Download Paternal Care in Canids1

AMER. ZOOL., 25:853-856 (1985)
Paternal Care in Canids1
JAMES R. MALCOLM
Department of Biology, University of Redlands,
Redlands, California 92374
SYNOPSIS. Paternal care has never been reported as absent in any canid species, and some
form of care has been seen in 18 of the 36 species in the family. Food provisioning, active
defense of the young, and protecting young by remaining at the den as the female forages
appear to be the commonest forms of male care. In addition males may groom, retrieve,
play and rest with young. Male canids are rarely involved in den selection or construction.
The effect on the fitness of the young of indirect forms of male care such as provisioning
the female and territory defense are hard to assess. Quantitative studies of male provisioning in seven species offer few generalizations. In two species (Canis aureus, C mesomelas)
females provided more food to the young than males; in one species (Alopex lagopus) the
pair contributed equally to feeding young, and in four species (Canis lupus, Vulpes vulpes,
Chrysocyon brachyurus, and Lycaon pictus), males provided more food than females. Much
more data are required, particularly from field studies, before patterns of variation can
be interpreted.
Indirect care involves those behaviors
Some form of male parental care, usually
in the context of biparental investment in which could benefit offspring even if there
monogamous pairs, is probably universal is no direct interaction between the male
in the family Canidae. Kleiman and Mal- and young. Den construction, giving alarm
colm (1981) reviewed literature up to 1979, calls, and caring for females fall into this
and found reports of male care in 17 of 35 category. In addition, the establishment
species. In no species was male care and maintenance of a territory may be a
reported to be completely absent. Paternal prerequisite for breeding but also benefits
care has subsequently been reported in two young.
more fox species {Vulpes bengalensis (John- In most canids, pairs or groups defend
singh, 1978], and V. macrotis [McGrew, multi-purpose territories (Kleiman and
1979] now recognized as distinct from V. Eisenberg, 1973) and all adults help directly
velox, in which male care was reported by in raising the young. The most commonly
Seton [1909]). Male care has, therefore, recorded forms of care are food provisionbeen seen in exactly half (18/36) of the ing (by regurgitation in larger species, and
species in the family.
carrying food items to the den in smaller
Kleiman and Malcolm (1981) distin- species), active defense of the young (often
guished between direct and indirect pat- reported in captive situations where males,
terns of care-giving behavior. Direct care with young, attack "keepers"), and care to
comprises those interactions between males the female, usually in the form of food proand their offspring which can reasonably visioning. However, in the species with the
be interpreted to increase offspring fitness. most extensive direct field observations, the
In canids, direct care can be classified into African wild dog {Lycaon pictus), all cateeight categories: resting and huddling with gories of male care were observed, with the
young, grooming and cleaning, carrying, exception of huddling with neonates, and
retrieving, providing food, defending den construction. Most categories of male
against predators, babysitting (remaining care probably occur in other species but
with young while others forage), and play- have not been reported. Den selection and
construction may be an exception, as it
ing or socializing.
appears to be a female perogative in canids.
(Data from Kleiman and Malcolm, 1981,
1
From the Symposium on Paternal Behavior pre- Table 2.)
sented at the Annual Meeting of the American SociSome recent quantitative studies of
ety of Zoologists, 27-30 December 1983, at Philaparental behavior in canids (Table 1) prodelphia, Pennsylvania.
853
854
JAMES R. MALCOLM
TABLE 1. Quantitative studies of parental care in canids.
Species
Location
Behaviors
Canis lupus
Field
Atten.
Wolf
Prov.
Canis lupus
Capt.
Canis aureus
Atten.; Prov.
Field
Golden jackal
Canis mesomelas
Atten.; Prov.
Field
Silver-back jackal
Vulpes vulpes
Prov.; Def.
Capt.
Red fox
Aloxpex lagopus
Atten.; Prov.
Field
Arctic fox
Chrysocycn brachyurus
Prov.
Capt.
Maned wolf
Lycaon pictus
Field
Atten.; Prov.;
African wild dog
Def.; Hunt
Key: Capt., captivity; Prov. provisioning young; Atten., attendance at
hunting.
vide some details on the distribution of care
giving behavior between the sexes. Five of
seven species (wolf, both jackals, the African wild dog and the red fox) on which
quantitative data have been reported, may
live in groups which are larger than the
nuclear family. In these species paternal
care can be directed to offspring from two
years, and care-giving behavior can also be
provided to younger offspring by males
other than the father. Some inter-species
comparisons can be made with respect to
attendance at the den and food provisioning.
Female canids usually spend more time
at the den than males. This seems to be
universally true in the first two weeks after
parturition when females seldom leave the
den. When young are more than a month
old, females in wolves, silver (black)-backed
jackals and Arctic foxes are reported to
spend more time than males in the vicinity
of pups (although the total time either adult
is present decreases with the pups' age).
Similarly, in African wild dogs, dominant
males (the probable fathers) are significantly less likely than mothers to remain
at the den when the pack hunts, and mothers actively chased more predators from
the vicinity of the den than did dominant
males (34 vs. 24). Conversely, dominant
males played a greater role in the pack's
Aulhor(s)
Harrington and Mech (1982)
Fentress and Ryon (1982)
Moehlman(1983)
Moehlman(1983)
Macdonald(1979)
Garrott (1980)
Rasmussen and Tilson (1984)
Malcolm and Marten (1982)
den; Def., defense of young; Hunt,
hunting than did mothers. (Data from Malcolm and Marten, 1982.)
In only two cases have males been
recorded spending more time at the den
than females. In silver (black)-backed jackals, the mother was with her pups almost
constantly for the first two weeks. However, in the third week her attendance
dropped sharply (to «35% of the time).
Simultaneously, the male's attendance rose
steadily from the second to the fourth week,
and in the third week the males spent almost
twice as much time at the den as females.
It appeared as if the female, after two weeks
confinement, foraged extensively in the
third week, and the male compensated by
babysitting. A preponderance of male
attendance has been recorded in bat-eared
foxes (Otocyon megalotis) (Malcolm, personal
observation; Nel, 1978). In this nocturnal
insectivorous species, the young are unusually dependent on milk as it is not economical to carry insects back to the den. Females
foraged through most of the night and the
male at two dens spent significantly more
time at the den than did the female.
Contributions of males versus females to
provisioning young are variable. In one
captive wolf pair (Fentress and Ryon, 1982;
litter of 1973), two captive pairs of maned
wolves (Rasmussen and Tilson, 1984;
Anon., 1983), and a family of red foxes in
855
PATERNAL CARE IN CANIDS
a pen (Macdonald, 1979) males provided
twice as much food to the pups as the
mother. In four packs of African wild dogs,
dominant males provided slightly more
food to pups than did mothers, but this
difference was mainly a result of mothers
staying with young pups. In Fentress and
Ryon's (1982) captive wolf pack (with multiple litters and uncertain paternity), males,
on a per capita basis, provided more food
than females in four offiveyears. The sexes
contributed almost exactly equally to young
of the year, but the males (and not the
females) also responded to begging by
yearlings which increased their total contribution.
In Arctic foxes there was no difference
in the food brought to the four dens by
males and females (Garrott, 1980). However, in both golden and silver (black)backed jackals females were recorded
regurgitating more frequently than males
(Moehlman, 1983).
The data on male care allow few definite
conclusions on the influence of male parental investment on canid social organization.
However, the results seem broadly in
agreement with Trivers' (1972) predictions that the sex providing the most
parental investment becomes a limiting
commodity for the other sex. Although
both sexes play an important role in raising
the young in canids, females play a special
role in choosing a den, lactating and
remaining with young pups. In quantitative field studies females also appear to provide as much or more food directly to the
young as males. It appears that, as predicted, most deviations from monogamy,
at least in smaller species, are in the direction of polygyny (e.g., Reynolds, 1977 for
Otocyon, von Schantz, 1981 for Vulpes
females. In African wild dogs (Frame et al,
1979), this seems to have led to a sexreversed species in which females compete
for access to groups of investing males.
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