Aseasonality of Bush Dog Reproduction and the Influence of Social Factors on the

Estrous Cycle
Author(s): Ingrid J. Porton, Devra G. Kleiman and Melissa Rodden
Source: Journal of Mammalogy , Nov., 1987, Vol. 68, No. 4 (Nov., 1987), pp. 867-871
Published by: American Society of Mammalogists

Stable URL: https://www.jstor.org/stable/1381569

REFERENCES
Linked references are available on JSTOR for this article:
https://www.jstor.org/stable/1381569?seq=1&cid=pdf-
reference#references_tab_contents
You may need to log in to JSTOR to access the linked references.

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide
range of content in a trusted digital archive. We use information technology and tools to increase productivity and
facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at
https://about.jstor.org/terms

is collaborating with JSTOR to digitize, preserve and extend access to Journal of Mammalogy

This content downloaded from

177.17.141.39 on Wed, 15 Nov 2023 18:29:09 +00:00
All use subject to https://about.jstor.org/terms
 November 1987 GENERAL NOTES 867

LITERATURE CITED

ALLEN, J. A. 1905. The mammalia of southern Pat- kins University Press, Baltimore, Maryland, 1 and
agonia. in Reports of the Princeton University Ex- 2:1-1362.
peditions to Patagonia. Vol III, pt. 1. Zoology. The OLROG, C. C., AND M. M. LUCERo. 1980. Guia de
New Era Printing Co., Lancaster, Pennsylvania, los mamiferos argentinos. Ministerio de Cultura y
210 pp. Educaci6n, Fundaci6n Miguel Lillo, San Miguel
CABRERA, A. 1957. Catalogo de los mamiferos de de Tucuman, 151 pp.
America del Sur. Rev. Mus. Arg. Cienc. Nat., Bs. OSGOOD, W. H. 1943. The mammals of Chile. Field
As., No. 1, 307 pp. Mus. Nat. Hist. Zool. Ser., 30:1-268.
CABRERA, A., AND J. YEPES. 1940. Mamiferos Sud PATTON, R. F. 1974. Ecological and behavioral re-
Americanos (vida, costumbres y descripci6n). His- lationships of the skunks of Trans Pecos, Texas.
toria Natural Ediar, Compafiia Argentina de Edi- Unpubl. Ph.D. dissert., Texas A&M University,
tores, Buenos Aires, Argentina, 370 pp. College Station.
FULLER, T. K., D. W. KUEHN, P. L. COY, AND R. K. PISANO, E. 1974. Estudio ecol6gico de la regi6n
MARKL. 1985. Physical characteristics of striped continental sur del irea Andino Patagonica. II.
skunks in northern Minnesota. J. Mamm., 66:371- Contribuci6n a la fitogeografia de la zona del Par-
374. que Nacional Torres del Paine. Anales del Instituto
MILLER, S., AND J. ROTTMANN. 1976. Guia para el de la Patagonia (Punta Arenas), 5:59-104.
reconocimiento de mamiferos chilenos. Editore RAMSDEN, R. O., P. F. COPPIN, AND D. H. JOHNSTON.
Nacional Gabriela Mistral, Santiago, 200 pp. 1976. Clinical observations on the use of ketamine
NOWAK, R. M., AND J. L. PARADISO. 1983. Walker's hydrochloride in wild carnivores. J. Wildl. Dis.,
mammals of the world. Fourth ed. The Johns Hop- 12:221-225.

Submitted 29 July 1986. Accepted 20 October 1986.

J. Mamm., 68(4):867-871, 1987

ASEASONALITY OF BUSH DOG REPRODUCTION AND THE INFLUENCE OF

SOCIAL FACTORS ON THE ESTROUS CYCLE

INGRID J. PORTON, DEVRA G. KLEIMAN, AND MELISSA RODDEN

Department of Zoological Research, National Zoological Park, Smithsonian Institution,

Washington, D.C. 20008
Present address of IJP: St. Louis Zoo, Forest Park, MO 63110

Temperate species of canids, both in captivity and in the wild, exhibit a rigid reproductive cycle
seasonal breeding and only one estrous period per year (Asdell, 1964; Ewer, 1973; Kleiman, 1968). Sc
accounts indicate that some tropical species of canids exhibit a greater degree of reproductive flex
Lycaon pictus, cerdocyon thous, and Canis aureus in tropical Asia, are reproductively aseasonal (Bra
Cullen, 1976; Frame et al., 1979; Nowak and Paradiso, 1983). C. thous (Brady, 1978) and Speothos ven
(Kleiman, 1968) exhibit two estrous cycles per year, and L. pictus (Brand and Cullen, 1976; Fram
1979) and Fennecus zerda (Koenig, 1970) have been reported to produce a second litter of pups w
months of losing the first.
The following observations of captive bush dogs (S. venaticus) suggest that this tropical canid po
more labile reproductive cycle than previously thought. Our research indicates that this species is a
and occasionally polyestrous with the reproductive cycle influenced, in part, by social factors.
The main subjects of this study were five pairs of captive bush dogs whose identities and ages, p
dates, social environment prior to pairing, and subsequent reproductive histories are presented in T
The animals were housed in 4.9 by 10.4 m or 8.5 by 10.4 m outdoor yards, with access to 2.4 by
indoor dens at the National Zoological Park's Conservation and Research Center (CRC) in Front
Virginia. They were exposed to ambient temperatures and natural light cycles except during f
temperatures, when they were locked indoors.

This content downloaded from

177.17.141.39 on Wed, 15 Nov 2023 18:29:09 +00:00
All use subject to https://about.jstor.org/terms
 TABLE 1.-The reproductive history of five pairs of bush dogs at the National Zoological Park's C
Pair

Individual his
Indi Age at Interestrous
vidual pairing interval inte
I.D. (months) Prior social condition Pairing date Estrous period (ties

6 102669 9 with brother 28 Aug 1979 2 Sept 1979 (1

9 102231 17 alone 21-24 Feb 1980 (*) 17
18-29 Dec 1980
28-30 June 198
22 Feb-3 Mar 19
1-7 April 1982 (
6 103330 48 alone 26 March 1980 19 April 1980 (1)
9 102671 15 with father and brother 26-29 Dec 1980 (6)
17-21 Aug 198
20-24 Feb 1982
a 102668 26 alone 13 Feb 1981 6-11 March 1981 (4)
9 103561 7.5 with parents and two sisters 24-28 April 19
24-27 Sept 198
7-10 March 1982 (
6 103462 11 with brother 13 April 1981 9-10 May 1981 (2)
Y 103560 9.5 with sister 10 June 1981 (*)
5 July 1981 (1
20 July 1981 (1
13-14 Aug 1981 (
19-25 Sept 19
5-9 April 1982
6 103461 11 with brother 13 April 1981 8 May 1981
9 103559 9.5 with sister

* Copulatory tie not observed, but male mounting, female lordosis, and vulval swelling and redn
** Pairing-to-estrus interval.
*** Copulation during pregnancy after pair moved to a new cage.

This content downloaded from

177.17.141.39 on Wed, 15 Nov 2023 18:29:09 +00:00
All use subject to https://about.jstor.org/terms
November 1987 GENERAL NOTES 869

Males and females were introduced in on

paired. After permanent cohabitation, pair
observed 12 to 14 h per week. Compara
(Chrysocyon brachyurus) are from CRC re
We defined estrus as the occurrence of (1
region concurrent with the female's assumpt
encompassed the days when we observed es
Captive bush dogs did not exhibit a fixed b
was as follows: February, 2; March, 4; Apri
1; November, 5; December, 3 (Dmoch, 198
recorded in every month at CRC with an a
crab-eatingfoxes, also born at the CRC facili
and October, suggesting aseasonality in this
maned wolves were born from December t
Estrous periods of bush dogs at CRC ran
young primiparous females (103560 and 1
condition never before reported for a canid
44 days. Within a 1-year period female 103
female 103561 had four estrous periods a
conception was 10 months (female 103561).
Additional observations suggest that in bu
alone, with sisters, or with one or both par
primiparous females (age 7.5 to 17 months)
estrous behavior within 4 weeks of pairing
Two family groups, one with three fem
offspring, were kept intact for 7.5 and 10
exhibited physical or behavioral signs of pu
littermates that regularly urine marked at
living with the parents. In one family, fem
into estrus 22 days after being paired with
103560) were separated from the parents an
itored daily for signs of vulval swelling an
female 103559 5 days after separation from
separation from the parents, the two siste
within 25 and 26 days respectively.
In another family group, the five 10-mon
together, and within 11 days one of the fem
a third family, female 102671 remained hous
death, when the female was 11 months old
within 26 days of being paired with an unf
Typically, females have a 238 +39.6 (ran
1981; Roben, 1974-1979; CRC records). Ge
occur until approximately 171 days postpartu
estrus was significantly shorter in five fe
compared with 11 females who raised yo
1980-1981; Roben, 1974-1979; CRC records
removed for medical reasons not related to
2, 5, and 21 days after the pups' removal
bred 79 days postpartum. Similarly, pups f
1981) died at 4 days of age and, based on th
days later. Thus, the interbirth interval ma
died.

The above data indicate that bush dog rep

factors. Moreover, reproductive suppression
for mammalian species that are monogamou
e.g., in marmosets and tamarins (Abbott an
(Frame et al., 1979; Zimen, 1976), and in
Mech, 1976) and in two callitrichid species

This content downloaded from

177.17.141.39 on Wed, 15 Nov 2023 18:29:09 +00:00
All use subject to https://about.jstor.org/terms
 870 JOURNAL OF MAMMALOGY Vol. 68, No. 4

parental influence results in reproduction at a younger ag

social organization in the wild, but captive juveniles and p
long as they are housed together suggesting strong family b
It appears that the reproductive strategy of the bush dog i
with the potential of a large number of young. However,
similar to other canids since bush dog litters are relativel
(CRC records; Dmoch, 1980-1981; Emerson, 1972; Jantsc
especially when compared to other group-living, pack-hunt
to 6.5 (range 1-9), African wild dog litters are usually s
alpinus) litters are typically eight (Johnsingh, 1979). Furth
and in the same direction as litter size; wolves have 10 te
whereas bush dogs have 8 teats (Ewer, 1973). Thus, small li
per year, and reproductive flexibility appear to be coevolv
All canids which have been reported to exhibit some deg
distribution. However, not all tropical canids are reproduc
to climate, seasonal variation in food availability most like
species. Bush dogs probably prey on medium-sized mamma
paca), capybaras (Hydrochoerus hydrochoeris), and smal
1975). Where they inhabit forest with permanent access to
numbers and density (Collett, 1981; Macdonald, 1981; Sm
have allowed year-round breeding by bush dogs and there
influenced by social factors.
There is an obvious need for more research on both the
dogs and other tropical canids. At a time when the existence
destruction, the need to understand their basic reproducti
This research was supported by NIMH grant 27241 aw
Lumpkin, H. Smith, C. Wemmer, J. F. Eisenberg, and two re
Reprint requests should be directed to D. G. Kleiman.

LITERATURE CITED

ABBOTT, D. H., AND J. P. HEARN. 1978. Physical, EWER, R. F. 1973. The carnivores. Cornell Univ.
hormonal, and behavioral aspects of sexual devel- Press, New York, 494 pp.
opment in the marmoset monkey, Callithrix jac-FRAME, L. H., J. R. MALCOLM, G. W. FRAME, AND H.
chus. J. Reprod. Fertil., 53:155-166. VAN LAWICK. 1979. Social organization of African
ASDELL, S. A. 1964. Patterns of mammalian repro- wild dogs (Lycaon pictus) on the Serengeti Plains,
duction. Cornell Univ. Press, Ithaca, New York, Tanzania (1967-1978). Z. Tierpsychol., 50:225-249.
670 pp. JANTSCHKE, F. 1973. On the breeding and rearing
BRADY, C. A. 1978. Reproduction, growth and pa- of bush dogs, Speothos venaticus at Frankfurt Zoo.
rental care in crab-eating foxes, Cerdocyon thous, Int. Zoo Yearb., 13:141-143.
at the National Zoological Park, Washington. Int. JOHNSINGH, A. J. T. 1979. Ecology and behaviour
Zoo Yearb., 18:130-134. of the dhole or Indian wild dog, Cuon alpinus
BRAND, D. J., AND L. CULLEN. 1976. Breeding the Pallas 1811, with special reference to predator-
Cape Hunting dog Lycaon pictus at Pretoria Zoo. prey relationships at Bandipur. Unpubl. Ph.D. dis-
Int. Zoo Yearb., 7:124-126. sert., Madurai Univ., Madurai, India, 308 pp.
COLLETT, S. F. 1981. Population characteristics of KITCHENER, S. L. 1971. Observations on the breed-
Agouti paca (Rodentia) in Columbia. Publ. Mus., ing of the bush dog Speothos venaticus at Lincoln
Michigan State Univ. (Biol. Ser.), 5:485-601. Park Zoo, Chicago. Int. Zoo Yearb., 11:99-101.
DEUTSCH, L. A. 1983. An encounter between bush KLEIMAN, D. G. 1968. Reproduction in the Canidae.
dog (Speothos venaticus) and paca (Agouti paca). Int. Zoo Yearb., 8:3-8.
J. Mamm., 64:532-533. . 1979. Parent-offspring conflict and sibling
DMOCH, R. 1980-1981. International studbook for competition in a monogamous primate. Amer. Nat.,
the bush dog, Speothos venaticus (Lund, 1842). 114:753-760.
Frankfurt Zoological Society. KOENIG, L. 1970. Zur Fortpflanzung and Jugen-
EMERSON, S. B. 1972. Notes on bush dogs. Ann. dentwicklung des Wustenfuchses (Fennecus zerda
Proc. Amer. Assoc. Zool. Parks Aquar. Conf., 48: Zimm. 1780). Z. Tierpshchol., 27:205-246.
79-86. LANGGUTH, A. 1975. Ecology and evolution in South
EPPLE, G., AND Y. KATZ. 1980. Social influences on American canids. Pp. 192-206, in The wild canids
first reproductive success and related behaviors in (M. Fox, ed.). Van Nostrand Reinhold, New York,
the saddle-back tamarin (Saguinus fuscicollis, 508 pp.
Callitrichidae). Int. J. Primatol., 1:171-183. MACDONALD, D. W. 1981. Dwindling resources and

This content downloaded from

177.17.141.39 on Wed, 15 Nov 2023 18:29:09 +00:00
All use subject to https://about.jstor.org/terms
 November 1987 GENERAL NOTES 871

the social behavior in of capybaras,

pair formation and maintenance.(Hydroc
Anim. Behav.,
hydrochaeris) (Mammalia). 31:1061-1069. J. Zool., London
171-391. ROBEN, P. 1974-1979. International studbook for
MECH, L. D. 1970. The wolf: the ecology and be- the bush dog, Speothos venaticus (Lund, 1842).
Frankfurt Zoological Society.
havior of an endangered species. Natural History
Press, New York, 384 pp. ROOD, J. P. 1980. Mating relationships and breed-
MEDJO, D. C., AND L. D. MECH. 1976. Reproductiveing suppression in the dwarf mongoose. Anim. Be-
activity in nine and ten month old wolves. J.hav., 28:143-150.
Mamm., 57:406-408. SMYTHE, N. 1978. The natural history of the Central
NOWAK, R. M., AND J. L. PARADISO. 1983. Walker's American agouti (Dasyprocta punctata). Smith-
mammals of the world. Fourth ed. The Johns Hop-sonian Contrib. Zool., 257:1-52.
ZIMEN, E. 1976. On the regulation of pack size in
kins Univ. Press, Baltimore, Maryland, 1 and 2:1-
1362. wolves. Z. Tierpsychol., 40:300-341.
PORTON, I. 1983. Bush dog urine-marking: its role

Submitted 7 November 1985. Accepted 31 October 1986.

J. Mamm., 68(4):871-873, 1987

PREMATURE REPRODUCTIVE ACTIVITY IN WILD WOLVES

L. DAVID MECH AND ULYSSES S. SEAL

U.S. Fish and Wildlife Service, Patuxent Wildlife Research Center, Laurel, MD 20708
V.A. Medical Center, Research Service 54th St. & 48th Ave. So., Minneapolis, MN 55417
Departments of Biochemistry and Fisheries and Wildlife, University of Minnesota, St. Paul, MN 55108
Mailing address for Mech: North Central Forest Experiment Station, 1992 Folwell Ave.,
St. Paul, MN 55108

This note discusses two female wolves that showed signs of reproductive activity despite anatomical
evidence that they were reproductively immature.
Some female wolves can conceive at 10 months of age (Medjo and Mech, 1976; Zimen 1976), but most
do not breed until at least 22 months old (Seal et al., 1979). The youngest wild female wolf known to have
produced pups was 3 years old, although evidence from nipple size and condition indicated that a 2-year-
old wolf produced pups that died before 4-months old (Mech, 1987). Captive wolves sometimes undergo
pseudopregnancy (Seal et al., 1979), a condition in which a non-pregnant female with a corpus luteum
sustains a hormonal and behavioral state indistinguishable from that of a pregnant animal at about 2 months
post-estrus (Johnston, 1986). Pseudopregnancy has not been reported from wild wolves.
On 9 May 1970 wolf 2202 of unknown age and weighing 38.6 kg was live trapped in St. Louis County,
Minnesota. Her nipples were flaccid and about 3 mm wide, an area of about 28 cm2 around each was barren
of guard hair (although underfur was present), and white, milk-like material was squeezed from each. The
wolf was tagged and released, and on 25 March 1971 she was killed; she weighed 36.8 kg. Her reproductive
tract was of mature size, but no placental scar, developing follicle, corpus luteum, or corpus albicans, was
apparent upon gross examination.
The second wolf, 6433, was born in April 1982 to 8-year-old radioed female 5176 and radioed male 5132,
who was at least 9-years old (Mech, 1987). Wolf 6433 was live trapped and radio-tagged as a pup on 19
September 1982 and was caught twice more in 1982 and twice in 1983. Luteinizing hormone (LH) levels
at each capture indicated that 6433's hypothalamic-pituitary axis was normal, that she was not a functional
castrate, and that in December 1982 she was approaching puberty. Her movements were monitored regularly
until her death about 8-14 February 1985.
On 22 November 1983, # 6433 was first found outside her natal pack's territory, and 53 of 55 subsequent
radio-locations in the next 6 months were outside her natal territory. She was first seen with another wolf
on 11 January 1984 and, of the next 15 times she was observed, she was with a companion 12 times. On 12

This content downloaded from

177.17.141.39 on Wed, 15 Nov 2023 18:29:09 +00:00
All use subject to https://about.jstor.org/terms