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Comparative social ecology of feral dogs and wolves
L. BOITANI and P. CIUCCI
Dipartimento di Biologia Animale e dell’Uomo, Università di Roma “La Sapienza”,
Viale dell’Università 32, 00185 Roma, Italy
Received 26 August 1994, accepted 29 November 1994
In this paper we compare some socio-ecological traits of feral dogs and
wolves in order to assess the social ecology of feral dogs in terms of its adaptive
value in the natural environment, and to evaluate to what extent the domestica-
tion process altered the wolf’s socio-ecological patterns. Referring to feral dogs
as those dogs living in a wild state with no food and shelter intentionally provid-
ed by humans, and showing a continuous and strong avoidance of direct human
contacts, we review the currently available information on feral dog ecology, and
particular reference is made to a 3-year term project on feral dog ecology in
Abruzzo, Italy. Through comparison of relevant behavioural and ecological fea-
tures of both wolves and feral dogs, we hypothesize that some aspects of the
feral dogs’ ecology, having escaped natural selection pressures, represent primar-
ily expression of “evolutionary inertia” or an epiphenomena of artificial selec-
tion. Fitness-related measures of sociality, demography, reproduction, space-use,
activity patterns, and feeding ecology in feral dogs tend to support our original
hypothesis: feral dogs are not reproductively self-sustaining, suffer from high
rates of juvenile mortality, depend indirectly upon humans for food, co-optable
individuals, and space, and their demography appears dominated by unpredict-
able mechanisms. However, further research is needed, especially concerning dif-
ferent ecological conditions and multi-generational time-scales, as well as the
role that dominant breed-types and cross-breeding history within feral dog
groups might play in the expression of the analyzed socio-ecological features.
KEY WORDS: Canis, wolf, canid evolution, social ecology, feralization, domestica-
tion.
Introduction . . . . . . . . . . . . . . . . . 50
Feral dogs and the feralization model . . . . . . . . . . 51
Demography and group composition . . . . . . . . . . 53
Social system . . . . . . . . . . . . . . . 53
Social unit . . . . . . . . . . . . . . . . 54
Group size and density . . . . . . . . . . . . . 54
Reproduction and life histories . . . . . . . . . . . . 56
Breeding . . . . . . . . . . . . . . . . 56
Denning and parental care . . . . . . . . . . . . 57
Litter size and pup survival . . . . . . . . . . . 58
Sex ratio . . . . . . . . . . . . . . . . 59
Ethology Ecology & Evolution 7: 49-72, 1995
50 L. Boitani and P. Ciucci
Space use patterns . . . . . . . . . . . . . . . 60
Home-range . . . . . . . . . . . . . . . . 60
Territoriality . . . . . . . . . . . . . . . 61
Activity patterns . . . . . . . . . . . . . . . . 62
Food sources and predation . . . . . . . . . . . . . 63
Concluding remarks . . . . . . . . . . . . . . . 65
Acknowledgments . . . . . . . . . . . . . . . 67
References . . . . . . . . . . . . . . . . . 67
INTRODUCTION
Feral dogs probably existed in Eurasia soon after the beginning of dog domes-
tication, as a result of the high integration of Mesolithic human cultures and the
natural environment, and of the many opportunities the dogs had to move in and
out of human settlements. Also on the North American continent feral dogs are
believed to have existed well before European contact (MCKNIGHT 1964). There are
at least two notable examples showing that the feralization process was already
going on several millennia ago: the dingo in Australia and its likely ancestor the
pariah dogs of all southern Eurasia (ZEUNER 1963; BRISBIN 1974, 1977; CLUTTON-
BROCK in press). In the 18th century, stray and feral dogs roaming many of the
large cities of the Mediterranean basin (Istanbul, Alexandria) were consistently
reported by many authors, and indeed described almost as separate subspecies (cf.
BREHM 1893). Warm climates and plenty of food resources have probably favoured
the subsistence of dog populations around villages and towns, shifting through the
house-stray-feral conditions. In particular, Mediterranean lifestyles and environ-
mental conditions appear very favourable to the support of stray and feral dog pop-
ulations (relatively warm climate, small game, free-ranging livestock, garbage
dumps, loose interests and controls on the “commons”) (BOITANI & FABBRI 1983). In
a nation-wide dog census conducted in Italy in 1981, BOITANI & FABBRI (1983)
revealed that feral dogs, i.e. those domestic dogs that live without any direct con-
tact with and dependence on humans, were estimated to be about 80,000, repre-
senting about 10% of the total free-ranging dog population, which includes also
stray dogs and all those left by their owner to move freely in and out of villages
into the surrounding areas. In spite of their significant impact on human and natu-
ral environments, free-ranging dogs have rarely been investigated until recently and
few studies of their ecology have been reported (BECK 1973; SCOTT &CAUSEY 1973;
NESBITT 1975; CAUSEY & CUDE 1980; BARNETT & RUDD 1983; DANIELS 1983a, 1983b;
GIPSON 1983; DANIELS & BEKOFF 1989a, 1989b; BOITANI et al. in press).
Wolves and dogs are often given separate names (i.e., Canis lupus for the
wolf, and Canis familiaris for the dogs), but they are, by all taxonomic criteria, the
same species, and it is now widely accepted that the wolf is the ancestor of all dog
types. Almost 12,000 years of selection by humans has amplified the phenotype
diversity of the dogs, as a result of natural and artificial selection. While the dog’s
fitness appears high when tested in its “natural” human context, little is known of
the dog’s performance when matched by natural selection forces alone.
In this paper we compare ecological and life-history traits of feral dogs and
wolves; analysis of their differences and similarities might help understand to what
extent the domestication process altered the wild progenitors’ behavioural and eco-
logical patterns, and affected the dog’s fitness in a natural environment. We focus
51Social ecology of feral dogs and wolves
primarily on feral dog groups whose existence in the wild is recent (i.e., a few gen-
erations) and who do not belong to populations which have completed a de-domes-
tication process in an evolutionary perspective (PRICE 1984). Dingoes and pariah
dogs are therefore excluded from the analysis having been exposed to natural selec-
tion for a number of generations significant enough to attain a rather stable “wild”
phenotype: dingoes, having totally lost any domesticity, are often no longer consid-
ered feral (PRICE 1984). However, given (i) relatively brief (in terms of generations)
exposure of feral dog groups to natural selection forces, and (ii) the degree of both
intra- and inter-group individual variability due to differences in breed-types and
cross-breeding history, we believe that one cannot reasonably expect any adaptive
value of feral dog socio-ecological traits. It is also obvious that the great variability
observed in the 350 and more different dog breeds might play a substantial role in
determining the degree of expression of some socio-ecological traits (e.g., social
attitudes, territorialism, etc.). However, even though information on breed influ-
ence on feral dogs’ biology is not yet available, “pure” modern breeds are rarely
found in well-established feral dog groups, where one might assume extreme breed
tendencies to be selected against.
The perceived differences in ecological strategies as revealed by feral dog
groups and wolf packs living in the natural environment, might be viewed as a test
of our understanding of the evolutionary and adaptive values of the ecological
traits in the wolf, as well as in other social, wild canids. Our underlying hypothesis
is that many aspects of feral dog ecology, being the results of relaxed natural selec-
tion forces, represent primarily expressions of “evolutionary inertia” and/or results
and epiphenomena of artificial selection in the dog.
In our comparative approach, we focus on those behavioural and ecological
features of both wolves and feral dogs for which data are available and for which
similar methodologies have been adopted. Particular reference is made to a re-
search programme carried out on a group of feral dogs from 1984 to 1988 by
means of radio-tracking in a mountainous area of the central Apennines (Abruzzo,
Italy) (ANDREOLI 1987, CIUCCI 1987, FRANCISCI et al. 1991), and whose results are
comprehensively reported by BOITANI et al. (in press). In particular we analyze
parameters such as demography and sociality, reproduction and life-history, space-
use patterns, activity patterns and feeding habits.
Although limited data on feral dog ecology are currently available, we have
attempted a critical review of the existing literature. However, the same problems
found in the study of domestication such as the geographic variation in biological
characteristics and the choice of what constitutes a “representative” study popula-
tion (PRICE 1984) might apply in our case, therefore limiting the generalization of
our conclusions.
FERAL DOGS AND THE FERALIZATION MODEL
Feral dogs are not a homogeneous category of animals. One of the critical dif-
ficulties encountered in conducting feral dog research is the determination of the
true status of the dog being investigated, and several different definitions have been
proposed (CAUSEY & CUDE 1980; BOITANI & FABBRI 1983; DANIELS & BEKOFF 1989a,
1989b). The distinction among feral, stray, and other free-ranging dogs is some-
times a matter of degree (NESBITT 1975). Dogs categories have been classified on
52 L. Boitani and P. Ciucci
the basis of behavioural or ecological traits (SCOTT & CAUSEY 1973, CAUSEY & CUDE
1980); the dog’s origins (DANIELS & BEKOFF 1989a, 1989b); the dog’s main type of
range (rural vs urban free-ranging: BERMAN & DUNBAR 1983; those having unre-
stricted access to public property: BECK 1973); the dog’s kind and level of human
dependency (WHO 1988). BOITANI et al. (in press) defined feral dogs as those living
in a wild and free state with no direct food or shelter intentionally supplied by
humans (CAUSEY & CUDE 1980), and that did not show any evidence of socialization
to humans (DANIELS & BEKOFF 1989a), but rather a strong continuous avoidance of
direct human contacts. Visual and radiotracking observations were used to distin-
guish feral dogs from other free-ranging dogs. This diversity of definitions contrib-
utes to the difficulties of comparing results from different studies. Further com-
plexity arises from considering feralization from the evolutionary perspective,
where it has been viewed as the domestication process in reverse (HALE 1969, BRIS-
BIN 1974, PRICE 1984), or as a behavioural ontogenetic (developmental) process
(DANIELS & BEKOFF 1989c): the two interpretations focus on different levels (popula-
tion and individual) and imply different temporal scales as well as different theo-
retical and research approaches (DANIELS & BEKOFF 1989c).
Indeed, most authors agree that “owned”, “stray” and “feral” dogs are not clo-
sed categories and that dogs may change their status throughout their life (SCOTT &
CAUSEY 1973, NESBITT 1975, HIRATA et al. 1987, DANIELS 1988, DANIELS & BEKOFF
1989a), supporting DANIELS & BEKOFF (1989c) view of feralization as a behavioural
ontogenetic process, sometime occurring within an individual lifetime. Only three
dogs out of the 11 adults studied by BOITANI et al. (in press) were certainly born in
the wild, while the others were recruited from the village populations, shifting from
a stray to a feral condition. Shifting status may depend upon several natural and
artificial causes (Fig. 1): a dog can become stray by escaping human control, by
abandonment or simply by being born to a stray mother (BECK 1975). A stray dog
can become feral when forced out of a human environment or when co-opted or
simply accepted by a feral group existing nearby (DANIELS 1988; DANIELS & BEKOFF
1989a, 1989c), like the majority of group members studied by BOITANI et al. (in
press). In the same study, it was found that some free-ranging dogs could display
behaviours and attitudes intermediate to those expected on the basis of the pro-
posed categories. This suggests that change of status in dogs is not always radical
and abrupt, but rather, depending on local stimuli and conditions, might require a
Fig. 1. — The feralization model.
53Social ecology of feral dogs and wolves
significant portion of an individual life-span. A change in local conditions might
force an individual dog to reverse its tendencies. The way back (i.e., toward the
“house” category) may be observed when a stray dog is adopted by humans. The
further step (i.e., from feral to stray or even owned condition), although generally
unlikely, has been observed by BOITANI et al. (in press), and more recently experi-
mentally demonstrated by one of us (P. CIUCCI unpubl.) by rehabilitating a feral dog
to a domestic status (both cases referring to individuals who, although living as
feral dogs, were not born in the wild). However, the evidence so far collected sug-
gests that, when feral dogs live in socially independent units (i.e., they are socially
bonded with other dogs), and in the absence of interference from humans, it is
highly unlikely for them to reverse their status (i.e., the feralization process is rein-
forced through new generations). From this perspective, our definition of feral dogs
(cf. BOITANI et al. in press) matches DANIELS & BEKOFF (1989c) view that feralization
occurs through the development of a fear response to humans, and does not neces-
sarily involves a significant genetic divergence from their domestic ancestors.
DEMOGRAPHY AND GROUP COMPOSITION
Social system
The typical social system found in wolves is a linear hierarchy structure
extended to all members of the social unit (males and females), where dominant-
subordinate distances are regulated on the basis of individual agonistic behaviour
(MECH 1970, ZIMEN 1982). The wolf social structure, where dominance is reflected
both in terms of privilege (e.g., “peck order”) and initiative (travelling, hunting, terri-
torial defence, reproduction, etc.) appears to allow for a higher functional integra-
tion among individual members than the mere aggregation of individual behaviours.
Indeed, forms of social control might affect individual tendencies to increase the
group’s functionality and survival. For example, social control on reproduction (i.e.,
deferred reproduction, PACKARD et al. 1983) regulates the number of reproducing
individuals within the group and, at the same time, might increase the chances of
pup survival through availability of non-reproducing adults (HARRINGTON et al. 1983).
In the case of feral dogs the social structure appears to be essentially an aggre-
gation of monogamous breeding pairs and their associates (pups and/or subadults of
pair members). Agonistic behaviour, which has been observed in ritualized forms
similar to those of wolves (L. BOITANI et al. unpubl. data), does not seem to extend
over the individual level and does not seem to translate into a higher social structure
(i.e., the hierarchical scale of wolves) that includes all individuals and exerts forms
of social control on group activity (e.g., in reproduction). It is not clear if the lack of
a higher social organization is related to the nature, the intensity and the frequency
of agonistic confrontations among individuals in feral dog group, or rather to the
lack in the dog of the same social susceptibility, in terms of behaviour and physiolo-
gy, which appears to be critical in wolf sociality, or to all these. Indeed, in the case of
feral dogs the observed intraspecific social behaviour might simply reflect the influ-
ence of human contacts during its early phases (SCOTT & FULLER 1965). This is even
more important in dogs living under feral conditions but who have been recruited
from the house and/or stray categories. If this were true we would expect, as the
54 L. Boitani and P. Ciucci
number of generations under feral conditions increases, for social attitudes in feral
dogs to resemble more closely those of wolves. Also, the extent different breeds influ-
ence the social behaviour should be carefully considered.
Social unit
Among canids, packs are social units that hunt, rear young and protect a
communal territory as a stable group (MECH 1970); and their members are usually
related individuals (BEKOFF et al. 1984).
Wolves, in particular, live in packs that are basically one-family units (MECH
1970, HABER 1977, PETERSON 1977) that form when two adults of opposite sex meet
on a vacant territory, and reproduce (ROTHMAN & MECH 1979, FRITTS & MECH 1981).
Formation and persistence of the pack as a functional unit is based on the social
bond among its members, or what has been referred to, in human terms, as a sort
of “affectional tie” (MECH 1970: 46). Although there are also cases of “non-family”
packs reported in the wild, all have in common a male-female pair (MECH & NEL-
SON 1990).
Feral dogs in Italy showed characteristics of the wolf pack only to a limited
extent and, most notably, members of the same social unit were not generally relat-
ed (BOITANI et al. in press), as in most cases of stray and feral dogs studied else-
where (SCOTT & CAUSEY 1973; NESBITT 1975; CAUSEY & CUDE 1980; BERMAN & DUN-
BAR 1983; DANIELS & BEKOFF 1989a, 1989b). However, even though the whole social
unit centred around stable breeding-pairs, the kinds of association and social bond
among feral dogs did not reflect the precise rules of pack living, as known for other
canids (KLEIMAN & EISENBERG 1973, BEKOFF et al. 1984, GITTLEMAN 1989). There-
fore, we propose the term “group” as more appropriate for a feral dogs’ social unit
rather than pack.
Group size and density
Group size was 2-5 individuals in SCOTT & CAUSEY (1973) and 2-6 individuals
in 14 groups in CAUSEY & CUDE (1980), both in Alabama. DANIELS & BEKOFF (1989b)
report 2-4 animals per pack in their feral population in Arizona. NESBITT (1975), in
his 5-years study of feral dogs in Illinois, found a mean group size of 5-6 animals.
BOITANI & RACANA (1984) also report feral dogs in Basilicata (southern Italy) seen
mostly in pairs. BOITANI et al. (in press) found group size ranging 3-6 adults. Urban
free-ranging dog studies report a majority of animals being seen alone or in pairs
(BECK 1975, BERMAN & DUNBAR 1983, DANIELS 1983a, HIRATA et al. 1986, DANIELS &
BEKOFF 1989b, MACDONALD & CARR in press). It is possible that the smaller group
sizes in urban and rural areas are due to the little advantage conferred on group-
living when food resources are scarce (BECK 1973, DANIELS & BEKOFF 1989b), or,
alternatively, to the fact that in urban areas food resources are plentiful and co-
operative “hunting” is not an advantage (BERMAN & DUNBAR 1983). Unfortunately
neither of these considerations are supported by accurate estimates of food
resources, in both spatial and temporal terms.
Pack size in wolves varies greatly, generally ranging from two to 15 individu-
als per pack in winter (MECH 1986), and packs of up to 36 individuals have been
reported from Alaska (RAUSCH 1967). However, average pack size from different
55Social ecology of feral dogs and wolves
areas appears to be seven or less (MECH 1970). Although information regarding
pack dynamics in wolves has been obtained both in captivity and in the wild (e.g.
ZIMEN 1982; MECH 1977a, 1986), the factors through which pack size are regulated
have not been clearly defined. Pack size is a function of mortality and recruitment
rates, as well as time of dispersal by pack members (PACKARD & MECH 1980). How-
ever, the ultimate and most important factor affecting pack size appears to be prey
abundance, as changes in prey availability correspond to proportional changes in
pack size: MECH (1977a) reported a decrease in mean pack size (midwinter) from
5.7-8.6 to 3.7 during a period of deer decline in Minnesota, and MESSIER (1985)
reported a mean pack size of 5.7 and 3.7, respectively in areas of high and low prey
density in Quebec. Pack size is not to be expected to be stationary year-round
(MECH 1977a, 1986), and major fluctuations (up to 12%) have been observed from
December through March in areas of low prey density (MESSIER 1985).
The group composition of feral dogs was found to be rather stable by BOITANI
et al. (in press), although in the absence of any apparent, intrinsic regulatory mech-
anism. In the 1984-1987 period, all events reducing or increasing the group num-
bers appeared to be related to external factors (human persecution and the avail-
ability of stray dogs), and without any predictable adaptive value. All deaths of sex-
ually mature individuals were accidental and caused by human interference, while
the newborn from feral parents contributed almost nil to the long term group
stability. Only by recruitment of new members from the village stray population
was the feral group able to maintain its size: at the end of the study, all but one
dog in the group were of stray origin. The accidental disruption of the breeding
pair appeared essential to trigger the recruitment mechanism: as a pair-bonded
adult died the adult left alone managed to co-opt a sexually mature individual of
the opposite sex. This new member became in turn socially accepted by the whole
feral group. Breeding periods in canids are accompanied by extensive social inter-
actions which, in turn, may contribute to stronger pair bonds (KLEIMAN & EISEN-
BERG 1973). Increased social interactions may facilitate co-option of external adults
(most interactions with dogs from the villages were observed during these periods),
while a strong pair bond may be the major cause of preventing further co-options.
Although these speculations might be a promising hypothesis, they do not foresee
any self-regulating mechanism for group size, and are weakened by the lack of data
on the behavioural responses of in-group individuals to the attempts of external
dogs to approach and join the group.
Quantity and distribution of food resources are often called as primary causes
of social groups and determinants of group size (MACDONALD 1983, VON SCHANTZ
1984, MACDONALD & CARR 1989 and in press). BOITANI et al. (in press) discussed the
role of the dumps at their study area, which provided a food supply in excess dur-
ing all seasons: group size was then referred more to social factors rather than eco-
logical ones. It is interesting to note that the marked philopatry of those dogs
would meet the general premises of the Territory Inheritance Hypothesis (LIND-
STROM 1986): this hypothesis on the evolution of group-living in carnivores gives
stronger importance to the attachment of individuals to the parents’ territories, and
it predicts an optimal group size which fits the results obtained by BOITANI et al. (in
press). The same authors, however, failed to report differential measures of fitness
of phylopatric sub-adults versus dispersers, which are critical to assess LINDSTROM’s
hypothesis. The feral dog ecology must undergo deeper and longer term research in
order to be reasonably tested against these theoretical generalizations. In addition,
we believe that theoretical analysis of canid evolutionary strategies (BEKOFF et al.
56 L. Boitani and P. Ciucci
1984) are of limited value when carried out on animals which have been living
under artificial and natural selection pressures.
It seems reasonable to assume that the lack of social structure and social
bonds, that is typical of the wolf pack, poses an upper limit to the number of feral
dogs that can effectively cooperate as a functional unit (in hunting, territorial
defence, offspring care, etc.). This might in part explain the smaller size of feral
dog groups (SCOTT & CAUSEY 1973, NESBITT 1975, CAUSEY & CUDE 1980, DANIELS &
BEKOFF 1989b, BOITANI et al. in press) as compared to wolf packs. However, the
situation appears obscured in Italy where, for both wolves and feral dogs, even
though to a different extent, human persecution and interference is perhaps the
most important (and unpredictable) factor affecting, directly and indirectly, the
pack and group demography. This could also explain the lower density values
observed for wolves in Italy (1.25/100 km2, ZIMEN & BOITANI 1975) and in Spain
(1.5-2/100 km2, VILA et al. 1993) as compared to those at the same latitudes in
North-America, even in periods of population decline (3.4/100 km2, MECH 1986). A
conservative density estimate for the feral dogs studied through intense field
research by BOITANI et al. (in press) accounts for 1.3-2.0 feral dogs/100 km2, which
appears closer to the wolf density in Italy (ZIMEN & BOITANI 1975). This could sug-
gest that feral dog groups in Abruzzo tend to be regulated by the same ecological
factors (including human persecution) that operate on wolves, although through
very different mechanisms (see BOITANI et al. in press).
Conversely, wolf densities in North-America appear to be regulated essentially
by the availability of the ungulate biomass (KEITH 1983) which, through the wolves’
social behaviour, has an effect on their numerical and functional responses.
REPRODUCTION AND LIFE HISTORIES
Breeding
Domestic dogs usually breed twice a year, although without seasonal patterns.
In feral dogs, BOITANI et al. (in press) found an average 7.3 months (range 6.5-10
months) between oestrus. As 50% of the parturitions occurred during the period
February-May, they indicate a breeding increase in spring, whereas the others were
scattered during the rest of the year. Although the spring concentration was signifi-
cant, it has not yet been possible to determine any real synchrony of breeding among
females. MACDONALD & CARR (in press) report a much more synchronized breeding in
their dogs and refer this to a period of group stability: this hypothesis is consistent
with data by BOITANI et al. (in press), and it deserves further observation. Increase in
breeding in spring and fall was reported by GIPSON (1972) and suggested as possible
by DANIELS & BEKOFF (1989b). Time of the year breeding occurs has a critical adap-
tive value, in terms of pup survival rates, and wild canids in Italy generally give birth
in April (Vulpes vulpes) and in April-May (Canis lupus) (BOITANI 1981). In this regard,
however, it is quite unlikely that the observed concentration in spring of oestrus
cycles in feral dogs might indicate a converging strategy, even at a very early degree,
with other wild canids. The life spans and poor viability of the dog population would
not allow sufficient time for a strategy to be successfully attained. It might rather
represent the remnant of an ancestral condition, i.e. the physiological potential to
synchronize endogenous reproductive rhythms to the photoperiod cycle, as can be
inferred from wolves (SEAL & MECH 1983, SEAL et al. 1987) and Cape hunting dogs
57Social ecology of feral dogs and wolves
(CUNNINGHAM 1905). On the other hand, the lack of synchronicity among the breed-
ing females of the studied group (BOITANI et al. in press), as well as the casual distri-
bution throughout the year of the off-spring oestrus cycles, might be expression of
modifications of the ancestral reproductive traits due to artificial selection: the loss
of susceptibility to a social control on reproduction and an escape from photoperiod
synchronization (PACKARD et al. 1985).
Although there are cases of two litters being raised successfully within the
same pack of wolves (MURIE 1944, VAN BALLENBERGHE 1983, MECH & NELSON 1989),
restriction of breeding to a single dominant female appears to be the general rule
(e.g. RABB et al. 1967, MECH 1970, ZIMEN 1976, PACKARD & MECH 1980, HARRINGTON
et al. 1982), and it is achieved through deferred reproduction of the subordinates
(PACKARD et al. 1983, 1985). Within the feral dog group studied in Italy (BOITANI et
al. in press) no indication was gathered of any form of social control on the sexual
behaviour of the (subordinate) adults. All females reproduced, giving the group its
full potential for demographic increase.
Denning and parental care
Only one case of group splitting was observed by BOITANI et al. (in press) when
denning and pup rearing by one of the feral dog females was observed to last more
than 5 months. In all other denning activities, females always reared their pups
without any care or threats by other group members, and located their dens within
or in close proximity to the group’s traditional core areas. Denning females spent
most of the time at the den, with frequent visits to the closest feeding sources. Den-
ning females who reared offspring within the group’s traditional home-range were
often visited by other group members, although there was no indication of commu-
nal care of the litters (BOITANI et al. in press). Pups in their first weeks of life were
often left unattended at the den while the mother was feeding, and this might
indeed account for a significant rate of infant mortality due to predation (BOITANI et
al. in press). Denning and rearing pups apart from the group has been reported in
feral dogs by DANIELS (1988) and DANIELS & BEKOFF (1989b). Group splitting during
denning has been suggested as an adaptive strategy for pack-living canids as a way
to reduce the burden of alloparental care on the pack, and to reduce the threats of
infanticide by the dominant female (DANIELS & BEKOFF 1989b); on the other hand,
group participation in rearing is adaptive for just the opposite reasons, i.e. it relie-
ves the female from the burden of caring alone for her pups, and it provides more
protection for the young from other predators (KLEIMAN & EISENBERG 1973). In
absence of a selective pressure for pack splitting, denning within the group’s territo-
ry increases protection from intruders and potential predators.
In wolves, dens are generally located within the pack territory (LOWHEAD 1983,
CIUCCI & MECH 1992), and all the pack movements during denning periods are
influenced by the den location, where the adults and the yearlings return periodi-
cally to regurgitate and attend the pups (MURIE 1944, MECH 1970, CARBYN 1974, VAN
BALLENBERGHE et al. 1975, HARRINGTON & MECH 1982). In wolves, the role of auxil-
iaries (non-breeding adults and yearlings) in the communal rearing of the relatives’
offspring has recently been interpreted in terms of individual selection (PACKARD &
MECH 1980, HARRINGTON & MECH 1982, HARRINGTON et al. 1983) where, rather than
a form of altruism, it is viewed as part of the optimal reproductive strategy of those
individuals who temporarily defer reproduction (PACKARD et al. 1983).
58 L. Boitani and P. Ciucci
In feral dogs, rearing pups without male assistance might be due to the
domestication process, where humans have provided assistance. In fact, dogs stand
alone, among all canids, for a total lack of paternal care (KLEIMAN & MALCOM 1981).
This could also explain the apparent lack of auxiliaries of any sex in feral dog
groups, even though it is mostly the absence of deferred reproduction in the adults,
i.e. absence of social control on subordinates’ sexual behaviour (PACKARD et al.
1985), that seems to affect the availability of potential helpers.
Litter size and pup survival
Dogs are known to have litter sizes up to 17, although up to 10 is the most
usual range (KLEIMAN 1968, KLEIMAN & EISENBERG 1973). For feral dogs, a litter of
five and a total of eight for two other litters have been reported by NESBITT (1975),
and DANIELS & BEKOFF (1989b) report a total of 10 pups from two litters. However,
estimates of litter size rely on pups first sightings, which rarely occur before they
are mobile (2-4 weeks old), when natal and post-natal mortality might have already
contributed to lower estimates. BOITANI et al. (in press) report a lower mean litter
size (3.63 pups/litter, n = 11) than previously reported for feral dogs elsewhere, and
also lower than the figure (5.5 pups/litter, n = 17) obtained by MACDONALD & CARR
(in press) in the same study area for a different period and group composition.
Along with factors such as nutritional status of the bitches, occurrence of epizoo-
tics and competition among pups, and predation, the different litter size values
might be explained by the great variability in age-related fecundity in different
breeds, as well as by different age-structures of the groups studied.
Wolf average litter sizes in the wild range from 4.0 to 6.5 (MECH 1970), esti-
mates being mostly based on the first observation of the litters after parturition, as
for the dogs. Based on active nipple counts of lactating females, and on mid-sum-
mer and winter pups observations, MECH (1977a) reported an average litter size
from 3.0 to 3.4 pups/litter, relating the lower than average estimate to the consider-
able nutritional stress that the population was facing. It remains unclear, however,
if food availability affects the natality rates of the breeding females or rather pup
survival rates during the early post-natal period, or both.
Survival rates for wolf pups are generally low in conditions of malnutrition
(MECH 1970, VAN BALLENBERGHE & MECH 1975, SEAL et al. 1975), and appear posi-
tively correlated with the number of in-group auxiliaries (HARRINGTON et al. 1983).
Survival rates are generally low in feral dogs as compared with wolves: BOITANI et
al. (in press) found that out of 40 pups, 28 (70%) died within 70 days of birth, 9
(22.5 %) died within 120 days, 1 (2.5%) within one year, and only 2 (5%) survived
the age of one year. These values compare with similar ones found by NESBITT
(1975), SCOTT & CAUSEY (1973), DANIELS & BEKOFF (1989b), MACDONALD & CARR (in
press). The low survival rate at 4 months of age (7.5%) found by BOITANI et al. (in
press) is a clear indication that the majority of mortality occurs during this period
of early independence, and may be due to four factors relevant for the purpose of
this paper: a) in the absence of communal helping, pups are often left at the den
unattended, thus increasing the risks of predation; b) from about 6-8 weeks of age
the pups begin to explore, without adult supervision, increasing portions of range
surrounding the den site and this, again, increases risks of predation; c) as their
mother enters her new oestrus cycle, she is most likely to lower her interest in her
offspring, and d) irregular breeding cycles occurring twice a year cause many a lit-
59Social ecology of feral dogs and wolves
ter to be born in periods of inclement weather. Thus, although early mortality
deserves further study, low efficiency in reproduction in feral dogs appears to be
mostly affected by two relevant aspects of their breeding biology: (i) the social envi-
ronment of the group which does not provide auxiliaries for the communal care of
the offspring, and (ii) the reproductive physiology of the females in terms of num-
ber and timing of their oestrus cycles. Such conditions, when transferred to natural
environment, determine high rates of reproductive failures and offspring/juvenile
mortality.
From a demographic standpoint, survival rates of only 5% to one year of age
contribute to understanding why free-ranging dogs are not able to maintain their
population levels. This was already stressed by urban dog studies (BECK 1973, DAN-
IELS 1983a) and it was left unanswered by DANIELS & BEKOFF (1989b) for feral dogs
in Arizona. In the area investigated by BOITANI et al. (in press), the feral dog group
could not have maintained the observed population level without a continuous
recruitment of new group members from outside (i.e., free-ranging and stray dogs).
Sex ratio
A critical component of the set of parameters contributing to the negative
demographic balance observed by BOITANI et al. (in press) in feral dogs is the
skewed sex ratio. Urban and rural/suburban dog populations generally show a
skewed sex ratio for males, ranging from 1.6:1 to 5:1 in favour of males (BECK
1973, DANIELS 1983a, BOITANI & RACANA 1984, DANIELS & BEKOFF 1989b, WHO
1988). MACDONALD & CARR (in press) found ratios of 4:1 and 2.6:1 in favour of
males in the villages of the same Italian study area where BOITANI et al. (in press)
reported a sex ratio of from 2:1 to 1.5:1 (group composition at various stages, and
considering only the adult members of the group) in favour of females. Possible
reasons for these findings have already been discussed (BECK 1973, DANIELS & BEK-
OFF 1989b): the sex ratios of urban dogs result from the direct selection of males as
pets and from selective removal of females from the population, either temporarily
to avoid unwanted pregnancies, or permanently by killing them as newborn. How-
ever, a differential mortality rate for the two sexes is not likely to occur outside
artificial human interference. DANIELS & BEKOFF (1989b) also reported a sex ratio
of 3.5:1 in favour of females in a feral dog population in Arizona, in an area adja-
cent to the previously cited urban areas: they explained the result by suggesting
that an important source of feral dogs was the abandonment of females removed
from urban and rural areas. This appeared the only logical, though weak, explana-
tion since no evidence could be obtained either on skewed sex ratios of new-borns,
or on differential survival between sexes.
These results are even more difficult to discuss when considering that overall
litter composition was found by BOITANI et al (in press), highly skewed in favour of
males (3.2:1). In wolves, MECH (1975) found a male biased pup sex ratio in condi-
tion of marginal nutrition or intense food competition among adults suggesting
this might contribute to the population regulation process. While the dog litter sex
ratio in favour of males might suggest retention in feral dogs of the similar, though
unknown, physiological mechanism (MECH 1975), higher female survival rates
appear the only explanation when litter sex ratio is compared to adult sex-ratio: it
might possibly be related to a) a stronger tendency of male pups to explore the
60 L. Boitani and P. Ciucci
areas surrounding the den site, and b) differential maternal care of pups of differ-
ent sexes. Further research is needed on the relation between survival rates of the
two sexes to nutritional status, philopatry tendencies and maternal care.
SPACE USE PATTERNS
Home-range
Space use in feral dogs is not dissimilar from most other wild canids, in that
for their life-history activities they utilize definite and traditional areas (home-rang-
es) which, to a varying extent, intensity and variability, tend to be defended against
intruders (i.e., territory) (SCOTT & CAUSEY 1973, CAUSEY & CUDE 1980, GIPSON 1983,
DANIELS & BEKOFF 1989a, BOITANI et al. in press). In addition to the several environ-
mental factors which determine home-range size and configuration, human activ-
ities can play a powerful role in affecting the biological answers of canids (KLEIMAN
& BRADY 1978), and of dogs in particular.
Within the overall home-range of 57.8 km2, BOITANI et al. (in press) found that
the dog group used smaller portions at a time, shifting core areas in response to
several factors: the finding of new food resources (i.e., a large livestock carrion),
disturbance by human presence, denning activities, previous spatial-use patterns of
newly recruited dogs, unpredictable fluctuation in food availability at dumps, pos-
sible interference by wolves. These factors had no seasonal predictability, and they
appeared randomly in the group history. DANIELS & BEKOFF (1989a) reported sea-
sonal variations related to the presence of dependent pups for one of the two
groups studied, while the other did not show any change in its core area use. Diffe-
rential energetic requirements were suggested as a possible reason for the two
groups behaviour (DANIELS & BEKOFF 1989a). SCOTT & CAUSEY (1973) also found a
change of core areas depending on the presence of pups.
Drifting ranges have been recorded for urban foxes as a consequence of social
instability entailed by abrupt changes to population structure and food availability
(DONCASTER & MACDONALD 1991). BOITANI et al. (in press) suggest that drifting of
seasonal ranges reflect not only direct environmental changes, but also the influ-
ence of previous knowledge of the area by a new member of the group. The ran-
dom changing of core areas was maintained within three main alternate areas, and
within the same long term boundaries, indicating that a tradition of area use pre-
vented more random movements.
Home-range estimates can yield very different results as a consequence of the
methods being adopted, and this should be taken into account when interpreting
data from different studies (MACDONALD et al. 1980). Furthermore, telemetry tech-
niques produce sets of data hardly comparable with other type of information (i.e.,
visual observation, tracks in snow, etc.) in assessing the extent of range and habitat
utilization. These considerations might explain a great part of the variability in
home-range sizes reported so far for feral dogs (see BOITANI et al. in press, for a
review). Limited to radio-telemetry studies, home-range sizes reported for feral dogs
range from 4.44-10.4 km2for three different groups in east-central Alabama (SCOTT &
CAUSEY 1973), to 18.72 km2in Alabama (CAUSEY & CUDE 1980) and 70 km2in Alaska
(GIPSON 1983). BOITANI et al. (in press) discussed the factors believed to play a signifi-
cant role in determining home range size, namely the relative distances of dens,
dumps and resting (refuge) sites, that are largely independent of group size.
61Social ecology of feral dogs and wolves
Urban and suburban dogs are reported to have much smaller home-ranges
(from 2-11 up to 61 ha) (BECK 1973, FOX et al. 1975, DANIELS 1983a, BERMAN &
DUNBAR 1983, SANTAMARIA et al. 1990). Food availability patterns, small group sizes
and reduced social contacts are probably the determinants of such behaviour, con-
firming the mechanism being suggested for the feral dog data.
Average wolf territory sizes may range from 78 km2, in areas where wolves
prey primarily on white-tailed deer (Odocoileus virginianus) (FULLER 1989), to 2.541
km2 at higher latitudes with moose (Alces alces) and caribou (Rangifer tarandus) as
the main prey (BALLARD et al. 1987). Much of the observed variation in territory
size has been interpreted as dependent on several factors among which the most
relevant appear to be pack size (PETERSON et al. 1984, MESSIER 1985, BALLARD et al.
1987), prey density (MESSIER 1985), and wolf population density (FRITTS & MECH
1981). The influence of all these factors on territory size appears to depend also on
the level of human exploitation of the local wolf populations (PETERSON et al. 1984,
BALLARD et al. 1987). Shifts in the use of the internal parts of wolf territories have
been reported: in summer as a result of pup presence at the den and at rendezvous
sites, and in winter to increase effectiveness of prey exploitation (FRITTS & MECH
1981, MESSIER 1985). Thus, it seems reasonable to assume that the flexibility obser-
ved in space use by feral dogs might substantially be the same strategy of their wild
ancestors where, depending on local conditions, territory utilization tend to opti-
mize resource utilization and survival. In fact, in the same area where feral dogs
have been studied by BOITANI et al. (in press), radiocollared wolves exhibited a simi-
lar pattern of territory use (CIUCCI 1994).
Territoriality
Territorial behaviour in feral dogs was observed by BOITANI et al. (in press)
more consistently than previously reported (SCOTT & FULLER 1965, BEKOFF 1979,
DANIELS & BEKOFF 1989a, BERMAN & DUNBAR 1983, BOITANI & RACANA 1984), as it was
displayed not only in the vicinity of den sites, but within the entire core areas and
during the whole year; this pattern is also reported by MACDONALD & CARR (in press)
for the same area. Territorial defence could be inferred through observation of direct
encounters (chasing and/or facing conspecific intruders), vocal advertisement (bark-
ing), and the absence for the study time-span of other stable groups of dogs in the
same territory. Even though there were signs of scent-marking activities, no attempt
was made to clarify the role of such marking in territory maintenance, as it has been
done for wolves (PETERS & MECH 1975, ROTHMAN & MECH 1979).
The higher frequency of territorial behaviour reported by BOITANI et al. (in
press) might be related to a higher level of integration within the group, a higher
degree of isolation from other dogs, and to the food resources being concentrated
mostly in localized patches at the dumps. Also, the partial overlap with two wolf
pack territories in the same study area might have increased the dogs’ general
alertness and territorial behaviour. In addition, breed influence on territorial
behaviour needs to be contemplated, since it is likely that the observed degree of
territoriality in part reflected the typical traits of the Maremma dog, one of the
dominant types in the feral group breeding history.
Although in the absence of any evidence for direct competition between feral
dogs and wolves (apart from one dog most probably being killed by wolves), partial
overlap of territories and the almost identical niche that they share in central Italy
62 L. Boitani and P. Ciucci
(BOITANI 1983) make competition for food and space highly likely. Thus, the pres-
ence of wolves might have been an important component in shaping the dogs’
home-range, and in determining its location and maintainance. The fact that feral
dogs’ core-areas were closer to human settlements than the wolves’ core-areas, and
located in the interstice between two neighbouring wolf territories (CIUCCI 1987,
BOITANI et al. in press), might indicate that the dog group was not competing effi-
ciently enough to succeed independently from human presence.
Territorial behaviour in wolves has been investigated (e.g., PETERS & MECH
1975, HARRINGTON & MECH 1979), related to intraspecific density and prey abun-
dance (e.g., MECH 1977a, 1986), and interpreted as a mechanism of population reg-
ulation (PACKARD & MECH 1980). Although the proximate causation of territorial
behaviour might be similar in dogs and wolves (i.e., the defence of resources within
the territory), it seems unlikely that in feral dogs it might work as a mechanism of
population regulation; in fact, for this to be true, one would expect feral dog popu-
lations to appear reproductively self-sustaining and their in-group sexual behaviour
susceptible to social control (PACKARD & MECH 1980). However, both these condi-
tions have not been met in the group studied by BOITANI et al. (in press). It would
be intriguing to investigate further the evolution of territorial patterns and mecha-
nisms in feral dogs, searching for the rules that minimize inter-group encounters
typical of saturated wolf populations (e.g., PETERS & MECH 1975, HARRINGTON &
MECH 1979). This would contribute to our understanding of the evolution scent-
marking and vocal behaviours, and the role that natural and artificial selection
plays in their shaping.
ACTIVITY PATTERNS
A tendency for nocturnal and crepuscular activity was first reported by BECK
(1973) for urban dogs: during the summer months, it was mainly restricted to two
periods, 07:00-10:00 p.m. and 05:00-08:00 a.m. A similar bimodal model of activity
distributions was found by BERMAN & DUNBAR (1983) for the dogs of Berkeley, Cali-
fornia. HIRATA et al. (1986) reported that dogs of several Japanese towns were most
active from 00:00 to 06:00 a.m., with a peak just before and around 06:00 a.m. The
predominant dawn peak had already been observed in free-ranging rural dogs in
Virginia (PERRY & GILES 1971), while the bimodal distribution was confirmed by
several studies on feral dogs (SCOTT & CAUSEY 1973, CAUSEY & CUDE 1980, BOITANI &
RACANA 1984, DANIELS & BEKOFF 1989a). NESBITT (1975), discussing similar tempo-
ral patterns, suggested that feral dogs could be active and travel all day but were
restricted to nocturnal and crepuscular hours in the attempt to avoid human con-
tact. BOITANI et al. (in press) presented similar results which seem to confirm
NESBITT’s hypothesis: when the human presence was low, a female dog moved most-
ly during daytime, while she later resumed nocturnal habits when visiting the more
“dangerous” dumps close to the villages. Avoiding humans may provide an explana-
tion for nocturnal activities, but it does not explain the bimodal pattern found for
all dogs and for all seasons. Nine out of 17 canids are strictly nocturnal (cf. BEKOFF
et al. 1981); bimodal activity regimes are known for a great variety of carnivores
(Vulpes fulva: ABLES 1975; Crocuta crocuta: KRUUK 1972; Chrysocyon brachyurus:
DIETZ 1984) and they are a common biological pattern. ASCHOFF (1966) called it the
“bigeminus pattern” and suggested that it is an innate behavioural trait, indepen-
63Social ecology of feral dogs and wolves
dent from any environmental pressure. ASCHOFF (1966) also pointed out that the
second (dawn) peak is usually lower, but this is not the case in the study by BOITANI
et al. (in press), where maximum activity levels have been always observed at
dawn.
Wolf activity patterns are extremely influenced by different environmental
conditions, including human interference. Activity in wolves in North-America has
been studied primarily through direct observation, and therefore most of data are
essentially limited to daylight hours. However, available radiotelemetry data (KOLE-
NOSKY & JOHNSTON 1967, BALLARD et al. 1991) and other direct observations (MURIE
1944, JOSLIN 1966, BALLARD et al. 1991) tend to confirm the nocturnal activity pat-
tern of wolves, with crepuscular times (dawn and sunset) being the most likely
periods of arrival at, and departure from, the den- and rendezvous-sites. Neverthe-
less, other authors reported that wolves tend to be active also during the daylight
hours, and that this proportion of activity (moving, travelling, hunting) tend to be
higher in winter months, as well as indirectly correlated with the nutritional status
of the wolves (MECH 1977b, PETERSON et al. 1984). West-European studies, based on
24-hr monitoring cycles, depict wolf activity as being almost nil during daylight
hours, with the wolves being primarily active from sunset to dawn (BOITANI 1982,
URIOS et al. 1993, VILA et al. 1993, CIUCCI 1994). In Spain, an analysis of wolf activ-
ity confirmed a nocturnal, bimodal pattern (URIOS et al. 1993, VILA et al. 1993),
whereas in highly populated areas of Italy nocturnal activity was mostly unimodal,
lacking periods of decreased activity in the night hours (CIUCCI 1994). This has to
be expected if the innate bimodal pattern (ASCHOFF 1966) has been selected against:
in areas of high anthropic interference, activity peaks at dawn and sunset would
increase the chance of wolf-human encounters, especially where the location of
wolf refuge-sites and feeding-sites appear to be interspersed with human settle-
ments (CIUCCI 1994).
Although the present studies on feral dog 24-hr activity patterns have been
essentially descriptive, and environmental correlates of activity (food abundance,
intra- and interspecific interferences, climatic and physiological factors, etc.) deser-
ve deeper investigation, they tend to confirm that the canids’ innate rhythmicity
has not been altered to a great extent through artificial selection, and that feral dog
activity patterns may just reflect their ancestors’ flexibility.
FOOD SOURCES AND PREDATION
Potential predation on wildlife and livestock has been the instigation for feral
dog studies: feral dogs have long been accused by the popular press of predation on
deer in North-America and on livestock in Italy, although with little supporting evi-
dence (cf. BOITANI 1983, for a short review on wolf-dog competition). BOITANI et al.
(in press) and SCOTT & CAUSEY (1973) did not find any evidence of predation on
livestock. NESBITT (1975) was unable to document a single case of livestock depre-
dation in his 5-years study. This last situation resembles that found by BOITANI et
al. (in press), where cattle were free-ranging over most of the area, yet no interfer-
ence was ever observed. On the other hand NESBITT (1975) reported that free-rang-
ing pet dogs killed three calves in that area during his study. This again compares
with severe damage by free-ranging owned dogs on livestock that one of us was
able to document in other areas of Italy (L. BOITANI unpubl.). Thus, some evidence
64 L. Boitani and P. Ciucci
would suggest that it is essentially free-ranging owned and stray dogs that are pri-
marily responsible for livestock depredation, although more research is needed on
this subject. As for predation on wildlife, BOITANI et al. (in press) report that wild
boar (Sus scrofa), the only ungulate significantly present in the area, was rarely
found in dogs’ faeces and, as yet, no evidence has been gathered for predation on a
live wild boar. Feral dogs have been reported to have little impact on wildlife by all
previous research on their feeding ecology (PERRY & GILES 1971, SCOTT & CAUSEY
1973, GIPSON & SEALANDER 1977, CAUSEY & CUDE 1980). Different experimental
hunting with radio-marked and trained dogs failed to prove successful hunting on
deer (PROGULSKE & BASKETT 1958, CORBETT et al. 1971, SWEENEY et al. 1971, OLSON
1974). On the other hand, HAWKINS et al. (1970) and LOWRY & MACARTHUR (1978)
reported a small percentage of successful hunting in Illinois and in Idaho, respec-
tively. Most importantly, DENNEY (1974) in Colorado and GAVITT et al. (1974) in Vir-
ginia reported deer being killed by feral dogs. As for livestock, these apparently
contradictory results are probably best explained by local conditions in terms of
the presence of alternate food resources (and their stability and predictability), dog
group size and breed-types, foraging tradition, livestock husbandry techniques,
wildlife distribution and density, and level of human persecution. It seems reason-
able to assume that all these factors somehow modulate to different extents an
innate propensity for predation, which also depends strongly on the attitude of the
dominant breed-types in the group (i.e., an effect of artificial selection). Other cas-
ual factors, such as the previous experience of some individual in the group, might
influence the feeding habits of the whole group (i.e., cultural tradition): exposure to
prey, successful predatory trials and cultural transmission are important compo-
nents in determining hunting techniques, attitudes and prey-type preference.
SCOTT & CAUSEY (1973) and BOITANI et al. (in press) reported brief, apparently
uncoordinated, and unsuccessful hunting chases during which the dogs were con-
stantly barking: one might assume that predatory tendencies in a feral dog group
are lowered in the long term by unsuccessful attempts due to lack of a coordinated
and effective hunting technique. If this were true, one might expect, in the presence
of alternative food sources, dog groups to have a more flexible diet, and to incorpo-
rate in their diet items easier for single individuals to catch. Alternatively, their diet
might just be more stable and predictable. This has been reported by several
authors who found that feral dogs hunt and feed on rodents, rabbits, and other
small game, although the details are not reported. Detailed accounts of a special
predatory situation were given for feral dogs on the Galapagos Islands who feeding
on marine iguanas (Amblyrhynchus cristatus) (KRUUK & SNELL 1981, BARNETT &
RUDD 1983), and for feral and semi-feral dogs predating capybaras (Hydrochoerus
hydrochaeris) in Venezuela’s llanos (MACDONALD 1981). BOITANI et al. (in press) and
MACDONALD & CARR (in press) stressed the importance for the feral dogs of refuse
found at the several garbage dumps located within the dogs’ home range.
The importance of parental-teaching, exposure to prey, and successful experi-
ence for the development of hunting efficiency in wolf pups has long since been
documented (MECH 1970). The same group-living in wolves has been interpreted as
a way to overcome prey with a larger body mass (MECH 1970), as also suggested for
other predators (e.g., hyenas: KRUUK 1972). Indeed, it is the same social structure
of wolf packs that allows each individual to be integrated within an efficient hunt-
ing unit. In feral dog groups, however, where leadership is more questionable and
social bonds among individuals are more flexible, one would expect predatory ten-
dencies, if any, to be less effective, mostly uncoordinated, and with severe limita-
65Social ecology of feral dogs and wolves
tions (e.g., kill rate, prey type and size, energetic cost and benefit balance, time to
functional response, etc.). Hence, given the feral dogs’ social structure, it seems
reasonable to expect diversified food habits which include smaller and easier-to-
catch items (see above). It is noteworthy to report that this also appears to be the
case for the dingo in Australia (CORBETT 1989). However, for those dog groups that
prey on wildlife (DENNEY 1974, GAVITT et al. 1974), the same mechanisms that regu-
late most wolf-prey systems (MECH 1977a, 1977b; PETERSON 1977; NELSON & MECH
1981; MESSIER 1985) cannot be expected to be effective, for the feral dog ecology
seems to be dictated by non self-regulating biological traits (e.g., productivity and
survival). Further research is especially needed in areas were dogs are known to
prey on wildlife and in the absence of more predictable food sources (i.e., garbage
dumps).
CONCLUDING REMARKS
The ecology of feral dogs, dominated by their behavioural tendency to avoid
contact with humans, appears to be shaped by a complex of biological traits which,
on the one hand, still resemble their ancestors’ flexibility, but on the other fail to
meet their original fitness requirements.
Feral dogs’ survival strategies still maintain, to some extent, some ancestral
traits, namely group-living tendencies, territorialism, predatory instincts, and a
large degree of ecological flexibility. Even though most of these traits appear void
of their adaptive value (just as “evolutionary inertia” or artificial selection epiphe-
nomena, as our hypothesis postulated), the dog’s ancestor flexibility seems to be
reflected mostly by food habits, space-use patterns, and activity patterns.
The dog flexibility, in both behavioural and morphological traits was enhan-
ced by the domestication process: in particular, the retention of juvenile characters
into adulthood (i.e., neoteny), having a significant adaptive value in domesticated
environments (e.g., lower aggressiveness, more dependence on humans, early sexu-
al maturation, easier submission), also provided greater plasticity in behavioural
development (GINSBURG & SCHOTTE 1978, FRANK 1980, FRANK & FRANK 1982, PRICE
1984). By escaping the full pressure of natural selection, dogs attained greater plas-
ticity also through relaxed selection. This enhanced flexibility of the dogs as com-
pared to wolves appears to be the critical mechanism which partly allow the dogs
to overcome the toll of natural selection and to survive in the natural environ-
ments. On the other hand, the domestication process is largely responsible for a
loss of fitness through altering the response thresholds of several behavioural traits,
lowering the observational capability and affecting the cognitive mechanisms, and
reducing the overall responsiveness to environmental change due to living for gen-
erations in “safe” environments (PRICE 1984). Feral dogs still show variable degrees
of these quantitative changes.
The limited data currently available on feral dogs do not allow for a thorough
testing of our original hypothesis, even though they tend to confirm the lack of
adaptive value of the most relevant socio-ecological traits: feral dogs are not repro-
ductively self-sustaining, suffer from high rates of juvenile mortality, depend indi-
rectly upon humans for food, co-optable individuals, and space, and their demogra-
phy appears dominated by stochastic and unpredictable mechanisms (BOITANI et al.
in press). It is noteworthy that feral dogs which are still dependent on humans for
66 L. Boitani and P. Ciucci
some aspects of their biology, may still be affected by some form of selection con-
nected to human activities (BECK 1973, BRISBIN 1977, DANIELS & BEKOFF 1989c).
Therefore, the feral dog ecology may be significantly different in various habitats
and ecological conditions (i.e., different levels of dependency of the dogs on
humans). In addition to this, the lack of an operational definition of feral status,
based on consistent criteria and referring to a definite temporal scale, makes the
comparison among different studies difficult. However, at a general level of analy-
sis, the characteristics of feral dogs and the adaptive value that they fail to match,
can be classified in terms of three main biological aspects:
Sociality. Feral dogs social units do not function as wolf packs, namely for the
lack of clear dominant-subordinate relationships throughout and of firm social
bonds among all group members. This directly affects the potential groups size,
the breeding system, the group’s efficiency as a functional unit (hunting, territory
defence, communal pup rearing, etc.). The most relevant consequences in terms of
fitness are: (i) potential limits in the social unit’s efficiency and functionality as
group size increases (i.e., potential upper limit to group size); (ii) potentially dras-
tic population fluctuations. The social structure and its expression in feral dog
groups does not allow for an efficient mechanism of population regulation in rela-
tion to environmental and ecological conditions; (iii) unavailability of potential
non-reproducing ‘auxiliaries’; (iv) negative energetic balance of breeding females;
(v) high pup/juvenile mortality rates; (vi) low recruitment rates.
Physiology. Among the main traits to be affected by artificial selection, repro-
duction in the dog, as well as in most other domestic species, has been strongly
influenced to increase reproductive potential and to shorten generation time. Con-
sequences of this process on physiology are obvious and may be inferred from the
frequency of breeding cycles in the females, the apparent escape from photo period
synchronization and social-control (i.e., deferred reproduction). In addition it has
been suggested, though without good evidence, that the immunodefence system of
the domestic dog appears limited in a natural environment, accounting for high
mortality rates among pups and subadults (FRANCISCI et al. 1991). All this directly
affects reproduction, survival and recruitment in feral dogs, and the most relevant
consequences in terms of fitness are: (i) negative energetic balance of breeding
females; (ii) high pup/juvenile mortality; (iii) low recruitment rates.
Behaviour. Both in terms of individual and social behaviour, the effectiveness
of group activities such as hunting, territorial defence, cultural transmission, and
reproduction appear limited to some extent. The most relevant consequences in
terms of fitness are: (i) low predatory attitude; (ii) low kill rate (efficiency of preda-
tion); (iii) potential limits in prey size; (iv) indirect food-dependency upon humans;
(v) indirect space-dependency upon humans; (vi) low efficiency or absence of allo-
parental care of pups.
In conclusion, we believe the present discussion offers a general framework
for an assessment of the effects of artificial selection on the dog as it returns to live
in the wild, analyzing the adaptive values of the most relevant socio-ecological fea-
tures of the wolf is natural history. The ultimate goal of assessing the adaptive
value of traits which are the results of a mixture of ancient and recent natural and
artificial selective pressure might prove impossible to achieve. It is also important
to emphasize that our discussion mostly applies to those feral dogs living in the
wild for a few generations: our conclusions, then, might not be valid for feral dog
populations which have been living under the effects of natural selection forces for
67Social ecology of feral dogs and wolves
hundreds of years (e.g., pariah dogs and dingoes), where longer reproductive isola-
tion from their domestic ancestors might result in lower phenotypic variability and
in behavioural differences which could be genetically determined.
An investigation of feral dog biology under different ecological conditions and
longer, multi-generational time-scales, might offer a deeper insight into the issues
herein discussed. Future research should also address the role that different domi-
nant breed-types in feral dog groups, as well as the cross-breeding history within
them, might play in the expression (nature, quantity, frequency) of the analyzed
socio-ecological traits. In this regard, we hope that future studies will benefit from
some of the working hypotheses highlighted by our discussion.
ACKNOWLEDGMENTS
We are greatly indebted to F. Francisci and G. Andreoli who contributed to the work on
the feral dogs in Central Italy. We are also grateful to F. Dessì-Fulgheri, J. Clutton-Brock and
an anonymous referee for substantial comments on an earlier version of this manuscript. The
work was partly supported by a grant from the Regione Toscana.
REFERENCES
ABLES E.D. 1975. Ecology of the red fox in North America, pp. 216-236. In: Fox M.W., Edit.
The wild canids. New York: Van Nostrand Reinhold.
ANDREOLI G. 1987. Analisi dei modelli di attività di un gruppo di cani inselvatichiti in Abruz-
zo. Laurea Thesis, University of Roma, Rome.
ASCHOFF J. 1966. Circadian activity patterns with two peaks. Ecology 47: 657-702.
BALLARD W.B., AYERS L.A., GARDNER C.L. & FOSTER J.W. 1991. Den site activity paterns of gray
wolves, Canis lupus, in South-Central Alaska. Canadian Field Naturalist 105: 497-504.
BALLARD W.B.,WITHMAN J.S. & GARDNER C.L. 1987. Ecology of an exploited wolf population in
South-Central Alaska. Wildlife Monograph 98: 1-54.
BARNETT B.D. & RUDD R.L. 1983. Feral dogs of the Galapagos Islands: impact and control.
International Journal of Studies on Animal Problems 4: 44-58.
BECK A.M. 1973. The ecology of stray dogs: a study of free-ranging urban animals. Baltimore:
York Press.
BECK A.M. 1975. The ecology of “feral” and free-roving dogs in Baltimore, pp. 380-390. In:
Fox M.W., Edit. The wild canids. New York: Van Nostrand Reinhold.
BEKOFF M. 1979. Scent-marking by free-ranging domestic dogs. Biological Behaviour 4: 123-139.
BEKOFF M., DANIELS T.J. & GITTLEMAN J.L. 1984. Life history patterns and the comparative
social ecology of carnivores. Annual Review of Ecology and Systematics 15: 191-232.
BEKOFF M., DIAMOND J. & MITTON J.B. 1981. Life history patterns and sociality in canids: body
size, reproduction, and behaviour. Oecologia 50: 388-390.
BERMAN M. & DUNBAR I. 1983. The social behaviour of free-ranging suburban dogs. Applied
Animal Ethology 10: 5-17.
BOITANI L. 1981. Lupo, Canis lupus, pp. 61-67. In: Pavan M., Edit. Distribuzione e biologia di
22 specie di mammiferi in Italia. Roma: Consiglio Nazionale delle Ricerche.
BOITANI L. 1982. Wolf management in intensively used areas of Italy, pp. 158-172. In: Harring-
ton F.H. & Paquet P.C., Edits. Wolves of the world. Perspectives of behavior, ecology
and conservation. New Jersey: Noyes.
BOITANI L. 1983. Wolf and dog competition in Italy. Acta Zoologica Fennica 174: 259-264.
BOITANI L. & FABBRI M.L. 1983. Censimento dei cani in Italia con particolare riguardo al feno-
meno del randagismo. Ricerche di Biologia della Selvaggina, INBS, Bologna 73: 1-51.
68 L. Boitani and P. Ciucci
BOITANI L., FRANCISCI F., CIUCCI P. & A NDREOLI, G. (in press). Population biology and ecology of
feral dogs in central Italy. In: Serpell J., Edit. The dog: its ecology, behaviour and evolu-
tion. Cambridge: Cambridge University Press.
BOITANI L. & RACANA A. 1984. Indagine eco-etologica sulla popolazione di cani domestici e
randagi di due comuni della Basilicata. Silva Lucana, Bari 3: 1-86.
BREHM A. 1893. Tierleben. Liepzig-Wien, 4 vols.
BRISBIN I.L. JR1974. The ecology of animal domestication: its relevance to man’s environmen-
tal crises - past, present and future. Association of Southeastern Biologists Bulletin 21: 3-
8.
BRISBIN I.L. JR1977. The pariah. Its ecology and importance to the origin, development and
study of pure-bred dogs. Pure-bred Dogs American Kennel Gazette 94: 22-29.
CARBYN L.N. 1974. Wolf predation and behavioral interactions with elk and other ungulates in
an area of high prey density. Ph.D. Thesis, University of Toronto, Edmonton.
CAUSEY M.K. & CUDE C.A. 1980. Feral dog and white-tailed deer interactions in Alabama.
Journal of Wildlife Management 44: 481-484.
CIUCCI P. 1987. Uso dello spazio e dell’habitat in una popolazione di cani inselvatichiti
nell’Appennino abruzzese. Laurea Thesis, University of Roma, Rome.
CIUCCI P. 1994. Movimenti, attività e risorse del lupo (Canis lupus) in due aree dell’Appennino
centro-settentrionale. Ph.D. Thesis, University of Roma, Rome.
CIUCCI P. & M ECH L.D. 1992 Selection of wolf dens in relation to winter territories in North-
eastern Minnesota. Journal of Mammalogy 73: 899-905.
CLUTTON-BROCK J. (in press). Origins of the dog: domestication and early history. In: Serpell
J., Edit. The dog: its ecology, behaviour and evolution. Cambridge: Cambridge University
Press.
CORBETT L.K. 1989. Assessing the diet of dingoes from feces: a comparison of 3 methods.
Journal of Wildlife Management 53: 343-346.
CORBETT R.L., MARCHINTON R.L. & HILL C.L. 1971. Preliminary study of the effects of dogs on
radio-equipped deer in mountainous habitat. Proceedings of the Annual Conference of
the Southeastern Association of State Game and Fish Commissioners 25: 69-77.
CUNNINGHAM D.J. 1905. Cape hunting dog (Lycaon pictus) in the gardens of Royal Zoological
Society of Ireland. Proceedings of the Royal Society of Edinburgh 25: 843-848.
DANIELS T.J. 1983a. The social organization of free-ranging urban dogs: I. Non-estrous social
behaviour. Applied Animal Ethology 10: 341-363.
DANIELS T.J. 1983b. The social organization of free-ranging urban dogs: II. Estrous groups
and the mating system. Applied Animal Ethology 10: 365-373.
DANIELS T.J. 1988. Down in the dumps. Natural History 97: 8-12.
DANIELS T.J. & BEKOFF M. 1989a. Spatial and temporal resource use by feral and abandoned
dogs. Ethology 81: 300-312.
DANIELS T.J. & BEKOFF M. 1989b. Population and social biology of free-ranging dogs, Canis
familiaris. Journal of Mammalogy 70: 754-762.
DANIELS T. J. & BEKOFF M. 1989c. Feralization: the making of wild domestic animals. Behav-
ioural Processes 19: 79-94.
DENNEY R.N. 1974. Impact of uncontrolled dogs on wildlife and livestock. Transactions of the
North-American Wildlife and Natural Resources Conference 39: 257-291.
DIETZ J.M. 1984. Ecology and social organization of the maned wolf (Chrysocyon brachiurus).
Smithsonian Contributions to Zoology 392: 1-51.
DONCASTER C.P. & MACDONALD D.W. 1991. Drifting territoriality in the red fox Vulpes vulpes.
Journal of Animal Ecology 60: 423-439.
FOX M.W., BECK A.M. & BLACKMAN E. 1975. Behaviour and ecology of a small group of urban
dogs (Canis familiaris). Applied Animal Ethology 1: 119-137.
FRANCISCI F., CIUCCI P., ANDREOLI G. & BOITANI L. 1991. Storia naturale di un gruppo di cani
inselvatichiti residenti nell’areale del lupo in Marsica (AQ). Final Report to the Italian
Institute of Wildlife Biology (INFS), Bologna.
FRANK H. 1980. Evolution of canine information processing under conditions of natural and
artificial selection. Zeitschrift für Tierpsychologie 53: 389-399.
69Social ecology of feral dogs and wolves
FRANK H. & FRANK M.G. 1982. On the effects of domestication on canine social development
and behaviour. Applied Animal Ethology 8: 507-525.
FRITTS S.H. & MECH L.D. 1981. Dynamics, movements, and feeding ecology of a newly pro-
tected wolf population in northwestern Minnesota. Wildlife Monograph 80: 1-79.
FULLER T.K. 1989. Population dynamics of wolves in North-Central Minnnesota. Wildlife
Monograph 105: 1- 41.
GAVITT J.D., DOWNING R.L. & MCGINNES B.S. 1974. Effects of dogs on deer reproduction in
Virginia. Proceedings of the Annual Conference of the Southeastern Association of State
Game and Fish Commissioners 28: 532-539.
GINSBURG B. E. & SCHOTTE C.S. 1978. Genetic differences determining the capacities for social
behaviour among wolves and dogs. Behavioural Genetics 8: 94 (abstract).
GIPSON P.S. 1972. The taxonomy, reproductive biology, food habits, and range of wild Canis
(Canidae) in Arkansas. Ph.D. Dissertation, University of Arkansas, Fayetteville.
GIPSON P.S. 1983. Evaluation and control implications of behaviour of feral dogs in Interior
Alaska, pp. 78-92. In: Kaukeinen D.E., Edit. Vertebrate pest control and management
materials: 4th Symposium. Philadelphia: ASTM Special Technical Publication.
GIPSON P.S. & SEALANDER J.A. 1977. Ecological relationships of white-tailed deer and dogs in
Arkansas, pp. 3-16. In: Philips R.L. & Jonkel C., Edits. Proceedings of 1975 Predator
Symposium. Bulletin of Montana Forest Conservation Experimental Station. Missoula:
University of Montana.
GITTLEMAN J.L. 1989. Carnivore behaviour, ecology and evolution. London: Chapman & Hall.
HABER G.C. 1977. Socio-ecological dynamics of wolves and prey in a subartic ecosystem.
Ph.D. Thesis, University of British Columbia, Vancouver.
HALE E.B. 1969. Domestication and the evolution of behaviour, pp. 22-42. In: Hafez E.S.E.,
Edit. The behaviour of domestic animals. London: Bailliere Tindall.
HARRINGTON F. H. & MECH L.D. 1979. Wolf howling and its role in territory maintenance.
Behaviour 68: 207-249.
HARRINGTON F.H. & MECH L.D. 1982. Patterns of homesite attendence in two Minnesota wolf
packs, pp. 81-105. In: Harrington F.H. & Paquet P.C., Edits. Wolves of the world: per-
psectives of behavior, ecology and conservation. New Jersey: Noyes.
HARRINGTON F.H., MECH L.D. & FRITTS S.H. 1983. Pack size and wolf pup survival: their rela-
tionship under varying ecological conditions. Behavioral Ecology and Sociobiology 13:
19-26.
HARRINGTON F.H., PAQUET P.C., RYON J. & FENTRESS J.C. 1982. Monogamy in wolves: a review
of the evidence, pp. 209-222. In: Harrington F.H. & Paquet P.C., Edits. Wolves of the
world: perpsectives of behavior, ecology and conservation. New Jersey: Noyes.
HAWKINS R.E., KLIMSTRA W.D. & ANTRY D.C. 1970. Significant mortality factors of deer on
Crab Orchard National Wildlife Refuge. Transactions Illinois State Academy of Sciences
63: 202-206.
HIRATA H., OKUZAKI M. & OBARA H. 1986. Characteristics of urban dogs and cats, pp. 163-175.
In: Obara H., Edit. Integrated studies in urban ecosystems as the basis of urban plan-
ning, I. Special research project on environmental science (B276-R15-3). Tokyo: Minis-
try of Education.
HIRATA H., OKUZAKI M. & OBARA, H. 1987. Relationships between men and dogs in urban eco-
system, pp. 113-120. In: Obara H., Edit. Integrated studies in urban ecosystems as the
basis of urban planning, II. Special research project on environmental science (B334-
R15-3). Tokyo: Ministry of Education.
JOSLIN P. W.B. 1966. Summer activities of two timber wolf (Canis lupus) packs in Algoquin
Park. M.A. Thesis, University of Toronto, Toronto.
KEITH L.B. 1983. Population dynamics of wolves, pp. 66-77. In: Carbyn L.N., Edit. Wolves in
Canada and Alaska. Their status biology and management. Canadian Wildlife Service
Report Series No. 45.
KLEIMAN D.G. 1968. Reproduction in the Canidae. International Zoo Yearbook 8: 1-7.
KLEIMAN D.G. & BRADY C.A. 1978. Coyote behaviour in the context of recent canid research:
problems and perspectives, pp. 163-188. In: Bekoff M., Edit. Coyotes. New York: Aca-
demic Press.
70 L. Boitani and P. Ciucci
KLEIMAN D.G. & EISENBERG J.F. 1973. Comparisons of canid and felid social systems from an
evolutionary perspective. Animal Behaviour 21: 637-659.
KLEIMAN D.G. & MALCOM J.R. 1981. The evolution of male parental investment in mammals,
pp. 347-387. In: Gubernik D.J. & Klopfer P.H., Edits. Parental care in mammals. New
York: Plenum.
KOLENOSKY G.B. & JOHNSTON D.H. 1967. Radio-tracking in timberwolves in Ontario. American
Zoologist 7: 289-303.
KRUUK H. 1972. The spotted hyena: a study of predation and social behaviour. Chicago: Uni-
versity of Chicago Press.
KRUUK H. & SNELL H. 1981. Prey selection by feral dogs from a population of marine iguanas
(Amblyrhynchus cristatus). Journal of Applied Ecology 18: 197-204.
LINDSTROM E. 1986. Territory inheritance and the evolution of group living in carnivore. Ani-
mal Behaviour 34: 1825-1835.
LOWHEAD B.D. 1983. Wolf den site characteristics in the Nelchina Basin, Alaska. MS Thesis,
University of Alaska, Fairbanks.
LOWRY D.A. & MACARTHUR K.L. 1978. Domestic dogs as predators on deer. Wildlife Society Bul-
letin 6: 38-39.
MACDONALD D.W. 1981. Dwindling resources and the social behaviour of Capybaras, (Hydro-
choerus hydrochaeris). Journal of Zoology 194: 371-391.
MACDONALD D.W. 1983. The ecology of carnivore social behaviour. Nature 301: 379-384.
MACDONALD D.W., BALL F.G. & HOUGH N.G. 1980. Evaluation of home range size and configu-
ration, pp. 405-424. In: Amlaner C.J. & Macdonald D.W., Edits. A handbook on biote-
lemetry and radio-tracking. Oxford: Pergamon Press.
MACDONALD D.W. & CARR G.M. 1989. Food security and the rewards of tolerance, pp. 75-99.
In: Standen V. & Foley R.A., Edits. Comparative socioecology: the behavioural ecology
of humans and other mammals. Oxford: Blackwell Scientific Publications.
MACDONALD D.W. & CARR G.M. (in press). Variation in dog society: between resource disper-
sion and social flux. In: Serpell J., Edit. The dog: its ecology, behaviour and evolution.
Cambridge: Cambridge University Press.
MCKNIGHT T. 1964. Feral livestock in Anglo-America. Berkeley: University of California Press.
MECH L.D. 1970. The wolf: the ecology and behaviour of an endangered species. New York:
Natural History Press.
MECH L.D. 1975. Disproportionate sex ratios of wolf pups. Journal of Wildlife Management 39:
737-740.
MECH L.D. 1977a. Productivity, mortality and population trends of wolves in North-Eastern
Minnesota. Journal of Mammalogy 58: 559-574.
MECH L.D. 1977b. Population trend and winter deer consumption in a Minnesota wolf pack,
pp. 55-83. In: Phillips R.L. & Jonkel C., Edits. Proceedings of the 1975 Predator sympo-
sium. Bulletin of Montana Forest Conservation Experimental Station. Missoula: Univer-
sity of Montana.
MECH L.D. 1986. Wolf population in the Central Superior National Forest, 1967-1985.
U.S.D.A., North Central Forest Experimental Station Research Paper NC-270: 1-6.
MECH L.D. & NELSON M.E. 1989. Polygyny in a wild wolf pack. Journal of Mammalogy 70:
675-676.
MECH L.D. & NELSON M.E. 1990. Non-family wolf, Canis lupus, pack. The Canadian Field Nat-
uralist 104: 482-483.
MESSIER F. 1985. Social organization, spatial distribution and population density of wolves in
relation to moose density. Canadian Journal of Zoology 63: 239-245.
MURIE A. 1944. The wolves of Mount Mckinley. U.S. National Park Service Fauna Series 5: 1-238.
NELSON M.E. & MECH L.D. 1981. Deer social organization and wolf predation in Northeastern
Minnesota. Wildlife Monograph 77: 1-53.
NESBITT W.H. 1975. Ecology of a feral dog pack on a wildlife refuge, pp. 391-395. In: Fox.
M.W., Edit. The wild canids. New York: Van Nostrand Reinhold.
OLSON J.C. 1974. Movements of deer as influenced by dogs. Indiana Department of Natural
Resources, Job Progress Report Project W-26-R-5: 1-36.
71Social ecology of feral dogs and wolves
PACKARD J.M. & MECH L.D. 1980. Population regulation in wolves, pp. 135-158. In: Klein A.G.
& Cohen M., Edits. Biosocial mechanisms of population regulation. New Haven: Yale
University Press.
PACKARD J.M., MECH L.D. & SEAL U.S. 1983. Social influences on reproduction in wolves, pp.
66-77. In: Carbyn L.N., Edit. Wolves in Canada and Alaska. Their status biology and
management. Canadian Wildlife Service Report Series No. 45.
PACKARD J.M., SEAL U.S., MECH L.D. & PLOTKA E.D. 1985. Causes of reproductive failure in
two family groups of wolves (Canis lupus). Zeitschrift für Tierpsychologie 68: 24-40.
PERRY M.C. & GILES R.H. 1971. Free running dogs. Virginia Wildlife 32: 17-19.
PETERS L. & MECH L.D. 1975. Scent-marking in wolves. American Scientist 63: 628-637.
PETERSON R.O. 1977. Wolf ecology and prey relationship on Isle Royale. National Park Service
Scientific Monographs Series 11: 1-210.
PETERSON R.O., WOOLINGTON J.D. & BAILEY T.N. 1984. Wolves of the Kenay peninsula, Alaska.
Wildlife Monograph 88: 1-52.
PRICE E.O. 1984. Behavioural aspects of animal domestication. The Quarterly Review of Biolo-
gy 59: 1-32.
PROGULSKE D.R. & BASKETT T.S. 1958. Mobility of Missouri deer and their harassment by
dogs. Journal of Wildlife Management 22: 184-192.
RABB G.B., WOOLPY J.H. & GINSBURG B.E. 1967. Social relationships in a group of captive wol-
ves. American Zoologist 7: 253-265.
RAUSCH R.A. 1967. Some aspects of the population ecology of wolves, Alaska. American Zoolo-
gist 7: 253-265.
ROTHMAN R.J. & MECH L.D. 1979. Scent-marking in lone wolves and newly formed pairs. Ani-
mal Behaviour 27: 750-760.
SANTAMARIA A., PASSANNANTI S. & DIFRANZA D. 1990. Censimento dei cani randagi in un quar-
tiere di Napoli. Acta Medica Veterinaria 36: 201-213.
SCOTT M.D. & CAUSEY K. 1973. Ecology of feral dogs in Alabama. Journal of Wildlife Manage-
ment 37: 253-265.
SCOTT J.P. & FULLER J.L. 1965. Genetics and the social behaviour of the dog. Chicago: Univer-
sity of Chicago Press.
SEAL U.S. & MECH L.D. 1983. Blood indicators of seasonal metabolic patterns in captive adult
gray wolves. Journal of Wildlife Management 47: 704-715.
SEAL U.S., MECH L.D. & VAN BALLENBERGHE V. 1975. Blood analyses of wolf pup and their eco-
logical and metabolic interpretation. Journal of Mammalogy 56: 64-75.
SEAL U.S., PLOTKA E.D., MECH L.D. & PACKARD J.M. 1987. Seasonal metabolic and reproduc-
tive cycles in wolves, pp. 109-125. In: Frank H., Edit. Man and wolf. The Netherlands:
Junk Publisher.
SWEENEY J.R., MARCHINTON R.L. & SWEENEY J.M. 1971. Responses of radiomonitored white-
tailed deer chased by hunting dogs. Journal Wildlife Management 35: 707-716.
URIOS V., VILA C. & CASTROVIEJO J. 1993. Foraging strategies of the Iberian wolf (Canis lupus
signatus). Simposium Internacional sobre el Lobo (Leon, 19-23 October, 1993, Spain)
(abstract).
VAN BALLENBERGHE V. 1983. Two litters raised in one year by a wolf pack. Journal of Mammal-
ogy 64: 171-173.
VAN BALLENBERGHE V., ERICKSON A.W. & BYMAN D. 1975. Ecology of the timber wolf in North-
eastern Minnesota. Wildlife Monograph 43: 1-43.
VAN BALLENBERGHE V. & M ECH L.D. 1975. Weights, growth, and survival of timber wolf pups
in Minnesota. Journal of Mammalogy 56: 44-63.
VILA C., URIOS V. & C ASTROVIEJO J. 1993. The iberian wolf: a persecuted species. Simposium
Internacional sobre el Lobo,(Leon, 19-23 October, 1993, Spain) (abstract).
VON SCHANTZ T. 1984. Carnivore social behaviour: does it need patches? Nature 307: 389.
WORLD HEALTH ORGANIZATION 1988. Report of WHO consultation on dog ecology studies relat-
ed to rabies control. WHO/Rab. Res./88.25.
ZEUNER F.E. 1963. A history of domesticated animals. London: Hutchinson & Co. Publishers.
ZIMEN E. 1976. On the regulation of pack size in wolves. Zeitschrift für Tierpsychologie 40:
300-341.
72 L. Boitani and P. Ciucci
ZIMEN E. 1982. A wolf pack sociogram, pp. 282-382. In: Harrington F.H. & Paquet P.C., Edits.
Wolves of the world. Perspectives of behavior, ecology and conservation. New Jersey:
Noyes.
ZIMEN E. & BOITANI L. 1975. Number and distribution of wolves in Italy. Zeitschrift für Säuge-
tierkunde 40: 102-112.
