Research Report Etosha National Park

June 2015

A study of the distribution and diversity of small mammal species within the different habitats of Etosha National Park

(M. Hauptfleisch)

Student Name: Matthew Walters Lecturer Name: M. Hauptfleisch Student Number: 213082969 Course of Study: Bachelor of Natural Resource Management Faculty: School of Natural Resources and Spatial Sciences Department: Agriculture and Natural Resource Management

2

Table of Contents

Introduction ............................................................................................................................................ 3

Aims and Objectives ................................................................................................................................ 3

Methods and Materials ........................................................................................................................... 4

Study Sites ........................................................................................................................................... 4

Dolomite Hills .................................................................................................................................. 5

Plains ............................................................................................................................................... 5

Okaukuejo ....................................................................................................................................... 5

Pan Edge .......................................................................................................................................... 5

Plain Steppes ................................................................................................................................... 5

Seasonal Water Depressions........................................................................................................... 5

Hillocks ............................................................................................................................................ 6

Terminalia Woodland ..................................................................................................................... 6

Methods .......................................................................................................................................... 6

Results ..................................................................................................................................................... 8

Overall Results .................................................................................................................................... 8

Trap success and abundance within each habitat .............................................................................. 9

Small mammal species richness within each of the habitats ........................................................... 10

Small Mammal Diversity ................................................................................................................... 11

Discussion.............................................................................................................................................. 12

Conclusions ........................................................................................................................................... 13

Recommendations ................................................................................................................................ 14

Acknowledgements ............................................................................................................................... 14

References ............................................................................................................................................ 15

3

Introduction

Important to the concept of rangeland management within the African context is the appreciation and understanding of small mammal distribution and population size. Small mammals within this study ranged from the three orders Rodentia, Insectivora, and Macroscelidea. Rodentia, by far, makes up the largest percentage of mammal species, with an estimated percentage of 40% (Danell & Aave-Olsson, 2002, as cited in Sizuka, 2014). Globally Rodentia, characterised by their continuously growing chisel-like incisor teeth (Apps, 2000), has a total of 33 families, 481 genera and 2277 species (Carleton & Musser, 2005). The orders Insectivora (aimed at Shrews) and Macroselidae (aimed at Elephant Shrews), though less numerous than the Rodentia are also important indicators of food availability due to their insectivorous eating requirements. In terms of Insectivora, Shrews are placed in the family Soricidae, with its three sub-families, which along with three other families creates a total of 385 shrew species within 26 genera worldwide (Wilson & Reeder, 2011). Elephant shrews, found only in Africa, are made up of 19 species within 4 different genera (Schlitter, 2005). Since the study will be conducted in the Etosha National Park (ENP) in Namibia, a brief overview of small mammals in Namibia is required. Within the overall Namibian regional context, small mammal species of order Rodentia comprises of 35 species, Insectivora comprises of 4 species, and Macroscelidea comprises of 3 species (Matson & Blood, 1994). However, past research and knowledge into the distribution and composition of small species remains limited due to the ENP’s greater focus on the bigger game species (H. Berry, 2013). However, there have been two noteworthy previous studies within ENP that have for a large part given a better understanding of the small mammal composition within the ENP. The first study was conducted through the Etosha Ecological Institute, by M. Griffin, over two periods of his stay in Etosha, in which small mammals were trapped and identified from Kaross in the west to Namutoni in the east (Griffin, 1993). A second study related to the study of the diet of owl species within ENP, mostly in the Okaukuejo area (Dilley, 1996). This study was able to partly determine what small mammal species occurred in the area based primarily on what was found in the owl pellets. The study of small mammals is the key to understanding the dynamics of veld condition, based primarily on the small mammal species’ immediate position as primary and secondary consumers (Avenant, 2000; David & Jarvis, 1983; Foresman & Badyaev, 2005). All data acquired within the study period will be included in the Small Mammal Atlas of Namibia Project.

Aims and Objectives Due to the lack of any current and updated small mammal species inventories within ENP, the aim of the project would be to contribute towards drawing up an inventory of all the small mammals that occur within the habitats of the Park. The research question posed and which the research project was meant to clarify is: what is the difference in species distribution and density within the various habitats of ENP? The null hypothesis (N0 Hypothesis) would be that there is no significant difference between the factors influencing diversity namely: species abundance and species richness observed between the various study sites. The alternative hypothesis (NA Hypothesis) is that there is a significant difference in abundance and species richness between the various study sites. As a means of achieving the overall aims, four objectives need to be realised:

To determine the species distribution and diversity of small mammals within the Park.

To determine the species richness of each sample site.

4

To determine the species diversity of the sample site according to individual species abundance and species richness.

To compare the study sites and to determine the reasons as to why each of them would have been significantly different.

Methods and Materials

Study Sites

Figure 1: Map of study sites at Okaukuejo

Figure 2: Map of the study sites at Namutoni

5

Dolomite Hills

As part of the February sampling period, a trapping site was placed on the mountain ranges 20 km south west of Okaukuejo on a road that led to a radio tower, that go by the name Ondundozonananandana (H. Berry, 1995). Overall, the geology is overwhelmingly dolomite typical of the Karst region with weakly developed shallow soils typical of arid regions (Barnard, 1998). Additionally, the vegetation is described as Mopane Savannah (Mannheimer & Curtis, 2009). The dominant tree species recorded were Colophospermum mopane, Sterculia africana, and several Commiphora species. The dominant grass species recorded were Aristida meridionales, Cenchrus ciliaris,and Enneapogon cenchroides. Trapping within this habitat zone took place 21-24 of February.

Plains

The open plains in which the study site is not a continuous open plain, rather it is open grassland plain surrounded by woodland. The sample site is situated near the road 9.3 km south west of Okaukuejo. Typical of the general Etosha area, the geology is mainly that of calcrete-limestone, and the soils are typically shallow (Barnard, 1998). The vegetation zone, same as that of the dolomite hills, is Mopane Savannah (Mannheimer & Curtis, 2009).The dominant tree species in and around the study site included: Colophospermum mopane, Catophractes alexandrii, and Acacia nebrownii. The dominant grass species within the study site included: Cenchrus ciliaris and Enneapogon cenchroides. Trapping within this habitat zone took place 22-25 of February.

Okaukuejo

The sample site at Okaukuejo was laid out in an area that was still within the town boundaries but was fairly free from the influence of human activity. Starting in the north-western corner of the town, the transect line ran parallel to the western fence in an area free from development, but with a noticeable amount of building rubble. The dominant grasses in Okaukuejo were Cenchrus ciliaris and Enneapogon cenchroides and the dominant tree species were Acacia tortilis, Acacia reficiens, Maerua schinzii and Aloe littoralis. Trapping within this zone took place 16-18 of March.

Pan Edge

The sample site at the Pan edge took place at a seasonal seepage point known as Wolfsnes, known only produce water in years of good rainfall, which is 15.2 km directly north of Okaukuejo. The soil type of the area is predominantly clay in structure (Capobianco Dondonna, personal com., 2015). Defined as a part of the saline desert with dwarf shrub savannah fringe, the dominant grass species Oddysea paucinervis and the dominant tree and shrub species are Acacia tortilis and Salsola tuberculatec (Cunningham & Jankowitz, 2011). Trapping within this zone took place 16-18 of March.

Plain Steppes

The plain steppes are an area typically found north of Okaukuejo and west of the Etosha Pan. The locality of the trapping sites were placed at the airstrip 3.31 km from Okaukuejo itself. In term of the vegetation zone, the steppe plains are within the margins of the dwarf shrub savannah that borders a saline desert. The typical vegetation found within this area consists mostly of Leucosphaera bainesii and Petalidium engleranum (H. H. Berry, 1980), though the grass species were not observed due to the dry nature of the season and the high grazing frequency in and around Okaukuejo. Trapping within this zone took place 23-25 of April.

Seasonal Water Depressions

The seasonal water depressions are a collective name for the depressions immediately west of Okaukuejo that accumulates water in the rainy season. The geology and the soil structure is consistent with that of Okaukuejo in terms of the calcrete-limestone and the shallow soils (Barnard, 1998). The vegetation is virtually Mopane woodland, with the dominant tree species being Colophospermum mopane and Terminalia prunioides. Once again, no grass species were able to be

6

identified as a dominant species due to the dry nature of the season and the high grazing frequency. Trapping within this zone took place 23-25 of April.

Hillocks

The hillocks are a group of hills just north of the Namutoni locality, and are also subsequently north of Fischer’s Pan. The trapping site was placed at the waterhole of Groot Okevi in a line that ran approximately 20 metres from the road. The geology of the Groot Okevi waterhole area is typical sandveld or sandy plains. The typical vegetation of this area includes Terminalia prunioides, Acacia reficiens, and Croton gratissimus. The typical grass species include Cenchrus ciliaris and Eragrostis echinocloidea. Trapping within this zone took place from the 26-27 of May.

Terminalia Woodland

The Terminalia/Spirostachys woodland is the typical woodland found south of the Namutoni and Klein Namutoni woodland. The trapping site at in this locality was place on the Dik-Dik Drive. The geology of the woodland was a typical of the calcrete plains that extend south of the Pan. The main vegetation of the woodland was Terminalia prunioides, Spirostachys africanum, and Boscia foetida. No particular grass species were observed due to the nature of the woodland and the high utilization by grazers. Trapping within this area took place 26-27 of May.

Methods

Small mammal distribution was measured by means of live trapping sessions. The traps used (Sherman traps) were 7.6x7.6x25.4 cm in dimension. Each trapping sites, placed within the various habitats consisted of one transect line made up of 100 Sherman traps; each were spaced five metres from each other within the transect line. The only instances where the transect was not always a straight line, was at the depressions trapping sites (necessitated by the lack of extensive space of the sampling area), in which case the transect line veered off to another clear depression zone, and where the transect line at the hillocks ran in a loop parallel to the road. For the purpose of noting the exact position of the trapping sites, a GPS was to mark the beginning and ending points of each transect line. The bait used for the traps was created to be suitable for both herbivorous and insectivorous species and consisted of peanut butter, oats, sunflower oil and Bovril (Avenant, 2000). Re-baiting took place every morning and every evening, for the possibility of diurnal and nocturnal species. Trapping periods were planned for the middle of each month starting in February and ending in May; and on average, the traps were set for three trap nights (a trap night being defined as a 24 hour period) (Avenant, 2000; Golley, Petrusewicz, & Ryszkowski, 1975). The traps were checked twice per trap night (morning and afternoon) due to the possibility of catching both diurnal and nocturnal species. The Dolomite hills and the open plains were the only two trap sites that had a period of 4 trap nights. Okaukuejo, the Pan edge, the plain steppes, and the water depressions had a total of 3 trap nights in which small mammals were trapped. The hillocks and the Terminalia woodland were the only two sites that received a total of two trap nights due to time constraints. Each small mammal captured was identified, sexed, and then marked by means of a fur trimming for the purposes of mark-recapture (Cameron, 1996; Seber, 1982). For the purposes of determining the diversity of small mammals at each site, the Shannon Information Index (Shannon & Weaver, 1948) was used to calculate diversity based on the species richness and the number of small mammals caught at each site. Using the Shannon Information Index (Shannon & Weaver, 1948), relative species diversity can be calculated for each site and can also be subsequently used to compare the habitat diversity of the each habitat. All calculations are based on the number of species and the abundance of each species. If one or more species of a particular dataset have the highest abundance and the other species tend to be more obscure and

7

rare then the calculations of the Index will bring the Shannon entropy closer to 0. If there is only one or no species in the data set then the Shannon entropy will equal exactly 0. For the purpose, of analysing the data statistically for any significant difference, the acquired data was tested for normality using the Shapiro-Wilks W Test, and as the data was not normally distributed, the Kruskal-Wallis Test for non-parametric data was used to test for significant differences between the sites based on the habitats and abundance or the habitats and species richness. A statistically significant level of 95% (p<0.05) was used for the data analysis. All the data gathered were analysed statistically (Shapiro-Wilks W Test and Kruskal Wallis Test) using Statistica for Windows version 10 (StatSoft Inc., 2011) (Hauptfleisch & D’Alton, 2015).

8

Results

Overall Results In total 259 individual small mammals from 11 species were captured across all habitats over the course of the study period. The highest number of total

trapped individuals was collected at the Dolomite hills, with a total of 105 individuals of 3 species (Table 1). The trapping site with the highest total species

richness was within the boundaries of the Okaukuejo rest camp, with a total of 9 species being recorded. The lowest number caught as well as the lowest

recorded species richness occurred during the sampling on the plain steppes, with nothing being caught in any of the trap nights. All species sampled were

nocturnal, except for Elephantulus intufi (crepuscular to diurnal) (De Graaff, 1981) which was only caught at the camp. The species and the habitats in which

they were caught are summarized in Table 1.

Table 1: Overall number of small species captured at each trapping site.

Habitat Aethomys chrysophilus

Aethomys namaquensis

Mastomys coucha

Malacothrix typica

Saccostomus campestris

Mus indutus

Thallomys paedulcus

Gerbilliscus leucogaster

Desmodillus auricularis

Crocidura fuscomurina

Elephantulus intufi

Total Number Trapped*

Species abunda-nce**

Species richness

Dolomite Hills

0 83 21 0 0 1 0 0 0 0 0 105 72 3

Open Plains 0 0 1 1 1 9 0 0 0 0 0 12 12 4

Okaukuejo 28 13 10 0 0 2 3 29 1 3 5 94 87 9

Pan Edge 2 0 0 2 0 0 0 1 8 0 0 13 12 4

Plain Steppes

0 0 0 0 0 0 0 0 0 0 0 0 0 0

Water Depressions

1 0 0 0 0 0 0 0 0 0 0 1 1 1

Hillocks 4 0 6 0 0 0 0 14 0 0 0 24 21 3

Terminalia Woodland

1 0 8 0 0 0 0 1 0 0 0 10 9 3

Total 259 214

*Number of individuals caught over the fixed trapping nights (retrapped individuals included) **Number of individuals caught over the fixed trapping nights (retrapped individuals excluded)

9

Trap success and abundance within each habitat

Figure 3: Number of small mammals trapped within the assigned trapping periods at each study site.

Figure 4: Species abundance between the habitats.

As can be seen in Figure 3, the Dolomite hills had the highest trap success of small mammals within the entire project period with a total of 105 being caught, but produced 33 retraps, making it the second most abundant habitat (Figure 4). Okaukuejo itself had the highest abundance where a total of 94 small mammals were trapped within the study period. Though the hillocks had a trapping period of only two nights, it had the third highest trap success with a total of 24 small mammals being caught, and even with 3 retraps, the hillocks still have the third highest abundance with a total of 21 small mammals. The Pan edge, open plains and Terminalia woodland were almost similar in terms of numbers trapped with a total of 13 (including 1 retrap), 12, and 10 (including 1retrap) being caught at each site respectively. The lowest species abundance and trap success came from both

0

20

40

60

80

100

120

Ind

ivid

ual

s ca

ugh

t

Habitats

0

20

40

60

80

100

120

Ab

un

dan

ce

Habitats

10

the water depressions and the plain steppes where only one small mammal caught at the water depressions and nothing was caught on the plain steppes.

Figure 5: Mean and confidence intervals of the habitats according to trap success based on the Kruskal-Wallis Test.

There was only a significant difference in trap success (p=0.031) between the plain steppes and

Okaukuejo. However, the statistical software, could not make a direct comparison between the

between half of the sites as some varied in the number of trapping nights. Since, the Dolomite hills

and the open plains had a total of four trap nights and since the hillocks and Terminalia woodland

had a total of two trap nights, they cannot be tested for significance with the rest of the habitats.

Small mammal species richness within each of the habitats

Figure 6: Species richness of each habitat.

As can be seen in the Figure 6, the highest species richness occurred in the Okaukuejo rest camp area with a total of 9 different species being trapped over the study period. The open plains and the

0123456789

10

Spe

cie

s ri

chn

ess

Habitats

11

Pan edge came second in terms of species richness as both had 4 species trapped over the study period. The hillocks, the Terminalia woodland and the Dolomite hills all had a species richness of 3. The plain steppes had the lowest species richness (0) and the water depressions only had a species richness of 1.

Figure 7: Mean and confidence intervals of the habitats according to species richness based on the Kruskal-Wallis Test.

Once again, as can be seen in Figure 7, only a significant difference (p=0.020) in the species richness between the plain steppes and Okaukuejo was found, influenced by the same limitations regarding trapping nights.

Small Mammal Diversity As can be seen in Table 2, the highest Shannon value is within Okaukuejo and the lowest in the plain steppes and the water depressions. Table 2: Calculated Shannon entropy for the habitats of the Park.

Habitat Shannon entropy

Dolomite Hills 0.55

Open Plains 0.88

Okaukuejo 1.7

Pan Edge 1.1

Plain Steppes 0

Water Depressions 0

Hillocks 0.96

Terminalia woodland

0.64

Species caught The most common species found across all eight study sites was Mastomys coucha, which although it did not dominate in any of the habitats except for the Terminalia woodland, was found in almost all of the habitats except for the plain steppes, the water depressions and the Pan edge. The second most common species caught were Aethomys chrysophilus which is the second abundant species in

12

Okaukuejo after Gerbilliscus leucogaster, and is found all the habitats except the Dolomite hills, the open plains, and the plain steppe. Another common species found within the study period was Gerbilliscus leucogaster which dominated in Okaukuejo and the hillocks. Aethomys namaquensis dominated on the Dolomite hills, which along with lower species richness and the lesser abundance of the other two species, contributed to the lower diversity within the Dolomite hills. The only insectivorous species found within the study period were Crocidura fuscomurina and Elephantulus intufi, which were only found in the Okaukuejo trapping site.

Discussion All species found were within their expected distribution range (Apps, 2000; Skinner & Smithers, 1990). However, species that were described previously in the area and that are recorded within the Park were not found within study period. Possibilities for these differences are most likely the result of seasonality, habitat variance, and the trapping methods (Golley et al., 1975) used during the course of the project. As indicated in previous studies (Baraoidan & McCleery, n.d.), the amount of Horizontal Visual Obstruction (HVO) plays a role in the distribution and abundance of small mammals. This is by providing shelter and protection from predators, but the abundance of grass in the study sites should not be overlooked as it too provides a considerable percentage of the HVO within the savannahs. However, most of the sites in which the sampling sites were placed had a considerable amount of grass coverage or relatively average tree canopy coverage. Though, the only sites that were observed to have effectively little to no grass cover were: the plain steppes, the water depressions and the Terminalia woodland. Species abundance that was determined within the study period has given a relatively good indication of the habitat health, but it has also been primarily due to the influence of specialised ecological niches(van As, du Preez, Brown, & Smit, 2012) at the sample sites. As a part of research undertaken in the Namib Desert the effect of the mountainous and rocky outcrops (the rock habitat itself provides shelter) can give rise to specialised niches which additionally increases the availability and predictability of food, improvement of climatic variables and, in some cases, a resource that can be defended by the mammals (Mares, 1997). This effect can be seen in the number of Aethomys namaquensis trapped at the Dolomite hills. Habitat preferences also formed an explanation to the presence of Gerbilliscus leucogaster at Okaukuejo, the Pan edge, the hillocks and the Terminalia woodland, as the presence of sandy soils creates a suitable environment for it to burrow (Apps, 2000; De Graaff, 1981), as can also be said for Desmodillus auriclaris (favouring sandy to clayey soils), the dominant species found at the Pan edge. The primary reason for the high species richness and high abundance (which is the reason for it also having the highest species diversity) for the small mammals found at Okaukuejo could mainly be due to the microhabitat diversification within in the boundaries of the rest camp. The habitat was ranged from bare sandy areas, to grassland, to rubble strewn areas, and finally to the verge of woodland. The sandy soils described in the northern part of the transect line accounts (Apps, 2000; De Graaff, 1981) for the habitat preferences of the habitat species: G. leucogaster, E. intufi , and D. auricularis. The more rocky terrain replicated by the rubble strewn areas created the environment suitable for A.namaquensis, as was seen in the Dolomite hills. Certain species of small mammals have a wide tolerance for a range of habitats, and were therefore found in more than one habitat. Species such as Mastomys coucha and Aethomys chrysophilus have

13

been described to having no particular habitat preference (Apps, 2000; De Graaff, 1981; Meester, Rautenbach, Dippenaar, & Baker, 1986), which is why each were found in 5 different habitats. In terms of the species diversity, the lower species diversity within the Dolomite hills, the open plains and the Terminalia woodland can be attributed to the high species abundance of just one species at each study site. The high abundance of A. namaquensis at the Dolomite hills, Mus indutus at the open plains, and M. coucha at the Terminalia woodland was the primary cause for the lower species diversity. The higher species richness in Okaukuejo, the higher species richness and relative abundance at the Pan edge and the relatively equal abundance of species at the hillocks contributed to a higher species diversity amongst the three sites. With regard to the diurnal species, it is of key interest that none were actually found in any of the sample sites. The study area is within the distribution range of the diurnal species E. intufi and Crocidura fuscomurina (Apps, 2000; Meester et al., 1986; Skinner & Smithers, 1990). Three reasons for the lack of diurnal species can be hypothesized. Firstly, as it has been discovered elsewhere, the presence of raptors, of which there was an abundance in ENP, can play a defining role in the presence of diurnal small mammals (Matos et al., 2015). Secondly, the poor vegetation coverage (Baraoidan & McCleery, n.d.), along with the presence of raptors, could have a difference on the small mammals presence and its ability to escape from predators. Thirdly, with the rainfall being of a particular drought-like nature (the rainfall amount recorded for Okaukuejo during the rainy season amounted to 258 mm opposed to the average rainfall of 350mm) (S. Kötting, personal communication, 2015), and with no respite from the daytime temperatures experienced in ENP, it reasonable to assume that diurnal small mammals would absent due to the difficulty of thermoregulation experienced in small mammals (McKechnie & Mzilikazi, 2011). Notwithstanding, the rainfall pattern of the year 2015 played a considerable role in the trapping of the small mammals. The rainfall that was below average and that was scattered in its distribution for most of the rainy season created a rather sparse or slowed regeneration of grass species for most of the Park (as above). This was felt mostly in areas such as the plain steppes and the water depressions (trapping sites that were set up just after the end of the rainy season), that due to the continued use of the Okaukuejo waterhole for the benefit of the tourism industry (S. Kötting, personal communication, 2015), continue to exist in an overgrazed state. This continued grazing impact (Grant, Birney, French, & Swift, 1982; Martin & Dickinson, 1985), along with the influences of the dry season, are hypothetically the primary reasons as to why there were hardly any small mammals discovered at the plain steppes and the water depressions. Furthermore the personal observations that were made when baiting of the trapping sites, were that the population of black-backed jackal in the Okaukuejo area were unsettling the sites by breaking into the traps to eat the bait inside. The importance of certain species found within the trapping sites cannot be overlooked. M. coucha, being the most widespread species within the Park, is considered an important prey species, and often falls prey to raptors and small carnivores (Apps, 2000). G. leucogaster, also being a seemingly common species within the Park adds a valuable ecological service in terms of the bioturbation (soil-reworking) it is responsible for (M. Hauptfleisch, personal communication, 2015). Small mammals also serve as a disease vectors, as is observed in M. coucha, a carrier of bubonic plague and bilharzia, and G. leucogaster, a carrier of bubonic plague (IUCN, 2015).

Conclusions The study provided an important update of the knowledge of small mammals in Etosha. With the last extensive study having been done in1992 (Griffin, 1993), the data will be submitted to the

14

Namibia mammal atlas project to add to the knowledge of small mammals in Namibia as a whole. The atlas is an initiative to update the current information regarding small mammals in Namibia and to explore the distributions of small mammals across Namibia. Small mammals are a vital component within the ecosystem of the ENP, based on their roles as primary and secondary consumers and their importance as a source of food for many predator species. As has been ascertained from the study period, each habitat sampled at each study site has a variance in distribution and abundance of small mammal species. Overall, there is a significant difference in the amount of small mammal species and their distribution within the various habitats of Etosha. Each habitat being variable to all the others creates an environment suitable for a wider variety of small mammals, which ultimately aids in increasing the chances of a higher biodiversity. As can be seen, Okaukuejo and the Dolomite hills support a high species abundance when compared with the other habitats, with Okaukuejo additionally having the highest species richness and diversity. An information poster can now be developed in the future for the EEI, detailing the results of the project and to create further awareness within the Park on the importance of small mammals.

Recommendations The project was a short term inventory, and in order for it to be more meaningful, it should be continued till a clearer and much more certain species representation of the Etosha area can be gathered. Due to the error of not being able to sample at certain habitats for the intended length of time and due to the large percentage of species that were not numerous enough for the purposes of mark-recapture, no comparisons were able to have been made in terms of the population sizes of all the habitats. Due to the nature of the year, most of the area around Okaukuejo did not specifically benefit from a particularly abundant rainy season. This year, effectively, is considered a below average rainfall year and this itself might have influenced the species found at the trapping sites at the time they were sampled. For a clearer representation of the species richness and distribution within the Park habitats to be made, it could be recommended that the research project be replicated within the timespan of a more favourable rainy season. Secondly, the research project should be inclusive of the Park on a larger scale. Different habitat areas, not specifically within the vicinity of the Okaukuejo rest camp, should be sampled across the Park.

Acknowledgements A special thanks to Dr Morgan Hauptfleisch for his recommendation in doing this research project and for his guidance and assistance over the course of the study period, and also for supplying the bulk of the live traps needed for the project. Also, thanks to Martin Kasaona for also supplying a considerable number of Sherman traps. A special thank you to the staff members Thomas Amadhila, Jonson Vejorerako, and Pieter Aukhumeb for providing the transport and for assisting with the actual setting up and sampling and for the protection provided. Also, thank you to Andrea Capobianco Dondona for allowing me to travel with him to set up the last trapping sites near Namutoni. A special thanks to my mentor Shayne Kötting for his assistance and support during the project period, and for his efforts to arrange for transport and for arranging that the project go through the correct procedures.

15

References

Apps, P. (Ed.). (2000). Smithers’ mammals of southern Africa: A field guide. Cape Town, South Africa:

Struik Nature.

Avenant, N. L. (2000). Small mammal community characteristics as indicators of ecological

disturbance in the Willem Pretorius Nature Reserve, Free State, South Africa. South African

Journal of Wildlife Resources, 30(1), 26–32.

Baraoidan, S. A., & McCleery, R. A. (n.d.). Habitat preferences and biodiversity of small mammals in

southern African savannahs. University of Florida, Gainesville.

Barnard, P. (Ed.). (1998). Biological diversity in Namibia: A country study. Windhoek, Namibia:

Namibian National Biodiversity Task Force.

Berry, H. (1995). Origin and meaning of place names the Etosha National Park, Namibia.

Berry, H. (2013, August 20). Etosha - Creatures great and small. Travel News Namibia. Retrieved

from http://travelnewsnamibia.com/news/etosha-creatures-great-and-small/#.VLfoGS5biyg

Berry, H. H. (1980). Behavourial and eco-physiological studies on blue wildebeest (Connochaetes

taurinus) at the · Etosha National Park (PhD). University of Cape Town.

Cameron, A. (1996). An investigation into small mammal community composition on Mount Elgon

with reference to habitat modification by humans (BSc. Thesis). Aberdeen University,

Scotland, UK. Retrieved from http://www.see.leeds.ac.uk/misc/elgon/mammal1.html

Carleton, M. D., & Musser, G. G. (Eds.). (2005). Order Rodentia. In Mammal species of the world: A

taxonomic and geographic reference (Vol. 12, pp. 745–752). JHU Press.

Cunningham, P. L., & Jankowitz, W. (2011). A review of fauna and flora associated with coastal and

inland saline flats from Namibia with special reference to the Etosha Pan. Sabkha

Ecosystems, 3, 9–17.

Danell, K., & Aave-Olsson, B. (2002). Endemic mammalian genera: are they really unique?, (29), 457–

464.

16

David, J. H. M., & Jarvis, J. U. M. (1983). Notes on two brief surveys of the small mammal fauna on

the Rooiberg, Ladismith, southern Cape Province. South African Journal of Zoology, 18(4),

370–377.

De Graaff, G. (1981). The rodents of southern Africa. Pretoria, South Africa: Butterworth & CO.

Dilley, D. (1996). The diet of a Barn owl, Tyto alba, in Okaukuejo Etosha National Park, Namibia. Cape

Town, South Africa: Cape Technicon.

Foresman, K. R., & Badyaev, A. V. (2005). Developmental instability and the environment: Why are

some species of shrews better indicators of stress than others? Special Publication of the

International Society of Shrew Biologists, 1, 265–272.

Golley, F. B., Petrusewicz, K., & Ryszkowski, D. W. (1975). Small mammals: their productivity and

population dynamics. International Biological Programme, 25–53.

Grant, W. E., Birney, E. C., French, N. R., & Swift, D. M. (1982). Structure and productivity of

grassland small mammal communities related to grazing-induced changes in vegetative

cover. Journal of Mammalogy, 63(2), 248–260.

Griffin, M. (1993). Notes on smaller mammals from the Etosha National Park. Windhoek, Namibia.

Hauptfleisch, M. L., & D’Alton, C. (2015). Arthropod phototaxis and its possible effect on bird strike

at two Namibian airports. Applied Ecology and Environmental Research, 13(4), 957–965.

IUCN. (2015). The IUCN red list of threatened species. Retrieved June 13, 2015, from

http://www.iucnredlist.org/details/21516/0

Mannheimer, C. A., & Curtis, B. A. (2009). Le Roux and Muller’s field guide to the trees and shrubs of

Namibia. Windhoek, Namibia: Macmillan Education.

Mares, M. A. (1997). The geobiological interface: Granitic outcrops as a selective force in mammalian

evolution. Journal of the Royal Society of Western Australia, (80), 131–139.

Martin, G. H. G., & Dickinson, N. M. (1985). Small mammal abundance in relation to microhabitat in

a dry sub-humid grassland in Kenya. African Journal of Ecology, 23, 223–234.

17

Matos, M., Alves, M., Ramos Pereira, M. J., Torres, I., Marques, S., & Fonseca, C. (2015). Clear as

daylight: analysis of diurnal raptor pellets for small mammal studies. Animal Biodiversity and

Conservation, 38(1), 37–48.

Matson, J. O., & Blood, B. R. (1994). A report on the distribution of small mammals from Namibia.

Zeitung Säugetierkunde, 59, 289–298.

McKechnie, A. E., & Mzilikazi, N. (2011). Heterothermy in Afrotropical mammals and birds: A review.

Integrative and Comparative Biology, 51(3), 349–363.

Meester, J. A. J., Rautenbach, I. L., Dippenaar, N. J., & Baker, C. M. (1986). Classification of southern

African mammals. Pretoria, South Africa: Transvaal Museum.

Schlitter, D. A. (2005). Order Macroscelidea. In Mammal species of the world (3rd ed., pp. 82–85).

USA: John Hopkins University Press. Retrieved from

http://www.google.com.na/books?id=JgAMbNSt8ikC&pg=PA82&redir_esc=y#v=onepage&q

&f=false

Seber, G. A. F. (1982). The estimation of animal abundance and related parameters (2nd ed.).

Caldwel, New Jersey: Blackburn Press.

Shannon, C. E., & Weaver, W. (1948). A mathematical theory of communication. The Bell System

Technical Journal, (27), 379–423 and 623–656.

Skinner, J. D., & Smithers, R. H. N. (1990). The mammals of the southern African subregion (2nd ed.).

Pretoria, South Africa: Univeristy of Pretoria.

Van As, J., du Preez, J., Brown, L., & Smit, N. (2012). The story of life and the environment: an African

perspective. Cape Town, South Africa: Struik Nature.

Wilson, D. E., & Reeder, D. M. (2011). Animal biodiversity: An outline of higher-level classification

and survey of taxonomic richness, (3148), 56–60.