Ecology and Social Organization
of the Maned Wolf
(Chrysocyon brachyurus)
JAMES M. DIETZ
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 392
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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 3 9 2
Ecology and Social Organization
of the Maned Wolf
(Chrysocyon brachyurus)
James M. Dietz
SMITHSONIAN INSTITUTION PRESS
City of Washington
1984
A B S T R A C T
Dietz, James M. Ecology and Social Organization of the Maned Wolf
(Chrysocyon brachyurus). Smithsonian Contributions to Zoology, number 392, 51
pages, 24 figures, 21 tables, 1984.?Although the maned wolf is prized by
zoological gardens and is considered endangered throughout its range, little
is known of the ecology and social behavior of the species. This monograph
presents the results of a 2-year field study conducted in the Serra de Canastra
National Park, Brazil, and of a 15-month behavioral study of maned wolves
captive at the Conservation and Research Center of the National Zoological
Park, Front Royal, Virginia, USA.
Results of the field study are based on radiotelemetric and visual observa-
tion of 11 individuals living in four discrete home ranges. Wolves are largely
nocturnal, with the majority of circadian activity occurring from dusk to
midnight. Males are active more often than females, and pair-members are
usually found in close association only during the breeding season. Solatium
lycocarpum, small mammals and miscellaneous fruits are the most important
dietary components, as determined by examination of feces. Consumption of
these food items is consistent with seasonal availability indicating that maned
wolves are trophic generalists. Males and females are monogamous and share
territories averaging 27 square kilometers. Vacancies created by the deaths
of resident wolves are filled presumably by unlanded loners. The greatest
source of recorded mortality was due to reprisals by ranchers for wolf
depredation on domestic fowl. Six of eight wolves tested positive for cystin-
una, a metabolic disease occasionally fatal in captivity.
Behavioral observations on eight captive pairs of maned wolves were
conducted during two breeding seasons. Males scent mark, vocalize, and
initiate activity more often than do females, suggesting a more prominent
role by males in territorial defense. Anestrus is characterized by mutual
avoidance and minimum levels of scent marking and vocalization. Proestrus
is a period of social approximation culminating in estrus. Distance-decreasing
behavior drops to a minimum during gestation. However, the continued high
level of mutual use of space suggests that females regulate the extent to which
males participate in parental care.
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is
recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral
Montastrea cavernosa (Linnaeus).
Library of Congress Cataloging in Publication Data
Dietz, James M.
Ecology and social organizaton of the maned wolf (Chrysocyon brachyurus).
(Smithsonian contributions to zoology ; no. 392)
Bibliography: p.
Supt. of Docs, no.: SI 1.27:392
I. Maned wolf?Ecology. 2. Maned wolf?Behavior. 3. Social behavior in animals. 4.
Mammals?Ecology. 5. Mammals?Behavior. I. Title. II. Series
QL1.S54 no. 392 [QL737.C22] 591s 83-600292 [599.74'442]
Contents
Page
Introduction 1
Acknowledgments 1
The Study Area 3
Literature Review 4
The Field Study 6
Methods 6
Capture Procedure 6
Handling Procedure 8
Location by Radiotelemetry 8
Determination of Food Habits 9
Scat Collection 9
Scat Analysis 9
Census of Available Food Items 10
Results 10
Physical Characteristics of Captured Maned Wolves 10
Pelage, Weights, and Measurements 10
Hematology and Blood Chemistry 10
Description of Home Ranges in the Study Area 11
West Range 11
East Range 11
Taperao Range 11
Other Ranges 11
Activity Patterns 13
Daily and Seasonal Variation 13
Observations on Individual Activity 13
Social Organization and Communication 14
Relationships of Individuals within Pairs 14
Vocalizations 17
Deposition of Feces and Urine 17
Relationships between Adjacent Pairs of Wolves 19
Loners or Nomadic Maned Wolves 20
Feeding Ecology 20
Seasonal Availability of Trophic Resources 20
Food Habits of Maned Wolves in the Study Area 21
Exploitation of Domestic Stock by Maned Wolves 23
Reproduction 23
Denning 23
Parturition and Litter Size 24
Parental Care of Pups 24
iii
IV SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Parasites, Diseases, and Mortality 25
Ectoparasites 25
Endoparasites 25
Diseases 26
Mortality in the Serra da Canastra 26
Relations of Maned Wolves with Humans 26
Effects of Human Disturbance on Maned Wolves 26
Attitudes of Area Residents toward Maned Wolves 26
Folklore 27
Study of Captive Maned Wolves 27
Methods 27
Facilities and Observation Schedule 27
Operational Definitions of Reproductive Seasons 28
Behavioral Observations 29
Results 30
Synchrony of Activity 30
Individual Distance 30
Distance-increasing and Distance-decreasing Behavior 30
Mount 30
Present 30
Investigation of Anogenital Region 30
Investigation of Muzzle 31
Tail-up, Tail-out 31
Yawn 31
Avoidance 31
Gape 31
Full Piloerection 31
Bite 32
Initiative Behavior 33
Scent Marking 33
Vocalization 35
Use of Space 36
Precedence in Feeding 36
Discussion 36
Evolution of Sociality in a Grassland Habitat 36
The Pair Bond in Maned Wolves 37
Conservation and Management 38
Maned Wolves in the Wild 39
Maned Wolves in Captivity 40
Appendix: Tables 1-21 41
Literature Cited 48
Ecology and Social Organization
of the Maned Wolf
(Chrysocyon brachyurus)
James M. Dietz
Introduction
Thousands of years before the advent of civi-
lization, the highlands of central South America
were covered with grass prairie and scrub forest.
These habitats generally lacked the large and
diverse herds of ungulates characterizing the Af-
rican plains, but they abounded with smaller
vertebrates?particularly rodents of the family
Cricetidae. The maned wolf (Figure 1), largest
and most distinct of the South American canids,
must have hunted in this environment and been
shaped morphologically and socially by it.
Since the arrival of modern man, the Brazilian
Central Highlands have been changing dramati-
cally. Grassland and cerrado forest are being
converted at an unprecedented rate to cattle
pasture and farmland. Because we knew little of
the ecology of free-living maned wolves, it was
impossible to predict the effects of these environ-
mental changes on this species. Thus, the general
objective of the present study is to document the
ecology and social organization of the maned
wolf so that rational decisions can be more effec-
tively made concerning the managment and con-
servation of this endangered species.
This research project consists of two distinct
James Af. Dietz, Conservation and Research Center, National
Zoological Park, Smithsonian Institution, Front Royal, VA 22630.
studies: a 2-year field study of free-living maned
wolves in the Serra da Canastra National Park,
Brazil (July 1978-June 1980), and a 15-month
study of captive maned wolves at the National
Zoological Park's Conservation and Research
Center (CRC) in Front Royal, Virginia (Septem-
ber 1981-December 1982). This monograph re-
ports the methods and results of both studies,
and the subsequent discussion deals largely with
the social organization and conservation of this
endangered species.
The specific objectives of my field study of the
maned wolves of the Serra da Canastra include
quantifying for the relative use of available hab-
itat types, diurnal and seasonal activity patterns,
social organization, seasonal food habits, repro-
duction, and mortality and relations with hu-
mans. The objectives of my research at CRC
were suggested by results obtained in the Serra
da Canastra (Dietz, 1981) and include an exami-
nation of the behavioral mechanisms maintaining
the pair bond in maned wolves, and of the
changes in these parameters that were related to
stages of the reproductive cycle.
ACKNOWLEDGMENTS.?The field study partly
satisfied the requirement for my doctoral degree
at Michigan State University, and it is with plea-
sure that I acknowledge the contributions of the
following individuals to that portion of the proj-
ect: to R.H. Baker, chairman of my advisory
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 1.?Adult maned wolf.
committee, I extend my warmest thanks for par-
ticipation in every aspect of this research; similar
thanks are also due advisory committee members
S.C. Bromley, J.H. Fitch, L. Gysel, R.W. Hill,
and G.A. Petrides.
Coordination of this research project at the
federal level in Brazil was not an easy task, but it
was handled smoothly and efficiently by Dra.
Maria Tereza Padua, Dr. Renato Petty Leal, Jose
Carlos Duarte, Eduardo Kunze Bastos, and the
other competent professionals at Instituto Brasi-
leiro de Desenvolvimento Florestal (IBDF), Bras-
ilia. The late Dr. Ivens P. Franqueira, past direc-
tor of IBDF in Minas Gerais State, invited me to
study maned wolves in the Serra da Canastra.
Without his zeal and enthusiasm for conserva-
tion, that national park would not have been
created. I am also indebted to park director
Oliveiro A. Soares, and his administrative per-
sonnel, work crew, and guards.
Dra. Rosa Vieira, director of the Clinical Lab-
oratory, School of Veterinary Medicine, Federal
University of Minas Gerais, performed the labo-
ratory analyses during the course of this research.
I also thank J.O. Whitaker and N. Wilson for
identification of ectoparasites; D.O. Straney, P.
Myers, and R. Pine for identification of small
mammals; R. Ramalho and J. Duarte for identi-
fication of botanical specimens.
The field study was partially funded by grants
from the following agencies: World Wildlife
Fund/International Union for Conservation of
Nature and Natural Resources, Instituto Brasi-
leiro para Desenvolvimento Florestal/Fundacao
Brasileira para Conservacao de Natureza, New
York Zoological Society, and Michigan State
University.
Finally, I would like to acknowledge the par-
NUMBER 392
ticipation of my wife, Lou Ann, in this study. It
was she who gave up two years of her career to
help me chase maned wolves in the Serra da
Canastra.
My research at the National Zoo's Conserva-
tion and Research Center was made possible by
Devra Kleiman. She deserves special thanks for
allowing me to conduct the behavioral observa-
tions largely absent in my field study. Thanks are
also due Chris Wemmer and his staff for making
the Conservation and Research Center an enjoy-
able and productive place to work. M. Ditton
and the personnel who work with her in manag-
ing and caring for the canid collection contrib-
uted to several aspects of this study. L. Hayek
advised on statistics. D. Kleiman, C. Wemmer
andj. Horn read later drafts of this monograph
and made suggestions for its improvement. My
research at the Conservation and Research Cen-
ter was supported by grants from the Friends of
the National Zoo and from the Smithsonian In-
stitution Scholarly Studies Program (to J. Eisen-
berg et al.). The use of brand names in this report
is for descriptive purposes only and does not
constitute endorsement by the Smithsonian In-
stitution.
The Study Area
"Serra da Canastra," loosely translated, means
"mountain range shaped like a foot-locker," and
is the regional name given to an area of hills and
ridges located 300 km west of Belo Horizonte,
capital of the state of Minas Gerais, Brazil (Figure
2). The hills rise abruptly from surrounding ag-
ricultural land to elevations of nearly 1500 m
and descend into valleys of about 800 m eleva-
tion. The areas of higher elevation are charac-
terized by poorly drained soils supporting sea-
sonal grasslands. Subject to heavy grazing and to
local pasture improvement techniques, these
grasslands are composed of a variety of species
of Gramineae, such as Aristida sp., Tristachya sp.,
and with the exotic Melinis minutiflora. Also pres-
ent are occasional shrubs and small trees such as
Vellozia sp. and Arnica montana. Grassland habitat
comprises perhaps 50 percent of the area of the
Serra da Canastra.
SERRA DA CANASTRA
NATIONAL RARK
FIGURE 2.?Location of the Serra da Canastra National
Park, Minas Gerais State, Brazil, South America.
Approximately 35 percent of the area is com-
posed of cerrado vegetation. This habitat type is
defined by well-spaced, thick-barked trees (e.g.,
Byrsonima sp. and Vellozia sp.), less than 5 m in
height. Trees of the cerrado are typically
drought- and fire-resistant with thick tortuous
stems, limbs, and branches. Understories are
composed of a variety of annual plant species
(e.g., Gramineae, Cyperaceae, and Leguminosae)
whose presence is often influenced by disturb-
ance factors such as annual burning or clearing
of brush to "improve" pasture for cattle grazing.
Many streams and rivers, including the Rio
Sao Francisco, originate in the highlands of the
Serra da Canastra. Along these waterways and at
their headwaters are found narrow strips of sea-
sonal tropical forest. This riparian habitat type
accounts for perhaps 15 percent of the area and
is composed of a wide variety of tree species. In
this habitat type, understories vary from rela-
tively open ones in drier areas to thick tangles of
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
vines, herbaceous and shrub species in more
mesic areas.
The climate of the region is subtropical with
well-defined wet and dry seasons (Figure 3). No-
vember through February are typically the wet-
test months and have a mean temperature of
about 22?C. June through August are the driest
months and have a mean temperature of approx-
imately 18? C. Frosts occur rarely in the Serra
da Canastra. Yearly rainfall varies from about
1300 to 1700 mm. Winds average 18.9 km/hr
and are predominantly from the East (Padua,
1978).
A final characteristic common to all regions of
the Serra da Canastra is the ubiquitous presence
of humans and their activities. Accessible parts
of each range have been annually burned and
heavily grazed by cattle during the past several
generations. Cowboys tending cattle ride or walk
over much of the area on a daily basis. Tempo-
rary and permanent human dwellings, as well as
dirt roadways, are scattered throughout the area.
Within this hill country (20? South latitude,
46? West longitude) the Serra da Canastra Na-
tional Park is located. The park was created by
Brazilian federal decree in 1972. At the time of
my study, however, the land had not yet been
purchased by the federal government and was
still being used for cattle grazing by local ranch-
ers. Today, the park is 715 km2 in area and is
fenced and regularly patrolled by guards in au-
tomobiles. A dirt road traverses the park from
the city of Sao Roque de Minas on the east to
Sacramento to the west. Upland seasonal grass-
land is the most common habitat type within the
park. The study area for my research consisted
of the eastern half of this park and the areas
adjacent to it to the east and north. My wife and
I lived in the park from July 1978 through May
1980.
Literature Review
The first detailed description of the maned
wolf and its habits was published by the famed
naturalist Don Felix D'Azara in 1801. He re-
ferred to the maned wolf by its Paraguayan name
Agouara Gouazou (his spelling) which he trans-
lated as "large fox." Simpson (1941) added that
this name originated in the language of the Tupi-
Guarani Indians and has been shortened to
guard. Rengger (1830) suggested that the name
WET SEASON ORY8EA8ON WET8EA8ON DRYSEA8ON
FIGURE 3.?Definition of wet and dry seasons: temperature (= the mean of daily minimum
temperatures for each calendar month); length of day (= the mean number of minutes of
daylight for the days in each month); rainfall (= total measureable precipitation per month).
NUMBER 392
Aguara-guazu was onomatopoeic for the vocali-
zation of the maned wolf. The animal is now
known throughout Brazil as lobo guard, in Bolivia
as boroche, aguard guazu in Paraguay and Argen-
tina, and lobo de crin in Bolivia and Peru.
The taxonomic relationships within the family
Canidae have been relatively unstable since Gray
(1825) coined the family name in its present
form. Classification of the South American forms
at the generic level has been the most unstable
(see Cabrera, 1931, and Clutton-Brock et al.,
1976, for reviews). The maned wolf has largely
escaped these problems of technical nomencla-
ture of the canids and has not been challenged
as a distinct genus since clarification of its current
scientific name by Osgood in 1919 and 1934.
The maned wolf is monotypic, and no geo-
graphic races are recognized.
The identification of the canid groups that are
the maned wolf's nearest relatives remains an
unsolved problem: Clutton-Brock et al. (1976),
using morphological character analysis, con-
cluded that the maned wolf is weakly related to
the Dusicyon group. Chiarelli (1975) writes that
the karyotype of Chrysocyon is very similar to that
of Canis. However, the diploid number of the
former is 76, while that of the latter is 78 (Newn-
ham and Davidson, 1966). Interbreeding be-
tween these two genera is not likely (Bastos,
1978). Gel electrophoresis has been used to dif-
ferentiate between the serum proteins of various
carnivores including the maned wolf and domes-
tic dog (Canis familiaris) (Durr and Schmitt,
1970), but a relatively low incidence of polymor-
phic loci has generally limited the effectiveness
of this technique in the Canoidea (Seal, 1975).
Pocock (1927), Langguth (1969), and Studer
(1904) have described the external characters
and the morphology of the maned wolf and have
compared these data with those from other South
American canids. Whereas the caeca of most
canids are convoluted, the caecum of the maned
wolf is short and cylindrical (Flower, 1879), as it
is in the bush dog (Speothos venaticus), the crab-
eating fox (Cerdocyon thous), and the short-eared
dog (Atelocynus microtis) (Langguth, 1969). Soko-
lowsky (1927), speculating on the adaptive sig-
nificance of the long legs of maned wolves, sur-
mised that this species is not adapted for swift
running, as stated by Stains (1975), but rather
for efficient vision and travel in tall vegetation.
Since no fossil remains of the maned wolf have
been found outside of the Brazilian Central
Highlands, it has been assumed that the maned
wolf evolved in that region (Langguth, 1975)
either during the beginning of the Pleistocene
Epoch (Simpson, 1980) or well before the Pleis-
tocene (Langguth, 1969). The present range of
the maned wolf includes the following areas: all
geographic regions of Brazil (excluding the Am-
azon Basin, parts of the arid Northeast, and the
Atlantic coast); the provinces of Formosa, Cor-
rientes, and Chaco in Argentina; most of Para-
guay; eastern Bolivia; and the Pampas del Heath
in Peru (Cabrera, 1958; Cabrera and Yepes,
1960; Paiva, 1973; Schaller, 1975; Hofmann et
al., 1975-1976; Schaller and Vasconcelos, 1976;
Dietz, in press; Breyer, in prep.).
Acosta (1972) described hand-rearing a litter
of maned wolf pups. These data were used by
Bekoff and Jamieson (1975) to compare the
maned wolfs physical development with that of
other canids. The growth parameters for pups
were established by Acosta (1972), Encke et al.
(1970), and by Seidel (1972). Information on the
type of enclosure and diet necessary successfully
to maintain maned wolves in captivity has been
summarized by Crandall (1964), C. Carvalho
(1976), and by Brady and Ditton (1979). The
longevity of maned wolves in captivity has been
estimated at from 12 to 15 years (Silveira, 1969).
Female maned wolves are monestrous and may
copulate several times during an estrus of about
five days (Lippert, 1973; Kuhme, 1975). Copu-
latory behavior has been described by Lippert
(1973). Copulation by captive maned wolves has
been observed during the months of October
through February in the Northern Hemisphere
(Encke, 1971; D. Altmann, 1972), and from Au-
gust through October in South America (Acosta,
1972). The birth of from two to five pups, a
normal litter size for this species, follows a ges-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
tation period of about 65 days (Faust and Scherp-
ner, 1968). A record of seven pups in one litter
was recorded in the Sao Paulo Zoo (C. Carvalho,
1976). Parturition has been described by Ham-
merling and Lippert (1975), and the studbook
record for the species has been published by
Roeben (1975) and by Matern (1980).
In addition to mediocre reproductive success,
maned wolves suffer from several parasitic and
disease problems, including giant kidney worm
(Dioctophyma renale) (Giovannoni and Molfi,
1960; Lamina and Black, 1966; Matera et al.,
1968). This parasite is commonly fatal in captive
and probably also in free-living maned wolves.
Silveira (1969) suggested that the Solanaceae
eaten by maned wolves might have an inhibiting
action on this parasite, but the methods used to
test this hypothesis were rightly questioned by C.
Carvalho (1976). Other endoparasites found in
maned wolves are mentioned by Encke et al.
(1970), and by C. Carvalho (1976). Bush (1980)
reviewed the diseases and medical management
of maned wolves in captivity; two recent medical
problems include susceptibility to parvovirus
(Fletcher et al., 1979) and cystinuria (Bovee et
al., 1981).
Maned wolves in captivity lead essentially soli-
tary existences during the reproductive period.
Pairs composed of animals of opposite sexes are
easier to house in the same enclosure than same-
sex animals. Individuals maintain separate pre-
ferred resting sites and do not usually rest in
close association (Encke, 1971; D. Altmann,
1972; Kuhme, 1975). Kleiman (1972) contrasted
the solitary behavior of the maned wolf with the
highly gregarious nature of the bush dog (Speo-
thos venaticus). The vocalizations of maned wolves
have been described by Brady (1981), and the
behavior of pups by Encke et al. (1970) and Biben
(1983).
The wolf feeds mainly on small vertebrates
and on fruits (Azara, 1801; Cabrera and Yepes,
1960; C. Carvalho, 1976). F.W. Miller (1930)
and Dennler de la Tour (1968) describe the stiff-
legged pounce used by maned wolves capturing
small prey. Roosevelt (1925) and Krieg (1948)
write that the wolf often preys on small domestic
stock, but this allegation is refuted by Cabrera
and Yepes (1960). Krieg (1928, 1940, 1948)
refers to the wolf as the "savanna stroller" and
comments on its solitary nature. Dennler de la
Tour (1968) also notes that the wolf is usually
seen alone and hints that individual territories
may be maintained.
The maned wolf is classified by the Interna-
tional Union for the Conservation of Nature and
Natural Resources as "vulnerable" (IUCN Red
Data Book, 1982), and classified by agencies of
the Brazilian government as "endangered" (J.
Carvalho, 1968; Coimbra Filho and Magnanini,
1968). Silveira (1968) estimated that only 1500-
2200 maned wolves remain in 650,000 km2 in
Brazil, and Meritt (1973) details pathways by
which wolves captured in Paraguay and northern
Argentina were exported out of Paraguay in
1971. However, in spite of what appears to be a
rather bleak future for the maned wolf, little
research is currently underway on this species,
and no objective suggestions have been formu-
lated concerning its conservation.
The Field Study
METHODS
Capture Procedure
Political considerations and the likelihood of
breaking the long and apparently fragile limb
bones of captured maned wolves precluded the
use of steel leghold traps for trapping. Interviews
with area residents produced several models of
live-traps successful in capturing wolves. From
these suggestions, I designed and had built seven
wooden live-traps (Figure 4). Each weighed
about 110 kg and was constructed of durable
native hardwoods. Bolts permitted disassembly
for transport. The trap design was an open-
ended box with a falling door released by a
treadle mechanism located at the back of the
trap. For bait, a live chicken (supplied with food
and water) was supported on a platform above
NUMBER 392
97 a*
FIGURE 4.?Wooden live-trap used to capture maned wolves. Trap is baited with a live
chicken.
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
the treadle; the chicken was separated from the
inside of the trap by a wall of light chicken-wire.
All captured wolves perforated the chicken wire
and consumed the chicken.
Initial attempts using random distribution of
traps were unsuccessful. Through interviews, I
established the locations of recent maned wolf
activity and set traps within a few meters of major
trails. Major trails were determined by the quan-
tities of urine, feces, and tracks found there. All
such traps were successful in capturing resident
maned wolves. Traps were placed in pasture, tall
grass, or scrub brush and were not camouflaged
or hidden. When a trap repeatedly captured the
same wolves, that trap was temporarily closed or
was moved to a new area of wolf activity.
Since the objective of trapping was to place
radiotransmitters on all resident maned wolves,
trap placement was determined by signs of wolf
activity and not by an area grid design selected a
priori. The size and topographical irregularity of
the area, and the general lack of roads prevented
examination of all traps daily. Each trap was
assigned to a resident of the study area and was
placed so that he could check that trap from a
distance. Residents were reimbursed for rapidly
reporting a wolf capture.
Handling Procedure
The first capture of a wolf involved the collec-
tion of a urine sample for use in testing for
cystinuria. Since trapped maned wolves urinated
when first approached, I was able to collect urine
samples from eight wolves by using one stick to
distract the wolf and simultaneously inserting a
second stick, with a piece of filter paper attached,
between the legs of the wolf to collect the urine
as it was voided. The urine absorbed onto the
filter paper was analyzed chromatographically by
K. Bovee, School of Veterinary Medicine, Uni-
versity of Pennsylvania.
Immobilization involved an intramuscular in-
jection by means of a syringe recessed into a hole
in the end of a broomstick, using 20 mg Sernylan
(Parke-Davis); 7.5 mg Rompun (Haver-Lock-
hart); 0.75 mg Acepromazine (Haver-Lockhart).
This dose usually rendered the wolf immobile
within 10 minutes and allowed about two hours
of working time before the animal began to
regain consciousness. All immobilized maned
wolves recovered physical control quickly and
apparently completely.
Whole blood was collected from a superficial
vein of a limb and a portion was mixed with
anticoagulant (KEDTA) for serological determi-
nations. Feces from the trap and blood samples
were immediately sent for analysis to the School
of Veterinary Medicine, Federal University of
Minas Gerais, Belo Horizonte.
Tooth wear (after Gier, 1957) was used to
classify each wolf according to a relative age class:
young, middle-aged, or old. All ectoparasites
found were counted and collected for future
identification. Minor wounds were treated with
a topical spray antiseptic (Topazone, Eaton
Labs.) and a prophylactic dose of 2 ml Longicil
Fortified Antibiotic (Penicillin G; Fort Dodge
Products) was applied subcutaneously. Chloro-
mycetin Ophthalmic Ointment (Fort Dodge
Products) was used to reduce drying of the eyes
during immobilization. A single Rototag (size no.
5; Dalton, England) was affixed to the lateral
margin of each pinna. A radiotransmitting collar
was fitted around the neck of each wolf. All
wolves were weighed (Figure 5) and measured.
Transmitters (Wildlife Materials) weighed
about 250 g, were equipped with bimodal activ-
ity-monitoring switches, and were fastened to
collars of nylon webbing, which were riveted
around the neck. The sizes and shapes of collars
were based on models of the necks of captive
maned wolves from the San Diego Zoo (courtesy
of Dr. P.T. Robinson, San Diego Zoo). A col-
lapsible three-element yagi antenna (Wildlife Ma-
terials) proved satisfactory under field conditions
in the Serra da Ganastra. Recapture of a wolf
was common and involved only testing the radi-
otransmitter and photographing the wolf prior
to its release.
Location by Radiotelemetry
Radiotelemetry permitted the determination
on a year-round basis of home ranges, activity
patterns, habitat preference, and social behavior.
NUMBER 392
FIGURE 5.?Weighing an immobilized maned wolf in the
Serra da Canastra.
Wolves were located on a daily schedule based
on four 6-hour time blocks: 0601-1200, 1201-
1800, 1801-2400, and 0001-0600 hours. Al-
though the accentuated topographic relief in the
area restricted my capacity to travel at night and
often reduced the radius of transmission of a
transmitter to a few hundred meters, an effort
was made to locate each wolf at least once every
24-hours. Attempts to radiolocate wolves were
dispersed over time so that each time-block was
represented in a given month.
I used triangulation based on two or more
vectors to determine the location of each wolf
(e.g., Craigheadand Craighead, 1972; Craighead
et al., 1973; Seidensticker et al., 1973). For each
location, I recorded the time of day, activity
level, and habitat type. After 30 seconds of mon-
itoring I classified a wolf as: inactive (no activity
recorded during the interval), intermittently ac-
tive (activity not continuous during the interval),
or constantly active. These activity classes were
established by testing radiocollars on domestic
dogs.
Radiolocated wolves were visually monitored
for individual and social behavior. Observation
was attempted periodically when a wolf was
known to be active or in a location when and
where visual observation was likely to be success-
ful.
If a wolf had not been observed for approxi-
mately a month, or if it was inactive and in the
same location for several days, radiotelemetry
was used to approach the animal to determine its
physical condition. The constant movement of
cattle and ranch hands throughout my study area
often displaced these maned wolves in a similar
manner. Wolves fled for short distances, and it
is unlikely that my interference had any signifi-
cant additional effects on their behavior or activ-
ity patterns.
Determination of Food Habits
SCAT COLLECTION.?Food habits were deter-
mined by the collection and analysis of fecal
droppings (scats). Only scats 25 mm or larger in
diameter were collected, based on a comparison
of known maned wolf scats and those of other
canid species in my study area. After several
months of field work, additional characteristics
of maned wolf scats, including odor, texture, and
site, became evident to me.
All maned wolf scats found were collected for
analysis, but scheduled collection was limited to
the three home ranges of the maned wolves that
I intensively studied. A goal for collection of at
least 50 scats per home range per month was
arbitrarily set and usually achieved. Scats col-
lected from the interface between two maned
wolf ranges were omitted from analysis relating
to home ranges but were included in other anal-
yses. All feces found in a single location (~0.5 m
in diameter) were classified as a single scat except
when obvious differences in dates of deposition
were evident.
SCAT ANALYSIS.?Scats were air-dried and
stored until analysis. All components of each scat
were identified and grouped into major food
types. A reference collection of fruits and animals
collected in the study area was used to identify
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
scat components. I measured the volume of the
major scat components by compacting the mate-
rial in a graduated cylinder using a close-fitting
dowel.
The relative importance of food items in col-
lected scats was calculated using relative volume
(total volume of that component as a function of
total volume of all components) and frequency
of occurrence (number of scats containing that
component as a function of the number of oc-
currences of all components). Data analyses were
performed using a series of SPSS programs (Sta-
tistical Package for the Social Sciences, Version
8, Vogelback Computing Center, Northwestern
University). The quantitative problems relating
to the use of scat analysis in determining food
habits have been documented (e.g., Floyd et al.,
1978; Meriwether and Johnson, 1980; OJ. Mu-
rie, 1946).
CENSUS OF AVAILABLE FOOD ITEMS.?Pub-
lished accounts of the food habits of maned
wolves indicated that small mammals and native
fruits were perhaps the items most commonly
exploited by this species (e.g., Cabrera and
Yepes, 1960; C. Carvalho, 1976). To determine
how maned wolves use available trophic re-
sources, the relative abundance of small mam-
mals (Dietz, 1983) and of fruits was estimated on
a monthly basis in each of the three major habi-
tats in the park.
The trap lines for small mammals were also
used as transect lines for the estimation of rela-
tive diversity and abundance of fruits available
to maned wolves. The 5 hectare area sampled in
each habitat was arbitrarily defined by a strip 10
m wide running the length of the trap line. Any
fruits larger than 1 cm in diameter and less than
1 m above the ground level were defined as
available to maned wolves. Relative abundance
of a fruit species was estimated by the number of
individual plants in fruit in the sampled area.
Interviews with area residents were used to
determine which fruit species might be eaten by
maned wolves. The months when these 43 spe-
cies were in fruit were noted and examples in-
corporated into the reference collection.
RESULTS
Physical Characteristics of Captured Maned Wolves
My observations are in general agreement with
previously cited descriptions of maned wolf phys-
ical attributes and, therefore, are not stressed.
However, as published data on the hematology
and blood chemistry of this species are unavaila-
ble for free-living individuals, I have included
information on these aspects in Tables 1 and 2.
PELAGE, WEIGHTS, AND MEASUREMENTS.?
Photographs of the nine adult maned wolves
captured in the Serra da Canastra revealed al-
most no observable individual variation in pelage
coloration. One male, Lambda, was slightly
darker on the cheeks and muzzle, but at distances
greater than about 50 m I found it impossible to
identify individuals without the aid of radiote-
lemetry.
The guard hair on the bodies of captured
wolves was about 8 cm in length, straight, and
golden-red in color. No underfur was present on
these adult wolves. The distal portion of the
muzzle was black, as was the antebrachium distal
from the elbow and the hindlimb distal from the
tarsus. Of the dorsal erectile mane, approxi-
mately the cranial half was black. Black from the
mane also radiated laterally over the scapulae.
The inner aspects of the pinnae were white as
were the intermandibular and throat regions.
The distal half of the tail was also white. The
length of white on the tail and of black on the
forelimb, hindlimb, and on the mane were meas-
ured on five of the captured wolves (Table 3).
Weights and selected measurements of nine
captured adult wolves are also presented in Table
3. All comparisons of these values among age
classes, and all but one comparison between sexes
were not significantly different (Student's t-test
for unpaired observations; Steel and Torrie,
1960; p > 0.05). However, sampled male wolves
had significantly longer pinnae than sampled fe-
males (p < 0.05).
HEMATOLOGY AND BLOOD CHEMISTRY.?I col-
lected whole blood for hematologic and serum
NUMBER 392 11
analysis from eight adult maned wolves in the
Serra da Canastra. In addition, the veterinarian
at the Belo Horizonte Zoological Garden contrib-
uted blood samples from two captive maned
wolves (Zoo 1 and Zoo 2) recently acquired by
that institution. One of the latter two wolves was
a female; the sex of the other was not recorded.
These data (Tables 1 and 2) may be compared
with values from other maned wolves, but small
sample sizes, difficulties during collection and
transportation of blood samples, and the lack of
information concerning environmental or physio-
logical conditions suggest that a degree of cau-
tion should be used.
Description of Home Ranges in the Study Area
During the two years of this study, I captured
eight maned wolves in the Serra da Canastra and
three wolves elsewhere. Radiotelemetric and vis-
ual observations of the eight wolves residing in
the study area allowed the definition of three
mutually exclusive home ranges (Figure 6). I
named these ranges the West Range, the East
Range, and the Taperao Range. Portions of each
range were located within the boundaries of the
Serra da Canastra National Park. Each range was
occupied by a single male-female bonded pair of
maned wolves.
WEST RANGE.?Connecting peripheral points
on a map of plotted radiotelemetric locations
MANED WOLF RANGES
FIGURE 6.?Locations of home ranges of maned wolves in
the study area.
yields a polygon with an area of 21.7 km2. How-
ever, because one series of ridges and valleys,
measuring about 4 km2, was difficult to reach
from my field headquarters, this estimate is prob-
ably smaller than the area actually used by the
resident pair of wolves.
EAST RANGE.?Directly northeast of my field
headquarters and adjacent to the West Range
was the East Range (Figures 6, 7). The pairs of
wolves that successfully used this range were
more easily located than the wolves of the other
two ranges. Therefore, my estimate of the area
of this range, 30.0 km2, is probably the most
accurate of the three.
TAPERAO RANGE.?The Taperao Range (Fig-
ure 6) was located about 2 km southwest of the
city of Sao Roque de Minas, and about 28 km
east of my field headquarters. Ranchers and
farmers declared that the movements of the res-
ident wolves had remained unchanged for many
years. With the help of these farmers, I first
trapped a male maned wolf, Beta, on 10 Septem-
ber 1978.
The Taperao Range was characterized by ex-
tremes of topographic relief. Only one road was
usually passable by automobile. Any of a number
of large ridges effectively blocked the transmitter
signal. Therefore, the area of this home range,
12 km2 as calculated from radiotelemetric data,
is an underestimation. A more accurate estimate,
based on single radiotelemetric vectors and visual
sightings, is about 24 km2. With the exception of
scat collection, data were collected only oppor-
tunistically in the Taperao Range.
OTHER RANGES.?In addition to the three de-
scribed home ranges, there were several other
ranges in the region (Figure 6). One such range
abutted the West Range to the north, and a
second range was adjacent to, and east of, the
East Range. Between this second range and the
city of Sao Roque de Minas was a third range,
while a fourth existed to the south of Sao Roque
and east of the Taperao Range. Access to these
additional ranges was difficult and I made no
attempts to capture their resident maned wolves.
Pairs of wolves were sighted in three of the four
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 7.?Part of the East Range, Serra da Canastra National Park.
ranges, and reproduction took place in all four
ranges. Reports by area residents of the numbers
of wolves vocalizing in these areas and my own
subjective estimates of the amounts of wolf-sign
present suggest that two wolves occupied each of
these additional ranges.
Radiolocations indicated that maned wolves
spent approximately 34 percent of their time in
grassland, 43 percent in cerrado, and 24 percent
in forest (Table 4). After transformation to re-
flect equal sample sizes for both sexes, a chi-
square test of goodness of fit (Steel and Torrie,
1960), suggested that males used grassland hab-
itat significantly more than did females (p <
0.05), and that females used cerrado habitat sig-
nificantly more than did males (p < 0.025). No
significant difference between sexes was found
in the use of forest habitat.
G-tests of independence (Sokal and Rohlf,
1969) indicated a significant relationship among
male-female pairs with respect to their use of
cerrado (p < 0.005) and of forest habitat (p <
0.005) (Table 5).
Seasonal differences in habitat use were deter-
mined by dividing the radiolocations into those
occurring during the wet and dry seasons (Table
6). Results of chi-square tests within seasons in-
dicate that grasslands were used more often than
expected during the wet season (p < 0.025), and
forest habitat was used more often than expected
during the dry season (p < 0.025). A possible
biological significance of these results is that the
grassland affords less cover and supports a lower
relative abundance of small vertebrates during
the dry season when pastures are burned by local
ranchers.
The preceding analyses are biased in two ways:
first, the majority of the radiolocations were dur-
ing the day. Thus, because these wolves were
largely nocturnal, habitat types associated with
daytime rest sites would be overrepresented. Sec-
ond, although the three defined habitat types did
not differ among home ranges with respect to
relative availability, those three habitat types
were not equally abundant in the study area.
Therefore, the contribution of smaller habitat
NUMBER 392 13
types would be overemphasized in the above
analysis. To correct for these biases, I divided
radiolocations into day (1601-1800 hours), and
night (1801-0600) observations. A correction
factor was applied to adjust for differences in
relative abundance of the three habitat types. I
placed a confidence interval using the Bonferroni
z-statistic (after Neu et al., 1974) on the observed
proportion of occurrence of wolves in each hab-
itat (Table 7); results indicate that maned wolves
of the Serra da Canastra were less likely to be
found in grassland habitat than in other types of
habitat at any time. During the day, wolves were
most often found in cerrado; at night, they were
most commonly located in forest habitat.
Activity Patterns
DAILY AND SEASONAL VARIATION.?Results
include 763 radiolocations of eight wolves over
20 months (Figure 8). A maximum of two loca-
tions per wolf per time block are included in the
analysis. Radiotelemetric data collected during
daylight hours (0601-1800) indicate that wolves
were active during 34.7 percent of the time, or,
35.0 percent of the six hours prior to noon and
IOOT
0001-0600 0601-1200 1201-1800 1801-2400
TIME OF DAY
FIGURE 8.?Activity of maned wolves during six-hour time
blocks. Total number of radiolocations for each time block
(n) are classified according to level of activity (shading =
inactive; hatching = intermittently active; white = constantly
active).
33.8 percent of the six hours following noon.
The majority (53.8 percent) of these active ob-
servations were classified as intermittently active,
and the remainder (46.2 percent) as constantly
active; that is, wolves spent approximately 115
minutes travelling from place to place during
daylight hours.
The maned wolves were more active at night
than during the day. Ninety-one percent of the
radiotelemetric observations between 1801 and
0600 hours were of constantly or intermittently
active wolves. Between 1801 hours and mid-
night, 96 percent of the radiolocations of active
wolves were classified as constantly active. Dur-
ing the six hours following midnight, 48 percent
of the radiolocations were of intermittently ac-
tive wolves and 52 percent were of constantly
active wolves. Thus, wolves spent about eight
hours (five hours before midnight and three
hours after midnight) moving from place to place
at night.
Although a chi-square test of independence
did not indicate a dependent relationship be-
tween level of activity and four male-female pairs
(p = 0.33; Table 8), the same test did indicate a
relation between activity and sex (p = 0.012;
Table 9). Further, when numbers of intermit-
tently active and of constantly active observations
were pooled to form a single class for each sex,
results indicated that males were more active
than females (G-test; p < 0.01). Again, using
combined numbers of observations of constantly
active and intermittently active wolves, females
were active 34.4 percent of the time and males
were active 44.0 percent of the time.
OBSERVATIONS ON INDIVIDUAL AcnvrrY.?
Data collected during 16 monitoring sessions of
from two to six hours in length added descriptive
detail to the picture of temporal relationships
among maned wolves.
During daylight hours, the wolves rested in
thick cover?often in riparian forest or in the
grass and shrubs at the headwaters of small
streams. Individuals usually remained in the
same patch of cover or moved only a few meters
during the day. At intervals of about 45 minutes,
however, each wolf could be expected to move
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
for a few seconds before again settling down.
The transmitted radio signals suggested postural
changes or perhaps standing up and turning
around before again lying down. Shortly after
sunset (between the hours of 1750 and 1845),
the wolves left the areas where they had been
resting during the day and began to hunt and
travel throughout their ranges. These move-
ments continued for much of the night with
occasional pauses of up to about 20 minutes.
Occasionally wolves travelled across a home
range with only brief stops. At other times, an
entire night might be spent hunting in a small
area. Usually by 0700 hours, but occasionally as
late as 0840 hours, the wolves again retired to
thick cover for the day. Wolves appeared to
remain active longer during rainy or foggy days
than during days of bright morning sunshine.
Social Organization and Communication
RELATIONSHIPS OF INDIVIDUALS WITHIN
PAIRS.?Results based on 1537 trap-nights (Ta-
ble 10), 828 precise radiotelemetric locations,
nearly as many single directional vectors, and
approximately 50 visual sightings of eight maned
wolves indicate that the maned wolves of the
Serra da Canastra displayed facultative monog-
amy (see Kleiman, 1977) characterized by a long-
term pair bond between male and female. Al-
though the male and female, as a pair, had com-
pletely overlapping ranges, very little time was
spent in close association. According to local
observers, family groups were rarely seen.
Beta: This wolf, a middle-aged male, was cap-
tured in the Taperao Range on 10 September
1978 (Figure 9). He was observed on numerous
GAMMA-'
LAMBDA-t
DELTA-I
EPSLON-I
KAPPA-!
ZETA-f
SIGMA-1
BETA-*
NotTrappad-
cl-
cl-
cl-
Cl-
East Range
cl-
? >
West Range
cl- ?JD
Taperao Range
FIGURE 9.?A chronological representation of the relationship among wolves inhabiting the
East, West, and Taperao ranges. (C = date of initial capture of a maned wolf; D = date by
which a maned wolf was known to have died; maned wolf Zeta died some time during the
period indicated by the dotted line.)
NUMBER 392 15
occasions in the company of a small female wolf
that apparently shared that range. These wolves
shared part, if not all, of the same home range
estimated to be 24 km2. During the 20 months
that Beta carried a transmitter, he was observed
either alone or with the same female wolf.
In May 1979, the signal of his transmitter
ceased. I thought that perhaps Beta had been
replaced by another wolf and set traps accord-
ingly. On 18 June 1979, Beta was retrapped and
fitted with a new transmitter. The external an-
tenna of his transmitter had been bitten off at its
base. The transmitter was also deeply gouged by
the teeth of a large carnivore, and one of the
rivets attaching the transmitter to its collar had
been pulled loose. This damage was probably
done during a social interaction with another
wolf.
Gamma and Delta: Gamma, a middle-aged
male wolf, was first captured on 15 December
1978 in the East Range (Figure 9). Although
severe seasonal rains initially complicated radio
telemetry, I learned the general outline of the
East Range, that the perimeter was well defined,
and that it was stable (Figure 10).
An old female wolf, Delta, was taken in the
same trap as was Gamma, but about one month
FIGURE 10.?East range occupied sequentially by maned
wolves Delta (dotted line; n = 18 radiolocations; female) and
Gamma (striped line; n = 20 radiolocations; male); Epsilon
(dashed line; n = 220 radiolocations; female); and Lambda
(solid line; 225 radiolocations; male). Polygons represent the
connected peripheral locations for each wolf.
later, 18 January 1979. At the time of her cap-
ture, she appeared weak and disoriented, often
shaking her head from side to side. Closer ex-
amination revealed that the left pinna and inner
ear of this wolf was severely infested with screw-
worm larvae (Cochliomyia hominovorax). A single
application of a lindane-based product (Myzin
Smear, Fort Dodge Products) was successful in
curing this infestation by the time of her second
capture on 4 February. On 5 February this wolf
suffered a compound fracture of the right radius
and ulna. She never recovered from this injury
and died of apparent starvation about 25 Feb-
ruary. Necrospy revealed that the screw-worm
infestation had caused erosion of the left tym-
panic bulla and surrounding temporal bone.
The number of radiolocations for Delta were
few (n = 18). In addition, during three of the six
weeks that this wolf carried a radio collar, her
locomotion was severely hampered by her bro-
ken leg. Nevertheless, the portion of her range
that I was able to identify almost completely
overlapped that of Gamma (Figure 10), and the
area she used corresponded to that most inten-
sively used by later occupants of this range.
These facts indicate that Delta was the mate of
Gamma and that both were residents of the East
Range.
Epsilon and Lambda: In late April, a young
adult female wolf, Epsilon, was captured in the
East Range in one of the traps in which Gamma
had been recaptured. Visual sightings of wolves
and the amount of sign present in the East Range
indicated that Epsilon was probably trapped
shortly after her arrival in this area. She carried
a radiotransmitter in this range for the duration
of my study.
About one month after the capture of Epsilon,
it became increasingly difficult to locate Gamma.
While Epsilon used the western half of the East
Range, Gamma travelled widely in the eastern
half of the range. During the month of June,
Gamma was often outside the range of my radio-
telemetric equipment. On 25 June 1979, he was
located for the last time. I believe that either he
was killed by ranchers or he permanently left the
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
study area and crossed a series of ridges to the
east of the East Range.
Ten days before the last location of Gamma in
the East Range, a young male wolf, Lambda, was
taken in the same trap that had captured both
Gamma and Epsilon (Table 10). Lambda was
captured an additional 21 times and radiolocated
225 times in the East Range (Figure 10). After
the captures of Epsilon and Lambda, no addi-
tional maned wolves were observed or trapped
in this range.
In summary, the East Range was occupied by
maned wolves Gamma (male) and Delta (female)
for an unknown period of time prior to their
respective captures in December 1978 and Jan-
uary 1979. About two months after her death,
Delta was replaced by Epsilon, a young female
new to the East Range. Gamma did not closely
associate with Epsilon. About 10 days before
Gamma's departure, a young male, Lambda, was
captured in the East Range and rapidly took over
most of the area formerly used by Gamma. Based
on this circumstantial evidence, Lambda may
have been responsible for the eviction of Gamma
from the East Range. The areas used by Lambda
and Epsilon, a total of 30 km2, overlapped almost
completely and differed little from that of former
inhabitants Gamma and Delta.
Kappa, Zeta, and Sigma: Reported wolf activ-
ity to the west of the East Range prompted me
to extend trapping efforts to that area. On 15
May 1979, Kappa, a young adult female wolf,
was taken in what I named the West Range. She
carried a radiotransmitter in this range for the
remaining 12 months of this study. Seven weeks
after the capture of Kappa, a middle-aged male
wolf, Zeta, was taken in the same trap. The home
ranges of these two wolves overlapped exten-
sively (Figure 11) for about 14 weeks. On 17
October 1979, Zeta was located for the last time.
Nearly six weeks later I followed the signal of his
transmitter to the bank of a swollen stream. The
nylon collar had been cut with a sharp instru-
ment. Zeta had probably been killed by an area
farmer and the removed transmiter thrown into
a deep pond. A flash flood washed the transmit-
FIGURE 11.?West Range occupied by maned wolves Kappa
(solid line; n = 106 total radiolocations; female), Zeta (dashed
and dotted line; n ? 25 radiolocations; male), and Sigma
(dashed line; n = 46 radiolocations; male). Sigma replaced
Zeta in this range after Zeta's death. Polygons represent the
connected peripheral locations for each wolf.
ter up onto the shore on the day that I received
the final radiosignal.
On 18 November, a middle-aged male wolf,
Sigma, was trapped in the West Range. Sigma
took over the range formerly occupied by the
missing Zeta (Figure 11), and on several occa-
sions was observed in close association with
Kappa.
The maned wolves of the Serra da Ganastra
led predominantly solitary existences. Wolves
were never observed resting together, and they
rarely travelled or hunted together. On only 25
occasions during this study were two adult wolves
observed in close association. Groups of more
than two adults were never seen. Lambda and
Epsilon were seen in close association on 17
occasions in the East Range (Figure 12). Other
data, such as the distribution of wolf feces and
tracks, indicate that social activities involving
both individuals were most common in Septem-
ber 1979 and were least common during March
through May of the following year.
During periods when the members of a dyad
were not often in close association, the location
of one maned wolf was not independent of that
of its mate. Both individuals used the same trails
and resting places, but not simultaneously. In all
NUMBER 392 17
FIGURE 12.?Percent of radiotelemetric locations per month
in which Lambda and Epsilon were found in close association
(n = number of such observations).
but the Taperao Range, both members of a pair
were taken in the same trap. Capturing one wolf
often facilitated the prompt capture of its mate
in the same trap. However, precedence in the
use of traps and favorite resting places was not
sex-specific.
VOCALIZATIONS.?High-amplitude vocaliza-
tions produced by maned wolves bear little re-
semblance to the variable and oscillating howls
of the gray wolf (Canis lupus) or of the coyote (C.
latrans). The vocalizations of maned wolves more
resemble an extended variation of the bark of a
large domestic dog. Each bark lasted approxi-
mately 0.7 second, was repeated at regular inter-
vals of from 2 to 4 seconds, and sounded some-
thing like "oo-wahh." Wolves repeated this call
from 2 to 23 times in succession with an average
of about 9 repetitions. Both sexes bark, but rec-
ognition by the author of an individual by its call
was not possible. Occasionally as many as three
wolves were heard barking simultaneously, and
in at least two instances, wolves of two adjacent
home ranges were heard exchanging vocaliza-
tions.
Wolves were heard barking on 46 occasions
(Figure 13) of which 29 occurred in the East
Range. These data indicate that maned wolves
vocalized at all times of the year, and that a peak
in frequency occurred during the months of June
through August 1979. This higher incidence of
vocalization might have been related to the re-
placement of Gamma by Lambda in June 1979
and to the subsequent reproductive behavior of
Lambda and Epsilon.
The only additional vocalization that I heard
was a low-volume growl produced when a
trapped wolf was first approached by a human.
When approached more closely, the trapped wolf
invariably began a series of extended barks. Ar-
tificial predator calls and my own attempts to
imitate the extended barks of maned wolves pro-
duced no discernible response in wolves in the
Serra da Canastra.
DEPOSITION OF FECES AND URINE.?Feces
were most commonly encountered in the vicinity
of favorite resting sites, at den sites, and on or
near prominent points such as rock outcrops,
fence lines, trail intersections, or termite
mounds. Scat grouping, elevation above sur-
rounding ground level, and substrate often dif-
fered between these location types.
I examined three den sites recently used by
maned wolves (see "Reproduction"); one had
scats deposited by one or more adults and by two
pups. Scats of 1 cm or less in diameter were
found within a few meters of the den, while
slightly larger scats were found farther from the
den. Apparently, as pups grew they wandered
and defecated farther from the den. All scats
deposited by adults were located within a radius
of 50 m from the mouth of the den, and all were
deposited at ground elevation. Outside of this
perimeter, I found no scats in the region of the
den. I often found one, and occasionally two or
three, concentrations of scats near vicinities of
frequently used resting places. In three cases,
these "latrines" were about 20 m from a favorite
resting place of a pair of maned wolves, consisted
of a low, flat rock, and contained between 10
and 20 scats. Several scats were also found on
trails and rocks near these concentrations. These
latrines were probably used by both wolves resid-
ing in a home range.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
East Range
(?=2S)
Al Ranges
(IMS)
MONTH
FIGURE 13.?Number of nights per month during which maned wolves were heard vocalizing
in the East Range and in all ranges in the study area (n = the total number of observations).
Scats on trails were not distributed in a random
fashion throughout a home range. When travell-
ing from place to place, a maned wolf not actively
hunting usually followed the path of least envi-
ronmental resistance. Often-used trails were cus-
tomarily located along open ridge tops, and scats
along these important routes were deposited in-
dividually at intervals along trails. When I col-
lected scats that had remained undisturbed along
a trail for several weeks, maned wolves often
replaced them with fresh scats within a few days.
Sixty-five percent of the 740 collected scats
were found on rocks, dirt piles, or termite
mounds elevated above the surrounding ground
level (Table 11). The mean elevation from sur-
rounding ground for all collected scats was 39.4
cm, and scats were collected from atop termite
mounds up to 170 cm in height.
A few additional observations imply that
defecation serves a defensive or alarm function.
When startled or confronted at close quarters,
those animals (a male and a fernale) sprang onto
rocks about 60 cm high and defecated before
fleeing. Finally, one night I observed the meeting
of two wolves coming from opposite directions
along the same trail. One of the two defecated
on the trail and one barked repeatedly.
Little information was collected concerning
urine-marking by maned wolves. The odor of
the urine of maned wolves was prevalent along
frequently used trails, indicating that urine-
marking may have been employed in the same
context as the elevated deposition of feces. No
scratch marks were observed at the urination
sites or with feces. In addition to urination by all
wolves within traps, two individuals urinated im-
NUMBER 392 19
mediately after being released from traps?again
suggesting a parallel between the alarm or defen-
sive function of feces deposition and urination.
All trapped wolves and a free female wolf were
observed to use a squatting posture during uri-
nation. A free male was observed on two occa-
sions to cock a hind leg when urinating.
In summary, maned wolves deposited feces
and perhaps urine selectively with respect to
location. Feces near a den site were deposited at
ground level and were restricted to the area
immediately surrounding the mouth of the den.
Latrines in the vicinity of resting locations were
used by both occupants of the home range and
contained large concentrations of scats. Finally,
scats found throughout the remainder of a home
range were physically elevated, individually lo-
cated, spaced at intervals and replaced after re-
moval. I presume that these elevated and spaced
feces served to advertise the presence of a maned
wolf and may have aided indirectly in the defense
of territories. Reduced dispersal of scats in the
area of the den site and resting sites perhaps
served to emphasize the ownership of those areas
to unfriendly conspecifics or other predators.
RELATIONSHIPS BETWEEN ADJACENT PAIRS OF
WOLVES.?The maned wolves studied main-
tained strict territoriality, i.e., boundaries of the
ranges of each bonded pair were fixed and were
almost never transgressed by wolves of an adja-
cent range. In most cases, the limits of these
ranges were composed of physically identifiable
landmarks, such as roads, ridges, or streams.
Where they could be identified, the interfaces
between adjacent territories were only a few
meters wide. For example, a north-south road
divided the West Range and the East Range. The
wolves of each range used the areas on their
respective sides of the road but neither pair used
the road per se or crossed to the opposite side.
Scats were liberally deposited along trails on
either side of the road, and wolves were occa-
sionally heard vocalizing back and forth across
the boundary.
Home range boundaries appeared to remain
constant over time. The two exceptions con-
cerned those of wolves that had died or left the
area. These two observations give some insight
into the type of territoriality maintained by these
maned wolves.
When Delta (female) died and Gamma (male)
died or abandoned the west portion of the East
Range, the interface between the West Range
and the East range was the Gameleira Road
(Figure 14). Within several weeks, these wolves
were replaced by Epsilon (female) and Lambda
(male). However, just prior to, or perhaps shortly
after, the arrival of the latter pair, one or both
wolves of the West Range extended their terri-
tory into the region referred to as Antonio's Area
in the East Range. Within six weeks, all four
resident wolves were observed at least once in
this area of about 0.8 km2. From early July until
October, only Zeta (male) was found using An-
tonio's Area, indicating that a portion of the East
Range had been annexed to the West Range.
When Zeta was killed in late October or early
November, Antonio's Area remained vacant for
about four weeks before being repossessed by
Lambda, the male of the East Range. He was the
only wolf known to use this area for the remain-
ing five months of the study.
A second region of contention along the bor-
der of the West and East ranges was a 0.75 km2
WEST RANGE
FIGURE 14.?Areas of dispute between maned wolves resid-
ing in the West Range and in the East Range. Antonio's
Area was successively occupied by maned wolves of the East
Range, West Range and East Range. Jose's Area was taken
over by Lambda (East Range) after the death of Zeta (West
Range). The Gameleira Road divides the two ranges.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
pasture referred to as Jose's Area. This region
was used occasionally by both wolves of the West
Range until November 1979. After the death of
Zeta, resident male of the West Range, Jose's
Area was taken over for the duration of the study
by Lambda, resident male of the East Range.
Subsequent to this appropriation, the only wolf
observed in this area was Lambda. Jose's Area
had apparently also been added to the East
Range.
The territoriality maintained between adja-
cent pairs of maned wolves was dynamic and
expansive in nature. An area left vacant in one
range by the death or departure of its owner
resulted in appropriation by a wolf of the adja-
cent range. All successful appropriations were
accomplished by male wolves; female wolves
were not subsequently observed in these areas,
suggesting a system in which males competed to
secure territory rather than mates. These new
additions to the territories of males may later be
acquired by their mates during periods of de-
creased social distance associated with the breed-
ing season.
LONERS OR NOMADIC MANED WOLVES.?On
three occasions, maned wolves other than radio-
collared individuals were observed in the vicinity
of the West and East ranges. One sighting took
place on the boundary between the West and
East ranges, and the other two sightings were
made along the southern boundary of the East
Range. A skilled dog trailed one of these wolves
north along the eastern periphery of the East
Range for several kilometers before losing the
scent at a river. Finally, on one occasion, a set of
wolf tracks not attributable to a radiocollared
wolf was followed from the northern extreme to
the southern extreme of the interface between
the West and East ranges. These observations
suggest that maned wolves without fixed ranges
travelled the peripheries of occupied ranges and
presumably filled vacancies created by death or
abandonment. Maned wolves Lambda, Epsilon,
and Sigma replaced previous resident wolves and
could have been loners prior to inheriting their
respective ranges. No general statement can be
made about the gender and age class of these
wolves (a young male, a young female, and a
middle-age male) as a group.
Feeding Ecology
SEASONAL AVAILABILITY OF TROPHIC RE-
SOURCES.?The numbers of available fruit spe-
cies determined by questionnaires (Figure 15,
Curve A), and monthly transect-lines (Figure 15,
Curve C) were greater during the wet season
than the dry season. The seasonality of 14 species
of fruits identified in maned wolf scats (Curve B
in Figure 15) suggests a similar seasonal fruiting
pattern between the fruits available to and con-
sumed by maned wolves.
The fruit lobeira (Figure 16) was considered
apart from other fruits. This species is common
in the Serra da Canastra but is not ubiquitous.
Since this species prefers more organic soils than
were prevalent in the national park, lobeira in
this region was commonly located in richer soils
and along horse-trails and near corrals. The spe-
cies is everbearing, and I noted no quantitative
differences in fruit production between seasons.
The results from the small-mammal trapping
study (Figure 17; Dietz, 1983) suggest that small
FIGURE 15.?Seasonal availability of fruit species in the study
area. (Curves: A = the months during which 43 plant species
reported consumed by maned wolves were in fruit; B = the
months of fruiting of 14 species known to be consumed by
maned wolves; c = the estimated total number of species
fruiting per month as determined by transect-lines.)
NUMBER 392 21
FIGURE 16.?Lobeira (Solatium lycocarpum), "fruit of the maned wolf."
WET SEASON DRY SEASON WET SEASON
J ' ' * ' * " 1*0
FIGURE 17.?Numbers of small mammals captured by
author.
mammals were more abundant, or at least were
more readily caught, during the dry season than
during the wet season (x2 = 46.5, 2 d.f.,
p < 0.005). Coleopteran beetles were extremely
common during the months of October and No-
vember. Large volant ants swarmed during the
month of November.
FOOD HABITS OF MANED WOLVES IN THE
STUDY AREA.?The diet of maned wolves of the
Serra da Canastra, based on a complete list of
food items identified in scats (Tables 12, 13), is
typical for a generalist omnivore; of 41 identifi-
able items, 20 were animal in origin, 20 were of
plant materials and one was inorganic. Approxi-
mately 49 percent of the 2056 occurrences of
food items in the analyzed scats were animal
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
material and 51 percent were plant material.
The components of the list of food items were
divided into categories based on taxonomic re-
lationships and on questions of interest. For ex-
ample, the endangered status of the giant antea-
ter (Myrmecophaga tridactyla) suggested that this
species be included as a separate category (Table
14).
In terms of volume and occurrence in scats,
the fruit Solatium lycocarpum was the single most
important food item in the diet of the maned
wolves of the Serra da Canastra. The regional
common name for this fruit, "lobeira," freely
translated means "fruit of the maned wolf."
Larger examples of this fruit weighed about 450
g and resembled large tomatoes: green before,
and yellow upon, ripening. Although plants of
this species produced ripe fruit in our corral, I
never observed ripe fruit in any other region.
This, I assume, was due to domestic and wild
fauna feeding on the fruits before or as soon as
ripening took place. Maned wolves apparently
did not await ripening before consuming it, as
evidenced by large pieces of the green fruit in
fresh feces.
Small mammals and birds occurred in 69 and
33 percent, respectively, of examined scats. The
majority of small mammals consumed were prob-
ably cricetid rodents; however, specific identifi-
cation was not attempted. The relatively high
incidence of grass in the examined scats cannot
be explained on nutritional grounds. Ingestion
of grass has been observed in captive maned
wolves (Kuhme, 1975; Brady and Ditton, 1979),
and D. Altmann (1972) reports that consumed
grass is defecated within seven minutes of inges-
tion. Lloyd (1980) suggested that grass is often
eaten accidentally by foxes catching small prey.
A. Murie (1944) noted roundworms in wolf (C.
lupus) scats composed largely of grass and spec-
ulated on the beneficial scouring action of grass.
Lever (1959) mentioned the diuretic and emetic
actions of some grasses on red foxes. On one
occasion, I observed a maned wolf investigate
and consume a clump of grass that had previously
been urinated upon by another wolf.
At least six species of armadillo were common
in the study area, and four were identified by
their dermal scutes in 8.6 percent of examined
wolf scats: Dasypus novemcinctus (in 31 scats), Da-
sypus septemcinctus (2), Dasypus sp. (2), and Eu-
phractus sexcinctus (5). In most cases, ants and soil
were also found in scats containing armadillo
scutes, indicating that the former two items were
perhaps ingested incidentally. Carcasses of ar-
madillo preyed upon by maned wolves usually
consisted of an intact but deeply scratched cara-
pace and head. The ventral musculature and
viscera, except the descending colon, were re-
moved by the wolf. Maned wolves apparently
surprise armadillos above ground and then use
teeth and claws to turn the prey onto its dorsum,
thereby exposing the more vulnerable venter.
Insect material was found in 14 percent of
examined scats but comprised only 0.3 percent
of total scat volume. This is in marked contrast
with scats of sympatric foxes Dusicyon and Cerdo-
cyon, which contained high percentages and vol-
umes of insect remains. Maned wolves apparently
do not forage as intensively for insects as do these
smaller canids.
Small amounts of the hair of giant anteaters
were found in 0.7 percent of wolf scats. These
long, fiat hairs were typical of the tail and dorsal
mane of the giant anteater and were never found
in sufficient quantity to suggest that a maned
wolf fed on an anteater. Maned wolves may have
removed a few hairs from the carcasses of dead
anteaters, or may have occasionally attacked live
anteaters. However, my observations indicate
that during apparently chance encounters be-
tween maned wolves and giant anteaters, neither
animal paid any noticeable attention to the other.
Seasonal changes in the food habits of the
maned wolves of the Serra da Canastra were
determined empirically by categorizing scats by
collection date: dry season or wet season. Weath-
ering and insects destroyed scats rapidly; there-
fore the differential survivorship of scats from
one season to the next was ignored for the pur-
poses of this analysis.
Lobeira, the main staple in the diet of these
NUMBER 392 23
maned wolves, was consumed consistently across
seasons (chi-square, p > 0.90), whereas small
mammal remains and foliage were encountered
more often in dry-season scats (Table 15). Con-
versely, fruits (except lobeira) and insect remains
were more prevalent in scats collected in the wet
season.
The seven food items most important in terms
of volume and of occurrence in scats were di-
vided according to home range to test if use of
trophic resources varied between wolf pairs.
There were no differences in the percent of
occurrences of these items between home ranges
(chi-square, p > 0.05).
In summary, the numerically most important
dietary components of the maned wolves of the
Serra da Canastra were lobeira, small mammals
and miscellaneous fruits. Consumption was ap-
parently consistent with relative availability for
all three of these classes of food items: lobeira
was available and consistently consumed during
both the wet and dry seasons; small mammals
were most numerous and were consumed in
greatest numbers during the dry season; fruits
other than lobeira were most abundant and most
often consumed during the wet season. Insects
appeared to be most available and consumed
most often during the wet season.
Maned wolves at the Serra da Canastra are,
therefore, opportunistic generalists in their for-
aging habits. They feed on those food types that
are most abundant, shifting to alternative re-
sources when they become abundant. Trophic
resource use did not differ between mutually
exclusive home ranges.
EXPLOITATION OF DOMESTIC STOCK BY MANED
WOLVES.?Beef carrion was usually available
throughout the study area in the form of cattle
dead from disease or injury. These carcasses
were not exploited by area ranchers and would
seemingly represent a large store of available
protein for resident maned wolves. However, I
neither located a radiocollared wolf, nor found
a track or scent post, in the vicinity of cattle
carcasses. Only one rancher reported having seen
a maned wolf feeding on carrion: the wolf was
seen with its head inside the abdominal cavity of
a recently dead steer and was tugging at the
viscera.
Interviews with area residents suggest that
maned wolves never preyed upon calves or the
adults of cattle, sheep, or pigs. Maned wolves
may very rarely take newborn lambs in this re-
gion. Wolves occasionally prey on young pigs
and frequently take domestic chickens.
The maned wolfs preference for chickens re-
sults in more human-related wolf mortality than
any other interaction. Most Brazilians in the
Serra da Canastra maintain flocks of chickens for
eggs and meat, but the flocks are not shut in at
night. A marauding wolf may kill one or many
chickens in a single or multiple visits. The wary
wolf becomes quite the opposite when approach-
ing a flock of chickens. Wolves attacking my
chickens advanced with bodies close to the
ground and their gaze fixed on the prey.
Farmers can effectively defend barnyard fowl
from predation by maned wolves by enclosing
the roosting birds at night and by the acquistion
of reliable watch dogs.
Radiotelemetric data suggest that some maned
wolves attacked chickens with greater frequency
than others. Of 23 reported incidents of preda-
tion on domestic stock, 14 (61 percent) took
place from September through November, the
time when maned wolf pups are most likely to be
at den sites. Increased energy requirements im-
posed by nourishing the young may force the
adult female and perhaps the male to exploit
domestic stock to a greater degree at this time of
year.
Reproduction
DENNING.?I examined three den sites, and
was informed of another, in which maned wolf
pups had recently been raised. These dens were
above ground, took advantage of natural fea-
tures, and were located in open grass pasture.
All were within a few hundred meters of roads
frequently used by ranchers in cars or on horses.
One den was situated in a clump of grass at the
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
base of a hillside; a second den was found in a
shallow crevice formed by several large rocks.
The third den, from which two pups were cap-
tured, was located in a grass-covered gully
formed by previous water erosion. The cavity of
this den was 30 to 100 cm deep and about 60 cm
wide. Near the opening of the den, grass had
been trampled and flattened. Wolf scats and
chicken feathers were strewn within a radius of
approximately 50 m. The fourth den was within
the hollow of a termite mound excavated perhaps
by a giant anteater. I obtained no evidence to
indicate that maned wolves either excavated or
enlarged abandoned underground dens.
PARTURITION AND LITTER SIZE.?None of the
radiocollared wolves succeeded in raising a litter
during my study. However, observations of Ep-
silon and Lambda during the last two weeks of
August 1979, suggest that Epsilon may have
whelped at that time. Radiotelemetry at this time
indicated that Epsilon rarely moved from a dry
creek bed near the west border of the East
Range. Lambda was usually within a few
hundred meters of her location and was never
found farther than 2 km from the creek bed.
Wolves were heard barking in this area on five
nights during this period. The area immediately
surrounding the possible den site was littered
with scats. I did not examine the den site at close
quarters until after the animals had abandoned
the site and, therefore, have no direct evidence
that whelping took place.
Interviews with area residents produced some
J F
FIGURE 18.?Estimated months of birth of 21 litters of
maned wolves seen by residents of the Serra da Canastra.
Stature of pups was used to estimate months of birth.
data on 27 litters, as well as on one pregnant
female near term and on one lactating female.
To estimate dates of birth, information concern-
ing the sizes of pups from 21 litters was compared
with a theoretical growth curve based on inter-
views, my own observations, and published infor-
mation (Acosta, 1972). Results (Figure 18) indi-
cate that whelping may begin as early as February
and reach a peak between June and September.
Seventy-one percent (n = 15) of observed litters
were estimated to have been born during these
latter four months. The mean number of pups
observed in 17 litters was 2.47, and the range
was one to five pups (Figure 19). Weights and
measurements of a litter of two female pups,
estimated to have been born during the last two
weeks of June 1978, are presented in Table 16.
PARENTAL CARE OF PUPS.?If Epsilon did pro-
duce a litter during the last two weeks of August
1979, inferences about the behavior of Lambda,
her mate, are perhaps warranted. Although he
was not located in the vicinity of the den site with
the same regularity as Epsilon, his persistence in
that area suggests that he may have played some
role in the early care of the pups. On the morning
of 25 August, my field assistant watched Lambda
hunting in an area about 500 m from the den
site and saw the wolf pounce on and seize a small
animal in its jaws. Lambda carried his prey di-
rectly to the den site where Epsilon was located.
Since Epsilon's pups would not yet have been old
enough to ingest solid food, Lambda probably
carried this food item to his mate. Beginning on
FIGURE 19.?Numbers 5
of pups in 17 maned wolf fc
litters observed by resi- tt 3.
dents of the Serra da I
Canastra. 2
1
1 2 3 4 5
PUPS MUTTER
NUMBER 392 25
about 26 August, this pair of wolves again began
to travel widely about their range and no longer
spent a disproportionate amount of time at the
den site. Although these wolves were observed
together several times during the month of Sep-
tember, thorough searches revealed no evidence
of pups at the den site.
Some inferences concerning parental care of
pups at the den can also be drawn from infor-
mation collected during interviews of area resi-
dents: of 24 litters of maned wolf pups observed
by area residents, 16 were unaccompanied by
adult wolves, 6 each had one adult wolf present,
and 2 litters had 2 adult wolves present. If the
only adult wolves participating in raising a litter
are the parents of that litter, then at least some
males associate with the female and her pups
before the pups disperse.
I recorded six reliable reports by area residents
who experienced aggressive attacks by single
adult maned wolves as a result of human inter-
ference with litters of pups. In four cases, the
advances were accompanied by threats and at-
tempts to bite the offending humans. Three of
the six attacks took place while cowboys were
burning pasture, which flushed the wolves. Sub-
sequently, the maned wolf pups were captured.
In one case a wolf (identified as a female) ad-
vanced to within 4 meters of a cowboy who then
released two pups. The adult wolf picked up both
pups simultaneously in its mouth and carried
them over a nearby hilltop. In another case, a
cowboy captured a wolf pup and carried it 6 km
to his ranch. An adult wolf followed him to the
ranch and barked in the vicinity until the pup
escaped.
A final observation concerns the probable re-
location of a litter of pups by an adult female
maned wolf. On 26 June 1977, an area resident
saw an adult female wolf and two pups at a den
site in the East Range about 500 m east of the
den site selected by Epsilon. The pups, which did
not yet have their eyes open, were located under
a bent-over thatch of grass. The following day,
the cowboy returned to the site and found nei-
ther the adult wolf nor the pups.
Parasites, Diseases, and Mortality
ECTOPARASITES.?On 11 occasions (involving
8 wolves), total ectoparasite loads were estimated
and representative parasites were collected for
identification. The most commonly observed ec-
toparasites were ixodid ticks of the genus Am-
blyomma. The majority were A. cajennense, but a
few A. tigrinum were also found (J. Whitaker per
N. Wilson, in litt.). From 10 to 50 ticks were
counted on each wolf (x = 24; SD = 12.1). Most
were located in and around the ears.
One individual flea was found during each of
3 of the 11 examinations. The scarcity of this
parasite may be related to the lack of underfur
in this host. The only other ectoparasites noted
were screw-worm larvae (Cochliomyia hominivorax)
infesting the pinna and inner ear of maned wolf
Delta. Chandler and Read (1961) mention that
these insects often infest wounds resulting from
tick bites; Delta had many ticks in the ears.
One ectoparasite conspicuous by its absence
on examined wolves was the cuterebrid skin bot
fly (Dermatobia hominis). Although the subcuta-
neous larvae of this parasite were extremely com-
mon in cattle and dogs in the study area, none
were observed infesting maned wolves.
ENDOPARASITES.?Fresh feces collected from
traps, after the release of six captured maned
wolves in the Serra da Canastra, and from the
cages of two recently captured wolves in the Belo
Horizonte Zoological Garden were sent to the
School of Veterinary Medicine, Federal Univer-
sity of Minas Gerais, for examination for endo-
parasitic eggs or larvae. Each examined wolf
carried from two to four genera of endoparasites
(x = 3, SD = 1; Table 18). Six genera of endo-
parasites were identified: four nematodes, one
cestode, and one coccidian protozoan. Three
nematodes, Trichurissp., Ancylostoma sp., and Tox-
ocara sp. were most frequently found, and were
detected, respectively, in eight, seven, and four
of the eight examined wolves.
Certain parasitic infestations, for example,
dioctophymiasis (Mace and Anderson, 1975)
probably would not be diagnosed by the sedi-
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
mentation method used in the above analyses. In
addition, because only one fecal sample was ex-
amined from most wolves, the results probably
underestimated the presence of parasites whose
eggs or larvae are found only intermittently. Two
additional genera of parasites were found in the
second sample of feces than in the first sample
from maned wolf Beta.
DISEASES.?The urine of six of eight wolves
tested positive for cystinuria (Table 17). Cystin-
uria is an inherited metabolic disease described
for some breeds of dogs and in humans. The
significance of the unexpectedly high incidence
of this disease in maned wolves is discussed by
Bovee et al. (1981). The relationship between
the high incidence of cystinuria and dibasic
amino-aciduria is not clearly understood. One
female wolf, Kappa, also showed abnormal
amounts of glucose in the examined urine sam-
pie.
MORTALITY IN THE SERRA DA CANASTRA.?I
saw the remains of only two wolves that had died
of apparently natural causes: a few bone frag-
ments from one subadult, and the nearly com-
plete skeleton of Delta.
I was informed of 19 adult wolves and of 4
subadults that were killed by residents of the
Serra da Canastra since approximately 1970.
These wolves were killed in the following man-
ners: seven were shot, five were run down and
killed by dogs, four were trapped, two pups were
taken from a den and subsequently died of dis-
ease in captivity, one pup was poisoned in captiv-
ity, one adult was killed by a thrown rock, one
was clubbed to death, and two died of unknown
causes. Sixteen of these 21 deaths took place
while wolves were attempting to steal chickens
from barnyards; no mortalities were attributed
to hunting for sport.
Relations of Maned Wolves with Humans
EFFECTS OF HUMAN DISTURBANCE ON MANED
WOLVES.?Published accounts (e.g., Walker,
1975) state that populations of maned wolves
tend to retreat before the advancement of hu-
man colonization. My observations do not sup-
port this assertion, because the home ranges oc-
cupied by maned wolves have also been occupied
by farmers and ranchers for generations. Dis-
turbance factors, such as annual burning, clear-
ing of brush, and overgrazing throughout the
entire area had no observable immediate effects
on the movements and activity patterns of adult
maned wolves. However, I noted three reports
of maned wolf pups captured after being driven
from dens by range fires. I observed no direct
effects of subsistence-level agricultural practices
on maned wolves in this area. Wolves made no
apparent attempt to avoid fences, planted fields,
roads, or the vicinities of uninhabited houses.
Hofmann et al. (1975-1976) stated that
maned wolves in the Peruvian Pampas del Heath
were diurnal in an area annually burned but
uninhabited by man. My observations indicated
that wolves were acutely sensitive to the physical
presence of a human on horseback or on foot.
Upon detecting a human in its vicinity, a wolf
would seek cover in nearby brush or when in
grassland would lie curled on the ground with
only its ears occasionally moving. When a person
approached to within about 2 to 3 m the wolves
flushed, would piloerect and often vocalize, and
bounded toward the nearest cover. In most in-
stances, retreating wolves stopped at least once,
turned broadside to the human, back arched and
piloerect, and displayed one or more of the fol-
lowing behaviors: defecation, urination, vocali-
zation, feigned hunting behavior, or olfactory
investigation of a nearby rock or clump of grass.
A close encounter with a maned wolf is an
extremely rare event in the Serra da Canastra.
In two years of field work, I saw only five wolves
without the aid of radiotelemetry. Three of these
encounters were with wolves stealing chickens
from my barnyard. Cowboys and farmers who
lived and worked within my study area saw
wolves only about once a year.
ATTITUDES OF AREA RESIDENTS TOWARD
MANED WOLVES.?The attitudes of the rural
residents of the Serra da Canastra toward maned
wolves were not confounded by emotional or
political conflicts and were based on a relatively
accurate evaluation of the ecological role of the
NUMBER 392 27
wolves in that area. Maned wolves were not
actively hunted for sport or for food but were
occasionally killed in retribution for damage
done to domestic stock. In these cases, individual
offending wolves were selectively eliminated by
the farmer or rancher suffering the loss. Except-
ing females defending pups, maned wolves are
generally thought of as timid and fearful of man,
and successfully coexist with him in the Serra da
Canastra.
FOLKLORE.?The maned wolf has become an
important part of Brazilian folk culture and is
considered to have more supernatural identifi-
cation with man than any other species of fauna.
Maned wolves are thought to possess inexplicable
powers, and a variety of parts of their anatomy
serve as ingredients in folk medicines. For ex-
ample, the right eye removed from a live maned
wolf is said to increase the sexual prowess of the
bearer after the wolf is released.
Another legend predicts that a large snake can
be found in the kidney of a freshly killed maned
wolf. The snake in this legend is likely to be the
giant kidney worm common in maned wolves.
Azara (1801) also mentions this legend for wolves
in Paraguay.
During two visits to a locally renowned spirit-
ualist healer (curador), I was informed of the
recipes for many medicinal preparations based
on maned wolf ingredients: A canine tooth from
a maned wolf is tied around the neck of a child
to cure or protect against dental problems. A
piece of hide from a wolf is attached to a saddle
or belt to cure the user of kidney or back-related
problems. Ingestion of two small pieces of the
heart of a freshly killed wolf will protect a snake-
bite victim from death. Finally, a chronic cough
is best cured by drinking hot tea made from the
dried feces of a maned wolf.
Study of Captive Maned Wolves
METHODS
Facilities and Observation Schedule
Behavioral observations were conducted on 15
maned wolves captive at the Conservation and
Research Center (CRC), National Zoological
Park, in Front Royal, Virginia. Wolves were
housed in male-female pairs in fenced outdoor
enclosures averaging approximately 725 m2 in
area (Figure 20). Each pair had access to one
heated den and in some cases to an additional
unheated den. Adjacent pairs were separated by
cinder-block walls. With the exception of daily
feeding and cleaning routines, wolves were not
often disturbed by humans. Management of
maned wolves at this facility was as described by
Brady and Ditton (1979).
Preliminary observations indicated crepuscu-
lar activity peaks in maned wolf activity. There-
fore, behavioral observations were conducted
during the period from one hour before sunset
to three hours after sunset, and during the period
from three hours before sunrise to one hour after
sunrise. Prior to each observation session I
switched on three to five red floodlights in the
enclosures where observations were to take
place. Observations were conducted using in-
frared binoculars from blinds mounted 3 m from
ground level adjacent to enclosures.
Scheduled observations were conducted (1)
throughout the breeding season of 1981-1982
(12 October 1981 to 23 December 1981) until
individuals of male-female pairs were separated
prior to whelping; (2) subsequent to the rejoining
of pair members following weaning of pups (8
February 1982 to 8 April 1982); and (3) through-
out the breeding season of 1982-1983 (13 Sep-
tember 1982 to 20 December 1982). The three
breeding pairs observed during these time pe-
riods produced five litters. One female appar-
ently aborted late in gestation of the 1981-1982
breeding season. In addition to information con-
cerning these breeding pairs, I recorded data
concerning one nonbreeding pair each year.
However, the same male was used in both non-
breeding pairs. Although these five pairs formed
the nucleus for my behavioral studies, I was able
to collect limited amounts of information on
three additional nonbreeding pairs also captive
at CRC.
I observed maned wolves from 1 to 4 hours
per day on an average of five days per week (total
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 20.?Maned wolves in an outdoor enclosure at the Conservation and Research
Center (photo taken from observation tower).
474 hours). Observation sessions with each pair
lasted 1 hour and were systematically scheduled
to represent all pairs equally. Sampling proce-
dures included recording all occurrences of cer-
tain behaviors and of scan sampling at 2-minute
intervals for other behaviors (J. Altmann, 1974).
Additional information on sampling procedures
is presented in the following sections.
Statistical analysis of the following behavioral
data generally concerns (1) comparison between
sexes of frequencies of behaviors and (2) com-
parisons between reproductive stages. In the for-
mer case I used the test for difference between
proportions (Bruning and Kintz, 1977:222). In
the latter case I applied a chi-square test for
interaction (Steel and Torrie, 1960:366) in which
expected values were calculated using means
weighted by the appropriate number of hours of
observation for compared stages. In both cases,
data from breeding pairs were analyzed indepen-
dently from those of nonbreeding pairs. Data
from all breeding pairs were summed for each
sex and for each reproductive stage, thus obscur-
ing individual variation.
Operational Definitions of Reproductive Seasons
In view of the lack of histological and physio-
logical information concerning the estrous cycle
in the maned wolf, behavioral information was
used to define four reproductive stages for
breeding pairs. Brady and Ditton (1979) state
that 30 days prior to copulation, adult maned
wolves show an increase in friendly interactions
and in urine marking. I have termed this period
of courtship as "proestrus." It as not possible to
NUMBER 392 29
record all incidences of copulation by breeding
pairs. Therefore, I defined the period (x = 7.8
days) during which a female accepted mounting
by her mate as "sexual receptivity." I refer to the
approximately 55 days following sexual receptiv-
ity and prior to separation of pair-members as
"gestation." Observations were not conducted
during the remaining days prior to birth when
male and female were housed separately. Finally,
the period subsequent to rejoining of male and
female, after weaning and removal of the pups
from the mother, is referred to as "anestrus."
Data concerning nonbreeding pairs were not di-
vided into reproductive stages.
Behavioral Observations
To test the hypothesis that synchrony of activ-
ity does not vary with the reproductive stage I
recorded at 2-minute intervals the activity states
of both wolves. Individuals on their feet were
classified as active; those lying prone were cate-
gorized as inactive. A wolf remaining in a den
for longer than 2 minutes was presumed to be
inactive. Synchrony of activity occurred when
pair-members were either both active or both
inactive.
Information from the field study indicates that
the individual distance maintained between pair-
members changes as a function of reproductive
stage. To establish the timing of these changes I
recorded at 2-minute intervals the distance (in
body-lengths) between wolves that were within
five body-lengths of one another. I reduced the
confounding effects of having only one den avail-
able to some pairs by deleting from the analysis
those observations in which both wolves were
inactive.
Throughout my observations I monitored all
occurrences of 30 specific behaviors used in the
regulation of individual distance. From these I
selected six behaviors that are distance-decreas-
ing and five behaviors that are distance-increas-
ing in function (pp. 30-32). I used the available
literature, interviews with personnel involved
with care and management of maned wolves, and
my own experience to select those behaviors least
ambivalent in function. The summed frequencies
of distance-increasing behaviors and of distance-
decreasing behaviors were compared between
sexes and among breeding stages.
To further understand the relationships be-
tween sexes and how these relationships change
with reproductive stages, I defined and sampled
on an all-occasion basis three types of initiative
behavior: "Initiation of approach" jvas defined
by one wolf approaching to within one body-
length of its mate. Cases in which one wolf did
not clearly initiate the approach were not scored.
"Initiation of activity" was defined by one of two
inactive wolves rising to a standing position. Con-
versely, "initiation of rest" occurred when one of
two active wolves lay down or remained in a den
for longer than 2 minutes.
Defecation, urination, and rubbing by maned
wolves observed in this study were similar to
descriptions of these activities by Kleiman (1967,
1972) and by Kuhme (1975). I recorded all oc-
currences of these behaviors and of straddling
(Brady, 1982). Frequencies of these scent-mark-
ing behaviors were compared between sexes and
among reproductive stages.
Information on free-living maned wolves sug-
gests that individuals of mated pairs are found in
close association only during the breeding sea-
son. However, because females radiocollared in
the Serra da Canastra did not produce viable
litters, it was not possible to determine the timing
of these changes with respect to reproductive
stages. Neither was it possible to identify differ-
ences between sexes in bringing about these spa-
tial changes. To address these questions each
enclosure was divided into from 76 to 102 quad-
rats measuring approximately 2.6 m by 3.3 m in
area. I assigned latitudinal and longitudinal co-
ordinates to each quadrat and recorded at 2-
minute intervals the locations of both animals in
the focal pair. These data were computerized
and the number of locations in each quadrat
determined for each wolf during each reproduc-
tive stage. To reduce the cumulative effects of
errors in location, and to correct for the differ-
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ences in numbers of quadrats per enclosure, I
used the following formula to estimate the per-
cent of an enclosure frequently used by a wolf
and also shared with its mate:
FQ, or the number of frequently used quad*
rats, is defined as those quadrats in which a wolf
of specified sex was located during at least 1
percent of the total number of locations for that
wolf. SFQ, or the number of shared frequently
used quadrats, is defined as those frequently used
quadrats with locations scored for both members
of a pair. Thus M is a sex-specific coefficient of
mutual use of space in the enclosure. An anala-
gous parameter in free-living maned wolves
would be range overlap.
Priority in feeding was recorded during the
period October 1981 through December 1982.
The staff caring for maned wolves noted whether
the male, female, or both wolves initiated feeding
when the afternoon ration was presented. No
score was recorded when feeding was not initi-
ated within 5 minutes after delivering the food.
for nonbreeding wolves was significantly less
(p < 0.001) than that for gestation, the repro-
ductive stage with the lowest relative number of
locations within five body-lengths.
In summary, individual distance as estimated
by the inverse of the frequency of approaches to
within five body-lengths, was at a minimum dur-
ing the period of sexual receptivity and at a
maximum during gestation in breeding pairs.
Individual distance was significantly greater for
nonbreeding pairs than for breeding pairs during
any reproductive stage.
Evaluation of the mean distance between
wolves (< five body-lengths apart) generally sup-
ports the above conclusions. Individuals of
breeding pairs were found closest together dur-
ing sexual receptivity and farthest apart during
gestation. However, the mean distance between
individuals of nonbreeding pairs was approxi-
mately equal to that for breeding pairs during
sexual receptivity. Thus it appears that although
members of nonbreeding pairs were rarely in
close proximity, the distance between them on
these occasions was small in comparison with that
for breeding pairs.
RESULTS
Synchrony of Activity
Results (Table 19) were analyzed for differ-
ences between reproductive stages. Excepting
sexual receptivity x anestrus, all pairwise com-
parisons were significant (p < 0.01). The percent
synchrony among nonbreeding wolves was less
(p < 0.001) than that for proestrus, the repro-
ductive stage with least synchrony in breeding
pairs (p< 0.001).
Individual Distance
The number of instances in which individuals
of breeding pairs were located within five body-
lengths (Table 20) was compared pairwise for the
four reproductive stages. Values for each stage
were significantly different from those of all
other stages (p < 0.001). In addition, the value
Distance-increasing and Distance-decreasing
Behavior
Occurrences of the following behaviors were
monitored throughout the study.
MOUNT.?Distance-decreasing. A wolf, usu-
ally the male, places one or both forepaws over
the back or hips of its mate. Observed most often
in a sexual context but occasionally during play.
PRESENT.?Distance-decreasing. The female
averts the base of the tail exposing the anogenital
region. During estrus this behavior may be ac-
companied by lordosis and/or by "trapping" of
the male. In the latter case the female positions
herself so that the male is forced to pass near
her, e.g., in an open gate. She then turns away
from him and presents. Presenting is rarely seen
outside proestrus and sexual receptivity.
INVESTIGATION OF ANOGENITAL REGION.?
Distance-decreasing. A pair-member sniffs and/
or licks the anogenital region of its mate. This
NUMBER 392 31
behavior was most common during proestrus and
sexual receptivity but was frequently seen at the
onset of activity bouts during all reproductive
stages. Males were observed performing this be-
havior significantly more often than were females
(n= 190; p < 0.001).
INVESTIGATION OF MUZZLE.?Distance-de-
creasing. A maned wolf sniffs and/or licks the
muzzle of its mate. This behavior was often used
in greetings and preceded investigation of the
anogenital region. I occasionally observed muz-
zle investigation following feeding and surmise
that this behavior may transmit information
about the recent feeding history of the recipient.
There was no significant difference between
sexes in the number of observed occurrences of
this behavior (n = 117; p > 0.05).
TAIL-UP, TAIL-OUT.?Distance-decreasing.
These actions form part of a behavioral continu-
um in which the sender elevates its extended tail
a minimum of about 30? to a maximum of about
180? from the pendent position (see Kleiman,
1972:798) for a duration of from one to several
seconds. These tail postures range in function
from low-level appeasement to a compelling in-
vitation for social contact. For example, a maned
wolf passing unusually close to its resting mate
might send a tail-out of 1-second duration. The
tail-up display by maned wolves does not appear
to be related to expression of dominance as in
Canis lupus (e.g., Zimen, 1975) and Canis fami-
liaris (e.g., Fox, 1978), but rather to the regula-
tion of individual distance. Tail-up and tail-out
were both performed significantly more often by
males than by females (tail-up: n = 229, p < 0.01;
tail-out: n = 376, p < 0.001).
YAWN.?Distance-increasing. The yawn was
one of the behaviors most frequently observed
in these maned wolves (n = 811). Yawning was
most often performed by males and was never
observed occurring synchronously or in sequence
by pairs. Thus, Ewer's (1968) hypothesis that
yawning in the Canidae is a social ceremony
serving to rouse and synchronize the moods of
group members is not substantiated by this study.
Yawning apparently functions as a low-level ag-
gressive display and was often illicited when a
pair-member approached unusually close to its
mate. In most cases the wolf receiving the yawn
altered its course to increase distance between
individuals.
AVOIDANCE.?Distance-increasing. I defined
avoidance as one wolf moving out of the way or
clearly altering its course to make way for its
mate. Males avoided females significantly more
often than females avoided males (p < 0.001).
However, these data are not homogeneous across
pairs. In the five observed pairs the percentages
of total occurrences of avoidance behavior con-
tributed by males were 95, 94, 61, 42, and 34.
Thus, in two pairs the majority of avoidance
behavior was exhibited by males, in two pairs
each sex behaved in roughly equal percentages,
and in one pair the female avoided the male
more often than conversely.
Avoidance behavior was at a maximum during
anestrus and at a minimum during sexual recep-
tivity (proestrus =1.15 occurrences/hour of ob-
servation; sexual receptivity = 0.32; gestation =
1.14; anestrus = 1.36).
GAPE.?Distance-increasing. This agonistic
display consisting of opening the jaws, was usually
accompanied by lowering the head, flattening
the ears laterally, and occasionally by growling.
The gape was often used in defense of food or a
favored resting place, or in retribution for an
invasion of individual space. I found no signifi-
cant difference between sexes in the frequency
of occurrence of gapes. Occasionally a wolf began
to gape at its mate and then terminated the
display in a yawn, usually when the stimulus
apparently inducing the gape was suddenly re-
moved, e.g., the approaching mate reversed its
direction. This grading of a gape into a yawn
suggests a motivational relationship between
these two behaviors.
FULL PILOERECTION.?Distance-increasing. A
maned wolf in full agonistic display can erect its
white throat-patch and its dorsal mane from be-
tween its ears to the tip of its tail. This impressive
piloerection may be accompanied by broadside
positioning and by cat-like arching of the back?
both presumably to maximize visual impact. Full
piloerection was defined as the raising of the
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
dorsal mane from the ears to at least the rump
and was observed on 68 occasions with no sig-
nificant difference in frequency detected be-
tween sexes.
BITE.?Distance-increasing. Biting occurred
in three contexts: copulation (never observed
during scheduled observations), intense play, and
intense agonistic encounters. Bites during play
were inhibited and were uncommon. Intense ag-
gressive interactions were usually ritualized and
involved the distance-increasing behaviors de-
scribed above. Uninhibited aggressive bites were
very rare and were usually directed to the recip-
ient's shoulder or flank. No bite wounds were
noted during the 1981-1982 breeding season,
but five were recorded during the following year:
a female with a wound at the base of the tail
during late anestrus; a female with a wound on
the left shoulder during proestrus, and another
on the left flank during early anestrus; a non-
breeding male with two wounds to the left shoul-
der. The increase in the number of adult maned
wolves housed at these facilities during the 1982-
1983 breeding season may have precipitated ag-
gressive interactions.
Total frequencies of distance-increasing and
distance-decreasing behavior for each sex (Fig-
ure 21) were not independent of reproductive
stage (p < 0.005). In addition, the frequency of
distance-increasing behavior by males was sig-
nificantly greater than that for females during
all reproductive stages (p < 0.05). The frequency
of distance-decreasing behavior for males was
also greater during proestrus (p < 0.05) and
anestrus (p < 0.001).
Examining Figure 21 it becomes apparent that
the rates of distance-increasing behavior re-
mained relatively more stable across reproduc-
tive stages than did those of distance-decreasing
behavior. The major variation from the mean
(for all reproductive stages) for distance-increas-
ing behavior was a 43 percent decrease by fe-
males and a 46 percent decrease by males during
sexual receptivity. However, distance-decreasing
behavior during gestation declined from 124
percent above the all-stages mean to 66 percent
below the mean for males, and from 62 percent
above the mean to 62 percent below the mean
for females.
Males of nonbreeding pairs performed pro-
portionally more distance-increasing and dis-
tance-decreasing behavior than did females
(p < 0.001). In addition, neither the percentages
of distance-increasing nor distance-decreasing
behavior exhibited by each sex in nonbreeding
wolves differed significantly (p < 0.01 from those
in breeding pairs (all-stages means). However,
the frequencies of distance-increasing and dis-
tance-decreasing behaviors in nonbreeding
wolves approximated those for the reproductive
oI4
K
Ul
S
2-
DISTANCE - INCREASING
BEHAVIORS
DISTANCE - DECRE ASI NG
BEHAVIORS
imfl
0
 PROESTRUS SEXUAL GESTATION ANESTRUS PROESTRUS SEXUAL GESTATION ANESTRUS
RECEPTIVITY RECEPTIVITY
FIGURE 21.?Total frequencies of selected distance-increasing and distance-decreasing
behavior for three pairs of maned wolves (striped bars = females; solid bars = males).
NUMBER 392 33
stage with the lowest frequencies in breeding
pairs (distance-increasing: nonbreeding male =
0.67 occurrences/hr observation, female = 0.37;
distance-decreasing: male = 0.91, female = 0.43).
In summary, although nonbreeding pairs did not
differ from breeding pairs with respect to relative
rates of behavior performed by each sex, in ab-
solute terms the rates for nonbreeding pairs were
lower than those of breeding pairs.
Initiative Behavior
There was no significant difference between
sexes with respect to the all-stages frequencies
for initiation of approach (p > 0.01; Figure 22).
However, approaches during gestation were ini-
tiated significantly less often by females than by
males (p < 0.001). Frequencies of approach ini-
tiation were highest during sexual receptivity
1.2
OS-
0J6-
0.3-
INITIATE REST
K I.2-.
UJ
? 0.9-
x 0.6-
8 0.3-
INITIATE ACTIVITY
2.0-
1.6-
1.2-
0.8-
0.4-
o
INITIATE
APPROACH
fM
PROESTRUS SEXUAL GESTATION ANESTRUS
RECEPTIVITY
FIGURE 22.?Frequencies of initiative behaviors for three
breeding pairs of maned wolves (striped bars = females; solid
bars = males).
(male = 128 percent above the all-stages mean;
female = 86 percent above the mean) and lowest
during gestation (male = 38 percent below the
mean; female = 76 percent below the mean).
The frequencies of approach initiation for non-
breeding pairs were significantly greater by
males (p < 0.05) and were approximately equal
to those for breeding pairs during gestation (male
= 0.33 occurrences/hr observation; female =
0.23).
Initiation of activity was performed significantly
more often by males than by females (p < 0.001)
irrespective of reproductive stage in breeding
and in nonbreeding pairs, and also for each stage
considered individually in breeding pairs (p <
0.05). Frequencies for nonbreeding pairs were
small in comparison with those for breeding pairs
(male = 0.35 occurrences/hr observation; female
= 0.12).
In contrast, initiation of resting was performed
significantly more often by females than by males
in breeding pairs (p < 0.05) and in nonbreeding
pairs (p < 0.001). The only exception to the
above was during anestrus, when males initiated
rest more often than did females (p < 0.05).
Frequencies for initiation of rest by nonbreeding
pairs were lower than those for breeding pairs
(male = 0.04 occurrences/hr observation; female
= 0.45).
Scent Marking
Straddling consists of the wolf swinging a hind
leg over a bush or tuft of grass taller than its
back. The wolf then advances until the bush
bends beneath its tail and anogenital region. This
behavior was often accompanied by alternately
elevating the hind limbs, by yawning, and by
shaking the mane. Straddling was usually main-
tained for several seconds and until the strad-
dling wolf was in the visual field of its mate. In
four instances I failed to find urine on or beneath
straddled vegetation. However, on various occa-
sions I observed wolves sniffing and/or rubbing
on shrubs recently straddled by their mates.
Thus, it is likely that some olfactory information,
34
perhaps from the anal glands or genitalia, is
deposited on straddled items. I found no signifi-
cant difference in frequency of straddling be-
tween sexes in breeding or nonbreeding pairs
(p > 0.1; Figure 23). There were, however, dif-
ferences in frequencies of straddling among re-
productive stages for both males and females
(p < 0.005). Straddling was most common during
proestrus and least common during anestrus in
breeding pairs. Straddling was very rare in non-
breeding pairs (male = 0.01 occurrences/hr ob-
servation; female = 0.01).
Rubbing by these maned wolves was always
accomplished from a standing position and usu-
ally included sliding the head and neck along an
elevated object or on the ground. I never ob-
served a maned wolf rubbing on feces, but rub-
bing on recently deposited urine, either its own
or that of its mate, was very common. The fre-
quency of rubbing behavior (Figure 23) did not
differ between sexes (p > 0.1), with all breeding
stages combined. However, during proestrus and
gestation females rubbed more than did males
(p < 0.005), and during anestrus males rubbed
more often than did females (p < 0.001). Fre-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
quencies of rubbing were not independent of
reproductive stage for males or females of breed-
ing pairs (p < 0.005). Males rubbed most during
anestrus and least during proestrus; females
rubbed most during gestation and least during
anestrus. Rubbing by nonbreeding wolves did
not differ between sexes (p > 0.01) and was
infrequent in comparison with rates for breeding
wolves (frequency for females = 0.11 occur-
rences/hr observation; male = 0.13).
Each wolf had two or three sites where it
deposited virtually all of its feces. These sites
were usually but not always shared by both pair-
members, and were sometimes used in excess of
two years. With two exceptions (on a table and
on a den roof), middens were located at ground
level. Placing five cross-sections of logs ranging
in height from 10 to 40 cm into enclosures
produced no observable changes in defecation
patterns. Defecation, often accompanied by an
extended-bark vocalization, was a common re-
sponse to the presence of an unfamiliar person
in an enclosure and suggests that defecation has
an aggressive or alarm function in maned wolves.
Also of interest is the observation that maned
f ?
6-
5-
4-
3-
2-
1-
0.7-
0.6
0.5
0.4
0.3
0.2-
0.1
DEFECATION
ri 11 n 1 it 11 il I I I 11 il
STRADDLING
_ _
1
PIT
1?
URINATION | 1
il1n
RUBBING
i
?i
PROESTRUS SEXUAL GESTATION ANESTRUS PROESTRUS SEXUAL GESTATION ANESTRUS
RECEPTIVITY RECEPTIVITY
FIGURE 23.?Frequency of scent marking by defecation, urination, straddling and rubbing by
breeding pairs of maned wolves (striped bars = females; solid bars = males).
NUMBER 392 35
wolf pups practiced locus-specific defecation as
soon as they became mobile, but maintained the
midden separate from that of their female par-
ent.
Defecation by males was more common than
by females for nonbreeding pairs. Males of
breeding pairs defecated more than females dur-
ing all reproductive stages except sexual recep-
tivity (p < 0.001). The lack of significance during
sexual receptivity (0.05 < p < 0.1) may be due
to small sample sizes (12 occurrences for males;
6 for females). Although defecation rates by fe-
males were independent of reproductive stage (p
> 0.1), those for males were not independent of
reproductive stage (p < 0.005). Defecation rates
for both sexes were slightly higher during gesta-
tion and lowest during anestrus in breeding pairs.
Rates for nonbreeding pairs were 0.11 occur-
rences per hour of observation for females, and
0.47 for males.
Urine was most often deposited on elevated
objects throughout the enclosure, but also on
level ground and on walls. Although some objects
were marked more often than others, I only
noted locus-specific urination practiced by fe-
males during gestation, and then usually along
the fences nearest other maned wolves. Se-
quences in which the urine mark of one wolf was
marked, or rubbed and then marked by the mate,
were common throughout the study and were
practiced with approximately equal frequency by
both sexes. With the exception of anestrus for
breeding pairs (p > 0.1), rates of urine marking
were greater for males than for females in non-
breeding and in breeding pairs, when reproduc-
tive stages were combined or when considering
each stage individually (p < 0.001). Rates of
urine marking for males and for females were
found not to be independent of reproductive
stage in breeding pairs (p < 0.005). Rates were
greatest during proestrus for both sexes, least
for females during sexual receptivity, and least
for males during anestrus. Rates of urine mark-
ing by nonbreeding wolves (female = 0.39 occur-
rences/hr observation; male = 1.5) were less
than those for breeding pairs.
Vocalization
Kleiman (1972) and Brady (1981) have de-
scribed the high-amplitude roar-bark or ex-
tended-bark of the maned wolf. The latter au-
thor postulated that this call functions primarily
as a spacing mechanism within sexes. During the
field study, I noted several exchanges of vocali-
zations between pairs and between pair-mem-
bers, indicating that the function of this call
might also be related to localization between and
within pairs.
I intended to determine at CRC the frequency
and duration of vocalization bouts and to identify
participants during each reproductive stage.
However, because my observations were mainly
at night, and I could not identify individual
wolves by their calls, it was often impossible to
identify individual vocalizing wolves. The small-
est identifiable unit was often of the focal pair.
Also, since the reproductive stages of breeding
females were not entirely synchronous, it was
necessary to combine data from the proestrous,
sexual receptivity, and gestation stages into a
single breeding season.
Results (Table 21) show that these wolves vo-
calized more often, engaged in more bouts in-
volving more than one wolf, and used more barks
per occurrence during the combined breeding
stage than during the single anestrus stage (p <
0.005). For those wolves whose sex could be
determined, males vocalized more often than did
females (p < 0.001). Occurrences of vocalization
were more common early in the breeding season
(Oct-Nov) than later in the breeding season
(Dec) suggesting that vocalization was more com-
mon during proestrus and sexual receptivity than
during gestation.
On five occasions a wolf inside a den ex-
changed vocalizations with its mate outside the
den. In three cases a wolf's vocalization outside
a den apparently caused its mate to leave the
den, and in two cases vocalization in one enclo-
sure apparently caused a wolf in an adjacent
enclosure to leave its den. These observations
support the hypothesis that during the breeding
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
season the long-distance call functions primarily
in communication between pair-mates.
Use of Space
Results (Figure 24) show a high coefficient of
mutual use of space (p. 30) by females during
proestrus. Combining information from studies
in the field and in captivity it appears that the
essentially solitary existences maintained during
anestrus are terminated by encroachment of
males into areas formerly used mainly by females.
Levels of mutual use of space by both sexes
remained high, at least until late gestation.
Precedence in Feeding
The results of this survey were not homoge-
neous with respect to initiating feeding by sex.
In two breeding pairs, the females were predict-
ably first to feed (97 and 80 percent of all re-
corded feedings). In the third breeding pair the
male fed first in 87 percent of recorded obser-
vations. Instances of male and female beginning
feeding simultaneously were rare (3, 3, and 10
percent in breeding pairs), and precedence did
not appear to vary as a function of breeding
60 1
5 0 -
4 0 -
3 0 -
2 0 -
10
PROESTRUS SEXUAL GESTATION
RECEPTIVITY
ANESTRUS
FIGURE 24.?Frequency of shared use of enclosure quadrats
by three breeding pairs of maned wolves (striped bars =
females; solid bars = males).
stage. In two of three nonbreeding pairs the male
fed first (86, 69 percent), and in the third pair
priority to feed was relatively evenly split (male
= 42 percent; female = 49 percent). Finally, in
one pair of hand-raised siblings one year of age
at the beginning of this survey, both male and
female initiated feeding together in 100 percent
of recorded feedings.
Caching of food has been described for female
maned wolves prior to parturition (Encke et al.,
1970; Brady and Ditton, 1979). During the pres-
ent study caching by both sexes was observed for
two breeding pairs late in gestation. The behav-
ior pattern was similar for both pair-members:
one pair-member carried a food item for several
minutes, buried it in a shallow depression exca-
vated with forepaws and nose, and moved a few
meters away. Occasionally, the same wolf then
dug up its cached food and repeated the se-
quence. In several cases, however, the wolf's
mate retrieved the cache, carried it for several
minutes and buried it again. This food sharing,
although accompanied by aggressive gaping, pilo-
erection, and short chases, represents a signifi-
cant reversal in the otherwise highly defensive
attitude concerning food possessed by maned
wolves.
Discussion
Evolution of Sociality in a Grassland Habitat
Generalized representatives of the dog family
arriving in South America were exposed to a
variety of selective pressures in their new envi-
ronment. One of these would have been the
necessity of locomotion through the tall grass
prairies covering much of the continent. There
were two evolutionary options open to cursorial
mammals living in this type of habitat: travel
through the grass at ground level or travel high
enough above ground level that resistance due
to body contact with stiff plant stems were re-
duced. Apparently, the maned wolf was the only
South American canid to evolve the latter spe-
cialization.
Long legs necessitate a relatively large body
NUMBER 392 37
size which in turn places increased energy de-
mands on the individual (Brown and Lasiewski,
1972). During the late Pleistocene, when the
maned wolf was dispersing across the savannas
of central South America, most of the large
native herbivores became extinct and were re-
placed ecologically by highly successful cricetid
rodents (Webb, 1969, 1978; Simpson, 1980).
Foraging on this dispersed trophic resource base
would not have been conducive to social living.
The fact that a perennial bond persists between
members of a pair of maned wolves supports the
assertion of Kleiman and Eisenberg (1973) that
the pair bond is a phylogenetically old trait within
the Canidae.
I suggest that the monogamy and strict terri-
toriality (sensu Burt, 1943) displayed by the
maned wolf are the results of long-term ecologi-
cal and genetic constraints. The energetic de-
mands imposed by the large body size of the
maned wolf and its morphological and physiolog-
ical dependence on a food source that is scarce,
dispersed, and relatively dependable may have
favored the selection of a social organization
based on solitary and mutually exclusive forag-
ing. Because of the lack of exposure to factors
promoting sociality, this social structure may now
be relatively inflexible. That is, the genome of
the maned wolf may limit the extent of cooper-
ative social interaction among individuals.
If territoriality promotes survival of maned
wolves, natural selection should favor character-
istics that emphasize the ownership of an area.
Several such morphological and behavioral traits
are displayed by this species. Extended-bark vo-
calizations are loud and far-reaching but not
complex or variable between individuals. This
type of signal is well suited for transmitting sim-
ple messages to all conspecifics at a long distance
from the sender. The urine of maned wolves has
a strong odor and is placed in a manner that
would increase detection by conspecifics at a
distance or long after it has been deposited.
Visual cues used by the maned wolf are discern-
ible at considerable distances: erection of the
dorsal mane, broadside arching of the back, and
contrasts in color of the regions of the body are
used in display. In monogamous species there
would be little need for sexual dimorphism, and
natural selection should favor monomorphism
(Kleiman, 1977) as is found in maned wolves.
The Pair Bond in Maned Wolves
Canid social systems are often highly flexible
in response to a number of factors, including
food availability, human persecution, and popu-
lation status (Macdonald, 1979, 1983). There-
fore, it would be premature to state that the
social organization of the maned wolves of the
Serra da Canastra is "species-typical." However,
based on my studies of free-living and captive
maned wolves I offer the following conclusions
concerning the pair bond in this species.
The majority of canids are monogamous (Klei-
man, 1977) with males of most of these species
assisting in provisioning of young (Wittenberger
and Tilson, 1980). Such direct male parental care
benefits the reproductive fitness of both sexes.
Male maned wolves also occasionally provision
their offspring in captivity (Kleiman and Mal-
colm, 1981). However, information from the
Serra da Canastra indicates that females are not
dependent on direct aid from their mates for the
care of offspring. There is no indication that
groups of maned wolves contribute to the welfare
of the offspring of a single female as in certain
other canid species (Macdonald, 1980; Moehl-
man, 1979). In only two instances during the
field study did local observers see two adult
wolves accompanying pups. All observations of
defense of pups by adults involved only a single
adult. In all cases in which it was possible to
determine the sex of an adult accompanying
pups, the adults were females.
The facultative monogamy (Kleiman, 1977)
seen in maned wolves apparently evolved as a
result of indirect selective factors, such as (1) the
mutual defense of an area large enough to sup-
port parents and offspring prior to dispersal of
the latter; (2) mutual defense of young from
conspecifics or other predators; and (3) reduced
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
levels of intersexual aggression, especially during
the breeding season. Higher rates of vocalization,
scant marking, and initiation of activity by cap-
tive males, and territory acquisition by males of
the Serra da Canastra, indicate that males play a
more prominent role in territorial defense than
do females. Because of the high level of intras-
pecific aggression, male territorial defense is
probably important in the indirect defense of
pups as they begin to travel farther from the
female parent.
In summary, territoriality in maned wolves is
associated with foraging on a dispersed but rela-
tively predictable trophic resource base. Male
and female share the same territory because so-
cial interactions, especially those during the
breeding season, are less costly in established
pairs. The amount of intrapair aggression would
be expected to decrease from year to year in
permanent pairs (Ewer, 1973). Facultative mo-
nogamy may have evolved under circumstances
in which the reproductive fitness of both parents
benefitted more from male territorial defense
than from direct male care of offspring. In ad-
dition, the association of a resident male with his
mate and her litter may increase his reproductive
fitness by decreasing the probability of another
male copulating with that female (Niewold,
1980).
Although the social organization of the maned
wolf appears relatively unflexible, the nature of
the pair bond changes with reproductive stages,
and allows female choice in the extent to which
males directly participate in parental care. Anes-
trus in captive females was characterized by mu-
tual avoidance by pair-members and by reduced
mutual use of space. The frequency of scent
marking, as well as the frequency, duration, and
synchrony of vocalization were at minimum
levels for both sexes. Males maintained maxi-
mum levels of distance-increasing behavior and
initiated rest more often than did females. Data
on vocalization and movements of wolves in the
Serra da Canastra are consistent with these con-
clusions.
During proestrus, urine-marking reached
maximum frequencies for both sexes, suggesting
increased exchange of information between
sexes and/or increased territorial defense. As
the females become more olfactorily attractive,
males exhibited a high level of distance-decreas-
ing behavior, and the coefficient of mutual use
of space increased to a maximum for females.
Thus, proestrus is a period of social approxima-
tion reversing the avoidance characterizing anes-
trus. Because of the important role of this ex-
tended courtship, I would expect cuckoldry to
be rare in free-living maned wolves.
As expected, sexual receptivity features a re-
duction in individual distance and in distance-
decreasing behavior. However, although urine
markings by males continued at a relatively high
rate, frequency of urination by females dropped
to a minimum level. Because in a monogamous
canid with a short estrus the male presumably
accompanies the female, her scent marking is
perhaps less useful than other more overt behav-
iors. In addition, a female's advertisement of
sexual receptivity to more than one male risks
intermale combat and potentially reduces the
likelihood of her conception.
Gestation shows marked changes in relations
between the pair. Individual distance is maxi-
mized and distance-decreasing behavior drops to
a minimum. However, food sharing occurs and
the mutual use of space remains high for both
sexes. The extent to which a female allows her
mate to care for offspring may depend on the
persistence of these traits after parturition.
There was no evidence of a social dominance
hierarchy in these captive pairs. The outcome of
social conflicts between male and female was
generally predictable for each pair of wolves, but
was not generalizable with respect to sex. Domi-
nance was specific to each situation and was based
on norms established by each pair. For example,
one member of a pair had precedence in feeding,
whereas its mate had priority in the use of a
favorite resting place.
Conservation and Management
Most, if not all species are proceeding toward
extinction at varying rates. John Terborgh
NUMBER 392 39
(1974:715) writes: "Our goal as conservationists
is not to enforce a stasis of nature by somehow
stopping the march of evolution; that is impossi-
ble. Rather, it is to prevent, over the period of
transition to a steady state society the process of
species loss from running too far ahead of the
process of species gain." When a species rapidly
approaches extinction, the cause is often a dis-
equilibrium with environmental factors related
to human activities.
Maned Wolves in the Wild
Several traits have been implicated in extinc-
tion-prone species. For example, large body size
and habitat specialization were proposed by
Brown (1971) and Willis (1974) as detrimental
to long-term survival. The maned wolf qualifies
in the first category but not in the second cate-
gory. The geographic distribution of the maned
wolf includes a variety of habitats including grass-
lands, chaco, pantanal, pampas, several types of
cerrado, as well as part of the caatinga of north-
eastern Brazil (Dietz, in press). In addition, the
maned wolf has extended its range into the Zona
de Mata of southeastern Brazil. In the last 100
years, the tropical forest that covered the latter
region has been cut and replaced by farms,
ranches, and commercial conifer and Eucalyptus
plantations. Maned wolves are often observed in
these commercial conifer stands where small
mammals are abundant (Dietz et al., 1975).
Factors such as den sites and resting areas do
not appear to be limiting population density in
this species. Although seasonal variation may re-
duce the availability of some food items, it is
likely that others would be favored and that
maned wolves would take advantage of those
items that are most abundant. The generalist
foraging habits of this canid would tend to buffer
it against population decline due to sudden
changes in food resource composition.
Several authors have stressed the negative ef-
fects of human colonization on population den-
sity of the maned wolf (Dennler de la tour, 1965;
Langguth, 1972, 1975). However, no evidence
is presented in support of these statements. The
wolves and humans of the Serra da Canastra
interacted very little. With the exception of an
occasional wolf killed for stealing poultry, farm-
ers and ranchers were indifferent to wolves. Con-
versely, maned wolves avoided being seen, but
did not avoid the areas inhabited, by humans. In
a few instances, wolves were reported using aban-
doned human habitations and hunting small ver-
tebrates dislodged by tractors. In general, direct
human disturbance, including my activities, ap-
peared to have little effect on the maned wolves
of the Serra da Canastra.
The long-term indirect effects of human activ-
ities on maned wolves are potentially significant.
Practices, such as extensive overgrazing by cattle,
pasture management by annual burning, and the
cutting of brush, and agricultural practices in-
ducing erosion, probably have significantly un-
favorable impact on the resource base available
to these wolves. These effects would be more
severe in regions where the percentage of tillable
land is greater than in the Serra de Canastra.
However, since agriculture has been much the
same in the Serra da Canastra for at least the
past 80 years and maned wolves are still present,
areas such as this will probably retain populations
of wolves for the present.
Populations of free-living maned wolves are
apparently not in immediate jeopardy over much
of their range in South America. However, this
situation could change radically as a result of
large-scale habitat modification taking place in
the cerrado of the Brazilian Highlands. With that
in mind, I recommend the following policies be
adopted throughout the species range of the
maned wolf.
The great majority of biological reserves, such
as state and national parks, will not be large
enough to support populations of maned wolves.
I estimate that no more than 10 maned wolves
live entirely within the confines of the Serra da
Canastra National Park. However, on the order
of 40 wolves may have ranges that overlap both
park and private land. I recommend the estab-
lishment of buffer strips 2 km wide around the
perimeter of parks and reserves within the spe-
cies range of the maned wolf. Landowners within
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
these protected zones could be given tax incen-
tives in return for performing prescribed habitat
management practices such as controlled burn-
ing, grazing, hunting, and clearing of brush. In
the case of the Serra da Canastra National Park,
this action would effectively increase by one-
third the area of the land protected by this park
and perhaps triple its carrying capacity for
maned wolves.
A campaign to increase public awareness con-
cerning the endangered status of the wolf might
significantly reduce mortality of wolves stealing
poultry. A campaign of this nature is likely to
succeed in Brazil for two reasons: first, the maned
wolf does not have the negative public image that
large canids have in the United States; second,
Brazilians are nationalistic and, therefore, likely
to preserve an animal that they perceive to be
part of their national heritage.
Maned Wolves in Captivity
Several facts indicate that the future of the
maned wolf in captivity may not be entirely se-
cure. First, fewer than 100 individuals are ex-
hibited in 12 zoos worldwide (Bush, 1980). Since
the United States Endangered Species Act of
1973, and the International Convention on En-
dangered Species, also in 1973, few wild-caught
maned wolves have entered captive populations
in countries outside South America. Second, suc-
cessful reproduction by maned wolves in captiv-
ity is improving but is still relatively rare (C.
Carvalho, 1976). Although 105 births were re-
corded in zoos during the period from 1969 to
1973 (Roeben, 1975), a high percentage of these
resulted in early mortality. Third, due to the
high incidence of diseases, parasitization, and
problems associated with the social stress found
in maned wolves in captivity, mortality of adults
is also relatively high in this species. Mortality
has been particularly high in some of the smaller
South American zoos.
I suggest that increased numbers of zoos at-
tempt to breed maned wolves in captivity using
techniques recommended by Brady and Ditton
(1979). The registration and cooperative ex-
change of maned wolves, especially in South
America, should be encouraged. A monitored,
low-level introduction of wild-caught individuals
into the captive population would aid in main-
taining genetic diversity and would not at present
be detrimental to free-living populations. It has
been suggested that introduction of as few as one
or two individuals per generation into a small
population can counter the effects of genetic
drift (Avery, 1978; Franklin, 1980). Finally, the
prevalence and potentially fatal nature of cystin-
uria in captive maned wolves (Bovee et al., 1981)
make this disease a possible serious threat to this
species in captivity. Whenever possible, breeding
wolves should be selected from among noncystin-
uric individuals. I also recommend additional
research on the genetic basis, treatment and fre-
quency of occurrence of this disease in free-living
populations.
Appendix
TABLE 1.?Hematologic values for 10 maned wolves (see p. 10; dashes indicate no data).
Determination
Hematocrit
Hemoglobin
Erythrocytes(10H)
Leucocytes (10s)
Myelocytes
Metamyelocytes
Band cells
Segmented cells
Eosinophils
Basophils
Lymphocytes
Monocytes
A
lp
ha
 
(9
-
-
_
-
0
0
2.0
74.0
3.0
0
19.0
2.0
D
el
t;
47.0
15.8
8.61
14.1
0
0
6.0
77.0
3.0
0
14.0
0
6*
?
Ep
sil
Zo
o
Zo
o
(P)
Be
ta
BU
I
G
am
PERCENT OF BLOOD VOLUME
41.5 32.0 37.5 36.5 37.0
GRAMS PER 100 MILLILITERS
14.0 10.2 12.0 11.2 10.8
CELLS PER CUBIC MILLIMETER
6.60
12.3
5.78
14.1
6.87
17.75
5.83
14.8
6.04
12.15
(P)
bd
a
La
m
37.0
11.8
6.14
15.4
PERCENT OF WHITE BLOOD CELLS
0
0
5.0
74.0
7.0
2.0
12.0
2.0
0
0
1.0
49.0
8.0
0
42.0
0
0
0
3.0
53.0
6.0
0
35.0
3.0
0
0
2.0
74.0
14.0
0
9.0
1.0
0
0
5.0
70.0
2.0
0
20.0
3.0
0
0
3.0
66.0
7.0
0
22.0
2.0
?
3
u
N
37.5
12.0
5.06
12.15
0
0
1.0
78.0
2.0
0
17.0
2.0
?
bo
37.0
11.6
5.90
14.8
0
0
2.0
66.0
8.0
0
21.0
3.0
X
38.1
12.2
6.3
14.17
0
0
3.0
68.1
6.0
0.2
21.1
1.8
SD
4.11
1.72
1.00
1.83
0
0
1.76
9.93
3.71
0.63
10.18
1.14
TABLE 2.?Biochemical serum values for seven maned wolves (the blood sample for maned
wolf Epsilon hemolyzed prior to analysis and therefore was excluded from calculations of
sample mean and standard deviation; dashes indicate no data).
Determination
Total Protein
Albumin
Globulin, <*?
Globulin, aa
Globulin, 0
Globulin, y
SGPT(IU/l;R-f)*
SGOT(IU/l;R-f)*
Delta
(?)
-
_
_
-
_
_
51.0
200.0
Epsilon
(?)
Beta
(<$)
GRAMS PER
14.0 8.8
Gamma
(<5)
Lambda
(<5)
100 MILLILITERS
- 7.6
PERCENT OF TOTAL PROTEIN
22.0
7.0
10.0
55.0
6.0
-
-
26.0
12.0
13.0
36.0
13.0
50.0
66.0
_
-
-
-
-
47.0
92.0
21.0
11.0
12.0
41.0
15.0
58.0
95.0
Zeta
<<5)
6.4
22.0
7.0
10.0
42.0
19.0
49.0
68.0
Sigma
(<5)
7.8
24.0
12.0
14.0
36.0
14.0
-
-
X
7.7
23.3
10.5
12.3
38.8
15.3
51.0
104.2
SD
0.98
2.22
2.38
1.71
3.20
2.63
4.18
55.18
* Reitman-Frankel procedure.
41
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 3.?Weights and measurements of captured maned wolves (M = middle-aged, O = old,
Y = young; dashes = no data).
Character
Weight (kg)
Length (cm)
Head and body
Tail
Ear
Hindfoot
Forelimb1
Hindlimb2
White on
tail
Black on
antebrachium
Black on
hindlimb
Black stripe
on mane
Alpha
(M9)
25.1
99.0
49.0
16.0
29.5
85.0
80.0
_
_
-
-
Beta
(MS)
22.4
95.0
45.0
17.0
29.0
87.0
85.0
_
_
-
-
Gamma
(MS)
25.8
107.0
s
17.0
30.0
91.0
96.0
_
_
-
-
Delta
(O9)
21.3
104.0
43.0
16.5
30.0
91.0
84.0
_
_
-
-
Epsilon
(YS)
20.5
105.0
38.0
17.0
29.0
86.0
81.0
18.0
34.0
37.0
49.0
Kappa
(Y9)
24.0
104.0
46.0
15.0
30.5
90.0
95.0
8.0
50.0
38.0
47.0
Lambda
(Y(J)
23.1
99.0
45.0
18.0
28.0
91.0
92.0
19.0
46.0
37.0
40.0
Zeta
(MS)
23.1
100.0
43.0
17.5
27.5
89.0
91.0
23.0
44.0
38.0
45.0
Sigma
(MS)
24.4
114.0
47.0
17.5
32.0
96.0
99.0
31.0
47.0
37.0
54.0
x ? SD
23.3 ? 1.73
103.0 ?5.57
44.5 ? 3.30
16.8 ?0.90
29.5 ? 1.35
89.6 ? 3.32
89.2 ? 6.92
19.8 ? 8.35
44.2 ?6.10
37.4 ? 0.55
47.0 ?5.15
1
 Measured from the manus to the dorsal border of the scapula.
2
 Measured from the pes to the cranial crest of the ilium.
3
 Extremity missing.
TABLE 4.?Use (%) of three habitats by maned wolves as determined by number (N) of
radiolocations.
Habitat
Grassland
Cerrado
Forest
Total
N
97
89
62
248
Males
%
39.1
35.9
25.0
100.0
Females
N
66
116
51
233
%
28.3
49.8
21.9
100.0
All wolves
N
163
205
113
481
%
33.9
42.6
23.5
100.0
TABLE 5.?Use (%) of three habitats by bonded pairs of maned wolves as determined by
number (N) of radiolocations.
Habitat
Grassland
Cerrado
Forest
Total
Pair
N
104
163
62
329
1
%
31.6
49.5
18.8
99.9
N
17
11
30
58
Pair 2
%
29.3
19.0
51.7
100.0
N
27
24
20
71
Pair 3
%
38.0
33.8
28.2
100.0
NUMBER 392 43
TABLE 6.?Seasonal use (%) of three habitats by maned wolves as determined by number (N)
of radiolocations.
Habitat
Grassland
Cerrado
Forest
Total
Wet seasons
N
66
104
69
239
%
27.6
43.5
28.9
100.0
Dry seasons
N
97
101
44
242
%
40.1
41.7
18.2
100.0
All seasons
N
163
205
113
481
%
33.9
42.6
23.5
100.0
TABLE 7.?Use of available habitats with respect to their relative proportions in the study
area (ns = not significant).
Habitat
(Proportion of
study area)1
Number of
radiotelemetric
observations
Proportion of
total
observations
Confidence
interval
on proportion
of occurrence2
Significant
deviation from
expected
proportion
Grassland (0.50)
Cerrado (0.35)
Forest (0.15)
Grassland (0.50)
Cerrado (0.35)
Forest (0.15)
108
172
80
21
21
33
0601-1800 HOURS
0.30
0.48
0.22
1801-0600 HOURS
0.28
0.28
0.44
0.22 < p < 0.38
0.39 < p < 0.57
0.15 < p < 0.29
0.20 < p < 0.36
0.20 < p < 0.36
0.35 < p < 0.53
less
greater
less
ns
greater
1
 Proportions of habitat types in the study area also represent expected proportions of use by maned wolves of those
habitats. The proportions are tested against confidence intervals placed on observed proportions of occurrence.
2
 Confidence intervals are based on the Bonferroni normal statistics (Miller, 1966); p = 0.01.
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 8.?Levels of activity (%) of eight maned wolves grouped by bonded pairs as
determined by number (N) of radiolocations.'
Activity
level
Inactive
Intermittent
Constant
Total
Pair
N
302
99
89
490
1
%
61.6
20.2
18.2
100.0
N
28
7
15
50
Pair 2
%
56.0
14.0
30.0
100.0
Pair
N
101
27
38
166
3
%
60.8
16.3
22.9
100.0
N
36
13
9
58
Pair 4
%
62.1
22.4
15.5
100.0
1
 Chi-square value = 6.91, 6 d.f., p = 0.33.
TABLE 9.?Levels of activity (%) of eight maned wolves (grouped by sex) as determined by
number (N) of radiolocations.1
Activity
level
Inactive
Intermittent
Constant
Total
Females
N
267
74
66
407
%
65.6
18.2
16.2
100.0
N
200
72
85
357
Males
%
56.0
20.2
23.8
100.0
All
N
467
146
151
764
wolves
%
61.1
19.1
19.8
100.0
1
 Chi-square test value = 8.80, 2 d.f., p = 0.012. When intermittently active and constantly
active observations are combined, G-test value = 7.35, 1 d.f., p < 0.01.
TABLE 10.?Capture dates for maned wolves in three home ranges of the Serra da Canastra.
Wolf
(sex)
Beta (6)
Gamma (S)
Delta (9)
Epsilon (9)
Lambda (<$)
Kappa (9)
Zeta (<$)
Sigma (<5)
Initial
capture
Last location
or date of
death
TAPERAO RANGE
lOSep 1978
15 Dec 1978
18 Jan 1979
22 Apr 1979
15Jun 1979
15 May 1979
5Jul 1979
18 Nov1979
24 May 1980
EAST RANGE
25Jun 1979'
25 Feb 19792
25 May 1980
25 May 1980
WEST RANGE
25 May 1980
17Oct 1979 to
26 Nov 1979s
25 May 1980
Number of
captures
7
4
2
5
22
4
1
2
1
 Radiotelemetric contact was lost with this wolf on this date. It was assumed that the animal
left the study area.
2
 This wolf died as a result of suffering a broken leg.
3
 This wolf was killed by a local farmer sometime during this period.
NUMBER 392 45
TABLE 11.?Substrate used by maned wolves for the
deposition of feces.
TABLE 13.?Plant material identified in 740 scats of
maned wolves.
Substrate
Rocks
Termite mounds
Paths or roads
Dirt mounds
Clumps of grass
Log
Wolf trap
Substrate
not recorded
Total
No. of
Locations
257
206
202
16
4
1
1
53
740
(%)
34.7
27.8
27.3
2.2
0.5
0.1
0.1
7.2
99.9
TABLE 12.?Animal material identified in 740 scats of
maned wolves.
Food item
Small mammals
Bird bones and feathers
Armadillos
Miscellaneous insects
Beetles (Coleoptera)
Grasshoppers
Rabbits
(Sylvilagus brasiliensis)
Ants
Chickens
(Gallus gallus)
Bird egg shells
Snakes
Giant anteaters
Maned wolf, hair
Fish
Bees or wasps
Collared anteaters
Pacas
Ticks
Spiders
Termites
Unidentified
Total
N
524
230
63
43
31
28
23
10
10
7
6
6
5
3
2
2
1
1
1
1
7
997
Proportion of
occurrence1
0.255
0.112
0.031
0.021
0.015
0.014
0.011
0.005
0.005
0.003
0.003
0.003
0.002
0.002
0.001
0.001
0.001
0.001
0.001
0.001
0.003
0.491
Food item
Lobeira
(Solanum lycocarpum,
Solanaceae)
Miscellaneous grass
(Gramineae)
Gravata
(Bromelia anthiacantha,
Bromeliaceae)
Coquinho
(Astrocaryum sp., Palmaceae)
Casaco
(Unidentified, Myrtaceae)
Melancia do campo
(Unidentified, Cucurbitaceae?)
Goiaba
(Psidium sp., Myrtaceae)
Mangue
(Callophylum brasiliense,
Guttiferae)
Articum rasteiro
(Annona sp., Annonaceae)
Gabiroba
(Campomanesia sp., Myrtaceae)
Corn
(Zea mays, Panicoideae)
Araca
(Psidium sp., Myrtaceae)
Aperta-mao
(Unidentified, Melastomaceae
or Piperaceae?)
Limaozinho
(Unidentified, Rutaceae)
Amouripi
(Byrsonima sp., Malpighiaceae?)
Amora
(Moms sp., Moraceae)
Cega olho
(Unidentified)
Pitanga
(Eugenia uniflora, Myrtaceae)
Araca grande
(Psidium sp., Myrtaceae)
Articum grande
(Annona sp., Annonaceae)
Unidentified
Total
N
670
228
44
25
16
12
10
6
6
5
5
4
3
2
2
2
1
1
1
1
20
1044
Proportion of
occurrence1
0.326
0.111
0.021
0.012
0.008
0.006
0.005
0.003
0.003
0.002
0.002
0.002
0.002
0.001
0.001
0.001
0.001
0.001
0.001
0.001
0.010
0.510
1
 Expressed as a function of total occurrences of all items
(2056) in all examined scats.
1
 Expressed as a function of total occurrences of all items
(2056) in all examined scats.
TABLE 14.?Proportions of volume and occurrence of
selected food items in 740 scats of maned wolves.
Food item
Sol an urn Ixcocarpmn
Small mammals
Birds
Foliage
Miscellaneous fruit
Insects
Armadillos
Rabbits
Unidentified material
Soil
Amphibians, reptiles
Giant anteaters
Fish
Collared anteaters
Pacas
Proportion of
total volume
0.576
0.281
0.023
0.013
0.068
0.003
0.016
0.009
0.004
0.006
0.000
0.001
0.000
0.001
0.000
Proportion of
occurrence1
0.326
0.254
0.122
0.108
0.063
0.052
0.032
0.011
0.007
0.005
0.003
0.002
0.001
0.001
0.000
1
 Expressed as a function of total occurrences of all items
(2056) in all examined scats.
TABLE 16.?Weights and measurements of two captured
female maned wolf pups.
Character
Weight (kg)
at capture
at release
Length (cm)
Total
Tail
Ear
Right hindfoot
Pup Al
4.5
5.9
81
21
14
17
25 Aug 1978
lOSep 1978
PupA2
3.9
5.4
69
19
12
16
TABLE 15.?Seasonal occurrence of selected food items in
740 scats of maned wolves (significantly larger frequencies
are underlined).
Food item
Lobeira
Small mammal
Bird
Foliage
Miscellaneous
fruit
Insect
Armadillo
Total
Dry season
N
375
306a
120
147h
50
43
29
1070
Proportion
of total
0.337
0.275
0.108
0.132
0.045
0.039
0.026
0.962
Wet season
N
304
204
124
70
76b
61"
35
874
Proportion
of total
0.340
0.228
0.139
0.078
0.085
0.068
0.039
0.977
a
 p < 0.025; b p < 0.005.
TABLE 17.?The results of selected laboratory examinations
performed on urine samples from eight captured maned
wolves (a plus indicates a positive result; minus indicates a
negative result; laboratory procedures were performed by
the School of Veterinary Medicine, University of Pennsyl-
vania).
Maned
wolf
Beu(d)
Gamma (6)
Delta (9)
Epsilon (9)
Kappa (9)
Lambda (<J)
Zeta (<5)
Sigma (<5)
Cystinuria
Dibasic
amino-
aciduria Glucosuria
TABLE 18.?Endoparasites observed in fecal samples from eight maned wolves (analyses were
performed using a sedimentation method at the Clinical Laboratory of the School of Veterinary
Medicine, Federal University of Minas Gerais; the degree of parasitic infestation, as estimated
by the relative number of eggs or cysts in the examined feces, is represented by from one to
three pluses).
Endoparasite Alpha Beta Beta
1
 Gamma
(<5) (<5)
Zoo 1 Zoo 2 Epsilon
(?)
Lambda Zeta
Trirhuris sp.
Toxocara sp.
Dipxlidium sp.
Ancxlostoma sp.
Oxxuris sp.
Eiweria sp.
' Second capture.
NUMBER 392 47
TABLE 19.?Synchrony of activity by males and females of paired maned wolves.
Reproductive
stage
Breeding Pairs
Proestrus
Sexual
receptivity
Gestation
Anestrus
Nonbreeding Pairs
Pairs
(N)
3
3
3
2
2
Reproductive
stages
(N)
5
6
6
2
-
Observations
(N)
1770
750
3540
2490
2280
Intervals with
synchronous
activity
(%)
65.0
76.4
68.6
75.3
58.0
TABLE 20.?Distance between males and females of paired maned wolves.
Reproductive
stage
Breeding Pairs
Proestrus
Sexual
receptivity
Gestation
Anestrus
Nonbreeding Pairs
Pairs
(N)
3
3
3
2
2
Reproductive
stages
(N)
5
6
6
2
-
Observation
intervals
(N)
1770
750
3540
2490
2250
Intervals when
wolves within
5 body-lengths
(%)
12.5
21.9
9.5
15.9
3.8
Mean distance
(body-lengths)
between wolves
that were within
5 body-lengths
2.8
2.5
3.7
3.5
2.6
TABLE 21.?Total rates of occurrence of extended-bark
vocalization per hour of observation (numbers in parenthe-
ses indicate total number of observed occurrences).
Character
Total instances of
vocalization
Bouts with more than
one wolf
Total barks
Males vocalizing
Females vocalizing
Vocalizing wolves of
undetermined sex
Breeding
season(N = 202 hours)
0.96
(193)
0.34
(68)
10.36
(2092)
0.26
(53)
0.08
(17)
0.61
(123)
Anestrus(N = 124 hours)
0.06
(8)
0
(0)
0.43
(53)
0.01
(1)
0.01
(1)
0.05
(6)
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