.1. United States of America in the rough country of Trans-Pecos, and on parts of Edwards Plateau (Jones & Jones According to Ogren (1965), the arrui apparently 1992). The total population in the wild exceeds has been present in the USA since about 1900; all 5 000 (Traweek 1985). known imports being from European zoos. And — Oklahoma: arrui sightings have been reported by 1965, nearly every sizeable zoo had acquired occasionally, probably as a result of dispersal the species. The species is present in six main re- from the populations already mentioned (Simp- gions: son & Krysl 1981, Gray 1985). — California: according to Barrett (1980), the — Colorado: sporadic reports of sightings (Ogren species, which is commonly known as aoudad 1965, Simpson & Krysl 1981, Gray 1985). in this state, has been a common in Gray (1985) doubted that any populations had zoos for decades. In 1953, the Hearst Zoo at become established. 158 Cassinello • ANN. ZOOL. FENNICI Vol. 35

8.2.2.2. Mexico any rugged terrain. They also evaluated that out of 49 desert bighorn food plants, 37 were recorded Rangel-Woodyard and Simpson (1980) document so far as eaten by arruis, and that if both species the release of arruis in three localities: in the Si- would happen to share their home ranges, in win- erra Morena Ranch, Sierra Pájaros Azules and in ter, when both are feeding predominantly on grasses, San Luis Potosí; these releases were not designed competition should be hard, to the detriment of to supplement the endemic fauna for sport-hunt- the desert bighorn (Simpson et al. 1978). On the ing purposes as it was in some areas of the USA, potential risk on native or endemic flora, arruis but they were made to fulfill the interests of indi- are generalists, and their wrong introduction in viduals in a private collection of exotics. On the valuable zones, such as the Caldera de Taburiente other hand, Gray (1985) was informed by the Uni- National Park, in La Palma island (Canary Islands; versidad Nacional Autónoma in Mexico on the see above), has shown the presence of endemic release of the species on Espíritu Santo Island, in plants in their diet (Luengo & Piñero 1987, Piñero the Sea of Cortez, but heavy hunting extirpated & Luengo 1992). An uncontrolled increase in arrui eventually not only this population but also the introduced populations has always been observed other three ones released in Mexico, so that cur- (e.g., Barrett & Beasom 1980, R. C. Bigalke, rently there seems to be no longer any free-rang- unpubl.), so that a proper monitoring should be ing population anywhere in the country. one of the priorities in any management project of the species. Concerning conservation prospects, apart from 9. Management and conservation pros- the management of already-introduced populations pects which are to be maintained, the re-introduction of the subspecies A. l. sahariensis in Western Africa African populations of Ammotragus are generally is desirable, as a viable population is kept in cap- threatened due to a combination of overgrazing tivity in Spain (e.g., Alados et al. 1988). Proper by domestic livestock, deforestation and habitat surveys urge to be tackled in arrui’s original sites destruction, and poaching (see Loggers et al. 1992, in Africa in order to know the actual distribution Shackleton 1997). The general conservation pro- and abundance of the species, which is mostly in- grammes of each country where the species oc- determinate (see Shackleton 1997), and to carry curs are summarized in Shackleton (1997). out pertinent management policies if necessary. But From a purely hunting perspective, the arrui is a series of logistical and even political problems of great interest (see e.g., Ogren 1965, Ortuño & make it particularly difficult to carry out proper De la Peña 1979, Christian 1980), but unfortu- protection actions in the native lands of Ammotra- nately the introductions carried out in the past did gus (see also Alados & Shackleton 1997). not take into account the potential costs for the environment that an opportunistic species like this Acknowledgements: The author wishes to thank Raul may cause. Careful management and control of Valdez for kindly giving permission to use his published material; Savino di Lernia for the information provided on introduced populations are needed in order to pre- his research in Libya; David M. Shackleton for kindly send- vent any potential risks on autochthonous species, ing me valuable information on the species current status in such as interspecific competition with other un- Africa; and Miguel Delibes and an anonymous referee for gulates (Simpson et al. 1978) or feeding on native revising an earlier version of the manuscript. The author is flora (Luengo & Piñero 1987, Piñero & Luengo currently supported by DGICYT (PB96-0880). 1992). There is no recorded instance of direct com- petition between arruis and other ungulate species; but, following Simpson et al.’s (1978) reasoning, References a real threat against the desert bighorn (Ovis cana- densis) might occur in those American sites where Alados, C. & Shackleton, D. M. 1997: Regional Summary. — In: Shackleton, D. M. (ed.), Wild sheep and goats both species show close home ranges. These au- and their relatives: status survey and conservation ac- thors point out the immense dispersal capacities tion plan for Caprinae: 47–48. IUCN, Gland, Switzer- of arruis and their ability to survive in virtually land. ANN. ZOOL. FENNICI Vol. 35 • Ammotragus lervia: a review 159

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