Notes and records - African Wildlife Conservation Fundafricanwildlifeconservationfund.org/wp-content/... · for four females in and near Masai Mara National Reserve, Kenya (Fuller

Notes and records

Long-distance transboundary dispersal ofAfrican wild dogs among protected areas insouthern Africa

Harriet T. Davies-Mostert1,2,*, Jan F. Kamler1, Michael

G. L.Mills3,4, CraigR. Jackson5, Gregory S.A. Rasmussen1,6,

Rosemary J. Groom4,7 and David W. Macdonald1

1Wildlife Conservation Research Unit, Recanati-Kaplan

Centre, Department of Zoology, Oxford University, Tubney

House, Abingdon Road, Tubney, Abingdon, OX13 5QL,

U.K., 2Endangered Wildlife Trust, Private Bag X11,

Modderfontain, 1645, Gauteng, South Africa, 3Tony and

Lisette Lewis Foundation, Private Bag X5890, Upington,

8800, South Africa, 4Mammal Research Institute,

University of Pretoria, Pretoria, 0002, South Africa,5Department of Biology, Norwegian University of Science and

Technology (NTNU), Realfagbygget, 7491, Trondheim,

Norway, 6Painted Dog Research, Natural History Museum,

Bulawayo, Zimbabwe and 7African Wildlife Conservation

Fund, 10564 NW 57th Street, Doral, FL 33178, U.S.A.

Introduction

African wild dogs (Lycaon pictus), hereafter wild dogs,

have exhibited severe declines in their distribution

throughout Africa, disappearing from at least 25 coun-

tries during the past 50 years (Fanshawe et al., 1997).

Wild dogs occur at low densities and even the largest

protected areas typically support low populations. The

species is highly susceptible to edge effects and problems

associated with small population sizes. Throughout their

range, wild dogs live mainly in protected areas in highly

fragmented populations, and few areas hold more than

100 individuals (Fanshawe et al., 1997), although a

number of large contiguous populations persist in parts

of east and southern Africa. In South Africa, wild dogs

historically occurred throughout the country (Skinner &

Smithers, 1990), but owing to widespread persecution

and prey loss, now occur naturally only in a single viable

population in Kruger National Park and some adjacent

farming areas (Fanshawe et al., 1997; Lindsey, Du Toit &

Mills, 2004). Because there is no suitable large protected

area to establish a second viable population of wild dogs in

South Africa, conservation efforts focused on creating a

managed metapopulation using a network of smaller

reserves (Mills et al., 1998). Consequently, beginning in

1997, packs were reintroduced into small (85–1100 km2),

isolated, fenced reserves with the goal of establishing a min-

imum of nine packs linked through active management by

2007 (Davies-Mostert, Mills & Macdonald, 2009; Gusset,

2010). Northern Botswana is an important stronghold for

the species; however, very few resident packs occur in the

south-east of the country bordering South Africa and

Zimbabwe. In Zimbabwe, the largest population occurs

in the Zambezi Valley, with secondary populations in

Hwange National Park and associated conservation

areas, as well as the lowveld region in the south (Ras-

mussen, 1997). Populations in southern Africa could

feasibly be linked by dispersal if wild dogs were capable

of navigating the intervening landscape.

This study describes eight long-distance (>80 km) dis-

persal events by wild dogs between isolated nature

reserves, and in one case on private land. All these

occurred between September 1992 and April 2011.

Wild dog dispersal typically involves same-sex groups

that leave their natal pack in search of mates (McNutt,

1996). Although dispersal distances of up to 250 and

195 km have been previously reported for males and

females, respectively (Fuller et al., 1992), the dispersal

events reported here are the longest recorded for wild

dogs and show that isolated subpopulations could

potentially be linked in a metapopulation via dispersing

individuals.

Materials and methods

Study species

Wild dogs are medium-sized (~25 kg) social canids that

live in packs of anywhere from 2 to 30 adults and year-

lings (Woodroffe, McNutt & Mills, 2004). New packs typi-*Correspondence: E-mail: [email protected]

500 © 2012 Blackwell Publishing Ltd, Afr. J. Ecol. 50, 500–506

cally form when small same-sex groups (usually siblings)

disperse from their natal pack to locate and join oppo-

site-sex groups from unrelated packs (McNutt, 1996).

Study areas

Data for this study were collected between 1992 and

2011 from three fenced reserves in South Africa that are

part of the metapopulation programme for wild dogs,

two reserves in Zimbabwe and one reserve in Botswana.

Sites included Pilanesberg National Park (Pilanesberg;

500 km2), Marakele National Park (Marakele; 740 km2)

and the De Beers Venetia Limpopo Nature Reserve (Vene-

tia; 320 km2) in South Africa; Hwange National Park

(Hwange; 14,650 km2) and Save Valley Conservancy

(Save; 3450 km2) in Zimbabwe; and the Northern Tuli

Game Reserve (Tuli; 720 km2) in Botswana (Fig. 1,

Table 1). All sites supported populations of wild dogs at

some point during the study period, either as naturally

occurring or as reintroduced packs (Table 1). All reserves

occur within the savannah biome that dominates this

region.

Methods

Identification of wild dogs was based on pelage charac-

teristics, as each wild dog has a unique coat pattern and

can be readily identified using photographs (Maddock &

Mills, 1994; Creel, Mills & McNutt, 2004). Within the

fenced reserves in South Africa, and unfenced reserve in

Botswana, wild dogs were monitored intensively by

researchers and staff, and data were recorded when wild

dogs broke out of reserves and went missing. Populations

in Zimbabwe were monitored less frequently, and there-

fore, dispersal dates and intervals provided here are only

an approximate estimation. When new wild dogs

appeared in reserves, they were easily identified using

photographs from the other reserves.

Wild dogs periodically break out of the fenced reserves

in South Africa and are usually recaptured <80 km

away, and returned (H.T. Davies-Mostert, pers. obs.;

Gusset et al., 2008a). However, this study focused only

on long-distance (>80 km) dispersal events that occurred

during the study. Dispersal distances were calculated

as the straight-line distance from the last confirmed

sightings in the source reserve to where they were first

re-sighted or recaptured. When these details were

unknown, the shortest straight-line distance between

reserve edges was used to obtain a minimum dispersal

distance. Wild dogs were classified as adult

(�24 months old) or sub-adult (12–23 months old).

Results

Eight long-distance (>80 km) dispersal events were

recorded. All involved movements to areas to which the

individuals had not been previously exposed (Fig. 1). The

details of these events, in order from shortest to longest,

with dispersal and arrival dates in parentheses are as

follows. (i) Five adult females dispersed 81 km from Mar-

akele (1 November 2005) to Pilanesberg (16 December

2005). These females dispersed and arrived as a group

and associated with a group of males in Pilanesberg. The

females were captured and moved back to Marakele

following concerns about potential inbreeding because of

related founder stock at both sites. (ii) Two sub-adult

males dispersed 169 km south-westwards from south–

western Venetia (6 September 2003) to private farmland

(20 September 2003). These males initially dispersed

from Venetia with two sub-adult females, their litter-

mates, but probably split into separate groups because

the females were not with them 14 days later. The males

were recaptured and returned to Venetia and later trans-

located to Marakele, but the females were never found

again. (iii) Two males dispersed 199 km from Hwange

(18 February 1992 in the evening) to the 40 km peg

near Bulawayo (15 March 1992). These males were later

observed with females and pups on 3 September 1992,

172 km from their last fix and 44 km from their original

dispersal site. (iv) Three adult males dispersed 255 km

from Marakele (8 May 2005) to Venetia (19 June 2005).

These males initially dispersed as a group of five, but two

males subsequently split from the group and dispersed

elsewhere (see case v). After arriving on Venetia, the

three males associated with a group of four females, and

then all seven dispersed soon thereafter. One animal from

this group was killed by a vehicle 40 km east of Venetia

(12 August 2005), a total straight-line distance of

304 km from the natal range. The others were never

found again. (v) Two adult males dispersed 262 km from

Marakele (8 May 2005) to NTGR (19 October 2005).

These males initially dispersed from Marakele with the

three males from above, but subsequently split from

them. They were never seen with females and their fate

remains unknown. (vi) One adult male dispersed 296 km

from Tuli (photographed last on 20 August 2006) to

© 2012 Blackwell Publishing Ltd, Afr. J. Ecol. 50, 500–506

Long-distance dispersal in African wild dogs 501

Marakele (15 November 2006). This male was seen prior

to dispersal within a group of three other males and one

female, but it is not known whether he dispersed alone

or with the group, as the group was not seen again on

Tuli. In Marakele, he associated with a lone female before

they both disappeared. (vii) Two sub-adult males

dispersed 447 km from Save (last seen in 29 April 2009)

to Tuli (photographed on 19 April 2011). These males

Table 1 Characteristics of the source and destination sites in southern Africa for which long-distance (>80 km) dispersal events were

recorded for African wild dogs between 2003 and 2011

Site Abbreviation Location

Year of

reintroduction

Maximum population

size during the study

Dispersal

distance (km)*

Venetia Limpopo

Nature Reserve

VLNR Limpopo Province,

South Africa

2002 30 169

Northern Tuli

Game Reserve

NTGR Central District,

Botswana

2008 27 296

476

Marakele National

Park

MNP Limpopo Province,

South Africa

2003 37 81

255

262

Pilanesberg

National Park

PNP North West Province,

South Africa

1999 30 –

Hwange

National Park

HNP Matebeleland North

Province, Zimbabwe

Resident 150 199

Save Valley

Conservancy

SVC Masvingo Province,

Zimbabwe

Resident 96 447

*Dispersal distances represent events originating from that population.

ZIMBABWE

BOTSWANA

SOUTH AFRICA Kruger National Park

Venetia LimpopoNature Reserve

MarakeleNational Park

PilanesbergNational Park

[7] 447 km

[6] 2

96 km

[5] 2

62 k

m

[1] 8

1 km

[2] 169 km

Northern TuliGame Reserve

HwangeNational Park

Save ValleyConservancy

[8] 476 km

[4] 2

55 km

[3] 199 km

0 150 30075 Kilometers

Fig 1 Long-distance (>80 km) dispersal of groups and individuals of African wild dogs among reserves in South Africa, Zimbabwe and

Botswana, 1992–2011. Solid arrows represent straight-line distances


502 Harriet T. Davies-Mostert et al.

joined up and sired pups with the resident alpha female.

(viii) Three adult males dispersed 476 km from Tuli (6

April 2008) to Hwange (20 October 2008) where they

were observed with an adult female. These males

dispersed from their natal pack in Tuli 24 h after the

pack was released from a holding boma following trans-

location from Marakele.

Discussion

Five of eight dispersal events we documented covered

255–476 km and are longer than any previously

reported for wild dogs. The previous record for dispersal

distance was 250 km for two males in Kruger National

Park, South Africa (Fuller et al., 1992). Maximum dis-

persal distances of wild dogs from other large continuous

populations are 77 km for four females in Selous Game

Reserve, Tanzania (Creel & Creel, 2002), 169 km for

males in northern Botswana (McNutt, 1996), 195 km

for four females in and near Masai Mara National

Reserve, Kenya (Fuller et al., 1992), and 150 km from

Hluhluwe-iMfolozi Park, South Africa (Somers et al.,

2008). Wild dogs can exhibit sex-biased dispersal, but

with different sexes dominating the dispersal cohort in

different populations: females in northern Tanzania

(Frame & Frame, 1976), males in northern Botswana

(McNutt, 1996) and neither sex dominating in northern

KwaZulu-Natal, South Africa (Somers et al., 2008). Our

results indicate that both sexes are capable of long-

distance dispersal, with both dispersing >80 km in our

study. That said, most of the events we describe involved

males, which dispersed farther than females, thus reinforc-

ing earlier findings that males are the less philopatric sex.

The long dispersal distances in our study are probably

due to the isolation of subpopulations and distance

between reserves. Among wolves (Canis lupus), the great-

est straight-line dispersal distances (1092 km) occurred

in low-density populations with disjunct pack distribu-

tions (Boyd et al., 1995; Wabakken et al., 2007).

Wabakken et al. (2007) concluded that the excessive dis-

persal distances of wolves occur across areas containing

suitable habitat and food resources, so the ultimate factor

for long-distance dispersals was finding a mate. The same

is true for wild dogs, particularly those in isolated or

low-density populations (Somers et al., 2008; Gusset

et al., 2009a). For example, private lands between fenced

reserves in South Africa often had similar habitat to

fenced reserves, and similar if not higher prey numbers

(H.T. Davies-Mostert, pers. obs.). Therefore, it seems the

ultimate factor for the long-distance dispersal of wild

dogs in our study was finding a mate. This might

especially be true for subpopulations in isolated reserves,

where wild dogs usually occurred in 1–2 packs of related

individuals. Wild dogs tend not to mate with relatives,

even if given frequent opportunities (McNutt, 1996);

thus, one of the only options for nonalpha members to

breed in isolated reserves was to emigrate and search for

nonrelated mates. That several of the dispersing wild

dogs found and associated with members of the opposite

sex in other reserves supports this assertion. In contrast,

in large continuous populations, the mean dispersal dis-

tance of wild dogs often was <50 km and the maximum

dispersal was never >250 km (McNutt, 1996; Creel &

Creel, 2002).

Finding pack mates is contingent on a combination

of well-timed dispersal (i.e. arriving when potential

mates are available), choosing the correct search direc-

tion, and surviving the risks associated with traversing

unprotected ranchland. Given the vast distances

covered and possible directions that could have been

followed, the likelihood of dispersing wild dogs locating

other packs by chance in the sparsely populated land-

scape of northern South Africa was extremely low. Of

six primary dispersal events recorded at Marakele

between 2003 and 2006, three groups found members

of the opposite sex (two cases are described above),

one group was captured and translocated to a different

population, and the outcome was unknown for the

remaining two (H.T. Davies-Mostert, unpublished data).

The frequency at which the dispersers found mates

was suggestive of long-range communication, such as

olfactory cues, which facilitates detection of other wild

dogs. A better understanding of how this mechanism

operates, and particularly its effective distance, would

be useful for landscape-level conservation planning for

the species.

All wild dogs that immigrated onto fenced reserves

were identified; thus, no immigrants were from the large

population in Kruger National Park (ca. 300 individuals;

Woodroffe, McNutt & Mills, 2004), which was well

within the dispersal ranges we recorded (Table 1, Fig. 1),

or other resident packs in Limpopo Province (Lindsey, Du

Toit & Mills, 2004), which may have been <50 km from

Venetia. Thus, wild dogs at the edge of large continuous

populations may disperse towards potential mates within

the main population, rather than crossing large vacant



areas where potential mates are absent. However, this

hypothesis needs to be tested.

Wild dogs dispersed through seemingly inhospitable

landscape, managing to cross a matrix of numerous pri-

vate lands (Fig. 2), many with high and electrical fencing

for game ranching purposes, and where predators were

not welcome (Lindsey, Du Toit & Mills, 2004). In fact,

within the reserves themselves, wild dogs often broke

through ‘predator-proof’ electrical fencing by digging

under or slipping through the strands, suggesting that

this type of fencing does not prevent wild dog move-

ments. This is in contrast to other large carnivores, such

as lions (Panthera leo), which can be effectively contained

in small reserves by high electrical fencing (Davies-Most-

ert, Mills & Macdonald, 2009). Anthropogenic mortality

of wild dogs is expected to be higher on private lands

than on reserves (Woodroffe, McNutt & Mills, 2004),

and we suspect that many of the dispersing wild dogs

that were never seen again were killed by humans.

Regardless, wild dogs successfully dispersed long dis-

SOUTH AFRICA

BOTSWANA

ZIMBABWE

0 30 6015 Kilometers

MarakeleNational Park

Venetia LimpopoNature Reserve

Fig 2 Private property lines between two reserves in South Africa. Darker areas in the south-east represent high density of fence lines.

Most private properties in South Africa are fenced, often with high and/or electrical fencing. African wild dogs successfully dispersed

across this landscapes several times, demonstrating this species is capable of dispersing across some of the most highly fragmented

landscapes in southern Africa (Source: ENPAT, 2001)



tances to other reserves across a matrix of private

lands, including well-fenced areas, indicating that wild

dogs are capable of navigating across some of the most

human-altered and fragmented landscapes in southern

Africa. Indeed, recent research showed that wild dogs

in an anthropogenic landscape altered their activity to

use moonlight periods more and thus decrease encoun-

ters with humans (Rasmussen & Macdonald, 2011).

Such behaviour, even though not necessarily optimal

for hunting, may thus help to explain how wild dogs

succeed to disperse across such human-dominated land-

scapes.

Management implications

The metapopulation conservation programme for wild

dogs in South Africa actively manages gene flow among

subpopulations, via transfer of wild dogs into subpopula-

tions every few years (Davies-Mostert, Mills & Macdonald,

2009). However, without human assistance, wild dogs

have contributed to this process themselves, by dispersing

among isolated subpopulations and finding groups of the

opposite sex; however, thereafter their fate is unknown.

Although many of our dispersers were prevented from

successfully reproducing (either because of the direct

management interventions or because of the absence of

suitable mates), our results cautiously suggest protected

areas for wild dogs need not be contiguous, and widely

spaced subpopulations can be connected via dispersing

individuals or groups; thus, active management may not

be necessary in all situations. Given the importance of

dispersal in wild dog conservation, dispersal can be

enhanced in innovative ways, if necessary (e.g. Graf

et al., 2006; McNutt et al., 2008). We also caution man-

agers of small reserves that traditional ‘predator-proof’

fencing may not be adequate to contain wild dogs,

especially for those individuals attempting to disperse.

Managers in other areas of Africa should consider our

results, especially where wild dog populations are small

and fragmented. Dispersers may be necessary to prevent

inbreeding and/or local extinctions of wild dogs in

small, isolated subpopulations (McNutt, 1996). There-

fore, if increasing the size of protected areas is not an

option, establishing a network of small subpopulations

on isolated reserves may be adequate to maintain a

metapopulation. However, it should be noted that when

wild dog populations become very fragmented, such as

those occurring in places like Senegal, long-distance

dispersals may cause elevated mortality as wild dogs

disperse from their natal home ranges with no hope of

finding mates.

Future research should investigate what minimal dis-

tance is necessary between reserves to maintain gene

flow among subpopulations, identify the success of these

movements and determine the dispersal levels necessary

to achieve metapopulation viability. Also, because wild

dogs can navigate across vast expanses of fragmented

and human-altered landscapes, several factors might

increase their survivorship while dispersing, such as

establishing corridors and improving human tolerance of

transient wild dogs, which probably will be key to their

survival. South Africa’s metapopulation project, for

instance, has invested heavily in the conservation and

management of wild dogs within reserves, but less so in

the linkage between reserves. The nature of the dispersal

events described here illustrates that future efforts may

additionally aim to facilitate inter-population movement

and, in this way, benefit population dynamics of the typi-

cally small populations.

Acknowledgements

Thanks to S. Dell, H. Fitchat, J. Gounaris, L. Hedges,

D. Hofmeyr, J.W. McNutt, K. Potgieter, B. Schroder,

V. Steyn and J. Sellier for providing identification data for

wild dogs on the various reserves. U. Stenkewitz assisted

with making the figures. Support for research on various

reserves was provided in South Africa by Land Rover

South Africa, De Beers Consolidated Mines, Masslift, Mar-

akele (Pty) Ltd, SANParks; in Botswana by Mashatu

Game Reserve, Tuli Safari Lodge and Northern Tuli

Game Reserve landowners; and in Zimbabwe by the

Director General of the Parks and Wildlife Management

Authority Zimbabwe for permission to conduct research,

National Geographic Conservation Trust and Columbus

Zoo. HDM was supported through a grant to DWM from

Fauna & Flora International. Matt Hayward, Markus

Gusset, Adrian Shrader, Frank van Manen and six anon-

ymous reviewers provided valuable comments on the

manuscript.

References

Boyd, D.K., Paquet, P.C., Donelon, S., Ream, R.R., Pletscher, D.H.

& White, C.C. (1995). Transboundary movements of a

recolonizing wolf population in the Rocky Mountains. In:



Ecology and Conservation of Wolves in a Changing World

(Eds L.N. CARBYN, S.H. FRITTS and D.R. SEIP). Canadian

Circumpolar Institute, University of Alberta, Edmonton,

Canada.

Creel, S. & Creel, N.M. (2002) The African Wild Dog: Behavior,

Ecology, and Conservation. Princeton University Press,

Princeton & Oxford.

Creel, S., Mills, M.G.L. & McNutt, J.W. (2004) Demography and

population dynamics of African wild dogs in three critical

populations. In: Biology and Conservation of Wild Canids (Eds

D.W. MACDONALD and C. SILLERO-ZUBIRI). Oxford University Press,

Oxford.

Davies-Mostert, H.T., Mills, M.G.L. & Macdonald, D.W. (2009)

A critical assessment of South Africa’s managed

metapopulation recovery strategy for African wild dogs. In:

Reintroduction of Top-Order Predators (Eds M. HAYWARD and

M. SOMERS). Blackwell Publishing, Oxford.

ENPAT (2001). Environmental potential atlas. Department of

Environmental Affairs. http://www.environment.gov.za/

Enviro-Info/info.htm (accessed on June 6 2008).

Fanshawe, J.H., Ginsberg, J.R., Sillero-Zubiri, C. & Woodroffe, R.

(1997) The status and distribution of remaining wild dog

populations. In: The African Wild Dog: Status Survey and

Conservation Action Plan (Eds R. WOODROFFE and J.R.

GINSBERG). IUCN, Gland.

Frame, L.H. & Frame, G.W. (1976) Female African wild dogs

emigrate. Nature 263, 227–229.

Fuller, T.K., Mills, M.G.L., Borner, M., Laurenson, M.K. & Kat,

P.W. (1992) Long distance dispersal by African wild dogs in

East and South Africa. J. Afr. Zool. 106, 535–537.

Graf, J.A., Gusset, M., Reid, C., Janse Van Rensburg, S., Slotow,

R. & Somers, M.J. (2006) Evolutionary ecology meets

wildlife management: artificial group augmentation in the

re-introduction of endangered African wild dogs (Lycaon

pictus). Anim. Conserv. 9, 398–403.

Gusset, M. (2010) The re-introduction of African wild dogs in

South Africa. In: Global Re-Introduction Perspectives:

Additional Case-Studies from around the Globe (Ed. P.S. SOORAE),

IUCN/SSC Re-introduction Specialist Group, Abu Dhabi.

Gusset, M., Ryan, S.J., Hofmeyr, M., Van Dyk, G., Davies-

Mostert, H.T., Graf, J.A., Owen, C., Szykman, M., Macdonald,

D.W., Monfort, S.L., Wildt, D.E., Maddock, A.H., Mills, M.G.

L., Slotow, R. & Somers, M.J. (2008a) Efforts going to the

dogs? Evaluating attempts to re-introduce endangered wild

dogs in South Africa J. Appl. Ecol. 45, 100–108.

Gusset, M., Jakoby, O., Muller, M.S., Somers, M.J., Slotow, R. &

Grimm, V. (2009a) Dogs on the catwalk: modelling

re-introduction and translocation of endangered wild dogs in

South Africa. Biol. Conserv. 142, 2774–2781.

Lindsey, P., Du Toit, J.T. & Mills, M.G.L. (2004) The distribution

and population status of African wild dogs (Lycaon pictus)

outside protected areas in South Africa. S. Afr. J. Wildl. Res.

34, 143–151.

Maddock, A.H. & Mills, M.G.L. (1994) Population characteristics

of African wild dogs Lycaon pictus in the Eastern transvaal

lowveld, South Africa, as revealed through photographic

records. Biol. Cons. 67, 57–62.

McNutt, J.W. (1996) Sex-biased dispersal in African wild dogs,

Lycaon pictus. Anim. Behav. 52, 1067–1077.

McNutt, J.W., Parker, M.N., Swarner, M.J. & Gusset, M. (2008)

Adoption as a conservation tool for endangered African wild

dogs (Lycaon pictus). S. Afr. J. Wildl. Res. 38, 109–112.

Mills, M.G.L., Ellis, S., Woodroffe, R., Maddock, A., Stander, P.,

Rasmussen, G. & Pole, A. et al. (1998). Population and Habitat

Viability Analysis for the African Wild Dog (Lycaon pictus) in

southern Africa. IUCN/SSC Conservation Breeding Specialist

Group workshop report, Pretoria.

Rasmussen, G.S.A. (1997).Conservation Status of the Painted

Hunting Dog (Lycaon pictus) in Zimbabwe. DNPWLM/Painted

Dog Research Project, Harare, Zimbabwe.

Rasmussen, G.S.A. & Macdonald, D.W. (2011) Masking of the

zeitgeber: African wild dogs mitigate persecution by balancing

time. J. Zool. (Lond.) 286, 232–242.

Skinner, J. & Smithers, R. (1990) The Mammals of the South

African Subregion. University of Pretoria, Pretoria, South

Africa.

Somers, M.J., Graf, J.A., Szykman, M., Slotow, R. & Gusset, M.

(2008) Dynamics of a small re-introduced population of wild

dogs over 25 years: Allee effects and the implications of

sociality for endangered species’ recovery. Oecologia 158,

239–247.

Wabakken, P., Sand, H., Kojola, I., Zimmermann, B., Arnemo, J.

M., Pedersen, H.C. & Liberg, O. (2007) Multistage, long-range

natal dispersal by a global positioning system-collared

Scandinavian wolf. J. Wildl. Manage. 71, 1631–1634.

Woodroffe, R., McNutt, J.W. & Mills, M.G.L. (2004) African

wild dog (Lycaon pictus). In: Canids: foxes, Wolves, Jackals

and Dogs: Status Survey and Conservation Action Plan (Eds

C. SILLERO-ZUBIRI, M. HOFFMAN and D.W. MACDONALD). IUCN,

Gland.

(Manuscript accepted 18 March 2012)

doi: 10.1111/j.1365-2028.2012.01335.x



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