MAMMALIAN SPECIES wo. 555, pp. 1-8, 3 figs. Pseudalopex culpaeus. By Andrés J. Novato Published 24 October 1997 by the American Society of Mammalogists Pseudalopex Burmeister, 1856 Peeudalopex Burmeister, 1856:24. Type species Canis magellan- HS subpeiesfF eulpans by sabeuent Paeudelyeos Pilpp 1908:157, No species mentioned Cerdocyon Allen, 1905:154 (not Cerdocyon Hamilton Smith, 1899). Angusticeps Hilaheime, 1906-114. Type species Canis (Angusti- eps) reisii Hilheimer, 1906 (a subspecies of P2 culpacus) by subsequent designation Microeyon Trovessart, 1906-1186. Type species Speothos rivet ‘ours 06 a syony af Be. Feil by subeuet lesignation. Dusieyon Kraplievie, 193055 (pat, not of Hamilton Smith, 1839). ora, Family Cav CONTEXT AND CONTENT, Onder Ca nidae, Subfamily Canine. The genus Peeudalopex includes five or Six species (Berta, 1987; Tedford et a, 1995: Worencrat, 1998; Yahnke eta, 1996; Zunino et al, 1995) Peulpacuss P peruanus (extinct) P griseus: P.gymnocercus: P.fudvipe: P sechurae. Zi- tino etal (1995) studied cranial measirements and pelage che aciers of P gymnocereus and P. griseus and concluded that they fare conspecific. A key to living species of Pacudalopex (modifies from Cabrera, 1982; Clutton-Brock et al, 1976: Crespo and De Carlo, 1963; Eisenberg, in press; Ginshers and Macdonald, 1990; Osgood, 1843; Redford and Eisenberg, 1992 follows 1. Til pot bushy and <506 of length of head nd body unis formly dark and rufous pages size smal, eng af bed and oly 490-542 rom P.falpes Tail bushy. and >50% of length of head and body: size ‘small to medium, length of head and bedy 446-1010 am 2 2, Short, course fur, overall pale coloration, lithe or no rufous ‘coloring onthe body: size small, length of head and body {530-590 mmm: skull without an interpavietal crest P.sechurae Fur neiier short nor coarse; coloration rot pales mall to ‘medium size, Length of head and body 446-010; skull ‘wth oF without interperital crest. 3 3. Whitish eli: brownish patch of pelage on back of thighs: skull with interparietal crest: medium size, length of head ‘and body’ 585-1010 num ® culparus Black chin; black patch of pelage on back of thighs: skull with or without interparital eres: size smal, length of head and body 446-722 ram 4 44 Length of eranium and hind foot ca. 130 mi Deng of Thea and body 520-722 min; color of pelage relatively sniform P. gymnocercus ‘Length of cranium <135 mon and length of hind foot <130 ‘mm: length of head and body #46600 mun: feet tawny fd head rust colored Previseus Pseudalopex culpaeus (Molina, 1782) Culpeo Fox Canis eulpaeus Molina, 1782:298. Type local (1931) suggested that it was restricted to Chile, now called Chilean Region V. llénica Gray. 18378578. Type loality “Magellan's (Port Famine: Gray 1843261), Magallanes, Chile. Canis (Pseudaloper) Iycoides Philipp 18563582. Type loeality "ine sulis Tierra del Fuego." Magallanes, Chil. Canis amblyodon Prilipp, 1908:188, Type loc paraiso,” Chi “Chili” Cabrera ntiago Province, ty “provincia Vale Canis albigula Philippi, 1903:159. Type locality “provinelis cen tealibas.” Chile Canis {Cerdocyon) prichardi Teowessae, 1904234. New name for Canis montanus Prichard, 1902:260 (preaccupie type lo ality “South-easter Patagonia.” Canis Angusticeps) ress: Milzheimer, 1906-116. Type locality “Quit.” Pichincha, Eouador. Cabrera (1958) suggested Yo: ‘in Cotopasi, Pichincha Province, Ecuador. peothos rvett Trouessrt, 1906-1185. Type lo province de Pichincha (Equateuy, altitude de 2101 tinder. Pseudalopex eulpacus Thomas, 19144357. First use of current Pseudalopex culpacolus Thomas, 19144359. Type local Elena,” Soriano, Uruguay. Considered a composite ‘ulpaens: skull oP gymnocereus) by Langguth (2967) who ‘elected the skin as lectotype Poeudalopes inca Thomas, 1914361. Type locality “Sumbay, Are “quips, Peru. Alt. 4000 m." Considered a composite (skin of P.gymnocercus; skull of P.culpaeus) by Lengguth (1967) who selected the akll a Ietotype Preudalopes smithersi Toor, 1914573. Type CCrdaba, Argentina. Cabreea (1958) suggested Pampa de Achala (2200 m). Ganis fergineus Huber, 1925:9. Type locality “la Conillera (de los Andes) ente los eos Mendoza, Atul, Neuguén y Calan Cr Argentina CONTEXT AND CONTENT: Contest noted in gensrie ac count. Six subspecies are recognized as follows (Cabrera, 1981): Pe. andina Thomas, 19140:357. P eulpaeolus Thomas anil P. inca Thomas are synonyms. Type locality “Esperanza, near Mt Sajama, Province of Oruro (La Paz). Bolivia. Alt. 4000 “Alehipihi Ee P. 6, eulpacus (Molina, 1782). See above. C. amblyadon Philip, 6. albgwtaPpi and Cf Her a 2 2. Iycoes (Philipp, 1896), See abo. Pc. magellaniea (Gray, 1880). See ave. C. prichardi Troves- saris a synonym Fic. 1. Adult male and female Pseudalopex eulpaets from Monumento Natural Bosques Peticsdos, Santa Caur Province, Ar ‘gentna. Photograph by F. Colavino.Fic. 2. Dorsal, venta, and lateral view of cranium and let cra view of mandible of Pseudaloper culpaeus trom Catan Lil, Neuquén Province, Argentina (Female, Museo Argentino de Cien cias Naturales specimen 15056). Greatest length of eranium is 174 tam Drawn by G- Carin Preis (Hileheimer, 1906). See above. S.riveti Trowessart isa “inithersi Thomas, 19148, See above. Lanaguth (1969) lists this sbepecies asa synonym of Pc andina, DIAGNOSIS. Pseudalopex culparus is distinguished from con severe by its lange size (length of heed and body, 585-1010 eam, ‘6-722 mm, spectively), rarower rostrum (strum width 24% of palate length in colpos, 27-825 in ether species) eddih cloration ‘tthe rad, neck, ears ad lps and white to Iighttawny chin (Car Tera and Yepes, 1940; Cliton Brock etal 1976 Crespo and De Carl, 1963; Eisenberg in press Ongon, 1943; Rediord and Eisen berg, 1992). Furthermore, the tall of flies is not bushy and shorter relative to the length of heed and Body: plage eslr of P vmnocercus is more uniform; P griseus and Psechuvae lack the fnterparcialexest and in the later hee litle or wo rufous coloring ‘on the bods, andthe palatine bones extend backwards beyond the Prsteror edge of M2 (Chtton-Brock et sl, 1976) GENERAL CHARACTERS, Preudalopex eulpacus (Fig, 1) is a large for. Ite skull has long and narrow facal region, fat frontal hones, slightly developed interparietal eres and a con tinuous sagital crest (Fig. 2). The palatine bones do not extend hackwate heyond the posterior enge of M2. Canines and premolars Re MAMMALIAN SPECIES 538 ae simple and “Yorke” and the metaconid of mie higher than the level of the talonid (Cluton-Broek et al, 1976: Langeuth, 1960), P. culpacus has a reduced m2 metaconid (Herta, 1987), ‘The dorsal part ofthe head, neck, outer aspect of the eas and legs, and flank tegions of Pculparns are tawny or rufous. The back ard shoulders are gray with agouti (barded black and white) fuant hair, underparts are pale, and underiur is fayn. Black hairs predominate along the center ofthe back, forming distinctive dark Tine that varies in width among, subepecies, The dorsal sides of the feet are light tawny. P eudpaens au bushy tall that is >05 of the length of head! and bods is black tipped. and has a black spot on the upper side near the hase (Citton-Brock otal, 1976: Lang rath, 1969), ‘Peeudaloper culpaeus increases significantly in body size sith increasing laitade inthe southern portion af is range Fuentes {nd Jaksie, 197%; Jiménez et al, 1995}. In nonh-central Chile (Aue, 31°30) adult mean body mass (in), length of body (in ‘mm), and total length {in mm; n= 9) are 432, 628, and 999, respectively. The samme measurements in southern Chile (Tomes del Paine, 5109'S: n = 7) are 10.16, 789, and 1,152, respectively Mean body mess (in kx) at anether site in northern-central Chile (Fray Jorge, 30°38) in 0.52 (n= 5 Sabai et al, 1993), and to the south in Neuguen Province, Argentina (39°83'S) itis 8.98 (n = 22—Novaro, 1991) ‘Adult male culpeos weigh significantly more than females in Torres del Paine (mean of 11.65 kg, n = 4, compared to 7.82 kg. n= 3-—Johnson and Franklin, 198) and in Newguén (11.02 kg, n= 11, compared to 884 kg, n = T—Novaro, 1991). Mean extemal measurements (in mm: range and n in parentheses) for tale and female culpoos, respectively, from Neuquen Province, are {otal lengthy 1150 (@20-1:5205 1 = 150), 1102 (000-1,1240, n= 107} length of tal, 412 (00-510, n= 130), 395 (315-455: n 108); length of hind fot, 168 (140-181; n = 13}, 152 (180-100; nn = 106}; length of eur, 88 (00-95: mn — 148), B¢ (26-90; 104—Creepo and De Carlo, 1963). Mean cranial and toath mea- surements (in mm: m in parentheses) for males and females. re spectively, from Nouquén Province, ar: length of cranium, 171.0 {oe = 139), 168.0 (n= 105}: basal length, 156.6 (n = 128), 199.4 (ir = 102); palatal length, 85.0 fv = 129), BL2 (n= 102) 2g0- ‘matic wid 85.0 (n= 129), 81.9 (n = 100), inerorbital width, 9 (n= 126), 27.0 (n = 101}; mastoid width, 493 (n ~ 136), 468 (n = 102); braincase width 49:9 (n= 131}, 498 in = 103): cl-m2 length, 73.7 (n= 134), 720 (n = 96), mandible length. 126.8 (n = 143), 121.7 (a = 108—Crespo and De Carlo, 1963), Subspecies of P culpaeus der in skull length, especially in the muzzle, in « north-south gradient along the Andean Cordillera (Thomas, 1914) The muzde and skull of Pc culpaeus are longer (condslobasal length, 148-174 mm) than in the northem subsp cies, Pees reisstiand Pc. andina (eondylobasal length c3. 155 ‘nme rostrum length ca. 70 mam), and shorter than in ‘2 magellanica(condyiobaca length, 155-175 mim). Pc. [ycoides, in Tierra del Fuego, han the longest muzale and skull ofall sub- species (condylabasal length 185-201 mms length of rostrum 86-98 ‘tim—Crespo and De Carlo, 1968; Kraglievien, 1930; Langguth, 1967, 1960), ?.c,culpacns has alight tawny chin, grayieh body and upper side of the tal, tawny head bright tawny feet and legs, and dull tawny underside of tai. Pelage of Pc. andina is similar, but is paler throughout (Osgood, 1048): the head, legs an fet ate ochra feous tawny rather than tawny (Osgood, 1983). Pc. andina is Similar to Pc react in skull characters, but coloration of Pe landing is more suffused with bull above, especially anteriony, the heavy black grialing starts forther back, ad pelage is whiter below (Thomas, 1914a). The two southeramont forms, Fc. Lycoides and Pc. magellanica, also differ im thatthe former han an overall ‘rayish coloration with gray throat and chest while the latter i more Fufous with white thoat and chest (Philippi 1896). Pc. Iyoides {the largest ofall subspecies, frequently reaching a total Henyth ‘of 1.500 mm (Cabrera and Yepes, 190), Pe. amithers is char- acterized by its smal size (condylobasal Length of skull, 143-148 mmm length of rostrum ca, 5 mm) and an intense ferroginows co ion (Kraglievich, 1980; Thomas, 1914) DISTRIBUTION. The culpeo fox inhabits semideser An dean plateau, mediterranean serub and grassland, aes of western and southern South Amerea (Fig. 3, nga, 1943). Ie range extends from northem Ecuador (and per:MAMMALIAN SPECIES 558, 75 io 15° Fic, 3. Geographic distribution of Pseudalopes culpacus in South America, Numbers correspond to approximate rages (bourd= lanes for subspecies 1, 2.3, and 5 are not well established) of Subspecies (Cabrera, 1981, 1058): 1, Pc, ress; 2. P.c. andina: 3, Pec. eulpacus; 4 P.c. smithersi: 5, Pc. magellaniew: 0. Pe Iyeoides. hhaps southern Colombia, Narifo Province—Jiménez et aly 1995) tw southem Chile and Argentina, along the foothill of the Andes, including Tierra de! Fuego, and thoughout the Patagonian steppe Gf Argentina (Redford and Eisenberg, 1992) In Eousdor ii ound {nthe Andean region, far north 8x Cotopa in Pichinch Prov ince (UCN, 1988, In Pert, culpeos also range throughout the An= dean region atleast between 1,000 and 4,500 m and in the high- Tans to the south (Grimmer. 1969), ‘Galpeo distribution may have expanded eastward in Argentina during the last century. According 10 early accounts (Pricha 1902) culpeos were found only along the Andean Covillera and foothills during the early 1900s, but curently their rango ex- rust the coast of Patagonia south of 43°S (Fig 8). The increase ‘culpeo range may be related to the iateduction of European hare (Eepus europarus) and sheep (Otis aries) in the eatly 1900s (Crespo and De Carlo, 1968; Grigera and Rapopst, 1988). Calpe Aistnbution in Chile also may be determined in par by the dist= bution and denaty “af intraduced lagomoephe and” may. have ‘changed signfiantly during the last century Johnson, 1992) Isolated ealpeo populations accu in the Crabs bill of een- tual Argentina and 0 the island of Tier del Fuego. The former (subspecies Pc. smitheri) is separated from the remaining pop= lations by the extensive, love plains of La Rioja Province (Cabrera, 1981). P-.Iyeoides in Tiera del Fuego is separated fom mainland populations By the Stats of Magellan. FOSSH. RECORD. South American canide originated i North and Central America and migrated south after emergence of the Panamanian Land Bridge ca. 3.0 % 10> years ago (Bers. 1981, 1988). Maximus diversification of eanids occured in South Amer 1, pethape due to the abeence of olher camivorvus placental mammals and the presence of only. few species of carivorons ne and O'Brien, 1987; Webb, 1985), The earliest ‘Pseudalopes ofthe Chapadinallan age (3.01.8 X10" years ago) in north and central Chile (Moreno etal, 1998) and P. gymnocercus (Fischer, 1814) of the Uquian age (25-1.5 3 10" years ago late Pliocene and early Pleistocene) in the Vorohu Formation, Buenos Aires Province, Argentina (Krolievich, 1952; Pascual, 1966) Fossil ofthe genus Pseudalopes are also recorded .5-0.3 % 10 years ago, mide Pleistocene) Buenos Aires Province, Argentina, and i north and earliest record for P culpacus is frm La Ca the Lujanian (3.0-0.1 % 10> Years ago; late Plestocene~-Berta, 1987; Moreno et al. 1998), FORM. AND FUNCTION. Psoudalopex culparus has 9 ental formula off 3/8, € 11, p W/4, m 2/3, total 42 (Berta, 1987 ‘The molars in the larger subspecies of P culpaeus are enlarged, but this is probably a consequence of lager body size rather than of specialization in food habits (Langu, 1969). The eulpen fos, fa im olher species of the gens, has a long and coiled cee (Langguth, 1969). The far af eulpaeus becomes longer and dens- fe duting the winter. particularly on the wil (Crespo and De Caro 1963). The front and hind feet have five and four tas, respectively, and the limb posture is digitigeade (Cabrera, 1932). ‘The increase in body size of culpeas in southern Chile may be the result of bioencrgetic adaptation to colder rogions Jiménce tal, 1995) To meet daily energy requirements (calelated using ‘theoretical basal metabolic rates), eulpeo foxes need OCI. Eur ropean hares or 26-70 rodents in Chilean Patagonia. In north-cen- tral Chile, where culpeos are smaller. they need 02-04 logo- morphs or 3-6 rodents. These energy requirements have implica {ions for eulpeo distribution in relation tothe distribution of ttro- ‘duced lagomorphs and mediutsized rodents auch as Octodon tdegus and Abracoma bennett (Joho, 1992), ONTOGENY AND REPRODUCTION. Culpeo foxes Neuquén Province, Argentina, have a period of serua acti both sexes fom June though October (Crespo ard De Carlo, 1963), ‘They mate mainly between August and Octaber and produce one lier per year. with a peak of biths from October to December. Females are in anestrus feam October to July, proestas fr end of July to mid-October, and estrus from eatly August to Oc- tober; estrus is followed by pregnancy or pseudopregnancy (males n= 5; females: n = 48—Crespo and De Carl, 1963). The ges- tation period is 55-60 days, and mean number of embryos is 5.2 (n= 6—Crespo and De Cari, 1963). In Chile the gestation period {i estimated at 65 days and Titer size ranges from three to five young (Housse, 1953) "Young eulpeos are bom with thei ayes closed. At 2 days of age a male weighed 166 g and a female 170 ¢ (Crespo and De aro, 1963), Young culpeus are weaned at 2 months of age and gow to adult size within 7 months (Crespo and De Carlo, 1963) Both male and female may care forthe young (Gitleman, 1980), By 1 year af age, developing males prodce mature sperm in bath testes, and females ovulate (Crespo and De Caro, 1963), ECOLOGY. Preudalopes eulpaeus lives in mountainous hab tats on the puna highlands up to 4.500 am, and on plains down to sea level on either side ofthe Andes mountain chain (Marquet et sl, 199%; Medel and Jaksic, 1988). 1 occupies habitats tha ‘ther forested or covered by shrubby or herbaceous vegetation. I fouthern Chile andthe Andean fothillsof northwest Argentinean Patagonia, culpens select Nothefigus thickets and matoral sheul- land habitats where prey are mere abundant and there is more cover for resting ad den sites Crespo and De Carlo, 1963; Johnson tnd Franklin, 1994) In steppe habita of noriwest Patagonia, fulpeos use humid valleys to forage and slopes and rugged reas to rest al den (Diuk Wasser, 1995). Paeudalopex culpaeus is gencrlly an opportunistic predator, although it can be leally celective for certain prey (Iriarte etal. 1980; Johnson and Franklin, 1994; Meserve etal, 1987; Rau et aL, 1987). Culpeos consume more vertebrate prey and les plant tmattor and invertebrates than any other South American canid (Redforl and Eisenberg, 1992), Main prey items of culpnos are small mammals wnd introduced lagomorphs. Small mammals and European rabbits (Oryetolagus cuniculus) are consumed in vay proportions in central Chile (Ebensperger etal 1991; Fuentes Jake 197 ate ot al 299% okie ad Yann 980 alse 1980 Yaner and Taksic, 1978) and on Tierra del Fuego faland (aksic and Yates, 1983) European hare Lzpus europacts) ‘dominate the eulpen diet in the Magallanes Region of southern Chile Johnson and Franklin, 1940) and in northwest Argent Patagonia (Crespo and De Carl, 1968; Novaro, 1991), Additional4 prey include arthropods and fruit at most sites, mainly in Chile: ‘sheep (Oris ares, party as earion) in Argentinean Patagonia (Bel- lati and von Thungen, 1990; Creapo and De Carlo, 1968: Novato, 1901) andl Tierra el Fuego (Jaksic et al. 1983) birds (tainly Chloephaga pieta and passerines) a guanacos (lama guanicoe, probably as carrion) in Southeen Chile Jaksic etal. 108%: Johason find Franklin, 19a; Yater and Ras, 1980); and reptiles in cent Chile (Ebensperger et al, 1991), In northern Chile eulpeo foxes prey mostly on small rodents Jaksie et al. 1093: Meserve etal, 9G), and ton lesser extent on cepts, ieds, arthropods, and Fruit (Dutdn etal, 1987: Jaksie otal, 1992: Jiménes, 1993), Functional and numerical” responses of eulpeo foxes to changes in prey availabilty vary among sites and type of prey. A functional responce of prey switching (Jaksic and Simonct, 187) was recorded ia ceneal snd southers Chile and Angentina. Doing sets nen ren declined, ex ‘and Fro 1991) diet to matked changes in Octodon degus abundance, maintsising 4 strong preference for this species. However, culpeos displayed 2 nificant numeral response fllowing fituations im abundance of 0. degus at Fray Jorge National Park in north-central Chile Uaksic etal, 1903}. Finaly, eulpeos did not show a functional oF ‘numerical response during decline of ther rodent prey (aay 1. degus and Abrocoma Bennett) at Aucé in north-central Chile laksie et al, 1992, 1906, Martinez etal, 1993). ‘Culpeo predation has significant elfects on populations of small rodent that ace overeepresented (relative to their wala) inthe for det (Meserve et al, 1996) Culpeo selectivity fr dent species has been related tothe prey's habitat use and body size (rare etal. 1989), body sie and time of aetivity Gaksi, 19865, Meserve etal, 1987}, a combination of size and abundance Uaksi 1960; Jaksic & a 1992), and higher vulnerability of cee cetine species beeause of morphology (Corley etal, 1995) Human disturbance may affect culpeo diets significantly. In areas of Chile and Argentina where sheep and lsopean hare and {abit were intsduced, up to 96% of eulpen diet hiomass is r= resented by these prey (Crespo and De Carlo, 1963: Johnson and Franklin, 1094; Novar, alteration by humans also may cause an increased consumption of O. degus, rodent abundant in disturbed habitats (Simonetti, 1998). How: fever, studies show that O. degus is the dominant fem in eulpr> se Aliets even where this rodent is not the most abundant speci faite etal, ISB; Meserve etal, 1987), suggesting that select plays a more important role than human disturbance in determin ulpeo fod habits (Meserve, 1588) “The role of the eulpeo fox as a disperser of ree seeds in central Chile is species-specific (Casto etal, 1994). Seeds of pa ‘no (Cryptocarya’ alba) and pimiento (Sehinus molle) defecate by fulpeos are viable and germinate at higher rates than those not pasted through digestive tacts, while sed of ie (Lthrea eats- fea) germinate at lower rates. Seeds of pimiento are defecated in sites where successful establishment of seedlings is possible, ‘whereas peumo seeds are not (Bustamante et al, 1902; Casio et 1994; Letin-Labos and Kalin-Arroyo, 1993), Culpoo population structure in Neuquén Province, Argentina was 77.9% for those OL year of age, 13.8% for those 1-2 years af ‘ge, and 8.7% for those 2 years of age (Crespo and De Carlo, 1908). Sex rato ofthe Neuqucn population was significantly biased towards males in 1959-61 (15s: n = 254——Crespo and De Carlo, 1968), but was not significantly different from a 1:1 ratio 30 years later (L081: n= 102—Novaro, 1991). This change in sex ratio ray be a response to increased hunting pressure (Novato, 1995). Population density was 0.72 ulpeos/km’ in Neuquén Province (e>- timated by removal—Crespo and De Carlo, 1968) kn in Torres del Paine Retional Pask in Chile (est ‘censts—Abell, 1979), Culpeo density in Ne 1915 when European hares and sheep were inineuced (Crespo and De Carly 1963}, and it may be declining in Salta Province, At renting (Mares etal, 1981). ‘The South American ray fox or chilla, P. griseus, may be a potential competitor of P eulpacus (Fuentes and Jaksic, 1979: Jak Sic otal, 1980, 1983) The two foxes are sympatric in many areas ‘of Argentina and Chile, but appear to use dflerent microhabitats. Cullptos use more wowed and densely vegetated habitats where MAMMALIAN SPECIES 558, their prefered prey are more abundant (Crespo and De Carlo, 1968; Jaksie etal, 1980; Jimenez, 1998; Jimener etal, 1995). In ropean ‘lant. The ptosis likely dictate! bythe higher seabolc eda ‘of culpeos and mediated by the aggressiveness of the larger culpeos and consequent avoidance by gray foxes Johnson and Franklin, 1904, although no dveetculpeo-geay fox interactions have been recorded Johnson and Franklin, 19940), Other potential compet tore of eulpeos are lesser grisons (Galictis cua), which prey on Eupean rabbits, rodents, and reptiles (Ebensperer etal. 1991), Geotlioys eat (Onefelis geafroyt) and paras cat (0. colocolo), which prey om small mammals and birds Uohnson et sl, 1992), nd owls (Speotyto cunicularia, Tyo alba, Bubo virginiana, Glaueilnm nani) av falconiform (Geranoaetus melanoe Busco polyosoma, Parabuteo unicinetus, and Falco sparverus), hich prey mainly on small mammals or lagomorphs Uaksic tal ToB1, 1092, 1003; Meseeve etal, 187) Parasite burdens in the digestive tracts of P eulpaeus from Neuquén. Argentina, are Jow. From one to five nematodes repre= senting three species (Physaloptera claus, Tosasears leonina, land Protosprura numidiea crveticola) were found io 5 of 129 culpeos studied. Arthropods and rodents are the main intermediate Iho: therefore, reduced predation hy eulpeos on these prey mi explain the low prevalence of nematodes in Neuguén (Sein et al 1994), The adult potastomid Linguatula serrata yas found inthe trachea of eulpeos from Chile. The larvae ofthis parasite ae fone in rodents and lagomorph (Alvarez, 1960), Fleas (Pale iritan) land biting lie (Trichodectes canis) arc ectoparasites of culpeos (Crespo and De Carl, 1963: Hopkins, 1989), Prevalence af epidemiologically signlieant Kchinococeus granulosus in culpeos from Neuguén was 12% in 1960 (6 of 50, culpeos—Seidat, 1960, 1963), 26% in 1971-1972 (9 of 3-— Schantz etal 1975), and 05% in 1989-1990 (0 of 129—Stein et al, 1908). The deeline in the prevalence of F. granulosus in cul ‘pes followed an intense eampaign in Neuquén Province to teat Alomestic dogs against this cestode (C. Rambeaud, pers. comm). CClpens ean become infected by eating infected sheep (Shantz #1 al, 1976), but lavae of B. granulosus azo also found in European hates from Newquén, which suggests the existence of cycle wi splvatic hosts (Schanty etal, 1972). However the low E. gran- Tosus burdens in culpeos from Neuguén and the absence of infec- tion in Chile (Alvaret, 1961; Blood and Leliveld, 1969) suggest that culpeos are of limited importance in the transmission ofthis paraiite (Schant et al, 1975) biants of the Andean region of Ecuador and Pent prac- iced hunts and burials of ealpeo and Sechura-decert foxes for cer- -monial purposes prior to 1750 B.C. (Wing. 1989). Culpeo foxes may have been domesticated by Indians on Tera del Fuego (Ham= fon Smith, 1839) Currently culpeos are intensively hunted it Ae eatinean Patagonia, bot Sheep and goats (Bell Macdonald, 1900}. Culpeos kill up 19736 of la in northwest Patagonia (Dellti, 1986). Commerc pos is dane mainly by rural workers, who supplement thet i Comes with sales of fox fur (Novara, 1098). Simulations of culpeo population dynamics indicate that the curcent harvest in Argentine an Patagonia may be sustainable because of heterogeneous istic bution of hunting, which is a consequence of topography and di ferent levels of hunting pressure on sheep and cattle ranches (N varo, 1995), This conclusion is supported by the results of moni= toring programs of culpeo density throughout southern Argentina {Novaro and Funes, 1994; von Thngea, 1991), except in Tierra pers. comm). Gulpeo hu hunting is common outside protected areas (iménex. 1993), The species is included im Appendis I of CITES (Medel and Jaksic. 1983, Researchers studying food habits of culpeos in areas of sym= patry with other eamivores may encounter difculties identifying theitfeees(liménez, 1993; Johnson and Franklin, 19944, Jiménez ct al. (1996) attempted to differentiate feces of eulpeo and chia foxes using thin-layer chromatography of thet ile acid contents, bout were notable to consistently profile bile acid standards, Ca- pura etal in press) successfully distinguished! feces of esipeos nd chills witha improved bile-aeid thislayer chromatographyMAMMALIAN SPEC 538 technique and reported bile-acd profiles of six other Neotropical carnivores potentially sympatie with culpeos BEHAVIOR. Pseudalopex eulpacus is primarily nocturnal in southern Chile and northwest Argentinean Patagonia, where it ‘was stuied with radiostelemetry(Diuke Wasser, 1995; Johnson and Franklin, 199). Culpeo eetvity in southern Chile increases dra- ‘matically within an hour or two of sunset and decreases near sin- ‘ice. In central Chile eulpews are ether erepuseular Jaksic et al, 1980) or active during day and night (rane tal, 1989), although these conclusions are inferred frm the activity palterns ofthe prey of. culpacus. Surplus killing of lambs by culpeos is reported in Rio Negro Province, A ‘Culpeos didnot feed on 48% of their hills (1 and von Thungen, 1990). “Annual and seasonal home range sizes of culpeos in southern Chile averaged 98 kv’ and 7-7 ki (x ~ 8), respectively. There were no significant differences in home range sizes among seasons and between sexes (Johnson and Franklin, 1994e), Culpeo. vocalizations in the Rld have not been described (Beanch, 1998). Captive culpeos make a mixed growi/scream sound (Cohen and Fox, 1970). GENETICS. The culpeo fox has a diploid numberof fundamental number of 76, and all tlocentie autowomes and Zuleta, 1992). The karyotype is identical to other species of the genus (Tedford et al. 1005). 7 culpaeus and P griseus have the southerumost range of species in South America and are likely lo represent the most recent speciation events in the radiation of Psoudalopes species (Wayne eal, 1989: Yahnke et al, 1990). Based on polymerphisms in mitocondrial DNA restriction sites, the two species dered apromtely 25-50% 10x ‘eranial morphology over this relatively short time span. Character divergence may have been fostered by low prey and predator d= ‘versity present in South America when the species evolved during the PliosPleistocene (Wayne etal, 1989) ‘The Samson mation is characterized by the absence of guard his and sYellowish-white appearance of far eaased by the under hair coloration (Voipio, 1950}. This mutation is present in 3.3% of the population (9 of 257 eulpeos) in northwestern Patagonia, At ‘genina (Crespo ann De Carlo, 1963), REMARKS. The assoc nat fhe genus of je fox in unresolved, Cabrem (1981, 1940) gave Parndafopex Burmeier, 56 the rank of gems, nla the eulgenfox in and con cred sopante om yealopes Burr, 51. Osgod 1508) ita isuficien evidence t apport evel spain of Pre dates from the ges Dnatson nite sty Cabra (1958) Scope We Inston of the geo Paeudolpe Stet gem ‘rateyon: Langu (1989 1910) considered Predalope sabe fens of ustojor, Ener (1973) sso asgted the grap to the emis Dae, mbereas Lange (1975) and Van Geer (1978) ve Pirudaloer the rank of seen Smeal elasticaon othe fail Ca {15%6) sgescd placing the eles fox inthe genta Dasion fd even dobied the recomton fille species previ ‘ihgued a Bcupacus an P gymnocecss Basal ont da Salas Sah American cant, ena (87 seoghaad Pate dates sa nepart taxon and reac as consi wh ‘lope, lng the fet tne forthe. gents, Wore (1903) Placed tte clpeo fox under the genus Deaton. together ih the treating on, Cdocyo thous Halon Shy TE39, bin Inter analsis(Werencra 198) aged the clpe ts Panda lope and peed with Bea (1987) in reaing yeolopeswcn- ftcrie" Zino oa (198) flowed ena and Wereaca trating Zcalopen and Peudaloper os congener bt sed 3 ‘lope fr the gens becue has 2 ear pry. Final ins ‘eee lai stay of living aver of ca, Teo! oo {1555 conned thal Pada en separate gen om y= "otier vemacalar names fr he species ae roo cola, seo sr de ilar Gia and Macon, 199) {tps ay dete fm the ave Chilean wor! eulpem, whieh team madncn esse the anal expones Hinsley Bring Killa by haters (Malin. 138 Dp, 1943, Tihank 1, FBsenberg. 3. Emenee, A Ber, and Pt Merve fr vtmable cn on Wis ene, Cari, skall figures and assisted with the li 41. Lombarlo, G. E. Zunino, 0. B, Vaccaro, P. Vuillermoz, RS. Walker, and'P. Colavino also provided valuable assistance. This per was written while the author held « Fulbright Fellows at the University of Foride and a graduate fellowship from the Ui ‘of Buenos Aires. This publication is No. R-05957 of the Florida Agricultural Experiment Station Series. LITERATURE CITED ‘Aweiio, 0. 1979. Densidad de una poblacida de 21s colorados Dusicyon culpacus, en el Parque Nacional Tores del Pat (Magallanes, Chile) Informe Corporacisn. 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