Rangeland Ecology & Management 79 (2021) 64–76 Contents lists available at ScienceDirect Rangeland Ecology & Management journal homepage: www.elsevier.com/locate/rama Local-Scale Variation in Land Use Practice Supports a Diverse Carnivore ✩ Guild on Namibian Multiple-Use Rangeland ∗ Stijn Verschueren 1,2, , Willem D. Briers-Louw 1,3, Pedro Monterroso 4, Laurie Marker 1 1 Cheetah Conservation Fund, Otjiwarongo, Namibia 2 Evolutionary Ecology Group, University of Antwerp, Wilrijk, Belgium 3 Zambeze Delta Conservation, Marromeu, Mozambique 4 CIBIO/InBIO, Biodiversity and Genetic Resources Research Center, University of Biodiversity and Genetic Resources Research Center, University of Porto, Vairão Campus, 4485-661 Vairão, Portugal a r t i c l e i n f o a b s t r a c t Article history: Many rangelands worldwide are threatened by human population growth, so there is an urgent need Received 19 October 2020 for understanding how we can preserve functional diversity across these systems. The conservation and Revised 7 July 2021 restoration of mammalian carnivores (order Carnivora) is critical because they impart important trophic Accepted 15 July 2021 cascading effects. Land use practice on rangelands may determine carnivore distributions and abun- dances; thus, to effectively facilitate coexistence between carnivores and humans, it is essential to under- Key Words: stand carnivore community functioning in human-dominated landscapes. We conducted a camera trap- camera trap ping survey on multiple-use rangeland in north-central Namibia to investigate the spatial ecology of free- coexistence ranging carnivores in a farming system that comprises both livestock farming activities and wildlife-based farmland land uses. We hypothesized that carnivore diversity and occupancy would be determined by farm type occupancy seasonality and predicted the associations of carnivore distributions with covariates related to resource availability, spatial ecology intraguild interactions, and anthropogenic influence. We considered single-season occupancy models and hypothesized that in this semiarid study system, seasonality had profound effects on the spatial ecology of carnivores. Our results show that Namibian multiple-use rangeland supported a diverse carnivore guild. Carnivore diversity and occupancy were generally similar across farm types, suggesting that the carnivore community assemblage in our study area was homogeneous. Local-scale variation in land use practices did not limit carnivore distributions, which could be key to maintaining ecological integrity of rangelands. The effect of seasonality suggested that carnivore space use on Namibian rangelands was influenced by availability of dry season resources. In addition, carnivores were dependent on natural resources, showed complex interactions with intraguild members, and had seasonally contrasting associations with anthro- pogenic activities. Namibian multiple-use rangelands may function as viable socioecological landscapes and could act as an important link between core conservation areas. ©2021 The Society for Range Management. Published by Elsevier Inc. All rights reserved. Introduction lacking ( Fjeldså et al. 2004 ), and many PAs perform poorly because of insufficient funding and poor management ( Lindsey et al. 2017 ). Although the establishment of protected areas (PAs) throughout In addition, edge effects reduce effective reserve size and enhance Africa has been instrumental for the conservation of biodiversity, isolation ( Woodroffe and Ginsberg 1998 ; Newmark 2008 ). Wide- many large mammal populations continue to decline ( Craigie et al. ranging species often extend their home ranges beyond the ad- 2010 ). A comprehensive and interconnected PAs network is often ministrative boundaries of PAs, so integrative management at the landscape-level is needed to target a more holistic approach to conservation ( Newmark 2008 ; Bowyer et al. 2019 ). ✩ The Go Green Fund, initiated by Nedbank Namibia and Namibia Nature Founda- Many unprotected lands primarily emphasize extensive grazing tion, provided funding for this research. P. M. was supported by UID/BIA/50027/2019 and pastoralism ( Lund 2007 ; Briske 2017 ). These rangelands are with funding from FCT/MCTES through national funds. This research was conducted common in arid and semiarid to dry subhumid areas and are usu- within the UNESCO Chair “Life on Land.” ally dominated by grasses and shrubs ( Briske 2017 ). Rangelands are ∗ Correspondence: Cheetah Conservation Fund, PO Box 1755, Otjiwarongo, keystone elements of these landscapes because they offer a liveli- Namibia. E-mail address: [email protected] (S. Verschueren). hood to millions of people while supporting important ecological https://doi.org/10.1016/j.rama.2021.07.007 1550-7424/© 2021 The Society for Range Management. Published by Elsevier Inc. All rights reserved. S. Verschueren, W.D. Briers-Louw and P. Monterroso et al. / Rangeland Ecology & Management 79 (2021) 64–76 65 regulatory services and biological diversity ( Lund 2007 ). Range- tions, and anthropogenic influence. We grouped carnivore species lands can support diverse communities of terrestrial vertebrates with shared characteristics (as defined by Jones et al. 2009 ) into ( Kinnaird and O’Brien 2012 ; Msuha et al. 2012 ; Kiffner et al. 2015 ; functional groups to detect prevailing drivers of occupancy of free- Drouilly et al. 2018a ), but livestock farming practices can be diffi- ranging carnivores in this mosaic rangeland system ( Table 1 ). We cult to reconcile with biodiversity conservation due to overstock- defined functional groups by adult body mass, intermediate-sized ing, large herbivore removal, and carnivore persecution ( Du Toit (1 −15 kg) versus large-sized carnivores ( > 15 kg) ( Buskirk 1999 ), and Cumming 1999 ; Ogutu et al. 2005 ; Fynn et al. 2016 ; Briske and by feeding strategy, specialists (i.e., carnivorous and insec- 2017 ). With continued human population growth, functional diver- tivorous [reliant on one dietary category: vertebrates or inverte- sity of rangelands becomes threatened, creating a strong demand brates]) versus generalists (i.e., omnivorous [reliant on multiple di- for multipurpose management of rangelands to provide sustainable etary categories: vertebrates, invertebrates, fruits and/or plants]) goods and services ( Bedunah and Angerer 2012 ). ( Jones et al. 2009 ). Throughout southern Africa, legislative changes in the past We hypothesized that the carnivore community would be more decades provided farmers with property rights over wildlife diverse and with higher species-specific occupancy rates on game occurring on their land, which encouraged farmers to invest in farms compared with mixed farms and livestock farms due to a wildlife as an alternative source of income to livestock ( Barnes suite of favorable characteristics, namely abundant natural prey, and de Jager 1996 ; Barnes et al. 2004 ; Clements et al. 2019 ). reduced disturbance, and high-quality habitat ( Boast and Houser The development of game farms, with land use practices such as 2012 ). However, as carnivores may move extensively in search for game meat production, ecotourism, trophy hunting, and/or wildlife resources, they may cross less favorable areas, possibly leading to a conservation (i.e., wildlife-based land uses), resulted in substantial homogeneous community assemblage across farm types ( Curveira- increases of wildlife numbers outside state PAs because more Santos et al. 2021 ). habitat became preserved ( Barnes et al. 2004 ). Game farming may We further hypothesized that seasonality would have profound be more profitable compared with livestock farming in semiarid effects on the processes governing the spatial ecology of carni- areas, which underpinned the transition toward multiple-use land- vores in semiarid systems such as ours ( Schuette et al. 2013 ; Rich scapes ( Lindsey et al. 2013b ; Holechek and Valdez 2018 ; Young et al. 2017b ). Therefore, we analyzed the strength and direction et al. 2018 ). This resulted in a mosaic distribution of livestock of seasonal covariate associations related to resource availability, farms, game farms, and mixed farms (both livestock production intraguild interactions, and anthropogenic influence. More specifi- and wildlife-based land uses) across southern African rangelands, cally, we expected resource scarcity during the dry season to re- and therefore it is vital to understand the effect of local-scale strict space use of free-ranging carnivores due to preferred use of variation in land use practices on patterns of biodiversity and space around resources of limited availability. ecosystem health. We also hypothesized that generalist species would be more Species of the order Carnivora (hereafter named “carnivores”) common than specialists because they are better suited to persist have a disproportionately large effect on ecosystems because of and thrive in human-dominated landscapes, given their higher eco- cascading effects across trophic webs and are therefore often re- logical plasticity ( Fedriani et al. 2001 ; Davis et al. 2018 ). Lastly, we ferred to as keystone species ( Roemer et al. 2009 ; Ripple et al. expected that the occupancy probability of large-sized carnivores 2014 ). They have been studied extensively worldwide and, despite would be lower than intermediate-sized ones because of their typ- regional variation, their main patterns of occurrence have been ically lower densities and higher vulnerability to local extinction found to be governed by 1) resource