ORIGINAL RESEARCH

 

Will woody plant encroachment impact the visitor experience and economy of conservation areas?

 

 

Emma F. Gray^{I, II}; William J. Bond^II

^ISouth African Environmental Observation Network Fynbos Node, Cape Town, South Africa
^IIDepartment of Botany, University of Cape Town, South Africa

Correspondence

 

 

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ABSTRACT

Woody plant encroachment into savannas is a globally prevalent phenomenon and impacts ecosystem goods and services such as biodiversity, carbon storage, nutrient cycling, grazing and hydrology. The direct ecological and economic consequences for rangelands have been fairly well studied, but, to our knowledge, the economic impact on conservation efforts has not been investigated. African savannas are important as conservation areas because they support large numbers of the world's remaining megafauna. This study used visitor surveys and long-term mammal distribution data to investigate how an increase in tree density might affect the visibility of animals in a conservation area, which could reduce the satisfaction of visitors to the area. We found that apparent herd sizes and density of animals were much reduced in woody areas, suggesting that visibility is negatively impacted. Visitor surveys determined that a large fraction (almost half) of potential future visitors to the park may be lost if animals became more difficult to see and that the majority of these would be the higher-spending visitors. Responses differed depending on the origin of visitors, with international visitors being more interested in seeing animals, whilst local visitors were more content with just being away from the city. The results suggest that woody plant encroachment may have significant impacts on visitor numbers to savanna conservation areas, whilst animal numbers and densities may also be significantly impacted.
Conservation implications: The results pointed to potentially significant economic consequences for conservation efforts as visitors become less satisfied with their experience. Perceptions of visitors are important for management decisions as park fees contribute significantly to conservation efforts. This could ultimately result in a reduced capacity for African conservation areas to conserve their biodiversity effectively. The results suggest that management may need to re-evaluate their approach to controlling woody plant encroachment.

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Introduction

Savannas and grasslands are economically important as rangelands and agricultural lands (Costanza et al. 1997; Sankaran, Ratnam & Hanan 2004). In Africa in particular, savannas have an intrinsic sentimental value because they are home to most of the world's last remaining megaherbivores (animals > 1000 kg). This translates into an economic importance because they attract visitors from all over the world, which contributes significantly to the gross domestic product of many African countries (Akama & Kieti 2003; Akama, Maingi & Camargo 2011; Di Minin et al. 2012). The experiences of these visitors are an important source of information to guide park managers and planners in appropriate measures to achieve both visitor satisfaction and conservation goals (Boshoff et al. 2007; Obua & Harding 1996).

Woody plant encroachment has been extensive in both farmed and conserved savannas all over the world over the last century (Archer, Schimel & Holland 1995; Asner et al. 2003; Britz & Ward 2007; Fensham, Fairfax & Archer 2005; Skarpe 1990; Van Auken 2000; Wigley, Bond & Hoffman 2010). The drivers are strongly debated, but recent literature recognises the importance of both local drivers, such as changes in land use (fire and herbivore stocking rates), and global drivers, such as increases in atmospheric carbon dioxide concentrations (Bond 2008; Bond & Midgley 2012; Bowman, Murphy & Banfai 2011; Wigley et al. 2010). Besides environmental repercussions such as loss of biodiversity (Bond & Parr 2010; Parr, Gray & Bond 2012; Ratajczak, Nippert & Collins 2012) and altered ecosystem goods and services such as water supply and carbon sequestration (Coetsee et al. 2013), woody thickening also has economic consequences on commercial rangelands. Stocking rates are likely to drop as trees increasingly restrict animal access and forage is reduced as grass cover declines. The consequence is a reduction in the overall carrying capacity of the land (Moleele et al. 2002; Roques, O'connor & Watkinson 2001). In Namibia, cattle numbers have been reduced to 36% of what they were in 1959, and woody plant encroachment is estimated to cause a loss of more than N$700 million per annum (De Klerk 2004). What is often overlooked is that in savannas under conservation there are likely to be direct consequences for wild animals, which may translate into unforeseeable indirect economic consequences for conservation areas.

Savannas support a diversity of habitats for mammals and birds (Sirami et al. 2009). Different animals show diverse preferences for certain habitats owing to direct (Sinclair, Mduma & Brashares 2003) and indirect (Brown 1988, 1992, 1999; Brown, Laundré & Gurung 1999; Laundré, Hernandez & Altendorf 2001; Ripple & Beschta 2004) effects of predation and the availability of grazing. This makes for a diversity of wildlife viewing experiences for visitors to conservation areas. This spatial heterogeneity is crucial for maintaining diverse species assemblages and may help to buffer against changes in resource availability due to a changing climate (Wang et al. 2006) by compensating for temporal variability in resource availability (Fryxell et al. 2005). Woody plant encroachment will reduce spatial heterogeneity in vegetation structure. This change in habitat structure is likely to affect animal distributions, herd sizes and densities, with the direction of change depending on the size and feeding habits of the species concerned (Du Toit & Owen-Smith 1989). However, whether actual animal numbers are reduced may be of little importance to visitors. An increase in tree cover will reduce visibility and, as more woody plants encroach, it is likely that animals will become more difficult to see (Marshall, Lovett & White 2008). An inability to see animals may lead to a reduced wildlife experience, which could lead to a decrease in visitor numbers in conservation areas. Regular visitors to a conservation area are likely to notice reduced animal visibility due to woody plant encroachment, which could translate to a reduction in their desire to return to the park.

Here we report how woody plant density affects wildlife viewing in a savanna park and how visitors are likely to respond to woody plant encroachment. Woody plant encroachment is widespread in African savannas and is well documented in our study area, which was far more open in the early and mid-20th century but has become heavily encroached in the last few decades (Watson & Macdonald 1983; Wigley et al. 2010). We addressed the following questions:

• Does denser vegetation result in fewer visible animals and therefore poorer game viewing opportunites?

• Do visitors go to parks primarily to see wildlife?

• If visitors' ability to view wildlife is reduced, will they be less likely to visit parks and, if so, could this lead to reduced visitor numbers in parks and subsequent loss of revenue?

 

Methods

Study area

This study was conducted in the Hluhluwe-iMfolozi Park (HiP) complex, South Africa (28°00'-28°26'S; 31°43'-32°09'E), which consists of the Hluhluwe Game Reserve (225 km²) in the north, the iMfolozi Game Reserve (447 km²) in the south and a corridor (227 km²) joining the two (Whateley & Porter 1983). HiP, which is managed by Ezemvelo KwaZulu-Natal (KZN) Wildlife, is a major contributor to the tourism industry of KZN (Aylward & Lutz 2003).

HiP contains the full complement of large mammal herbivores characteristic of the region (including black and white rhinoceros) and is the oldest conservation area in South Africa. The park is characterised by high habitat heterogeneity, with vegetation ranging from open grassland and savanna to closed woodland and forest (Whateley & Porter 1983). The vegetation in the park has thickened significantly over the last century (Watson 1995; Watson & Macdonald 1983) and a study by Wigley et al. (2010) found an increase in woody cover from 14% in 1937 to 58% in 2004 in a significant portion of the northern Hluhluwe section of the park. Visitors to HiP go there predominantly to see wildlife (Di Minin et al. 2012) and are an important source of income for the park, surrounding communities and smaller game reserves (Aylward & Lutz 2003).

The annual economic contribution of HiP to Ezemvelo KZN Wildlife is disproportionately large for its area and expenditures (Table 1) and therefore any loss in revenue that HiP suffers is likely to have repercussions for Ezemvelo KZN Wildlife parks in general. Reduced revenue generation from tourism may ultimately influence operational costs and conservation goals of the parks authority.

Visibility of animals in different habitats

To determine the impact of vegetation on animal visibility we used census data collected by the Earthwatch Institution in 2008, 2010 and 2012. During censuses, observation teams walked a total of 26 fixed-line transects of between 3.9 km and 10.4 km (Cromsigt et al. 2009). Teams of two observers walked transects just after sunrise over a three-month period in the dry season (from July to October). All sightings of herbivore species larger than a hare within 500 m of both sides of a transect were recorded. The species and abundance, the GPS coordinates of the observer and the distance and bearing of the animal(s) from the observer were recorded (Cromsigt et al. 2009). The vegetation in each transect was classified into five different categories according to increasing woodiness: grassland, open savanna, closed savanna, thicket (dense, impenetrable woody vegetation) and forest.

We used these data to make inferences about whether animal visibility is significantly reduced in habitats with dense tree cover. Besides analysing the visibility of all species in combination, we also looked at nyala (Tragelaphus angasi), kudu (Tragelaphus strepsiceros) and impala (Aepyceros melampus) in isolation. Nyala and kudu are predominantly browsers and so would be expected to be found in forested areas. Impala are predominantly grazers and so would be expected to be found mainly in savanna areas. We hypothesised that regardless of feeding preferences, larger herd sizes and higher animal densities would be recorded in open grassland and savanna areas because animals would more likely have been counted in these habitats, as visibility is not obstructed by trees (Marshall et al. 2008). We used the statistical software package R to build a generalised linear model to predict the response variable density of animals per area (km²). We used habitat as an explanatory variable to determine if it explained a significant amount of deviance in the model. We included transect length as an offset to account for different sizes of each habitat. This produced a response variable of animal density. In this step, we were interested in the apparent rather than the actual density of animals in each habitat, so there was no need to standardise our estimates using correction factors.

Visitor surveys

We used structured questionnaires to survey 220 visitors to the park in July 2009 and May 2010. We approached visitors in accommodation camps, at the entrances to the park and in recreational areas throughout the park. The park has two main accommodation areas, with Hilltop being a luxury camp in the Hluhluwe section of the park and Mpila a budget camp in the iMfolozi section of the park. All visitor groups present were approached, with one representative per group completing the questionnaire. The questionnaire included both open-ended and multiple-choice questions (see Online Appendix 1).

Vegetation change and desirability

We asked repeat visitors to describe any change in vegetation they may have noticed since their first visit, without explaining the woody plant encroachment phenomenon to them. Visitors were also shown photographs depicting three different vegetation types (open savanna, heterogeneous landscapes with savanna and forest elements, and encroached thicket) and asked which vegetation they most enjoyed driving through. The responses to these questions were used to establish the main reasons for visiting the park and which aspects of the park and its wildlife visitors were most attracted to. Visitors were also asked to identify, from the same pictures, in which vegetation they had seen the most wildlife.

Visitor demography and reasons for visiting

At the onset of the survey we established where visitors were from, where they were staying, whether they had visited the park before and, if so, how often. Visitors were asked why they had chosen to visit HiP and to indicate which animals they most wanted to see in the park. This allowed us to determine whether responses differed depending on visitor origin and accommodation choice using contingency tables and chi-square tests.

Effect of encroachment on visitor satisfaction

We then explained the woody plant encroachment phenomenon. Visitors were asked to record, on a scale between 1 and 5, how strongly they agreed with a series of statements (5 = strongly agree; 1 = strongly disagree; also see Online Appendix 1) to establish whether they would return to visit the park despite a reduction in the visibility of wildlife. The results were subsequently related to the demographic profiles of visitors using contingency tables and chi-square tests.

 

Results

Visibility of animals in different habitats

Results from a generalised linear model of the form:

suggested that mean density was significantly higher in grasslands and open habitats than in closed habitats (see results of model summarised in Table 2). Forests had the lowest observed animal densities (0.38 animals/km²). Some 30% of the deviance in the model was explained by habitat. The open habitats had much larger animal groups, with a maximum of 300 individuals in one group in grassland, and a maximum of just 50 in forest (Figure 1a). This pattern was repeated regardless of feeding ecology when animals were grouped according to species (Figure 1b-d). These results suggest that an increase in woody plants would likely cause a reduction in the apparent density of animals.

 

 

 

 

Visitor perceptions

Before woody plant encroachment was explained to visitors, they were asked how long they had been visiting the park and whether they had noticed a change in vegetation since they had first visited. Of the 64.7% of returning visitors (n = 66) who answered that they had noticed a change, 80% (n = 53) described a vegetation change that could be identified as woody plant encroachment. Figure 2 shows the percentage of visitors who had observed vegetation change as a function of how long ago their first visit was. Time since first visit increased the likelihood of having noticed vegetation change (χ² = 22.30, df = 6, p = 0.001). Of the 20 visitors who had been visiting the park for more than 30 years, 80% (n = 16) had noticed woody encroachment. A further 10% had noticed some change in the vegetation but did not describe it. This was in contrast to responses from visitors who had been visiting for less than 10 years (n = 48), with only 31.2% having noticed woody plant encroachment and 12.5% having noticed some change in vegetation.

 

 

When visitors were asked to identify, using photographs, the habitat they thought they had seen most animals in, 63% chose photographs showing open savannas or landscapes with both forest and savanna elements. Only 37% of visitors indicated that they had seen more animals in closed thicket and forest landscapes. We used the same photographs to ask visitors which scenery they enjoyed most when driving through the park. According to the responses, 48% of visitors preferred driving through landscapes with forest and savanna elements (i.e. heterogeneous landscapes), 27% preferred open savanna landscapes and 25% preferred closed habitats.

Demographic profiles of visitors

The demographic profiles of visitors are shown in Table 3 and Table 4. Although the majority of visitors to the park were local travellers (i.e. from KZN or elsewhere in or close to South Africa¹) a large proportion (45.5%) of visitors were international guests. Most guests were day visitors (58.1%). Of the overnight visitors (n = 92), 65.2% stayed at the Hilltop camp and the majority of these (60%) were international visitors. A chi-square test confirmed that local overnight visitors were less likely to stay at Hilltop, whilst international visitors were more likely to stay there than would be expected by chance (χ² = 13.04, df = 4, p = 0.011).

 

 

Of the surveyed visitors, 47.3% had been to the park before. More than half of the overnight visitors (n = 48) were return visitors. Most of the day visitors (56.1%, n = 72) had not visited the park before. Likelihood of staying in park accommodation was not different for first-time or regular visitors (χ² = 3.62, df = 2, p = 0.16) (Table 4).

Reasons for visiting Hluhluwe-iMfolozi Park

This was an open-ended question, so answers were combined into categories. The most commonly cited reason for choosing to visit HiP was that it was close or convenient (Figure 3). For international visitors, the most commonly cited reason for visiting was that it was recommended by someone or because it formed part of a tour. Tranquillity or being in nature was most commonly cited by local visitors as a reason for choosing the park. The park's heritage and its well-known rhino conservation efforts appeared to be more important to international than local visitors. A chi-square test confirmed that the origin of visitors (i.e. local vs international) influenced their reasons for choosing to visit HiP (χ² = 26.8, df = 10, p = 0.003). Surprisingly, only 17.7% of visitors cited seeing animals in general, and the 'big five' (lion, leopard, elephant, white rhino and buffalo) in particular, as the most important reason for visiting the park. However, in response to the statement: 'The main reason I am visiting this park is to see animals,' 58.6% strongly agreed, whilst only 5.58% strongly disagreed. Visitor demography strongly influenced how they answered this question (χ² = 38.9, df = 8, p < 0.0001): 46.0% of visitors who strongly agreed with the statement were international guests, whilst 83.3% of respondents who strongly disagreed with the statement were local visitors, specifically from KZN. Visitors from elsewhere in Africa were more likely to have a neutral view, although the majority of them still strongly agreed with the statement.

 

 

The 'big five' tended to be the most desirable animals (Figure 4) for visitors to see whilst in the park, although giraffe were more popular than buffalo. Animal preferences were affected by visitor origin (χ² = 51.90, df = 26, p = 0.002), with giraffe and zebra being more popular sightings amongst international visitors, and buffalo and birds being more popular sightings amongst local visitors. Whether visitors had been to the park before or not also affected their desire to see certain animals (χ² = 78.7, df = 52, p = 0.0098): the desire to see leopard, impala, nyala and cheetah increased if people had been to the park before, but the desire to see zebra decreased.

 

 

Visitor response to changing vegetation

Although the vast majority of visitors (87.4%) indicated that they would like to return to the park given the opportunity, 42.4% of visitors agreed or strongly agreed with the statement that they would not return to the park if it became more difficult to see animals (Table 5). Accommodation choice and whether they had been to the park before affected their response to this statement (χ² = 36.86, df = 22, p = 0.025). Day visitors or visitors staying at Hilltop were less likely to return if game became more difficult to spot than those staying at Mpila. First-time visitors were also less likely to return than regular visitors if animals became more difficult to see.

A chi-square test found that visitors whose responses indicated that they were unlikely to return if encroachment made wildlife more difficult to see were likely to agree with the statement that encroachment should be controlled (χ² = 50.18, df = 16, p < 0.0001). Day visitors were also more likely to agree with controlling encroachment than overnight guests staying in park accommodation (χ² = 19.46, df = 8, p = 0.01).

 

Discussion

Woody plant encroachment will impact on many savanna ecosystem goods and services such as fodder availability in rangelands (De Klerk 2004), biodiversity (Parr et al. 2012), hydrology and nutrient cycling (Archer, Boutton & McMurtry 2004). It is also likely to have regional-scale feedbacks to the earth-atmosphere system because of changes in albedo and gas exchange (Asner et al. 2004; Beerling & Osborne 2006).

This study set out to determine whether woody plant encroachment would have additional economic impacts due to a reduction in game viewing capacity in HiP, where evidence of extensive woody plant encroachment has been seen over the last century (Wigley et al. 2010). Specifically, we set out to investigate (1) whether woody encroachment would reduce visibility of animals, (2) whether guests visited the park predominantly to see animals and (3) whether reduced visibility of animals could therefore result in reduced visitor numbers and, consequently, reduced park revenue.

Our study showed clear evidence that wildlife visibility is compromised by woody plant encroachment (Figure 1), regardless of the species of animal. In addition, most visitors indicated that they had seen more animals in open and heterogeneous habitats. Visitors also found heterogeneous landscapes with typical savanna characteristics, such as open plains and single large trees, more aesthetically pleasing than encroached areas. Besides the aesthetics, heterogeneous vegetation is ecologically important in increasing the carrying capacity of populations, because it provides a diversity of resources (Fryxell et al. 2005; Wang et al. 2006).

In a previous study undertaken in HiP, Cromsigt, Prins and Olff (2009) suggested that habitat heterogeneity might facilitate herbivore diversity in savanna ecosystems.

Whilst the conclusion that denser vegetation will result in reduced visibility of animals may seem an obvious one, it is likely to have numerous less obvious knock-on effects, one of which is the potential reduction in visitor numbers to game reserves. Our results established that whilst wildlife viewing may not be the main reason for visitors choosing HiP (Figure 3), the majority of visitors were indeed intent on seeing wildlife. This was, however, affected by visitor origin, with international visitors being more interested in seeing animals than local visitors. Di Minin et al. (2012) found a similar trend when investigating the relative importance of the 'big five' in HiP, with more wealthy and less experienced international visitors most interested in seeing these animals.

Whether visitors will stop visiting the park because of encroachment was not clear from our results, but the responses to our questions have led us to make certain inferences. People who visit the park because it is close or convenient (Figure 3) are unlikely to be much affected by a change in woody vegetation as the location of the park and its proximity to other areas will not change. Those who visit the park because of its heritage and rhino conservation efforts, as well as those who have come to escape city life and experience the tranquillity of nature, are also unlikely to stop visiting the park as a results of encroachment. For international visitors, however, recommendations and choice of tour operator were important reasons for visiting HiP. Mmopelwa, Kgathi and Molefhe (2007) found that in Botswana international visitors were more willing to pay higher rates for wildlife tourism, a finding mirrored by our observations that international tourists often stayed in the more expensive accommodation. Tour operators are unlikely to take tourists to a game reserve where wildlife viewing success is poor, and the same may be expected from guide book recommendations. This suggests that the park could lose a significant fraction of their international visitors (and therefore highest spenders) because of woody plant encroachment.

Although the majority of visitors indicated they had enjoyed their experience and would like to return to the park, a large portion of visitors confirmed that reduced visibility of animals would impact their decision to the return to the park (with nuances depending on whether they were regular visitors and choice of accommodation). Based on visitor responses we estimated that Hilltop could lose at least half of its potential future visitors and about 40% of its regular visitors if animals became more difficult to see. If we consider return visitors as definite future visitors, the park would lose about 40% of its definite future visitors and about one third of these would be visitors to Hilltop (i.e. the visitors who are likely to spend the most money whilst in the park). In addition, with an estimated return rate of 47% for first-time visitors, Hilltop could lose nearly half of potential future visitors. This combines to a possible total loss of about 40% of potential future visitors to the park if animals became more difficult to see. These results suggest that park managers should consider suitable encroachment control measures if they are to ensure return visits. On the whole, visitors did not object to the suggestion of controlling encroachment, so park managers are not likely to face public outcry if such measures were put in place.

 

Conclusion

Our study has shown strong evidence that woody plant encroachment influences visitor experience in HiP and suggests that visitor numbers might diminish in heavily encroached areas. Woody plant encroachment is not merely a debate on conservation objectives for park management, but could negatively influence the economy of the park. Our results suggest that the park may lose a significant portion of visitors if woody encroachment continues unchecked.

However, studies have shown that visitor demand for large wildlife viewing is unlikely to disappear and the overall number of wildlife tourists in Africa is unlikely to drop (Boshoff et al. 2007; Goodwin & Leader-Williams 2000; Lindsey et al. 2007; Mmopelwa et al. 2007; Obua & Harding 1996). Rather, we may see a shift in visitor numbers to different parks as visitors choose parks where they are more likely to have their desired experience. To ensure success of their parks in the future, managers need to make decisions based on conservation principles, but with an understanding that conservation will be facilitated by the revenue generated by visitors (Boshoff et al. 2007; Kiss 2004; Lindsey et al. 2007; Mmopelwa et al. 2007). As HiP contributes significantly to the revenue of Ezemvelo KZN Wildlife as a whole (Aylward & Lutz 2003), a decrease in revenue may affect the organisation significantly. In addition, Ezemvelo KZN Wildlife employs many members of the local community (Aylward & Lutz 2003) and many livelihoods could therefore be affected by a drop in visitor numbers, as well as the many smaller businesses and private game reserves and local ecotourism ventures that benefit from being in close proximity to such a large tourist attraction (Bookbinder et al. 1998; Kiss 2004). Our conclusions on the impact of habitat change on park visitors are not restricted to HiP, as woody encroachment is a widespread phenomenon in many savannas in Africa. It will be prudent for managers to consider strategic management of the phenomenon using fire and other management tools.

 

Acknowledgements

This study was made possible through funding from the University of Cape Town Woolworths Fellowship and the German Academic Exchange Programme (DAAD). Thanks to the Earthwatch Institute for mammal census data, Geoffrey Clinning for providing data, and Amy Rose Wilkinson for assistance in the field. Thanks also to field and administrative staff at Hluhluwe-iMfolozi Park for allowing us to undertake the study and providing us with assistance at every stage.

Competing interests

The authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article.

Authors' contributions

W.J.B. (University of Cape Town) was the project leader. E.F.G. (University of Cape Town) and W.J.B. were both responsible for project design. E.F.G. undertook all fieldwork and analyses and wrote the manuscript. W.J.B. made conceptual contributions to the manuscript.

 

References

Akama, J.S. & Kieti, D.M., 2003, 'Measuring tourist satisfaction with Kenya's wildlife safari: A case study of Tsavo West National Park', Tourism Management 24(1), 73-81. http://dx.doi.org/10.1016/S0261-5177(02)00044-4        [ Links ]

Akama, J.S., Maingi, S. & Camargo, B., 2011, 'Wildlife conservation, safari tourism and the role of tourism certification in Kenya: A postcolonial critique', Tourism Recreation Research 36, 1-11.         [ Links ]

Archer, S., Schimel, D.S. & Holland, E.A., 1995, 'Mechanisms of shrubland expansion: Land use, climate or CO2', Climatic Change 29, 91-99. http://dx.doi.org/10.1007/BF01091640        [ Links ]

Archer, S., Boutton, T. & McMurtry, C., 2004, 'Carbon and nitrogen accumulation in a savanna landscape: Field and modeling perspectives', in M. Shiomi, H. Kawahata, H. Koizumi, A. Tsuda & Y. Awaya (eds.), Global Environmental Change in the Ocean and On Land, pp. 359-373, Terra Scientific, Tokyo.         [ Links ]

Asner, G.P., Archer, S., Hughes, R.F., Ansley, R.J. & Wessman, C.A., 2003, 'Net changes in regional woody vegetation cover and carbon storage in Texas drylands, 19371999', Global Change Biology 9(3), 316-335. http://dx.doi.org/10.1046/j.1365-2486.2003.00594.x        [ Links ]

Asner, G.P., Elmore, A.J., Olander, L.P., Martin, R.E. & Harris, T., 2004, 'Grazing systems, ecosystem responses, and global change', Annual Review of Environment and Resources 29, 261-299. http://dx.doi.org/10.1146/annurev.energy.29.062403.102142        [ Links ]

Aylward, B.A. & Lutz, E. (eds.), 2003, Nature tourism, conservation, and development in KwaZulu-Natal, South Africa, World Bank Publications, Washington.         [ Links ]

Beerling, D.J. & Osborne, C.P., 2006, 'The origin of the savanna biome', Global Change Biology 12(11), 2023-2031. http://dx.doi.org/10.1111/j.1365-2486.2006.01239.x        [ Links ]

Bond, W.J., 2008, 'What limits trees in C4 grassland and savannas?', Annual review of Ecology, Evolution and Systematics 39, 641-659. http://dx.doi.org/10.1146/annurev.ecolsys.39.110707.173411        [ Links ]

Bond, W.J. & Midgley, G.F., 2012, 'CO2 and the uneasy interactions of trees and savanna grasses', Philosophical Transactions of the Royal Society Biological Sciences 367, 601-612. http://dx.doi.org/10.1098/rstb.2011.0182, PMid:22232770        [ Links ]

Bond, W.J. & Parr, C.L., 2010, 'Beyond the forest edge: Ecology, diversity and conservation of the grassy biomes', Biological Conservation 143(10), 2395-2404. http://dx.doi.org/10.1016/j.biocon.2009.12.012        [ Links ]

Bookbinder, M.P., Dinerstein, E., Rijal, A., Cauley, H. & Rajouria, A., 1998, 'Ecotourism's support of biodiversity conservation', Conservation Biology 12(6), 1399-1404. http://dx.doi.org/10.1046/j.1523-1739.1998.97229.x        [ Links ]

Boshoff, A.F., Landman, M., Kerley, G.I.H. & Bradfield, M., 2007, 'Profiles, views and observations of visitors to the Addo Elephant National Park, Eastern Cape, South Africa', South African Journal of Wildlife Research 37(2), 189-196. http://dx.doi.org/10.3957/0379-4369-37.2.189        [ Links ]

Bowman, D.M.J.S., Murphy, B.P. & Banfai, D.S., 2011, 'Has global environmental change caused monsoon rainforests to expand in the Australian monsoon tropics?', Landscape Ecology 25, 1247-1260. http://dx.doi.org/10.1007/s10980-010-9496-8        [ Links ]

Britz, M.L. & Ward, D., 2007, 'Dynamics of woody vegetation in a semi-arid savanna, with a focus on bush encroachment', African Journal of Range and Forage Science 24(3), 131-140. http://dx.doi.org/10.2989/AJRFS.2007.24.33.296        [ Links ]

Brown, J.S., 1988, 'Patch use as an indicator of habitat preference, predation risk, and competition', Behavioral Ecology and Sociobiology 22(1), 37-47. http://dx.doi.org/10.1007/BF00395696        [ Links ]

Brown, J.S., 1992, 'Patch use under predation risk: I. Models and predictions', Annales Zoologici Fennici 29, 301-309.         [ Links ]

Brown, J.S., 1999, 'Vigilance, patch use and habitat selection: Foraging under predation risk', Evolutionary Ecology Research 1(1), 49-71.         [ Links ]

Brown, J.S., Laundré, J.W. & Gurung, M., 1999, 'The ecology of fear: Optimal foraging, game theory, and trophic interactions', Journal of Mammalogy 80(2), 385-399. http://dx.doi.org/10.2307/1383287        [ Links ]

Coetsee, C., Gray, E.F., Wakeling, J., Wigley, B.J. & Bond, W.J., 2013, 'Low gains in ecosystem carbon with woody plant encroachment in a South African savanna', Journal of Tropical Ecology 29(1), 49-60. http://dx.doi.org/10.1017/S0266467412000697        [ Links ]

Costanza, R., d'Arge, R., De Groot, R., Farber, S., Grasso, M., Hannon, B. et al., 1997, 'The value of the world's ecosystem services and natural capital', Nature 387(6630), 253-260. http://dx.doi.org/10.1038/387253a0        [ Links ]

Cromsigt, J.P., Van Rensburg, S.J., Etienne, R.S. & Olff, H., 2009 'Monitoring large herbivore diversity at different scales: Comparing direct and indirect methods', Biodiversity and Conservation 18(5), 1219-1231. http://dx.doi.org/10.1007/s10531-008-9506-1        [ Links ]

Cromsigt, J.P., Prins, H.T.H. & Olff, H., 2009, 'Habitat heterogeneity as a driver of ungulate diversity and distribution patterns: Interaction of body mass and digestive strategy', Diversity and Distributions 15, 513-522. http://dx.doi.org/10.1111/j.1472-4642.2008.00554.x        [ Links ]

De Klerk, J.N., 2004, Bush encroachment in Namibia: Report on Phase 1 of the Bush Encroachment Research, Monitoring, and Management Project, Windhoek, Namibia, Ministry of Environment and Tourism, Directorate of Environmental Affairs, Windhoek.         [ Links ]

Di Minin, E., Fraser, I., Slotow, R. & MacMillan, D.C., 2012, 'Understanding heterogeneous preference of tourists for big game species: Implications for conservation and management', Animal Conservation 16(3), 249-258. http://dx.doi.org/10.1111/j.1469-1795.2012.00595.x        [ Links ]

Du Toit, J.T. & Owen-Smith, N., 1989, 'Body size, population metabolism and habitat specialization among large African herbivores', American Naturalist 133, 736-740.         [ Links ]

Fensham, R.J., Fairfax, R.J. & Archer, S., 2005, 'Rainfall, land use and woody vegetation cover change in semi-arid Australian savanna', Journal of Ecology 93(3), 596-606. http://dx.doi.org/10.1111/j.1365-2745.2005.00998.x        [ Links ]

Fryxell, J.M., Wilmshurst, J.F., Sinclair, A.R.E., Haydon, D.T., Holt, R.D. & Abrams, P.A., 2005, 'Landscape scale, heterogeneity, and the viability of Serengeti grazers', Ecology Letters 8, 328-335. http://dx.doi.org/10.1111/j.1461-0248.2005.00727.x        [ Links ]

Goodwin, H.J. & Leader-Williams, N., 2000, 'Tourism and protected areas - distorting conservation priorities towards charismatic megafauna', in A. Entwhistle & N. Dunstone (eds.), Priorities for the conservation of mammalian diversity: Has the panda had its day?, pp. 257-275, Cambridge University Press, Cambridge.         [ Links ]

Kiss, A., 2004, 'Is community-based ecotourism a good use of biodiversity conservation funds?', Trends in Ecology and Evolution 19(5), 232-237. http://dx.doi.org/10.1016/j.tree.2004.03.010, PMid:16701261        [ Links ]

Laundré, J.W., Hernandez, L. & Altendorf, K.B., 2001, 'Wolves, elk, and bison: Reestablishing the "landscape of fear" in Yellowstone National Park, USA', Canadian Journal of Zoology 79(8), 1401-1409. http://dx.doi.org/10.1139/z01-094        [ Links ]

Lindsey, P.A., Alexander, R., Mills, M.G.L., Ramanach, S. & Woodroffe, R., 2007, 'Wildlife viewing preferences of visitors to protected areas in South Africa: Implications for the role of ecotourism in conservation', Journal of Ecotourism 6(1), 19-33. http://dx.doi.org/10.2167/joe133.0        [ Links ]

Marshall, A.R., Lovett, J.C. & White, P.C.L., 2008, 'Selection of line-transect methods for estimating the density of group-living animals: Lessons from the primates', American Journal of Primatology 70, 452-462. http://dx.doi.org/10.1002/ajp.20516, PMid:18240143        [ Links ]

Mmopelwa, G., Kgathi, D.L. & Molefhe, L., 2007, 'Tourists' perceptions and their willingness to pay for park fees: A case study of self-drive tourists and clients for mobile tour operators in Moremi Game Reserve, Botswana', Tourism Management 28(4), 1044-1056. http://dx.doi.org/10.1016/j.tourman.2006.08.014        [ Links ]

Moleele, N.M., Ringrose, S., Matheson, W. & Van der Post, C., 2002, 'More woody plants? The status of bush encroachment in Botswana's grazing areas', Journal of Environmental Management 64(1), 3-11. http://dx.doi.org/10.1006/jema.2001.0486, PMid:11876072        [ Links ]

Obua, J. & Harding, D.M., 1996, 'Visitor characteristics and attitudes towards Kibale National Park, Uganda', Tourism Management 17(7), 495-505. http://dx.doi.org/10.1016/S0261-5177(96)00067-2        [ Links ]

Parr, C.L., Gray, E.F. & Bond, W.J., 2012, 'Cascading biodiversity and functional consequences of a global change-induced biome switch', Diversity and Distributions 18(5), 493-503. http://dx.doi.org/10.1111/j.1472-4642.2012.00882.x        [ Links ]

Ratajczak, Z., Nippert, J.B. & Collins, S.L., 2012, 'Woody encroachment decreases diversity across North American grasslands and savannas', Ecology 93(4), 697703. http://dx.doi.org/10.1890/11-1199.1, PMid:22690619        [ Links ]

Ripple, W.J. & Beschta, R.L., 2004, 'Wolves and the ecology of fear: Can predation risk structure ecosystems?', BioScience 54(8), 755-766. http://dx.doi.org/10.1641/0006-3568(2004)054[0755:WATEOF]2.0.CO;2        [ Links ]

Roques, K.G., O'connor, T.G. & Watkinson, A.R., 2001, 'Dynamics of shrub encroachment in an African savanna: Relative influences of fire, herbivory, rainfall and density dependence', Journal of Applied Ecology 38(2), 268-280. http://dx.doi.org/10.1046/j.1365-2664.2001.00567.x        [ Links ]

Sankaran, M., Ratnam, J. & Hanan, N.P., 2004, 'Tree-grass coexistence in savannas revisited - insights from an examination of assumptions and mechanisms invoked in existing models', Ecology Letters 7, 480-490. http://dx.doi.org/10.1111/j.1461-0248.2004.00596.x        [ Links ]

Sinclair, A.R.E., Mduma, S. & Brashares, J.S., 2003, 'Patterns of predation in a diverse predator-prey system', Nature 425(6955), 288-290. http://dx.doi.org/10.1038/nature01934, PMid:13679915        [ Links ]

Sirami, C., Seymour, C., Midgley, G. & Barnard, P., 2009, 'The impact of shrub encroachment on savanna bird diversity from local to regional scale', Diversity and Distributions 15(6), 948-957. http://dx.doi.org/10.1111/j.1472-4642.2009.00612.x        [ Links ]

Skarpe, C., 1990, 'Structure of the woody vegetation in disturbed and undisturbed arid savanna, Botswana', Plant Ecology 87(1), 11-18. http://dx.doi.org/10.1007/BF00045650        [ Links ]

Van Auken, O.W., 2000, 'Shrub invasions of North American semiarid grasslands', Annual Review of Ecology and Systematics 31, 197-215. http://dx.doi.org/10.1146/annurev.ecolsys.31.1.197        [ Links ]

Wang, G., Hobbs, N.T., Boone, R.B., Illius, A.W., Gordon, I.J., Gross, J.E. et al., 2006, 'Spatial and temporal variability modify density dependence in populations of large herbivores', Ecology 87(1), 95-102. http://dx.doi.org/10.1890/05-0355, PMid:16634300        [ Links ]

Watson, H.K. & Macdonald, I.A.W., 1983, 'Vegetation changes in the Hluhluwe-Umfolozi Game Reserve Complex from 1937 to 1975', Bothalia 14(2), 265-269.         [ Links ]

Watson, H.K., 1995, 'Management implications of vegetation changes in Hluhluwe-iMfolozi Park', South African Geographical Journal 77(2), 77-83. http://dx.doi.org/10.1080/03736245.1995.9713595        [ Links ]

Whateley, A. & Porter, R.N., 1983, 'The woody vegetation communities of the Hluhluwe-Corridor-Umfolozi Game Reserve Complex', Bothalia 14(3), 745-758.         [ Links ]

Wigley, B.J., Bond, W.J. & Hoffman, M.T., 2010, 'Thicket expansion in a South African savanna under divergent land use: Local vs. global drivers?', Global Change Biology 16(3), 964-976. http://dx.doi.org/10.1111/j.1365-2486.2009.02030.x        [ Links ]

 

 

Correspondence:
Emma Gray
Private Bag X7
Claremont 7735, Africa
emmafionagray@gmail.com

Received: 05 Aug. 2012
Accepted: 29 Apr. 2013
Published: 02 Aug. 2013

 

 

Note

Additional supporting information may be found in the online version of this article as an Online Appendix 1 http://dx.doi.org/10.4102/koedoe.v55i1.1106-1
1. Very few African visitors were from outside South Africa and therefore all African and South African visitors were grouped together.