Speothos venaticus

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Speothos venaticus
Author(s): Beatriz de Mello Beisiegel and Gerald L. Zuercher
Source: Mammalian Species, :1-6.
Published By: American Society of Mammalogists
DOI: http://dx.doi.org/10.1644/783.1
URL: http://www.bioone.org/doi/full/10.1644/783.1
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M
AMMALIAN
S
PECIES
No. 783, pp. 1–6, 3 figs.
Speothos venaticus.
By Beatriz de Mello Beisiegel and Gerald L. Zuercher
Published 20 December 2005 by the American Society of Mammalogists
F
IG
. 1. A captive adult male
Speothos venaticus
at Parque
Estoril, Sa˜o Bernardo do Campo, SP, Brazil, 1999. Photograph by
B. M. Beisiegel.
Speothos
Lund, 1839
Speothos
Lund, 1839:224. Type species
Speothos pacivorus
Lund,
1839, by monotypy.
Cynogale
Lund, 1842:203. Type species
Cynogale venatica
Lund,
1842, by monotypy; preoccupied by
Cynogale
Gray, 1837.
Icticyon
Lund, 1842(1843):80. Replacement name for
Cynogale
Lund, 1842.
Abathmodon
Lund, 1843:74. Type species not given.
Cynalicus
Gray, 1846:293. Type species
Cynalicus melanogaster
Gray, 1846, by monotypy.
Cynalius
Gray, 1847:18. Incorrect subsequent spelling of
Cynali-
cus
Gray, 1846.
Melictis
Schinz, 1848:177. Type species
Melictis beskii
Schinz,
1848, by monotypy.
Melictes
Gray, 1868:498. Incorrect subsequent spelling of
Melictis
Schinz, 1848.
Cynalycus
Gray, 1869:183. Incorrect subsequent spelling of
Cyn-
alicus
Gray, 1846.
CONTEXT AND CONTENT. Order Carnivora, suborder
Caniformia, family Canidae, subfamily Caninae (Stains 1984). The
genus
Speothos
is monotypic.
Speothos
was previously placed in
the subfamily Simocyoninae (Simpson 1945) with
Cuon
and
Ly-
caon
.
Speothos venaticus
(Lund, 1842)
Bush Dog
Cynogale venatica
Lund, 1842:203. Type locality ‘‘Lagoa Santa,
Minas Gerais, Brazil.’’
Icticyon venaticus
: Lund, 1842(1843):80. Name combination.
Cynalicus melanogaster
Gray, 1846:293. Type locality ‘‘Brazils.’’
Melictis baskii
Schinz, 1848:177. Type locality ‘‘Nova Friburgo,
Estado do Rio de Janeiro, Brazil.’’
Speothos venaticus
: Thomas, 1903:236. First use of current name
combination.
Speothos wingei
Ihering, 1911:222. Type locality ‘‘Estado de Santa
Catarina, Coloˆnia Hansa, Brazil.’’
Icticyon panamensis
Goldman, 1912:14. Type locality ‘‘Cerro
Pierre, Panama.’’
CONTEXT AND CONTENT. Context as above. Three sub-
species are recognized:
S. v. panamensis
(Goldman, 1912:14); see above.
S. v. venaticus
(Lund, 1842:203); see above;
baskii
(Shinz) and
melanogaster
(Gray) are synonyms.
S. v. wingei
Ihering 1911:222; see above.
DIAGNOSIS.
Atelocynus microtis
and
Speothos venaticus
overlap geographically in the Amazonian regions of Brazil, Colom-
bia, Ecuador, and Peru.
S. venaticus
is distinguished from
A. mi-
crotis
by absence of M3 and absence of metaconid and entoconid
on M1 and M2.
S. venaticus
has interdigital webbing and a car-
nassial 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.
microtis
is fox-like (Vieira 1946).
GENERAL CHARACTERS.
Speothos venaticus
(Fig. 1) is
a small canid (body mass, 4–7 kg) with short legs and tail, inter-
digital membranes, elongated body, and small rounded ears (Vieira
1946). Color ranges from uniform dark brown to yellowish red, with
lighter color on neck, nape of neck, and ears (Sheldon 1992) and
darker color on limbs, tail, and ventrum (Coimbra-Filho 1972).
Young animals are uniformly black (Darling and Whitehead 1991).
Bush dogs lack facial markings. Skull is short (Fig. 2; Vieira 1946).
S. v. panamensis
is smaller than
S. v. venaticus
and
S. v. wingei
,
which are similar in size (Linares 1967).
S. v. panamensis
and
S.
v. wingei
are light-colored compared with
S. v. venaticus
(Linares
1967).
Measurements (ranges, in mm) of
S. v. wingei
are: length of
head and body, 730–740 (
n
5
3); length of tail, 110–150 (
n
5
3);
total length of skull, 132–133 (
n
5
2); palatal length, 61–63 (
n
5
2); zygomatic width, 80–81 (
n
5
2); braincase width, 46–47 (
n
5
2); interorbital width, 24 (
n
5
2); length of mandible, 102–102.5
(
n
5
2—Vieira 1946). Measurements (in mm) of 1 adult male
S.
v. panamensis
are length of head and body, 730; length of tail,
105; length of hind foot, 110; length of ear, 35; length of skull,
120; condylobasal length, 124; palatal length, 61.5; basal length,
116.5; nasal length, 29.5; interorbital width, 31.6; supraorbital
width, 39.8; postorbital width, 26.5; zygomatic width, 75.8; length
of mandible, 91 (Linares 1967).
DISTRIBUTION.
Speothos venaticus
occurs from Panama´
to southern Brazil, Paraguay, and northern Argentina, extending
west to Bolivia, Peru, and Ecuador (Fig. 3; Cabot et al. 1986; Ca-
brera and Yepes 1940; Linares 1967; Yahnke et al. 1998).
S. ven-
aticus
is widely distributed in northern South America (Eisenberg
1989), with fewer reports from southeastern (Sa˜o Miguel Arcanjo–
Beisiegel 1999; Sa˜o Joa˜ o da Boa Vista and Lorena—Carvalho 1980)
and southern Brazil (Cimardi 1996; Ihering 1911).
S. v. panamen-
sis
occurs in northwestern South America;
S. v. venaticus
occurs
in the Amazon River basin, central Brazil, eastern Bolivia, northern
Paraguay, northeastern Peru, eastern Ecuador and the Guyanas; and
S. v. wingei
occurs in southeastern Brazil (Linares 1967). Bush
dogs occur from low elevations (Eisenberg 1989) to 1,900 m alti-
tude (R. Wallace, pers. comm.).
FOSSIL RECORD.
Speothos
originated from canids that
evolved in the Brazilian highlands (Berta 1984; Langguth 1975).
Speothos pacivorus
(extinct species) and
Speothos venaticus
occur
in the late Pleistocene (300,000 years ago)—Recent deposits from
Lagoa Santa Caves, Brazil (Berta 1984, 1987).
FORM AND FUNCTION. Bush dogs are a highly special-
ized lineage of South American canids that evolved for dwelling in
forests (Langguth 1975). They have partially webbed feet (Macdon-
ald 1996) adapted for walking on soft soil near streams. Bush dog
tracks and stride in the field are as follows: forefeet, 5.7 by 5.4
cm; hind feet, 5.0 by 4.4 cm; stride, 28 cm (Villalba and Yanosky
2000). Bush dog dentition, specifically the reduced number of mo-
lars, indicates a highly carnivorous diet (Flower 1880). Dental for-

2 MAMMALIAN SPECIES 783—
Speothos venaticus
F
IG
. 2. Dorsal, ventral, and lateral views of cranium and lat-
eral view of mandible of
Speothos venaticus
(specimen 19743 from
Museu de Zoologia da Universidade de Sa˜ o Paulo, Brazil). Greatest
length of skull is 130 mm.
F
IG
. 3. Geographic distribution of
Speothos venaticus
. Sub-
species are 1,
S. v. panamensis
;2,
S. v. venaticus
;3,
S. v. wingei
.
Subspecies boundaries follow Cabrera (1957), Carvalho 1980, Iher-
ing (1911), Linares (1967), and Massoia et al. (1987). Circles (open
and solid) show sites of occurrence for which precise locations are
reported in the literature. Sources are Barnett et al. (2001), Beisie-
gel (1999), Calouro (1999), Carvalho (1980), Cimardi (1996), Dal-
ponte (1995), Eisenberg (1989), Massoia et al. (1987), Peres (1991),
Redford and Eisenberg (1992), Vieira (1946), Wallace et al. (2002).
Distribution map modified from Eisenberg (1989), Eisenberg and
Redford (1999), Emmons (1997), Redford and Eisenberg (1992),
and Zuercher et al. (2004).
mula is i 3/3, c 1/1, p 4/4, m 1/2, total 38 (Vieira 1946). Lower
carnassial differs from canid pattern: inner cusp of talonid is miss-
ing, resulting in this part of the tooth forming a subsidiary blade
and not a basin. M1 is modified to form a basin into which fits the
single hypoconid cusp of lower carnassial.
Bush dogs have 4 paired mammae (over the ribs ca. 5 cm
behind elbows, on abdomen, opposite knee joints, 4.9 cm in front
of vulva—Flower 1880). Stomach has a subglobular cardiac portion
and a narrower pyloric portion, separated by a constriction. Diges-
tive canal has a reduced caecum (ca. 3.2 cm—Berta 1984, 1988).
Anal glands are large (2.3 cm long and 1.8 cm wide), oval, and
open with a single orifice at lateral margin of anal aperture (Flower
1880).
Speothos venaticus
has a massive frontal pole of brain (Lyras
and van der Geer 2003). A complete sulcus does not separate the
upper curve of the gyrus immediately surrounding the Sylvian fis-
sure on left side from the sulcus above it (Flower 1880).
ONTOGENY AND REPRODUCTION. Captive bush dogs
do not reproduce seasonally (Porton et al. 1987). Aseasonality of
bush dog reproduction in the wild has been largely confirmed by
survey respondents from 7 range countries (DeMatteo 2004). Births
typically occur during the rainy season in Suriname (Husson 1978).
Estrus in captivity lasts 1–12 days (Porton et al. 1987). Two
primiparous females had interestrous intervals of 15–44 days. One
female had 4 and the other 7 estrous periods in 1 year. Earliest
age at conception was 10 months (Porton et al. 1987) and earliest
age of reproduction in males was 12 months (Bekoff et al. 1981).
Bush dogs have copulatory lock, but not back-to-back posture (Dru¨-
wa 1982, 1983; Kleiman 1972). In captive bush dogs, copulatory
locks occurred 1–3 times per day during estrus (DeMatteo 2004;
Dru¨ wa 1983; Kitchener 1971; Kleiman 1968; Porton et al. 1987).
Pseudopregnancy occurred in captive bush dogs after an infertile
mating or after an ovulation without mating (DeMatteo 2004).
In captivity, mean (
1
SE,
in days) interestrous interval is 238
days
6
39.6 (range, 179–301;
n
5
11), but may be shorter if the
litter is lost (Dmoch 1995; Porton et al. 1987). Gestation averages
67 days (Porton et al. 1987), ranging from 65 to 83 days (Moehlman
1986). Bush dog litters typically have 3–6 pups, although litters of
at least 10 occur (Dmoch 1995). Neonatal mass ranges from 125
to 190 g. Sex ratio at birth (males : females) is 1:1.4 (Moehlman
1986). Eyes of young open in 14–19 days (Collier and Emerson
1973; Darling and Whitehead 1991). Pups 1st emerge from den at
2.2 weeks and weaning occurs at 4 weeks (Bekoff et al. 1981).
A litter of 6 bush dogs (3 males and 3 females) was born in
1971 at the Los Angeles Zoo (Collier and Emerson 1973). One
female was dead when the pups were retrieved from the mother
and 1 died of bacterial septicemia 10 days later. Three males and
1 female were successfully hand raised. These surviving pups

MAMMALIAN SPECIES 3783—
Speothos venaticus
weighed 170 g at birth, lost weight during the 1st week, but reg-
ularly gained weight afterward, weighing 476–546 g at 40 days of
age.
Females raised by 1 or both parents, alone or with their sisters,
do not cycle, but do so when paired with a male (Porton et al.
1987). Presence of a male was not required for ovulation but short-
ened interestrous intervals and increased frequency of estrous cy-
cles (DeMatteo 2004).
ECOLOGY. Bush dogs are extremely rare over most of their
distribution, but are common at some sites in Suriname (Husson
1978).
S. venaticus
typically occurs in lowland habitats including
gallery forests, forest edge, wet savannas, and riparian areas (Aqui-
no and Puertas 1997; Defler 1986; Eisenberg 1989; Emmons 1997;
Strahl et al. 1992; Zuercher et al. 2005). Bush dogs also occur in
open habitats such as cerrado (savanna—Silveira et al. 1998;
Zuercher and Villalba 2002), ranchlands (T. Oliveira, pers. comm;
L. Silveira, pers. comm.), and altered habitats (da Fonseca and
Redford 1984). Home range estimates are 3.8–10.0 km
2
(minimum
convex polygon—Beisiegel 1999). Bush dogs have semi-aquatic
habits (Langguth 1975), and most sightings occurred near rivers,
streams, and watercourses (Aquino and Puertas 1997; Barnett et
al. 2001; Deutsch 1983; Linares 1967; Peres 1991; Wallace et al.
2002). A female swam across the Negro River with 2 pups (Coim-
bra-Filho 1972), and a pair swam in the Caura River in Venezuela
(Strahl et al. 1992). Bush dogs pursue and kill pacas (
Agouti paca
)
in water (Strahl et al. 1992; Tate 1931). At Emas National Park, a
cerrado (savanna) area of central Brazil, 3 of 9 sightings occurred
at sites within 200 m of water; the other 6 were from 2,600 to 5,700
m from water (Silveira et al. 1998). At the Mbaracayu´ Reserve in
eastern Paraguay, most bush dog feces, tracks, and sightings
(
.
70%) were detected
,
1,000 m from water (Zuercher 2001). A
captive female spent a great deal of time in a water pond where
she dived and swam with ease (Bates 1944).
Bush dogs use dens dug by large armadillos (Coimbra-Filho
1972) and sometimes tamanduas (Sanderson 1949). Other den sites
are fallen tree trunks (Aquino and Puertas 1997) and rock shelters
(Linares 1967). Den sites occur with scattered dry and fresh feces
(Aquino and Puertas 1997) or free of fecal material (Linares 1967).
Agouti (
Dasyprocta
), capybara (
Hydrochoerus hydrochaeris
),
and paca are the main prey of bush dogs (Aquino and Puertas 1997;
Cabrera and Yepes 1940; Deutsch 1983; Peres 1991; Silveira et
al. 1998; Strahl et al. 1992; Tate 1931; Zuercher et al. 2005). One
group of 6 bush dogs hunted a tapir (
Tapirus terrestris
) that weighed
250 kg (Wallace et al. 2002). Near Serra das Araras, State of Mato
Grosso, Brazil, bush dogs also hunt armadillos (Dasypodidae), col-
lared peccaries (
Tayassu tajacu
), and deer (
Mazama
—Beisiegel
1999). Indigenous groups from Ecuador and Peru report that
S.
venaticus
successfully hunts collared peccaries and tapirs (Descola
1994). Bush dogs also hunt rheas (
Rhea americana
—Santos 1945).
In Paraguay, bush dog feces contained remains of agouti, paca,
tinamou (Tinamidae), an unidentified snake, several unidentified
small rodents, and seeds from
Cecropia
(Zuercher et al. 2005). In
the Peruvian Amazon, a single bush dog’s feces contained hair of
agouti (
Dasyprocta fuliginosa
), coati (
Nasua nasua
), smaller ro-
dents (
Myoprocta pratti
or
Proechimys
), and feathers similar to
those of tinamous (Aquino and Puertas 1997). Captive bush dogs
readily accepted 9-banded armadillos (
Dasypus novemcinctus
),
opossums (
Didelphis
), and rabbits (
Sylvilagus brasiliensis
—Van
Humbeck and Perez 1998). Bush dogs individually hunt small prey
(Silveira et al. 1998). In Paraguay,
S. venaticus
and other carni-
vores have minimal dietary overlap (Zuercher 2001).
Only indirect evidence of predation exists on bush dogs. In
Peruvian Amazonia, the carcass of an adult bush dog was surround-
ed by tracks of jaguar (
Panthera onca
) or puma (
Puma concolor
—
Aquino and Puertas 1997).
Speothos venaticus
is hunted for human
consumption in some areas of Amazonia (Calouro 1999).
Speothos venaticus
hosts
Echinococcus vogeli
, a cestode with
pacas as the intermediate host (D’Alessandro et al. 1979; Rausch
and Bernstein 1972). In Venezuela, bush dogs host
Lagochilas-
caris
, an ascaridian nematode with rodents, such as
Dasyprocta
leporina
, as intermediate hosts (Volca´n and Medrano 1991). Cap-
tive bush dogs have succumbed to canine parvovirus (Janssen et
al. 1982; Montali and Kelly 1989). Captive animals may have suc-
cumbed to a vaccine-induced canine distemper virus (McInnes et
al. 1982). Other pathogens from captive bush dogs include bacteria
(
Escherichia coli
,
Klebsiella
,
Proteus vulgaris
,
Shigella
,
Staphy-
lococcus aureus
, and
S. epidermis
), fungi (
Candida
—Van Hum-
beck and Perez 1998), and protozoans (
Giardia
).
HUSBANDRY. One captive bush dog lived for 10 years and
4 months (Jones 1982). At the Los Angeles Zoo, a suckling re-
sponse was difficult to elicit from pups during the first 8 days. Pups
were fed a formula of 1 part Esbilac and 2 parts boiled distilled
water; concentration was increased until full Esbilac by day 19,
and bottle feeding continued until day 39. From day 7 to day 25,
a vitamin supplement was added and, from day 40 on, the young
were given Esbilac in a shallow dish and puppy chow in small
fragments (Collier and Emerson 1973). At Twycross Zoo, United
Kingdom, bush dogs are fed raw meat, live rats, and bone meal
with vitamin supplements. A more varied diet of commercial dog
food, fruit, and eggs caused vomiting, dehydration, and loss of ap-
petite, mainly due to the commercial dog food (Darling and White-
head 1991). Other zoos feed only carcasses and live animals (Mac-
donald 1996).
Bush dogs can be immobilized with 10 mg/kg Telazol supple-
mented, if necessary, with 10 mg/kg ketamine, or with 20 mg/kg
ketamine plus 0.2 mg/kg acepromazine (Kreeger et al. 2002). Bush
dogs at the St. Louis Zoo were immobilized with 50 mcg/kg med-
etomidine combined with 5–6 mg/kg ketamine (DeMatteo and Ko-
channy 2004).
BEHAVIOR. A summary of behavior of
S. venaticus
is avail-
able (Beisiegel and Ades 2002).
S. venaticus
is diurnal/crepuscular
(Dalponte 1995; Defler 1986; Deutsch 1983; Linares 1967; Peres
1991; Silveira et al. 1998; Strahl et al. 1992; Zuercher and Villalba
2002), although individuals have been observed at night between
0100 and 0400 h (Wallace et al. 2002). In captivity, bush dogs are
active during the day and retire to dens at night (Kleiman 1972).
Play behavior has been described for young
S. venaticus
(Biben
1983). Seventy-one percent of object play by a captive group oc-
curred in water (Macdonald 1996).
The basic social unit of
S. venaticus
is a monogamous pair
and extended family. In captivity, bush dogs form pair bonds (Dru-
wa 1983; Macdonald 1996; Porton et al. 1987) and parents are
dominant over all group members (Macdonald 1996). Only the al-
pha female reproduces and other group members display allopar-
ental behaviors, such as guarding, carrying, and licking pups (Mac-
donald 1996). Fathers help during parturition by grooming their
mate’s anogenital region and removing the afterbirth (Porton 1983).
Group living occurs in the field (Aquino and Puertas 1997; Barnett
et al. 2001; Defler 1986; Strahl et al. 1992). In the Peruvian Am-
azon, 2 adult bush dogs explored an area of ca. 60 m diameter
around a pile of branches. Fifteen minutes later, a young bush dog,
followed by a pup, emerged from the pile (Aquino and Puertas
1997). Although most sightings of bush dogs are of groups, sight-
ings in the cerrado of Emas National Park were mostly of single
individuals (Silveira et al. 1998).
Members of bush dog groups hunt cooperatively (Aquino and
Puertas 1997; Barnett et al. 2001; Dalponte 1995; Defler 1986;
Linares 1967; Peres 1991; Strahl et al. 1992). When a bush dog
group is hunting a paca, part of the group chases it on land and
part waits for the paca in the water (Cabrera and Yepes 1940).
Communal hunting occurs in captivity (Macdonald 1996). Captive
pups are fearful when they individually encounter prey, but attack
prey when near their parents. During ingestion of large prey, par-
ents position themselves at the extremes of the carcass, thus facil-
itating dismemberment of prey by pups (Biben 1982b).
Bush dogs have 10 distinct vocalizations: short whines, ex-
tended whines, repetitive whines, pulsed vocalizations, short
screams, long screams, barks, growls, infant whines, and infant
grunts (Brady 1981; Villa 2001). Frequency ranges for several vo-
calizations are short whines, 450–1,790 Hz; extended whines, 540–
1,660 Hz; repetitive whines, 520–1,570 Hz; pulsed vocalizations,
1,140–1,590 Hz; short screams, 800–1,720 Hz; long screams, 690–
1,550 Hz; barks, 410–1,720 Hz; growls, 450–1,590 Hz; infant
whines, 670–880 Hz; infant grunts, 820–1,590 Hz (Villa 2001).
Vocalizations facilitate communication among group members in an
environment that obstructs visual contact (Brady 1981; Villa 2001).
Recorded vocalizations and deposition of urine have attracted and
elicited vocal responses from wild bush dogs (DeMatteo et al.
2004).
Female bush dogs have a specific urinary posture (the raised-
hindquarters posture, or handstand) in which they climb a vertical

4 MAMMALIAN SPECIES 783—
Speothos venaticus
surface backward with their hindlegs and urinate while standing on
their forelegs. They then slide their ano-genital region down the
vertical surface (Biben 1982a; Kleiman 1966). Male bush dogs
slightly extrude their penis and move laterally to create a spray
when they urinate (Kleiman 1972).
GENETICS.
Speothos venaticus
has 2n
5
74 chromosomes
with 36 acrocentric chromosomes (Schreiber and Dmoch 1994;
Wayne et al. 1987). Fundamental number is 76. X chromosome is
a large metacentric and Y is a small subacrocentric (Chiarelli 1975;
Schreiber and Dmoch 1994).
CONSERVATION STATUS.
Speothos venaticus
is consid-
ered vulnerable by the IUCN and appears in CITES, Appendix I.
Studies of its distribution and abundance are considered a priority
by the IUCN Canid Action Plan (Sillero-Zubiri et al. 2004).
REMARKS.
Speothos
means cave wolf, referring to location
of 1st fossil specimens; living animals do not commonly frequent
caves (Simpson 1980). The species name
venaticus
means hunter
(Ihering 1968). Vernacular names include cachorro do mato vina-
gre, janauı´, and janauı´ra (Brazil); zorro vinagre, perro vinagre, and
perro de monte (Bolivia, Ecuador, and Venezuela); perrito venadero
(Colombia); perro de la selva, perro de agua, and guanfando (Ec-
uador); and jagua yvyguy (Paraguay).
Speothos
was originally placed in the subfamily Simocyoninae
with
Cuon
and
Lycaon
based on the unicuspid lower carnassial
(Simpson 1945; Stains 1975). However, this character evolved in-
dependently in other carnivore groups and convergence is possible
for
Cuon
,
Lycaon
, and
Speothos
. Morphological characters suggest
that
Speothos
is a sister taxon to
Atelocynus
(Berta 1988; Lyras
and van der Geer 2003; Tedford et al. 1995). Based on molecular
characters,
Speothos
forms a monophyletic clade with
Chrysocyon
(Wayne et al. 1997). The
Chrysocyon
–
Speothos
clade supports mul-
tiple canid invasions of South America from North America (Wayne
et al. 1997). Phylogenetic placement of
Speothos
, based on com-
bined analyses of 3 genes (cytochrome
b,
cytochrome
c
oxidase
subunit I, and cytochrome
c
oxidase subunit II), behavioral, de-
velopmental, ecological, and morphological characters supports the
monophyletic clade of
Speothos
and
Chrysocyon
(Zrzavy and Ri-
ca´nkova´ 2004).
B. Beisiegel thanks Drs. C. Ades and M. de Vivo, M. Sears,
P. Monticelli, and R. S. Tokumaru for institutional or scientific sup-
port, . Cavallete and M. Miretzki for helping with pictures, and
´
E
FAPESP postdoctoral fellowship #00/14591-0. G. Zuercher thanks
Drs. P. Gipson, A. Yanosky, R. Klemm, and R. Owen, and L. Aqui-
no, R. Farin˜ a, C. Mercolli, N. Cardozo, O. Carrillo, M. Zuercher,
D. Koch, and T. Livingston for support and assistance. The authors
thank D. Kleiman, R. Wallace, and an anonymous reviewer for
improvements in this manuscript.
LITERATURE CITED
A
QUINO
, R.,
AND
P. P
UERTAS
. 1997. Observations of
Speothos
venaticus
(Canidae: Carnivora) in its natural habitat in Peru-
vian Amazonia. Zeitschrift fu¨r Sa¨ugtierkunde 62:117–118.
B
ARNETT
, A., R. S
HAPLEY
,
AND
M. E
NGSTROM
. 2001. Records of
the bush dog,
Speothos venaticus
(Lund, 1842) from Guyana.
Mammalia 65:232–237.
B
ATES
, M. 1944. Notes on a captive
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Associate editors of this account were K
RISTOFER
H
ELGEN
,P
AMELA
O
WEN
, and B
ETTY
M
C
G
UIRE
. Editor was V
IRGINIA
H
AYSSEN
.
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EISIEGEL
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