Status of terrestrial mammals at the Kafue-Zambezi interface: implications for transboundary connectivity

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Status of terrestrial mammals at the Kafue-Zambezi interface: implications for transboundary connectivity
Status of terrestrial mammals at the
             Kafue–Zambezi interface: implications for
             transboundary connectivity
                                                     R O B I N L I N E S , J O S E P H T Z A N O P O U L O S and D O U G L A S M A C M I L L A N

             Abstract The Kavango–Zambezi Transfrontier Conservation                               Introduction
             Area Programme promotes landscape-level connectivity
             between clusters of wildlife management areas in five neigh-
             bouring countries. However, declining regional biodiversity                           W       ildlife management areas are frequently clustered
                                                                                                           along international borders, with ecosystems divided
                                                                                                   by arbitrarily drawn political boundaries (Zbicz, ;
             can undermine efforts to maintain, expand and link wildlife
             populations. Narratives promoting species connectivity                                Hanks, ). Where fences and physical barriers are com-
             should thus be founded on studies of system and state                                 bined with expanding human settlements and intensifying
             changes in key resources. By integrating and augmenting                               agro-pastoralist activities, overexploitation and decline of
             multiple data sources throughout eight wildlife management                            wildlife populations can occur (Ogutu et al., ).
             areas, covering . million ha, we report changes during                              Additionally, invasion, disease, pollution and climate
             – in the occurrence and distribution of  mammal                             change (Pachauri et al., ; Maxwell et al., ) interact
             species throughout a landscape linking the Greater Kafue                              with intrinsic species traits (Cardillo et al., ) to inhibit
             System to adjacent wildlife management areas in Namibia                               or sever wildlife movement patterns, isolating core wildlife
             and Botswana. Results indicate species diversity is largely un-                       management areas (Margules & Pressey, ; Newmark,
             changed in Kafue National Park and Mulobezi and Sichifulo                             ). Together these drivers are exposing wildlife popula-
             Game Management Areas. However, % of large carnivore                               tions to escalating edge-effects and ecological traps, threa-
             and % of prey diversity have been lost in the Simalaha                              tening species persistence within and outside protected
             areas, and there is no evidence of migrational behaviour or                           areas (Woodroffe & Ginsberg, ; Battin, ).
             species recolonization from adjacent wildlife areas.                                      Conversely, intact species assemblages have wide-ranging
             Although temporal sampling scales influence the definition                            implications for sustainable and resilient social-ecological
             of species occupancy and distribution, and data cannot elu-                           systems (Cumming, ). Heterogeneity and functional
             cidate population size or trends, our findings indicate an                            diversity drive productivity and a system’s capacity to ab-
             emerging connectivity bottleneck within Simalaha.                                     sorb, resist and respond to shocks, perturbations and other
             Evidence suggests that at current disturbance levels the                              stressors that negatively affect its structure and function
             Greater Kafue System, Zambia’s majority component in                                  (Fischer et al., ). Cumulatively, threats to species per-
             the Kavango–Zambezi Transfrontier Conservation Area, is                               sistence undermine habitat integrity, ecosystem services,
             becoming increasingly isolated at the trophic scale of large                          food security, the development of sustainable wildlife-based
             mammals. Further investigations of the site-specific, inter-                          land uses and human well-being (Millennium Ecosystem
             acting drivers influencing wildlife distribution and occur-                           Assessment, ; Lindsey et al., b).
             rence are required to inform appropriate conservation                                     Acknowledging the limitations imposed by these con-
             interventions for wildlife recovery in key areas identified to                        straints, stakeholders in southern Africa are increasingly
             promote transboundary connectivity in the Kavango–                                    embracing Transfrontier Conservation Areas as a new con-
             Zambezi Transfrontier Conservation Area.                                              servation paradigm (Hanks, ), considered to be an evo-
                                                                                                   lution of previous community-based natural resource
             Keywords Connectivity, Kafue, Kavango–Zambezi Transfron-                              management approaches that yielded mixed results
             tier Area, mammal loss, transboundary conservation, wildlife                          (Andersson, ). Enticing narratives include the integra-
             management area                                                                       tion of biodiversity conservation with the promotion of sus-
                                                                                                   tainable socio-economic development and a culture of peace
                                                                                                   and cooperation at the ecosystem level, linked to the re-
                                                                                                   moval of fences and other barriers inhibiting the free move-
                                                                                                   ment of wildlife across vast interconnected landscapes (van
             ROBIN LINES (Corresponding author), JOSEPH TZANOPOULOS and DOUGLAS                    der Linde et al., ; Hanks, ).
             MACMILLAN Durrell Institute of Conservation and Ecology, Marlow Building,                 The Kavango–Zambezi Transfrontier Conservation Area
             The University of Kent, Canterbury, Kent, CT2 7NR, UK
             E-mail rl291@kent.ac.uk                                                               is intended to capitalize on the region’s unique diversity and
             Received  February . Revision requested  June .                             distribution of wildlife assets by advocating shared natural
             Accepted  October . First published online  May .                         resource management and development goals across a

                                                                                        Oryx, 2019, 53(4), 764–773 © 2018 Fauna & Flora International doi:10.1017/S0030605317001594
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. https://doi.org/10.1017/S0030605317001594
The Kafue–Zambezi interface                  765

           network of protected areas spanning . , km at the                              the Zambezi River. This landscape is promoted as a key link-
           interface of Angola, Botswana, Namibia, Zambia and                                    age to the central cluster of wildlife management areas in
           Zimbabwe (KAZA TFCA Secretariat, b; Hanks &                                       Namibia and Botswana, at the heart of the Kavango–
           Myburgh, ). Stated objectives to integrate conservation                           Zambezi Transfrontier Conservation Area (KAZA TFCA
           and development, promote peace and cooperation, and fa-                               Secretariat, ). Through integration, harmonization
           cilitate connectivity of wildlife populations between clusters                        and triangulation of data we were able to determine changes
           of wildlife management areas have become popular and                                  in species occurrence and distribution by wildlife manage-
           compelling programme narratives driving north–south fi-                               ment area and designation.
           nance initiatives, NGO engagement, and state buy-in
           (PPF, ; KAZA TFCA Secretariat, a; WWF, ).
               Notwithstanding evolving conservation and develop-                                Study area
           ment narratives, the success of the Kavango–Zambezi
           Transfrontier Conservation Area Programme is constrained                              The boundaries of the Kavango–Zambezi Transfrontier
           by many existing and emerging challenges. Mounting                                    Conservation Area are imprecise (Andersson, ) but
           anthropogenic pressures combined with poor land-use                                   Cumming () characterized the area as comprising a
           planning, institutional conflicts and stakeholder disenfran-                          matrix of .  wildlife management areas, from National
           chisement (Andersson, ) are driving encroachment into                             Parks under strict state control to multiple-use areas
           wildlife areas, habitat loss and fragmentation (Newmark,                              under community management. These wildlife manage-
           ; Simukonda, ; Watson et al., ), and unsustain-                           ment areas are grouped into three major clusters and five
           able harvesting of wildlife, threatening many of the                                  periphery sub-clusters, with Kafue National Park and sur-
           Conservation Area’s iconic natural assets (Lindsey et al.,                            rounding wildlife management areas constituting the
           a). With the region’s human population expected to                                major northern cluster (Fig. ).
           double by  (United Nations, Department of                                             Kafue National Park (, km) is Zambia’s oldest and
           Economic and Social Affairs, Population Division, )                               largest protected area, the largest National Park in the
           and likely impacts of climate change exacerbating challenges                          Kavango–Zambezi Transfrontier Conservation Area and the
           to socio-economic development (Bellard et al., ;                                  second largest National Park in Africa (UNEP/WCMC, ).
           Pachauri et al., ), even moderately optimistic scenarios                          Together with nine surrounding IUCN category VI Game
           imply regional biodiversity loss will accelerate significantly                        Management Areas and multiple Forest Reserves, the effective
           this century (Biggs et al., ).                                                    unfenced wildlife management area, referred to as the Greater
               Collectively these challenges raise important questions                           Kafue Landscape or System, covers , km and includes
           about the scope, scale and ambition of narratives promoting                           % of Zambia’s land mass and . % of the Kavango–
           landscape-level linkages, the interventions required to                               Zambezi Transfrontier Conservation Area estate (Moss, ).
           maintain or expand connectivity, and the purposes these                                   Most of the Greater Kafue System lies at –, m
           proposed linkages may serve in the long term (Cumming,                                altitude. Mean annual rainfall is  mm in the south and
           ). There is thus a clear imperative to promote evidence-                          , mm in the north, falling predominantly during
           based socio-economic and environmental policies and in-                               November–April. Vegetation is characterized by the
           terventions built around the application of conservation sci-                         Zambezian Miombo Woodland ecoregion, typical of large
           ence (Sutherland et al., ), including research and                                areas throughout southern and eastern Africa, dominated
           monitoring of changes in the state of sites and systems,                              by Brachystegia spp., Combretum spp., Mopane spp.,
           and their response to factors driving connectivity at the                             Terminalia spp. and Baikaea spp. Woodlands are inter-
           scale of interest. The process of informed decision-making                            spersed with open floodplain grasslands and dambos
           is data hungry, and local, regional and transboundary data                            (ZAWA, ). Species records include  mammals, 
           sources are disparate and inconsistent, undermining attempts                          birds,  reptiles,  amphibians and  fish, with the great-
           to understand complex social-ecological systems such as the                           est diversity of antelopes in Africa ( species), an intact car-
           Kavango–Zambezi Transfrontier Conservation Area. Data                                 nivore guild and a full complement of Zambia’s large
           deficiencies ultimately constrain effective decision-making                           mammals, with the exception of the giraffe Giraffa giraffa,
           and appropriate interventions to promote biodiversity con-                            black rhinoceros Diceros bicornis and tsessebe Damaliscus
           servation and development.                                                            lunatus (Moss, ).
               Here we interrogate and synthesize existing data sources,                             The Greater Kafue System is Zambia’s majority component
           and supplement them with additional research to document                              within the Kavango–Zambezi Transfrontier Conservation
           the historical and contemporary status of the African ele-                            Area (KAZA TFCA Secretariat, ), with connectivity to
           phant Loxodonta africana, five large carnivores, one meso-                            the broader Kavango–Zambezi landscape contingent on the
           predator and  prey species throughout eight wildlife                                maintenance of a landscape-level linkage routing south–south-
           management areas between the Greater Kafue System and                                 west through a mosaic of nominally, potentially and possibly

           Oryx, 2019, 53(4), 764–773 © 2018 Fauna & Flora International doi:10.1017/S0030605317001594
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. https://doi.org/10.1017/S0030605317001594
766         R. Lines et al.

                                                                                                                                             FIG. 1 Clusters of wildlife
                                                                                                                                             management areas in the
                                                                                                                                             Kavango–Zambezi
                                                                                                                                             Transfrontier Conservation
                                                                                                                                             Area landscape in southern
                                                                                                                                             Africa (adapted from KAZA
                                                                                                                                             TFCA Secretariat, ; PPF,
                                                                                                                                             ).

             protected wildlife management areas, including Mulobezi and                           indicate that Kafue National Park (operating with only
             Sichifulo Game Management Areas, Nachitwe, Martin and                                 –% of recommended protected area budgets) has re-
             Machili Forest Reserves, the Nyawa communal areas, and the                            ceived the greatest level of long-term support for biodiver-
             Simalaha Communal Conservancy, proclaimed in  (Fig. ).                           sity conservation throughout the study area, followed by
             Together these wildlife management areas extend the Greater                           Mulobezi and then Sichifulo Game Management Areas,
             Kafue System to c. . million ha.                                                    which receive minor budget allocations from the State
                A secondary (south-westerly) linkage passing through                               Wildlife Authority, augmented by finance and in-kind op-
             Mulobezi to Sioma National Park (bordering Namibia and                                erational support from resident safari hunting operators and
             Angola) has been proposed, although our focus is on the                               conservation NGOs. Nachitwe, Martin and Machili Forest
             linkage that broadly follows the Machili stream catchment                             Reserves have received minor budgets intermittently from
             basin from the border of Kafue National Park to the northern                          the State Wildlife Authority and Forestry Department
             bank of the Zambezi River, adjacent to Kasika and Salambala                           (ZAWA, ; C. Chifunte, pers. comm. .). The
             Communal Conservancies of the eastern Zambezi region in                               Simalaha Communal Conservancy started receiving formal
             Namibia, and through to Chobe National Park in Botswana.                              wildlife resource protection as recently as , having had
                The proposed landscape linkage is – km. The                                  no formal biodiversity conservation budgets since before
             human population is c. , and increasing by .% an-                              (HRH Chief Inyambo Yeta, pers. comm. ). We
             nually, with a population density of c.  people per km                              were unable to ascertain if the Nyawa communal areas receive
             (CSO, ). Communities are centred on a few larger set-                             any formal wildlife management budget. In addition, a
             tlements of ,–, residents, and otherwise in clus-                            , ha fenced Wildlife Recovery Sanctuary to the south
             ters of scattered villages typically concentrated along water                         of Simalaha, with an extensive open border along the
             courses, seasonal waterholes and a few pumped ground-                                 Zambezi River, has received .  head of game of eight spe-
             water supplies. Subsistence agro-pastoralists dominate this                           cies since : a significant investment promoted as a justifi-
             landscape, with residents largely dependent on exploiting                             cation for restocking the broader Simalaha Communal
             a wide range of natural resources to meet their basic liveli-                         Conservancy (PPF, ).
             hood needs (Musgrave, ). Formal employment oppor-
             tunities beyond a few distant urban settlements are
             negligible. Customary law within the Lozi, Nkoya and                                  Methods
             Tonga ethnolinguistic groups constitutes the de facto re-
             gional governance system (Brelsford, ; Musgrave, ).                           Data sources
                Biodiversity conservation budgets have varied signifi-
             cantly throughout this landscape, both spatially and tem-                             The earliest records of occurrence and distribution of
             porally. Although precise figures are unavailable, sources                            terrestrial mammals in the vicinity of the proposed

                                                                                        Oryx, 2019, 53(4), 764–773 © 2018 Fauna & Flora International doi:10.1017/S0030605317001594
Downloaded from https://www.cambridge.org/core. IP address: 46.4.80.155, on 14 Mar 2021 at 03:55:34, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms
. https://doi.org/10.1017/S0030605317001594
The Kafue–Zambezi interface                  767

                                                                                                 FIG. 3 Boundaries of contemporary wildlife management areas
                                                                                                 (in white) and the study area (dotted line) projected onto
                                                                                                 Ansell’s () map of the known (filled squares), possible
                                                                                                 (half-filled squares) and former range (unfilled squares) of the
           FIG. 2 Wildlife management areas within the study area.                               blue wildebeest Connochaetes taurinus.

           Kafue–Zambezi linkage are limited to disparate notes and                                  Ansell () reported on  terrestrial mammals .  kg
           reports in the grey literature from early explorers, hunters,                         in weight, from  families, but we restricted the contempor-
           traders and missionaries, dating back to the late th century                        ary list to  readily detected species from nine families, omit-
           (e.g. Sampson, ; Holub, ), with approximate                                   ting seven species considered to be at the edge of their known
           location data variously reported in relation to key landscape                         range and/or habitat specialists requiring species-specific
           features. The first published checklists for Zambia (Pitman,                          survey techniques beyond the scope of this study.
           ; Lancaster, ; Ansell, , , ) indicate there                           Boundaries of contemporary land-use classifications
           were no changes to the assemblage of large mammals in and                             (UNEP-WCMC, ) were projected over Ansell’s ()
           around Kafue National Park prior to the extirpation of the                            maps using QGIS (QGIS Development Team, ; Fig. ),
           black rhinoceros in the mid s, although there are unre-                           to facilitate extraction of historical species distribution data
           solved questions regarding anecdotal records of a relic gir-                          at comparable spatial scales, for Kafue National Park
           affe population (P. Moss, pers. comm. ). Data for these                           (Kaunga and Nanzhila management blocks, , ha),
           checklists were collected ostensibly through ad hoc and op-                           Mulobezi Game Management Area (hereafter Mulobezi,
           portunistic sightings by government staff and expert obser-                           , ha), Sichifulo Game Management Area, including
           vers reporting from their travels throughout the country,                             Nachitwe, Martin and Machili Forest Reserves (hereafter
           augmented by trading records and hunting ledgers kept                                 Sichifulo, , ha), and the Nyawa/Simalaha areas
           by District Commissioners.                                                            (c. , ha).
               The first systematic collation of species occurrence and dis-                         In compiling contemporary data sets (Fig. ) we con-
           tribution data was published by Ansell (), superseding                            strained data gathering to three broadly comparable
           previous literature. Amalgamated checklist data were mapped                           ground-based survey approaches. We omitted aerial survey
           within quarter-degree grid squares, based on  : ,                               data (e.g. DNPW, ) because of limitations to detection
           Ordnance Survey map sheets. Given the absence of reports                              rates for many species of primary interest in forested areas
           from many inaccessible and largely unmapped periphery                                 (Jachmann, ).
           areas, the data reflect minimum regional species ranges; none-                            Firstly, the resident safari hunting operator, operational
           theless, much of the study area can be considered to be well                          throughout Mulobezi and Sichifulo during the preceding
           mapped because of the established network of access routes                            decade, was asked to provide sighting reports for  terres-
           developed alongside the nascent teak logging and safari hunt-                         trial mammals of interest through a questionnaire survey
           ing industries (Musgrave, ).                                                      following the  hunting season. Multiple groups of

           Oryx, 2019, 53(4), 764–773 © 2018 Fauna & Flora International doi:10.1017/S0030605317001594
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. https://doi.org/10.1017/S0030605317001594
768         R. Lines et al.

                                                                                                   three times by RL and two experienced local trackers from
                                                                                                   the safari hunting industry, amounting to , km of spoor
                                                                                                   transects during the dry season of May–October .

                                                                                                   Data analysis

                                                                                                   A confirmed sighting from any of the three selected expert
                                                                                                   contemporary sources was considered to be sufficient to de-
                                                                                                   tect species presence at the scale of interest. Given the atyp-
                                                                                                   ical nature of ongoing ungulate reintroductions and
                                                                                                   management in the fenced Simalaha Wildlife Sanctuary,
                                                                                                   we restrict reporting to the detection of the carnivore
                                                                                                   guild for this subset of the Simalaha Communal
                                                                                                   Conservancy.
                                                                                                      Data for each of the four composite wildlife management
                                                                                                   area blocks, from the three data sources, were compiled
                                                                                                   against historical data to determine if any changes in species
                                                                                                   occurrence and distribution had occurred. Outputs reflected
                                                                                                   species persistence, loss or colonization at the composite
                                                                                                   wildlife management area scale.
                                                                                                      As survey methods were optimized for resident large car-
                                                                                                   nivores and their principal prey species, the risk of non-
                                                                                                   detection was elevated for species that exhibited significant
             FIG. 4 Sources of data within the study area for contemporary                         seasonal (migration) or non-resident (emigration and im-
             analyses of terrestrial mammals at the Kafue–Zambezi interface                        migration) movement patterns, or where surveys did not
             (Fig. ).                                                                             cover the restricted ranges of habitat specialists. These con-
                                                                                                   straints are acknowledged in Table  and subsequent
                                                                                                   analyses.
             guides, hunters and skilled trackers traverse both Mulobezi
                                                                                                      An amalgamated distribution map was generated for the
             and Sichifulo on and off road, covering . , km each
                                                                                                   five extant large carnivores, indicating their historical range
             dry season (R. Kraljic, pers. comm. ). This was consid-
                                                                                                   within the survey area, and current known range within
             ered to amount to sufficient survey effort and expertise to
                                                                                                   studied wildlife management areas.
             detect target species.
                 Secondly we collected patrol data from the local State and
             Community Wildlife Police Officers responsible for wildlife                           Results
             protection in southern Kafue National Park, Mulobezi and
             Sichifulo. We amalgamated data for the Kafue patrol blocks                            Table  indicates few non-detections recorded against any
             adjacent to Mulobezi and Sichifulo to cover a single area in-                         data sources since  throughout southern Kafue
             cluding the border area north of both Mulobezi and                                    National Park, Mulobezi or Sichifulo. However, the hippo-
             Sichifulo Game Management Areas. These data comprised                                 potamus Hippopotamus amphibius appears to be no longer
             , georeferenced wildlife sightings from , man-days                           resident in any of the waterways along the Machili stream
             of foot patrols during – (ZAWA, unpubl. data).                                and catchment area, and the klipspringer Oreotragus oreo-
                 In  we undertook a systematic randomized spoor and                            tragus appears to be absent from Mulobezi, although core
             sightings survey of large carnivores and their principal prey                         habitat for this species went unsurveyed. The steenbok
             throughout  -km survey blocks in Mulobezi, Sichifulo                            Raphicerus campestris is considered to be at the extent of
             and the Nyawa/Simalaha areas. Detection probability and                               its north-east range approaching Kafue National Park,
             survey effort were optimized for large carnivores following                           with a single sighting recorded in Mulobezi.
             Funston et al. () and Thorn et al. (). In addition, a                            The absence of confirmed sightings of the caracal
             site-specific calibration process was undertaken during                               Caracal caracal and serval Leptailurus serval by Wildlife
             July–September , conducted at various spatio-temporal                             Police Officer patrols in southern Kafue National Park ap-
             scales, to establish the survey effort required to detect large                       pear to be an anomaly, given detections of the species in ad-
             carnivores and sample the landscape in a single season                                jacent Game Management Areas. Although it is likely this
             (MacKenzie & Royle, ; D.I. MacKenzie, pers. comm.                                 anomaly represents non-detection error versus absence,
             ). In total,  -km transects were surveyed on foot                             we discarded these species from the final checklist.

                                                                                        Oryx, 2019, 53(4), 764–773 © 2018 Fauna & Flora International doi:10.1017/S0030605317001594
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. https://doi.org/10.1017/S0030605317001594
. https://doi.org/10.1017/S0030605317001594
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                                                                                                                                                                                                                                                                                                   TABLE 1 Species detected in various studies conducted in wildlife management areas at the Kafue–Zambezi interface (Fig. ), their IUCN Red List status, and whether or not there was a change
                                                                                                                                                                                                                                                                                                   in distribution between  and –.
                                                                                                                                                                                                     Oryx, 2019, 53(4), 764–773 © 2018 Fauna & Flora International doi:10.1017/S0030605317001594

                                                                                                                                                                                                                                                                                                                                                                               R. Kraljik (unpubl.       Zambia Wildlife Authority
                                                                                                                                                                                                                                                                                                                                         Ansell (1978)                          data, 2013/2014)         (unpubl. data, 2014/2015)       This study (2014/2015)     Distribution change 1978–2014/2015
                                                                                                                                                                                                                                                                                                                              Red List   Kafue NP/                                                Kafue NP/                                                Kafue NP/
                                                                                                                                                                                                                                                                                                   Species                    status1    South     Mulobezi Sichifulo Simalaha Mulobezi Sichifulo South     Mulobezi Sichifulo Mulobezi Sichifulo Simalaha South     Mulobezi Sichifulo Simalaha
                                                                                                                                                                                                                                                                                                   Cheetah Acinonyx           VU         ✓          ✓        ✓        ✓        ✓         ✓           ✓                               ✓          ✓                 No        No        No        Yes
                                                                                                                                                                                                                                                                                                      jubatus
                                                                                                                                                                                                                                                                                                   Lion Panthera leo          VU         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓                 No        No        No        Yes2
                                                                                                                                                                                                                                                                                                   Leopard Panthera           VU         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓        ✓        No        No        No        Yes2
                                                                                                                                                                                                                                                                                                      pardus
                                                                                                                                                                                                                                                                                                   Spotted hyaena             LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓                 No        No        No        Yes2
                                                                                                                                                                                                                                                                                                      Crocuta crocuta
                                                                                                                                                                                                                                                                                                   Side-striped jackal        LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓                               ✓          ✓        ✓        No        No        No        No
                                                                                                                                                                                                                                                                                                      Canis adustus
                                                                                                                                                                                                                                                                                                   African wild dog           EN         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓                 No        No        No        Yes
                                                                                                                                                                                                                                                                                                      Lycaon pictus
                                                                                                                                                                                                                                                                                                   African elephant           VU         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓                 No        No        No        Yes2
                                                                                                                                                                                                                                                                                                      Loxodonta africana
                                                                                                                                                                                                                                                                                                   Burchell’s zebra           NT         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓                 No        No        No        Yes3
                                                                                                                                                                                                                                                                                                      Equus quagga
                                                                                                                                                                                                                                                                                                   Warthog Phacochoerus       LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓                 No        No        No        Yes
                                                                                                                                                                                                                                                                                                      africanus
                                                                                                                                                                                                                                                                                                   Bushpig Potamochoerus      LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓                 No        No        No        Yes
                                                                                                                                                                                                                                                                                                      larvatus
                                                                                                                                                                                                                                                                                                   Hippopotamus               VU         ✓          ✓        ✓        ✓                              ✓                                                            No        Yes2      Yes2      Yes2
                                                                                                                                                                                                                                                                                                      Hippopotamus
                                                                                                                                                                                                                                                                                                      amphibius
                                                                                                                                                                                                                                                                                                   Hartebeest Alcelaphus      LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓                 No        No        No        Yes
                                                                                                                                                                                                                                                                                                      lichtensteinii
                                                                                                                                                                                                                                                                                                   Blue wildebeest            LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓                 No        No        No        Yes3
                                                                                                                                                                                                                                                                                                      Connochaetes taurinus
                                                                                                                                                                                                                                                                                                   Klipspringer Oreotragus    LC         ✓                   ✓                 ✓         ✓           ✓                                                            No        No        No        No
                                                                                                                                                                                                                                                                                                      oreotragus
                                                                                                                                                                                                                                                                                                   Oribi Ourebia ourebi       LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓        ✓        No        No        No        No
                                                                                                                                                                                                                                                                                                   Steenbok Raphicerus        LC         ✓          ✓        ✓        ✓                                                              ✓                            UK4       No        Yes       Yes
                                                                                                                                                                                                                                                                                                      campestris
                                                                                                                                                                                                                                                                                                   Sharpe’s grysbok           LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓        ✓        No        No        No        No
                                                                                                                                                                                                                                                                                                      Raphicerus sharpei
                                                                                                                                                                                                                                                                                                   African buffalo            LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓                 No        No        No        Yes
                                                                                                                                                                                                                                                                                                      Syncerus caffer

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                         The Kafue–Zambezi interface
                                                                                                                                                                                                                                                                                                   Common eland               LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓                            No        No        No        Yes
                                                                                                                                                                                                                                                                                                      Tragelaphus oryx
                                                                                                                                                                                                                                                                                                   Bushbuck Tragelaphus       LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓                 No        No        No        Yes
                                                                                                                                                                                                                                                                                                      scriptus
                                                                                                                                                                                                                                                                                                   Sitatunga Tragelaphus      LC         ✓                                                                                                                        Yes       No        No        No
                                                                                                                                                                                                                                                                                                      spekii
                                                                                                                                                                                                                                                                                                   Greater kudu               LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓        ✓        No        No        No        No
                                                                                                                                                                                                                                                                                                      Tragelaphus
                                                                                                                                                                                                                                                                                                      strepsiceros
                                                                                                                                                                                                                                                                                                   Common duiker              LC         ✓          ✓        ✓        ✓        ✓         ✓           ✓           ✓          ✓        ✓          ✓        ✓        No        No        No        No
                                                                                                                                                                                                                                                                                                      Sylvicapra grimmia

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                         769
770          R. Lines et al.

                                                                                                                                                                                                                                                                                                                                                                                  Significant losses have occurred in the Simalaha

                                                                                 Kafue NP/ Mulobezi Sichifulo Simalaha Mulobezi Sichifulo Kafue NP/ Mulobezi Sichifulo Mulobezi Sichifulo Simalaha Kafue NP/ Mulobezi Sichifulo Simalaha
                                            Distribution change 1978–2014/2015
                                                                                                                                                                                                                                                                                                                                                                              Communal Conservancy, where  of  terrestrial mammals

                                                                                                                                                                                                                                                       Yes3

                                                                                                                                                                                                                                                              Yes3

                                                                                                                                                                                                                                                                     Yes3
                                                                                                                                                                                                                                                                     No3
                                                                                                                                                                                                                                           Yes

                                                                                                                                                                                                                                                 Yes

                                                                                                                                                                                                                                                                     No

                                                                                                                                                                                                                                                                             No
                                                                                                                                                                                                                                                                                                                                                                              were undetected (Fig. ) and the side-striped jackal Canis adu-
                                                                                                                                                                                                                                                                                                                                                                              stus was the only widespread carnivore detected. The spotted
                                                                                                                                                                                                                                           No                                                                                                                                 hyaena Crocuta crocuta and the leopard Panthera pardus

                                                                                                                                                                                                                                                 No

                                                                                                                                                                                                                                                       No

                                                                                                                                                                                                                                                              No

                                                                                                                                                                                                                                                                     No
                                                                                                                                                                                                                                                                     No
                                                                                                                                                                                                                                                                     No

                                                                                                                                                                                                                                                                             No
                                                                                                                                                                                                                                                                                                                                                                              were the only large carnivores detected within  km of the
                                                                                                                                                                                                                                                                                                                                                                              Zambezi River in the Nyawa communal areas (Fig. ). The re-
                                                                                                                                                                                                                                           No

                                                                                                                                                                                                                                                 No

                                                                                                                                                                                                                                                       No

                                                                                                                                                                                                                                                              No

                                                                                                                                                                                                                                                                     No
                                                                                                                                                                                                                                                                     No
                                                                                                                                                                                                                                                                     No

                                                                                                                                                                                                                                                                             No
                                                                                                                                                                                                                                                                                                                                                                              mainder of the large carnivore guild appears to be extirpated
                                                                                                                                                                                                                                                                                                                                                                              from the Simalaha/Nyawa area, along with all ungulates
                                                                                                                                                                                                                                                                                                                                                                              .  kg except the southern reedbuck Redunca arundinum
                                                                                                                                                                                                   South
                                                                                                                                                                                                                                           No

                                                                                                                                                                                                                                                 No

                                                                                                                                                                                                                                                       No

                                                                                                                                                                                                                                                              No

                                                                                                                                                                                                                                                                     No
                                                                                                                                                                                                                                                                     No
                                                                                                                                                                                                                                                                     No

                                                                                                                                                                                                                                                                             No
                                                                                                                                                                                                                                                                                                                                                                              and the greater kudu Tragelaphus strepsiceros. Kudu were
                                                                                                                                                                                                                                                                                                                                                                              also the only herding ungulate to be detected in Simalaha, al-
                                            This study (2014/2015)

                                                                                                                                                                                                                                                                         ✓
                                                                                                                                                                                                                                                                                                                                                                              though no aggregations of more than three individuals were
                                                                                                                                                                                                                                                                             ✓                                                                                                detected. Neither the warthog Phacochoerus africanus nor the
                                                                                                                                                                                                                                                                                                                                                                              bushpig Potamochoerus larvatus, both habitat and feeding
                                                                                                                                                                                                                                                                                                                                                                              generalists with high reproductive rates, were detected in
                                                                                                                                                                                                                                           ✓

                                                                                                                                                                                                                                                 ✓

                                                                                                                                                                                                                                                       ✓

                                                                                                                                                                                                                                                              ✓

                                                                                                                                                                                                                                                                         ✓

                                                                                                                                                                                                                                                                             ✓

                                                                                                                                                                                                                                                                                                                                                                              Simalaha. Although .  head of game, comprising seven
                                                                                                                                                                                                                                                                                                                                                                              species, have been introduced into the , ha Simalaha
                                                                                                                                                                                                                                                                                                                                                                              Wildlife Recovery Sanctuary since , only the side-striped
                                                                                                                                                                                                                                           ✓

                                                                                                                                                                                                                                                 ✓

                                                                                                                                                                                                                                                       ✓

                                                                                                                                                                                                                                                              ✓

                                                                                                                                                                                                                                                                         ✓

                                                                                                                                                                                                                                                                             ✓

                                                                                                                                                                                                                                                                                                                                                                              jackal was detected inside the (non-predator-proof) area.
                               Zambia Wildlife Authority
                               (unpubl. data, 2014/2015)

                                                                                                                                                                                                                                                                                                                                                                              There was no evidence of any species range extension or re-
                                                                                                                                                                                                                                           ✓

                                                                                                                                                                                                                                                 ✓

                                                                                                                                                                                                                                                       ✓

                                                                                                                                                                                                                                                              ✓

                                                                                                                                                                                                                                                                         ✓

                                                                                                                                                                                                                                                                                                                                                                              colonization throughout any of the sampled areas.
                                                                                                                                                                                                                                                                                                                                                                                  Although no long-term or landscape-level survey pro-
                                                                                                                                                                                                                                                                                                                                                                              gramme is in place for systematic monitoring of changes
                                                                                                                                                                                                                                           ✓

                                                                                                                                                                                                                                                 ✓

                                                                                                                                                                                                                                                       ✓

                                                                                                                                                                                                                                                              ✓

                                                                                                                                                                                                                                                                         ✓

                                                                                                                                                                                                                                                                                                                                                                              in species occurrence, distribution or abundance, much ex-
                                                                                                                                                                                                                                                                                                                                                                              isting expertise and anecdotal evidence implies there have
                                                                                                                                          South

                                                                                                                                                                                                                                                                                                                                                                              been large-scale population declines throughout the
                                                                                                                                                                                                                                           ✓

                                                                                                                                                                                                                                                 ✓

                                                                                                                                                                                                                                                       ✓

                                                                                                                                                                                                                                                              ✓

                                                                                                                                                                                                                                                                      ✓
                                                                                                                                                                                                                                                                      ✓

                                                                                                                                                                                                                                                                                                                                                                              Greater Kafue System and beyond since  (C. Chifunte,
                               R. Kraljik (unpubl.
                                data, 2013/2014)

                                                                                                                                                                                                                                                                                                                                                                              L. Daka, J. Hanks, HRH Chief Moomba, P. Moss & HRH
                                                                                                                                                                                                                                                                                                                                                                              Chief Inyambo Yeta, pers. comms). Contemporary data in-
                                                                                                                                                                                                                                           ✓

                                                                                                                                                                                                                                                 ✓

                                                                                                                                                                                                                                                       ✓

                                                                                                                                                                                                                                                              ✓

                                                                                                                                                                                                                                                                         ✓

                                                                                                                                                                                                                                                                             ✓

                                                                                                                                                                                                                                                                                                                                                                              dicate Kafue National Park, the region’s prime wildlife area,
                                                                                                                                                                                                                                                                                                                                                                              is maintaining the majority of terrestrial mammals at sig-
                                                                                                                                                                                                                                                                                    Non-resident individuals moving through the area, reported periodically by local people

                                                                                                                                                                                                                                                                                                                                                                              nificantly below carrying capacity (Simukonda, ).
                                                                                                                                                                                                                                           ✓

                                                                                                                                                                                                                                                 ✓

                                                                                                                                                                                                                                                       ✓

                                                                                                                                                                                                                                                              ✓

                                                                                                                                                                                                                                                                         ✓

                                                                                                                                                                                                                                                                             ✓

                                                                                                                                                                                                                                                                                                                                                                              Nonetheless, with few historical survey data available for
                                                                                                                                                                                                                                                                                                                                                                              direct comparison, we restricted our analyses to species di-
                                                                                                                                                                                                                                                                                   LC, Least Concern; NT, Near Threatened; VU, Vulnerable; EN, Endangered
                                                                                                                                                                                                                                           ✓

                                                                                                                                                                                                                                                 ✓

                                                                                                                                                                                                                                                       ✓

                                                                                                                                                                                                                                                              ✓

                                                                                                                                                                                                                                                                     ✓

                                                                                                                                                                                                                                                                         ✓

                                                                                                                                                                                                                                                                             ✓

                                                                                                                                                                                                                                                                                                                                                                              versity at the scale of interest, versus any interpretation of
                                                                                                                                                                                                                                                                                                                                                                              spatio-temporal changes to community structure and abun-
                                                                                                                                                                                                                                                                                    Reintroduced in fenced , ha breeding camp during –

                                                                                                                                                                                                                                                                                                                                                                              dance, which is beyond the scope of this article.
                                                                                                                                                                                                                                           ✓

                                                                                                                                                                                                                                                 ✓

                                                                                                                                                                                                                                                       ✓

                                                                                                                                                                                                                                                              ✓

                                                                                                                                                                                                                                                                         ✓

                                                                                                                                                                                                                                                                             ✓

                                                                                                                                                                                                                                                                                                                                                                              Discussion
                                                                                                                                                                                                                                           ✓

                                                                                                                                                                                                                                                 ✓

                                                                                                                                                                                                                                                       ✓

                                                                                                                                                                                                                                                              ✓

                                                                                                                                                                                                                                                                         ✓

                                                                                                                                                                                                                                                                             ✓
                                            Ansell (1978)

                                                                                                                                                                                                                                                                                                                                                                              Formal historical records accounting for species loss in the
                                                                                 South

                                                                                                                                                                                                                                                                                                                                                                              Simalaha and Nyawa areas are unavailable, although local
                                                                                                                                                                                                                                           ✓

                                                                                                                                                                                                                                                 ✓

                                                                                                                                                                                                                                                       ✓

                                                                                                                                                                                                                                                              ✓

                                                                                                                                                                                                                                                                      ✓
                                                                                                                                                                                                                                                                      ✓

                                                                                                                                                                                                                                                                             ✓

                                                                                                                                                                                                                                                                                                                                                                              Traditional Authorities (HRH Chief Inyambo Yeta &
                                                                                                                                                                                                                                                                                                                                                                              HRH Chief Moomba, pers. comms ) emphasized the
                                                                                 Red List
                                                                                 status1

                                                                                                                                                                                                                                                                                                                                                                              impact of the Angolan Bush War (–) as a key dri-
                                                                                                                                                                                                                                                                     NT
                                                                                                                                                                                                                                           LC

                                                                                                                                                                                                                                                 LC

                                                                                                                                                                                                                                                       LC

                                                                                                                                                                                                                                                              LC

                                                                                                                                                                                                                                                                     LC

                                                                                                                                                                                                                                                                     LC

                                                                                                                                                                                                                                                                             LC

                                                                                                                                                                                                                                                                                                                                                                              ver, describing encampments of foreign combatants in
                                                                                                                                                                                                                                           Sable antelope Hippotragus

                                                                                                                                                                                                                                                                                                                                                                              Simalaha being used as a base to exploit the area’s wildlife
                                                                                                                                                                                                                                           Cape porcupine Hystrix

                                                                                                                                                                                                                                                                                    Extent of known range
                                                                                                                                                                                                                                             Hippotragus equinus

                                                                                                                                                                                                                                             Redunca arundinum

                                                                                                                                                                                                                                                                                                                                                                              for rations and profit. Following the cessation of hostilities
                                                                                                                                                                                                                                           Puku Kobus vardonii
                                                                                                                                                                                                                                           Lechwe Kobus leche

                                                                                                                                                                                                                                           Southern reedbuck
                                                                                                                                                                                                                                           Defassa waterbuck

                                                                                                                                                                                                                                                                                                                                                                              there was a proliferation of small firearms in the area, and in
             Table 1 (Cont.)

                                                                                                                                                                                                                                             africaeaustralis
                                                                                                                                                                                                                                           Impala Aepyceros

                                                                                                                                                                                                                                             Kobus defassa
                                                                                                                                                                                                                                           Roan antelope

                                                                                                                                                                                                                                                                                                                                                                              conjunction with an increasing human population and lim-
                                                                                                                                                                                                                                             melampus

                                                                                                                                                                                                                                                                                                                                                                              ited funding for law enforcement and natural resource man-
                                                                                                                                                                                                                                             niger
                                                                                                                                                                   Species

                                                                                                                                                                                                                                                                                                                                                                              agement, unsustainable harvesting of wildlife continued.
                                                                                                                                                                                                                                                                                                                                                                              Given these circumstances we hypothesize that wildlife
                                                                                                                                                                                                                                                                                  
                                                                                                                                                                                                                                                                                                  
                                                                                                                                                                                                                                                                                  
                                                                                                                                                                                                                                                                                  

                                                                                                                                                                                                                                                                                                                                           Oryx, 2019, 53(4), 764–773 © 2018 Fauna & Flora International doi:10.1017/S0030605317001594
Downloaded from https://www.cambridge.org/core. IP address: 46.4.80.155, on 14 Mar 2021 at 03:55:34, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms
. https://doi.org/10.1017/S0030605317001594
The Kafue–Zambezi interface                  771

           FIG. 5 Changes in carnivore and herbivore diversity in wildlife
           management areas at the Kafue–Zambezi interface (Fig. ) during
           –.

           management areas closer to Kafue National Park were
           spared much of this pressure, having also received more pol-
           itical and revenue support for wildlife management in the
           long term (L. Daka, pers. comm. ).
               Surveys at the Kafue–Zambezi interface have employed a
           range of ad hoc methodological approaches that failed to
           detect the majority of resident species throughout this land-
           scape. The absence of a reliable baseline undermines efforts
           to evaluate the effectiveness of large-scale conservation in-
           terventions required to deliver key programme objectives
           within and between clusters of wildlife management areas.                             FIG. 6 Distribution of the spotted hyaena Crocuta crocuta,
                                                                                                 cheetah Acinonyx jubatus, African wild dog Lycaon pictus, lion
               Acknowledging non-detection error, we confirm that the
                                                                                                 Panthera leo and leopard Panthera pardus at the Kafue–Zambezi
           diversity of terrestrial mammals (.  kg) in southern Kafue                          interface (Fig. ) in –.
           National Park remains unchanged since . Mulobezi and
           Sichifulo retain largely intact mammalian diversity, with the                         of wildlife occurrence and distribution south of the Zambezi
           notable exception of the hippopotamus. No new data could                              River throughout the wildlife management areas of the east-
           be provided for the presence of free-ranging giraffes in any                          ern Zambezi region in Namibia, and the potential effects of
           of these wildlife management areas.                                                   ecological traps/attractive sinks at transboundary scales on
               A single-season survey design increases non-detection                             wildlife management interventions in Simalaha and other
           error associated with species dispersal or seasonal wildlife                          neighbouring wildlife management areas of Zambia.
           movement patterns; nonetheless, widespread losses, includ-                               Broader scale implications of species loss and ecological
           ing three of six carnivore species and  of  prey species,                         traps within the Kavango–Zambezi Transfrontier Conser-
           were detected in the Simalaha Communal Conservancy/                                   vation Area relate to dominant narratives about the
           Nyawa areas, which link wildlife management areas at the                              connectivity of wildlife management areas. The extent to
           interface of the Greater Kafue System and adjacent wildlife                           which existing and emerging drivers of species loss are
           management areas in Namibia and Botswana.                                             severing biological linkages between the Greater Kafue
               These data emphasize the challenges associated with the                           System and adjacent wildlife management areas remain
           scope and scale of conservation interventions required to                             unquantified and subject to speculation. However, data sug-
           limit factors driving species loss from seven of nine taxo-                           gest there is a connectivity bottleneck for large mammals in
           nomic families, representing a wide range of species traits.                          the Simalaha Communal Conservancy, with only  of 
           If drivers of species loss continue to limit population recov-                        species known from historical records detected throughout
           ery in the Simalaha/Nyawa areas then source–sink dynam-                               this area in –.
           ics and edge effects can have a negative impact on the                                   The long-distance dispersal capabilities of large carni-
           population viability of vulnerable species in peripheral wild-                        vores imply scope for gene flow between the Greater
           life management areas at local and transboundary scales.                              Kafue System and adjacent wildlife management areas in
               Wide-ranging species are particularly susceptible to                              the Kavango–Zambezi Transfrontier Conservation Area,
           source–sink dynamics and edge effects, and therefore the                              and therefore the extent to which connectivity bottlenecks
           absence of large carnivores from the Simalaha and the                                 impact processes of immigration and emigration in highly
           Simalaha Wildlife Recovery Sanctuary indicates the need                               mobile species is a priority area of research for regional con-
           for additional research to understand the status and drivers                          nectivity conservation management.

           Oryx, 2019, 53(4), 764–773 © 2018 Fauna & Flora International doi:10.1017/S0030605317001594
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. https://doi.org/10.1017/S0030605317001594
772         R. Lines et al.

                 We focused on ascertaining changes to the occurrence and                                         A N S E L L , W.F.H. () Some mammals from Northern Rhodesia.
             distribution of  terrestrial mammals .  kg known from                                                Journal of Natural History, , –.
                                                                                                                  A N S E L L , W.F.H. () Further data on Northern Rhodesian
             four composite wildlife management areas between the
                                                                                                                      ungulates. Mammalia, , –.
             Greater Kafue System and a central cluster of wildlife manage-                                       A N S E L L , W.F.H. () Mammals of Northern Rhodesia. The
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           Oryx, 2019, 53(4), 764–773 © 2018 Fauna & Flora International doi:10.1017/S0030605317001594
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