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  • 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: harrietd@ewt.org.za

    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) 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 k

    m

    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

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

    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 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

  • 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)

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

    504 Harriet T. Davies-Mostert et al.

  • 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.

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    doi: 10.1111/j.1365-2028.2012.01335.x

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

    506 Harriet T. Davies-Mostert et al.

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