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M S AMMALIAN PECIES No. 783, pp. 1–6, 3 figs. Speothos venaticus. ByBeatriz de Mello Beisiegel and Gerald L. Zuercher Published 20 December 2005 by the American Society of Mammalogists Speothos Lund, 1839 Bushdogslackfacialmarkings.Skullisshort(Fig.2;Vieira1946). S. v. panamensisissmallerthanS.v.venaticusandS.v.wingei, SpeothosLund,1839:224.TypespeciesSpeothospacivorusLund, which aresimilar in size(Linares1967).S. v. panamensisandS. 1839, by monotypy. v. wingeiarelight-coloredcomparedwithS.v.venaticus(Linares CynogaleLund,1842:203.TypespeciesCynogalevenaticaLund, 1967). 1842, by monotypy;preoccupiedbyCynogaleGray,1837. Measurements (ranges, in mm) of S. v. wingei are: length of Icticyon Lund, 1842(1843):80. Replacement name for Cynogale headandbody,730–740(n53);lengthoftail,110–150(n53); Lund, 1842. totallengthofskull,132–133(n52);palatallength,61–63(n5 AbathmodonLund,1843:74. Typespeciesnotgiven. 2); zygomatic width, 80–81 (n 52); braincase width,46–47(n5 Cynalicus Gray, 1846:293. Type species Cynalicus melanogaster 2); interorbital width, 24 (n 5 2); length of mandible, 102–102.5 Gray,1846, bymonotypy. (n 5 2—Vieira 1946). Measurements (in mm) of 1 adult male S. CynaliusGray, 1847:18. IncorrectsubsequentspellingofCynali- v. panamensis are length of head and body, 730; length of tail, cusGray,1846. 105; length of hind foot, 110; length of ear, 35; length of skull, Melictis Schinz, 1848:177. Type species Melictis beskii Schinz, 120; condylobasal length, 124; palatal length, 61.5; basal length, 1848, by monotypy. 116.5; nasal length, 29.5; interorbital width, 31.6; supraorbital MelictesGray,1868:498.IncorrectsubsequentspellingofMelictis width, 39.8; postorbitalwidth, 26.5; zygomatic width,75.8;length Schinz,1848. ofmandible,91 (Linares1967). CynalycusGray, 1869:183. IncorrectsubsequentspellingofCyn- alicusGray,1846. DISTRIBUTION. Speothos venaticus occurs from Panama´ to southern Brazil, Paraguay, and northern Argentina, extending CONTEXT AND CONTENT. Order Carnivora, suborder westto Bolivia, Peru,andEcuador(Fig.3;Cabotetal.1986;Ca- Caniformia,familyCanidae,subfamilyCaninae(Stains1984).The breraandYepes1940;Linares1967;Yahnkeetal.1998).S.ven- genus Speothos is monotypic. Speothos was previously placed in aticusis widely distributed in northernSouthAmerica(Eisenberg the subfamily Simocyoninae (Simpson 1945) with Cuon and Ly- 1989), with fewer reports from southeastern (SaõMiguelArcanjo– caon. Beisiegel1999;SaõJoaõdaBoaVistaandLorena—Carvalho1980) andsouthernBrazil(Cimardi1996;Ihering1911).S.v.panamen- Speothos venaticus (Lund, 1842) sis occurs in northwestern South America; S. v. venaticus occurs intheAmazonRiverbasin,centralBrazil,easternBolivia,northern Bush Dog Paraguay,northeasternPeru,easternEcuadorandtheGuyanas;and Cynogale venatica Lund, 1842:203. Type locality ‘‘Lagoa Santa, S. v. wingei occurs in southeastern Brazil (Linares 1967). Bush MinasGerais,Brazil.’’ dogs occur from low elevations (Eisenberg 1989) to 1,900 m alti- Icticyonvenaticus:Lund, 1842(1843):80.Namecombination. tude(R.Wallace,pers.comm.). Cynalicus melanogasterGray,1846:293. Typelocality‘‘Brazils.’’ FOSSIL RECORD. Speothos originated from canids that Melictis baskii Schinz, 1848:177. Type locality ‘‘Nova Friburgo, evolved in the Brazilian highlands (Berta 1984; Langguth 1975). Estadodo RiodeJaneiro,Brazil.’’ Speothospacivorus(extinctspecies)andSpeothosvenaticusoccur Speothos venaticus: Thomas, 1903:236. First use of currentname inthelatePleistocene(300,000yearsago)—Recentdepositsfrom combination. LagoaSantaCaves,Brazil(Berta1984, 1987). SpeothoswingeiIhering,1911:222.Typelocality‘‘EstadodeSanta Catarina,ColoˆniaHansa,Brazil.’’ FORM AND FUNCTION.Bush dogs are a highly special- Icticyon panamensis Goldman, 1912:14. Type locality ‘‘Cerro izedlineageofSouthAmericancanidsthatevolvedfordwellingin Pierre,Panama.’’ forests(Langguth1975).Theyhavepartiallywebbedfeet(Macdon- ald 1996)adapted for walkingonsoftsoilnearstreams.Bushdog CONTEXTANDCONTENT.Contextasabove.Threesub- tracks and stride in the field are as follows: forefeet, 5.7 by 5.4 speciesarerecognized: cm; hind feet,5.0by4.4cm;stride,28cm(VillalbaandYanosky S. v. panamensis(Goldman,1912:14);seeabove. 2000).Bushdogdentition,specificallythereducednumberofmo- S. v. venaticus (Lund, 1842:203); see above; baskii (Shinz) and lars,indicatesahighlycarnivorousdiet(Flower1880).Dentalfor- melanogaster(Gray)aresynonyms. S. v. wingeiIhering1911:222; seeabove. DIAGNOSIS. Atelocynus microtis and Speothos venaticus overlapgeographicallyintheAmazonianregionsofBrazil,Colom- bia, Ecuador, and Peru. S. venaticusisdistinguishedfromA.microtis by absence of M3 and absence of metaconid and entoconid on M1 and M2. S. venaticus has interdigital webbing and a carnassial ridge that A. microtis lacks. S. venaticus has shorter legs and tail than A. microtis and has paler pelage around head and neck. Head of S. venaticus is mustelid like, whereas head of A. microtisisfox-like(Vieira1946). GENERAL CHARACTERS.Speothos venaticus(Fig.1)is a small canid (body mass, 4–7 kg) with short legs and tail, inter- digitalmembranes,elongatedbody,andsmallroundedears(Vieira 1946).Colorrangesfromuniformdarkbrowntoyellowishred,with lighter color on neck, nape of neck, and ears (Sheldon 1992) and FIG. 1. A captive adult male Speothos venaticus at Parque darker color on limbs, tail, and ventrum (Coimbra-Filho 1972). Estoril, Saõ Bernardo do Campo, SP, Brazil, 1999. Photograph by Younganimalsareuniformlyblack(DarlingandWhitehead1991). B. M. Beisiegel. 2 MAMMALIANSPECIES 783—Speothosvenaticus FIG. 3. GeographicdistributionofSpeothosvenaticus.Sub- speciesare1,S.v.panamensis;2,S.v.venaticus;3,S.v.wingei. SubspeciesboundariesfollowCabrera(1957),Carvalho1980,Iher- ing(1911),Linares(1967),andMassoiaetal.(1987).Circles(open andsolid)showsitesofoccurrenceforwhichpreciselocationsare reportedintheliterature.SourcesareBarnettetal.(2001),Beisie- gel(1999), Calouro (1999),Carvalho(1980),Cimardi(1996),Dal- ponte(1995),Eisenberg(1989),Massoiaetal.(1987),Peres(1991), RedfordandEisenberg(1992),Vieira(1946),Wallaceetal.(2002). Distribution map modified from Eisenberg (1989), Eisenberg and Redford (1999), Emmons (1997), Redford and Eisenberg (1992), and Zuercheretal.(2004). surveyrespondentsfrom7rangecountries(DeMatteo2004).Births typicallyoccurduringtherainyseasoninSuriname(Husson1978). Estrus in captivity lasts 1–12 days (Porton et al. 1987). Two FIG.2. Dorsal,ventral,andlateralviewsofcraniumandlat- primiparousfemaleshadinterestrousintervalsof15–44days.One eralviewofmandibleofSpeothosvenaticus(specimen19743from female had 4 and the other 7 estrous periods in 1 year. Earliest MuseudeZoologiadaUniversidadedeSaõPaulo,Brazil).Greatest ageatconception was10 months(Porton et al. 1987)andearliest lengthofskullis130 mm. age of reproduction in males was 12 months (Bekoff et al. 1981). Bushdogshavecopulatorylock,butnotback-to-backposture(Dru¨- wa 1982, 1983; Kleiman 1972). In captive bush dogs, copulatory mula is i 3/3, c 1/1, p 4/4, m 1/2, total 38 (Vieira 1946). Lower locks occurred 1–3 times per day during estrus (DeMatteo 2004; carnassialdiffersfromcanidpattern:innercuspoftalonidismiss- Dru¨wa 1983; Kitchener 1971; Kleiman 1968; Porton et al. 1987). ing, resulting in this part of the tooth forming a subsidiary blade Pseudopregnancy occurred in captive bush dogs after an infertile andnotabasin.M1ismodifiedtoformabasinintowhichfitsthe matingorafteranovulationwithoutmating(DeMatteo2004). singlehypoconidcuspoflowercarnassial. Bush dogs have 4 paired mammae (over the ribs ca. 5 cm Incaptivity,mean(1SE,indays)interestrousintervalis238 behind elbows, on abdomen, opposite knee joints, 4.9 cm in front days6 39.6 (range, 179–301; n 5 11), but may be shorter if the ofvulva—Flower1880).Stomachhasasubglobularcardiacportion litter islost(Dmoch 1995; Portonetal.1987).Gestationaverages andanarrowerpyloricportion,separatedbyaconstriction.Diges- 67days(Portonetal.1987),rangingfrom65to83days(Moehlman tivecanalhasareducedcaecum(ca.3.2cm—Berta1984,1988). 1986).Bushdoglitterstypicallyhave3–6pups,althoughlittersof Anal glands are large (2.3 cm long and 1.8 cm wide), oval, and at least 10 occur (Dmoch 1995). Neonatal mass ranges from 125 openwithasingleorificeatlateralmarginofanalaperture(Flower to 190 g. Sex ratio at birth (males:females) is 1:1.4 (Moehlman 1880).Speothosvenaticushasamassivefrontalpoleofbrain(Lyras 1986). Eyes of young open in 14–19 days (Collier and Emerson 1973;DarlingandWhitehead1991).Pups1stemergefromdenat and van der Geer 2003). Acompletesulcusdoes notseparatethe 2.2 weeksandweaningoccursat4 weeks(Bekoffetal.1981). upper curve of the gyrus immediately surrounding the Sylvianfis- A litter of 6 bush dogs (3 males and 3 females) was born in sureon leftsidefromthesulcusaboveit(Flower1880). 1971 at the Los Angeles Zoo (Collier and Emerson 1973). One ONTOGENYANDREPRODUCTION.Captivebushdogs female was dead when the pups were retrieved from the mother do not reproduce seasonally (Porton et al. 1987). Aseasonality of and 1 died ofbacterialsepticemia10dayslater.Threemalesand bush dog reproduction in the wild has been largely confirmed by 1 female were successfully hand raised. These surviving pups 783—Speothosvenaticus MAMMALIANSPECIES 3 weighed 170 g at birth, lost weight during the 1st week, but reg- lococcus aureus, and S. epidermis), fungi (Candida—Van Hum- ularly gained weight afterward, weighing476–546 g at 40 daysof beck andPerez1998),and protozoans(Giardia). age. Femalesraisedby1orbothparents,aloneorwiththeirsisters, HUSBANDRY.Onecaptivebushdoglivedfor10yearsand do not cycle, but do so when paired with a male (Porton et al. 4 months (Jones 1982). At the Los Angeles Zoo, a suckling re- 1987).Presenceofamalewasnotrequiredforovulationbutshort- sponsewasdifficulttoelicitfrompupsduringthefirst8days.Pups ened interestrous intervals and increased frequency of estrouscy- were fed a formula of 1 part Esbilac and 2 parts boiled distilled cles(DeMatteo2004). water; concentration was increased until full Esbilac by day 19, and bottle feeding continued until day 39. From day 7 to day 25, ECOLOGY.Bushdogsareextremelyrareovermostoftheir a vitamin supplement was added and, from day 40 on, the young distribution, but are common at some sites in Suriname (Husson were given Esbilac in a shallow dish and puppy chow in small 1978). S. venaticus typically occurs in lowland habitatsincluding fragments (Collier and Emerson 1973). At Twycross Zoo, United galleryforests,forestedge,wetsavannas,andriparianareas(Aqui- Kingdom, bush dogs are fed raw meat, live rats, and bone meal noandPuertas1997;Defler1986;Eisenberg1989;Emmons1997; with vitamin supplements. A more varied diet of commercial dog Strahl et al. 1992; Zuercher et al. 2005). Bush dogs alsooccurin food,fruit,andeggscausedvomiting,dehydration,andlossofap- open habitats such as cerrado (savanna—Silveira et al. 1998; petite,mainlyduetothecommercialdogfood(DarlingandWhite- Zuercher and Villalba2002),ranchlands(T.Oliveira,pers.comm; head1991).Otherzoosfeedonlycarcassesandliveanimals(Mac- L. Silveira, pers. comm.), and altered habitats (da Fonseca and donald1996). Redford1984).Homerangeestimatesare3.8–10.0km2(minimum Bushdogscanbeimmobilizedwith10mg/kgTelazolsupple- convex polygon—Beisiegel 1999). Bush dogs have semi-aquatic mented, if necessary, with 10 mg/kg ketamine, or with 20 mg/kg habits (Langguth 1975), and most sightings occurred near rivers, ketamineplus0.2mg/kgacepromazine(Kreegeretal.2002).Bush streams, and watercourses (Aquino and Puertas 1997; Barnett et dogs at the St. Louis Zoo were immobilized with 50 mcg/kg med- al. 2001; Deutsch 1983; Linares1967; Peres1991;Wallaceetal. etomidine combined with5–6mg/kgketamine(DeMatteoandKo- 2002).AfemaleswamacrosstheNegroRiverwith2pups(Coim- channy2004). bra-Filho1972),andapairswamintheCauraRiverinVenezuela (Strahletal.1992).Bushdogspursueandkillpacas(Agoutipaca) BEHAVIOR.AsummaryofbehaviorofS.venaticusisavail- inwater(Strahletal.1992;Tate1931).AtEmasNationalPark,a able(BeisiegelandAdes2002).S.venaticusisdiurnal/crepuscular cerrado (savanna) area of central Brazil, 3 of 9 sightingsoccurred (Dalponte 1995; Defler 1986; Deutsch 1983; Linares 1967; Peres atsiteswithin200mofwater;theother6werefrom2,600to5,700 1991;Silveiraetal.1998;Strahletal.1992;ZuercherandVillalba m from water (Silveira et al. 1998). At the Mbaracayu´ Reservein 2002), although individuals have been observed at night between eastern Paraguay, most bush dog feces, tracks, and sightings 0100and0400h(Wallaceetal.2002).Incaptivity,bushdogsare (.70%) were detected ,1,000 m from water (Zuercher 2001). A active during the day and retire to dens at night (Kleiman 1972). captive female spent a great deal of time in a water pond where Play behavior has been described for young S. venaticus (Biben shedivedandswamwithease(Bates1944). 1983). Seventy-one percent of object play by a captive group oc- Bush dogs use dens dug by large armadillos (Coimbra-Filho curredinwater(Macdonald1996). 1972)andsometimestamanduas(Sanderson1949).Otherdensites The basic social unit of S. venaticus is a monogamous pair arefallentreetrunks(AquinoandPuertas1997)androckshelters andextendedfamily.Incaptivity,bushdogsformpairbonds(Dru- (Linares1967).Densitesoccurwithscattereddryandfreshfeces wa 1983; Macdonald 1996; Porton et al. 1987) and parents are (AquinoandPuertas1997)orfreeoffecalmaterial(Linares1967). dominant over all group members(Macdonald 1996). Onlytheal- Agouti (Dasyprocta), capybara (Hydrochoerushydrochaeris), pha female reproduces and other group members display allopar- andpacaarethemainpreyofbushdogs(AquinoandPuertas1997; entalbehaviors,suchasguarding,carrying,andlickingpups(Mac- Cabrera and Yepes 1940; Deutsch 1983; Peres 1991; Silveira et donald 1996). Fathers help during parturition by grooming their al.1998;Strahletal.1992;Tate1931;Zuercheretal.2005).One mate’sanogenitalregionandremovingtheafterbirth(Porton1983). groupof6bushdogshuntedatapir(Tapirusterrestris)thatweighed Grouplivingoccursinthefield(AquinoandPuertas1997;Barnett 250kg(Wallaceetal.2002).NearSerradasAraras,StateofMato etal. 2001; Defler1986; Strahletal. 1992).InthePeruvianAm- Grosso,Brazil,bushdogsalsohuntarmadillos(Dasypodidae),col- azon, 2 adult bush dogs explored an area of ca. 60 m diameter lared peccaries (Tayassu tajacu), and deer (Mazama—Beisiegel aroundapileofbranches.Fifteenminuteslater,ayoungbushdog, 1999). Indigenous groups from Ecuador and Peru report that S. followed by a pup, emerged from the pile (Aquino and Puertas venaticussuccessfullyhuntscollaredpeccariesandtapirs(Descola 1997). Although most sightings of bush dogs are of groups, sight- 1994).Bushdogsalsohuntrheas(Rheaamericana—Santos1945). ings in the cerrado of Emas National Park were mostly of single In Paraguay, bush dog feces contained remains of agouti, paca, individuals(Silveiraetal.1998). tinamou (Tinamidae), an unidentified snake, several unidentified Members ofbush doggroupshuntcooperatively(Aquinoand smallrodents, and seedsfromCecropia(Zuercheretal.2005).In Puertas 1997; Barnett et al. 2001; Dalponte 1995; Defler 1986; the Peruvian Amazon, a single bush dog’s feces contained hair of Linares 1967; Peres 1991; Strahl et al. 1992). When a bush dog agouti (Dasyprocta fuliginosa), coati (Nasua nasua), smaller ro- group is hunting a paca, part of the group chases it on land and dents (Myoprocta pratti or Proechimys), and feathers similar to part waits for the paca in the water (Cabrera and Yepes 1940). those of tinamous (Aquino and Puertas 1997). Captive bush dogs Communal hunting occurs in captivity (Macdonald1996).Captive readily accepted 9-banded armadillos (Dasypus novemcinctus), pupsarefearfulwhentheyindividuallyencounterprey,butattack opossums (Didelphis), and rabbits (Sylvilagus brasiliensis—Van prey when near their parents. During ingestion of large prey, par- HumbeckandPerez1998).Bushdogsindividuallyhuntsmallprey entspositionthemselvesattheextremesofthecarcass,thusfacil- (Silveira et al. 1998). In Paraguay, S. venaticus and other carni- itatingdismembermentofpreybypups (Biben1982b). voreshaveminimaldietaryoverlap(Zuercher2001). Bush dogs have 10 distinct vocalizations: short whines, ex- Only indirect evidence of predation exists on bush dogs. In tended whines, repetitive whines, pulsed vocalizations, short PeruvianAmazonia,thecarcassofanadultbushdogwassurround- screams, long screams, barks, growls, infant whines, and infant edbytracksofjaguar(Pantheraonca)orpuma(Pumaconcolor— grunts(Brady1981; Villa2001).Frequencyrangesforseveralvo- AquinoandPuertas1997).Speothosvenaticusishuntedforhuman calizationsareshortwhines,450–1,790Hz;extendedwhines,540– consumptioninsomeareasofAmazonia(Calouro1999). 1,660 Hz; repetitive whines, 520–1,570 Hz; pulsed vocalizations, SpeothosvenaticushostsEchinococcusvogeli,acestodewith 1,140–1,590Hz;shortscreams,800–1,720Hz;longscreams,690– pacas as the intermediate host (D’Alessandro et al. 1979; Rausch 1,550 Hz; barks, 410–1,720 Hz; growls, 450–1,590 Hz; infant and Bernstein 1972). In Venezuela, bush dogs host Lagochilas- whines, 670–880 Hz; infant grunts, 820–1,590 Hz (Villa 2001). caris, an ascaridian nematode with rodents, such as Dasyprocta Vocalizationsfacilitatecommunicationamonggroupmembersinan leporina, as intermediate hosts (Volcań and Medrano 1991). Cap- environmentthatobstructsvisualcontact(Brady1981;Villa2001). tive bush dogs have succumbed to canine parvovirus (Janssen et Recorded vocalizationsand depositionofurinehaveattractedand al.1982;MontaliandKelly1989).Captiveanimalsmayhavesuc- elicited vocal responses from wild bush dogs (DeMatteo et al. cumbed to a vaccine-induced canine distemper virus (McInneset 2004). al.1982).Otherpathogensfromcaptivebushdogsincludebacteria Femalebushdogshaveaspecificurinaryposture(theraised- (Escherichia coli, Klebsiella,Proteus vulgaris,Shigella,Staphy- hindquartersposture,orhandstand)inwhichtheyclimbavertical 4 MAMMALIANSPECIES 783—Speothosvenaticus surfacebackwardwiththeirhindlegsandurinatewhilestandingon versityofCaliforniaPublicationsinGeologicalSciences132: their forelegs. They then slide their ano-genital region down the 1–132. vertical surface (Biben 1982a; Kleiman 1966). Male bush dogs BIBEN, M. 1982a. Urine-marking during agonistic encounters in slightly extrude their penis and move laterally to create a spray thebushdog (Speothosvenaticus).ZooBiology1:359–362. when theyurinate(Kleiman1972). BIBEN,M. 1982b. Ontogenyofsocialbehaviourrelatedtofeeding in the crab-eating fox (Cerdocyon thous) and the bush dog GENETICS. Speothos venaticushas 2n 5 74 chromosomes (Speothosvenaticus).JournalofZoology196:207–216. with 36 acrocentric chromosomes (Schreiber and Dmoch 1994; BIBEN, M. 1983. Comparative ontogeny of social behaviour in Wayneetal. 1987). Fundamentalnumberis76.Xchromosomeis threeSouthAmericancanids,themanedwolf,crab-eatingfox alargemetacentricandYisasmallsubacrocentric(Chiarelli1975; andbushdog:implicationsforsociality.AnimalBehaviour31: Schreiberand Dmoch1994). 814–826. BRADY,C.A. 1981. Thevocalrepertoiresofthebushdog(Speo- CONSERVATION STATUS. Speothos venaticus is consid- thosvenaticus),crab-eatingfox(Cerdocyonthous)andmaned ered vulnerable by the IUCN and appears in CITES, Appendix I. wolf (Chrysocyon brachyurus). Animal Behaviour 29:649– Studiesofitsdistributionandabundanceareconsideredapriority 669. by theIUCNCanidActionPlan(Sillero-Zubirietal.2004). CABOT, J., P. SERRANO, C. IBAN˜EZ, AND F. BRAZA. 1986. Lista preliminardeavesymamiferosdelareserva‘‘EstacionBiol- REMARKS.Speothosmeanscave wolf,referringtolocation ogicadelBeni.’’EcologiaenBolivia8:37–44. of 1st fossil specimens; living animals do not commonly frequent caves (Simpson 1980). The species name venaticus means hunter CABRERA, A. 1957. Catalogo de los mamiferos de America del Sur.RevistadelMuseoArgentinodeCienciasNaturales‘‘Ber- (Ihering 1968). Vernacular names include cachorro do mato vina- nardinoRivadavia’’4:1–307. gre,janauı´,andjanauı´ra(Brazil);zorrovinagre,perrovinagre,and perrodemonte(Bolivia,Ecuador,andVenezuela);perritovenadero CABRERA, A., AND J. YEPES. 1940. Mamı´feros sudamericanos (Colombia); perro de la selva, perro de agua, and guanfando (Ec- (vida,costumbresydescripcion).HistoriaNaturalEdiar,Com- uador);andjaguayvyguy(Paraguay). paniaArgentinade Editores,BuenosAires,Argentina. SpeothoswasoriginallyplacedinthesubfamilySimocyoninae CALOURO,A.M. 1999. Riquezademamiferosdegrandeemedio with Cuon and Lycaon based on the unicuspid lower carnassial porte do Parque Nacional da Serra do Divisor (Acre,Brasil). (Simpson 1945; Stains 1975). However, this character evolvedin- RevistaBrasileirade Zoologia16(supplement2):195–213. dependentlyinothercarnivoregroupsandconvergenceispossible CARVALHO,C.T. 1980. Mamı´ferosdosparquesereservasdeSaõ forCuon,Lycaon,andSpeothos.Morphologicalcharacterssuggest Paulo.SilviculturaemSaõ Paulo,13/14:49–72. that Speothos is a sister taxon to Atelocynus (Berta 1988; Lyras CHIARELLI, A. B. 1975. The chromosomes of the Canidae. Pp. and van der Geer 2003; Tedford etal. 1995). Based on molecular 40–53 inThewildcanids:theirsystematics,behavioralecol- characters,SpeothosformsamonophyleticcladewithChrysocyon ogy and evolution (M. W. Fox, ed.). Van Nostrand Reinhold (Wayneetal.1997).TheChrysocyon–Speothoscladesupportsmul- Company,NewYork. tiplecanidinvasionsofSouthAmericafromNorthAmerica(Wayne CIMARDI,A.V. 1996. Mamı´ferosdeSantaCatarina.Fundaçaõdo et al. 1997). Phylogenetic placement of Speothos, based on com- MeioAmbiente-FATMA,Floriano´polis,Brazil. bined analyses of 3 genes (cytochrome b, cytochrome c oxidase COIMBRA-FILHO, A. F. 1972. Mamı´feros ameaçados de extinçaõ subunit I, and cytochrome c oxidase subunit II), behavioral, de- no Brasil. Pp. 13–98 in Espećies da fauna brasileira amea- velopmental,ecological,andmorphologicalcharacterssupportsthe çadasdeextinçaõ(AcademiaBrasileiradeCieˆncias,ed.).Ac- monophyletic clade of Speothos and Chrysocyon (Zrzavy and Ri- ademiaBrasileirade Ciencias,ed., Riode Janeiro,Brazil. cańkova´ 2004). COLLIER, C., AND S. EMERSON. 1973. Hand-raising bush dogs B. Beisiegel thanks Drs. C. Ades and M. de Vivo, M. Sears, SpeothosvenaticusattheLosAngelesZoo.InternationalZoo P.Monticelli,andR.S.Tokumaruforinstitutionalorscientificsup- Yearbook13:139–140. port, E´. Cavallete and M. Miretzki for helping with pictures, and DAFONSECA,G.A.B.,ANDK.H.REDFORD. 1984. Themammals FAPESPpostdoctoralfellowship#00/14591-0.G.Zuercherthanks ofIBGE’sEcologicalReserve,Brasilia,andananalysisofthe Drs.P.Gipson,A.Yanosky,R.Klemm,andR.Owen,andL.Aqui- roleofgalleryforestsinincreasingdiversity.RevistaBrasileira no, R. Farinã, C. Mercolli, N. Cardozo, O. Carrillo, M. Zuercher, de Biologia44:517–523. D.Koch,andT.Livingstonforsupportandassistance.Theauthors D’ALESSANDRO, A., R. RAUSCH, C. CUELLO, AND N. ARITZABAL. thank D. Kleiman, R. Wallace, and an anonymous reviewer for 1979. 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