Dissertations & Theses

Temminck’s ground pangolin is a myrmecophagous mammal and the only pangolin speciespresent in South Africa. The ground pangolin employs a bipedal gait and can roll up into aball when threatened by predators. The thoracic limbs display powerful claws that are usedto dig for prey species and excavate previously occupied burrows. All eight pangolin species, including Temminck’s ground pangolin, are listed on the International Union forConservation of Nature (IUCN) Red List of threatened species and are cited in Appendix I ofthe Convention on International Trade in Endangered Species (CITES) of Wild Fauna andFlora. The scientific literature on pangolin species is modest when compared to many other mammalian species and has focused mainly on their physiology and ecology. Anatomicalstudies on pangolins have mainly focused on their peculiar gastro-intestinal tract and verylittle information is available on the locomotor system. This study describes the radiologicalanatomy as well as the gross osteology, myology and arthrology of the thoracic limb ofTemminck’s ground pangolin. The aim of this study is to address the scarcity of informationavailable on the locomotor system of this threatened species. Five Temminck’s ground pangolins (3.0 kg; 3.2 kg; 7.0 kg; 7.3 kg; 8.3 kg) that had died from anthropogenic threats such as poaching or electrocution from electrical fences were collected and frozen. The frozen bodies were thawed to facilitate manipulation of the thoraciclimbs for radiographic imaging in situ. The specimens were skinned and the extrinsicmuscles described. The extrinsic muscles were transected and the thoracic limbs removedand immersion-fixed in 10% neutral-buffered formalin. The bodies were returned to the African Pangolin Working Group. Some of the disarticulated, skinned thoracic limbs wereradiographed before further investigation. The muscles of the thoracic limbs were identified and described using basic gross anatomical techniques, photographed and then removed to reveal the different joints of the thoracic limbs which were also dissected, described and photographed. After most of the soft tissue had been removed, the limbs were placed in a muslin cloth bag and boiled for at least 72 hours. The individual bones were then studied, described and photographed. The sites of origin and insertion of the various muscles associated with the thoracic limbs were identified on the individual bones. The skeleton of the thoracic limbs revealed a similar number and arrangement of bones compared to that of domestic carnivores. The bones were robust, numerous epiphyseal lines were open and many of the tubercles were well-developed. The scapula was broad and triangular-shaped. The humerus displayed a massive Epicondylus medialis. The ulna and radius were similarly sized, and displayed a large Tuber olecrani and broad Trochlea radii, respectively. The radius was shorter in length compared to the humerus. The Manus was composed of seven carpal bones, five short metacarpal bones and five digits of which the three central digits were the best developed. The Processus unguicularis of the distal phalangeal bones was bifid and elongated. The thoracic limb myology of Temminck’s ground pangolin revealed well-developed muscles with fleshy bodies and short tendons. The extrinsic muscles of the thoracic limb, in particular the M. cutaneus trunci, were prominent. Two novel muscles, which function as protractors of the thoracic limb, the M. omocephalicus and M. omomastoideus, were described. The extensor muscles of the elbow joint were well-developed. The largest intrinsic muscle of the thoracic limb, the M. triceps brachii, presented a Caput longum with four bellies which attached extensively to the scapula. A novel elbow extensor, the M. anconeus medialis was named and described. Rotators of the Antebrachium, the M. pronator teres and M. supinator, were well-developed and displayed extensive sites of insertion on the radius only. The M. flexor digitorum superficialis displayed a variation in the site of origin of the four individual muscle bellies that each inserted on a separate digit (II-V), except for digit I. The M. flexor digitorum profundus displayed an extensive area of origin as well as additional muscle bellies. The muscle bellies fused into a hard tendon plate over the palmar carpal area. Distal to the carpus the tendon divided into five individual tendon slips that each inserted on an individual digit. The extensors of the shoulder joint, flexors of the elbow joint and extensors of the carpal and digital joints followed the basic mammalian pattern. The thoracic limb joints of Temminck’s ground pangolin displayed shallow articular surfaces for the shoulder, elbow, carpal and carpometacarpal joints. The articular surfaces of the metacarpophalangeal, proximal and distal interphalangeal joints, however, displayed interlocking ridges and grooves. A number of the joint capsules acted as either sites of origin or insertion for a variety of muscles. Additionally, some muscles spanning joints were fused to the joint capsule. Articular ligaments were generally not well-developed and were represented mainly by thickenings of the joint capsule. The osteological characteristics indicated that Temminck’s ground pangolin is specifically adapted to protraction and retraction of the thoracic limb, strong elbow extension, flexion of the carpus and digits as well as pronation and supination of the antebrachium. The morphology of the muscles supported the observation that Temminck’s ground pangolin is non-weight bearing in the thoracic limbs, is adapted to roll into a ball if threatened by predators and is adapted to perform a variety of functions. These functions include powerful retraction of the thoracic limbs, extension of the elbow joint, simultaneous flexion of the carpal and digital joints as well as moderate supination and pronation of the Antebrachium. The morphology of the joints and ligaments suggested that Temminck’s ground pangolin does not experience major weight-bearing forces on its thoracic limbs but rather rotational forces of the proximal limb and impact forces in the distal limb due to digging in hard substrates. The thoracic limb of Temminck’s ground pangolin therefore displayed adaptations to both non-weight bearing and to scratch-digging.

A rise in African pangolin confiscation during trafficking prompts the need for knowledge of parasite loads, especially tick species, and this can contribute to treatment during rehabilitation. Ticks are external parasites that live on the skin of animals thus it is important to identify the species because of the different effects they can have on animals. To investigate the latter identification based on morpho-anatomical characteristics confirmed the presence of three tick species, Ornithodoros compactus and Rhipicephalus theileri identified from Temminck’s Ground Pangolins, while Amblyomma compressum was identified from samples collected from the White-bellied and Black-bellied Pangolins. The ticks collected from confiscated scales included an admixture of different species which were identified as Amblyomma species, Amblyomma compressum, Amblyomma hebraeum, and Rhipicephalus simus. To validate identification using molecular techniques, various molecular markers targeting different mitochondrial and nuclear gene regions were optimised for the identification of ticks in the study. The 16S gene region was used as the target gene for tick identification as this region had a large number of corresponding registered sequences on GenBank and demonstrated a high variation among species. In addition, the phylogenetic analysis based on this gene region could distinguish between the soft and hard ticks in the collection.

Pangolins (Pholidota: Manidae) form an important part of local cultures and traditions in the majority of central and west African countries. The use of whole pangolins or body parts for medicinal purposes in Africa is long-established and they are also hunted as a source of bushmeat across the continent. However, very little is known about the cultural importance of pangolin body parts that are used to treat, alleviate or cure particular ailments and diseases as well as the level of trade for bushmeat purposes in west Africa. In addition, the level of local knowledge relating to pangolins and how it can be integrated with scientific ecological knowledge to contribute to implementation plans and conservation management actions for pangolins in the forests of west Africa has not been determined. The aim of this study was to determine the traditional medicinal importance of pangolins to local communities, pangolin level of trade for bushmeat purposes and local ecological knowledge amongst collectors of pangolins. Using purposive and snowball sampling for the selection of participants, data were gathered on traditional medicinal use, level of trade for bushmeat purposes as well as level of local ecological knowledge of hunters using semi-structured interviews. Ethnozoological data on the use of pangolins for traditional medicinal purposes were collected from 63 and 48 traditional healers in Sierra Leone and Ghana, respectively. Data were gathered from 153 stakeholders comprising chopbar operators, farmer hunters and wholesalers to determine the level of trade in Ghana. Information on the level of local ecological knowledge of hunters was gathered from 126 and 153 farmer hunters in Sierra Leone and Ghana, respectively. Ethnozoological data were analysed using use value, cultural importance and informant agreement ratio indices. A total of 21 pangolin parts and tissue oil was found to be used to treat various ailments in Sierra Leone while 13 body parts were used to treat various ailments in Ghana. Pangolin scales were the most culturally important body part according to the rank at which this body part was used in both countries. The results revealed that pangolin body parts play a vital role in the primary healthcare of the people in both countries. A total of 341 pangolins were recorded to have been traded during the study period and, of this number, 98 pangolins were personally observed. Three species of African pangolin are known to occur in both countries. Of these 82% of pangolins traded were white-bellied pangolin (Phataginus tricuspis) while 18% where black-bellied pangolin (Phataginus tetradactyla), with no trade observed for the giant ground pangolin (Smutsia gigantea). Local ecological knowledge of farmer hunters revealed that they were familiar with two of the three African pangolin species that occur in Ghana and Sierra Leone, i.e. the arboreal black-bellied pangolin (P. tetradactyla) and white-bellied pangolin (P. tricuspis) but none reported hunting the giant ground pangolin (S. gigantea). Although in existence, current legislation has been unable to combat harvesting of pangolins for traditional therapeutic and dietary purposes. The Wildlife Conservation Act in both Ghana and Sierra Leone prohibits any person from hunting or being in possession of pangolins, however, enforcement is lacking. In addition, pangolins are listed in the Convention on International Trade in Endangered Species (CITES) as Appendix II and were up-listed as vulnerable on the recently revised International Union for Conservation of Nature Red List. It is highly likely that the unregulated harvesting and poaching of this threatened group of mammals for medicinal purposes and the bushmeat market, is completely unsustainable. A study to determine the population status and ecology of the three species of African pangolins occurring in both countries is urgently required in order to assess the impact of harvesting on the populations which is currently unmonitored and unsustainable. Within this context, there is an urgent need to determine pangolin population abundance within the region, and Africa in general, to ensure their survival.

Pangolins are medium-sized mammals, characterised by keratinous scales covering their entire body. There are currently eight species of pangolins left in the world with four occurring in Africa and four in Asia. All species of pangolin are currently considered vulnerable to extinction, with their numbers rapidly declining due to the excessive demand for pangolin scales from the illegal trade. This trade is often highly organised and lucrative, making it extremely problematic to counteract. These factors contribute to pangolins now being regarded as the most illegally trafficked mammals on Earth. Within the trade, pangolins are often kept in dire conditions and in cases where they have been recovered, they rarely survive the ordeal. For those that do survive, many assumptions are made when releasing them back into the wild, with only a few cases where actual monitoring has taken place post-release. This study aims to investigate: the demographics of Temmick’s pangolin which has been confiscated from the illicit trade in South Africa, the condition of the animals retrieved, survival between release strategies, and their distribution following release subsequent to veterinary treatment. Study animals were fitted with appropriate tracking equipment and monitored post-release to monitor movement and survival. Living pangolins confiscated in South Africa increased from 8 in 2016 to, 40 in 2018, but declined to 25 in 2019, with confiscations peaking during the cold and dryer seasons of winter and spring. Of the pangolins that were successfully confiscated, 63.24% were adult, 14.71% were sub-adult and 22.06% were juveniles. 68% of these pangolins were found in a compromised state. The soft-release approach has become the preferred method of release in South Africa, and current evidence suggest that it has a positive effect on pangolin survival, but more research is still required. Trade related stress was the main root of mortalities and a great cause for concern. Sex of animal played no significant role in susceptibility to poaching, or in survival. Post release monitoring revealed that released pangolins can travel vast distances, and this needs to be taken into consideration when selecting release sites and tracking equipment. Facilitated release procedures have proven to be effective in improving survival probability, but improvements can still be made in all aspects of recovery, treatment and release protocols.

Pangolins are evolutionarily distinct in that they are the only representatives of an entire mammalian order, the Pholidota. Temminck’s ground pangolin (Smutsia temminckii) is one of the four extant African pangolin species. It is a medium-sized mammal commonly called a scaly anteater and is of the family Manidae. They are considered to be very rare in South Africa and believed to be declining in most parts of their range and this species was recently up-listed to Vulnerable from Least Concerned by the International Union for the Conservation of Nature (ZSL & IUCN, 2014; IUCN, 2015). Pangolins are solitary animals that are revered in most South African communities because of their believed mystical powers and economic values. It is further believed that there is an extremely high demand for the species for traditional medicine purposes throughout southern Africa based on their perceived ability to cure a diverse number of ailments making use of the various pangolin body parts. It is therefore believed that the species is threatened as a result of this exploitation for traditional medicine purposes and other cultural beliefs. However, no studies have been carried out to document the various uses of the animal within tribal communities throughout its range in Africa. The aim of the study is to investigate the uses of the species for traditional, cultural and medicinal purposes and the potential these may have on the prevalence and distribution of this vulnerable species within its known distributional range within South Africa. In conjunction with this, this study aims to estimate the current need for the species within South African tribal communities and to document the associated uses of pangolin body parts to try and quantify this use as a potential threat to the species. Individuals, elders and traditional healers within tribal communities throughout the species known distributional range in South Africa were interviewed, namely in the Sepedi, Zulu, Tsonga, Venda, Tswana, SiSwati and Ndebele tribes. It was found that the use of the animal in the communities rode basically on the belief system of the people. The animals’ scales were the most frequent body part utilised to treat a wide array of ailments. Following this, the blood, fat, bones and meat were often prescribed for a particular disease or an ailment. Not only were pangolin body parts prescribed for medicinal purposes but often prescribed for protection against harm or to ward off evil spirits by either having a scale and/or blood on person or including the fat and/or blood into the building material of houses and livestock pens. In some instances, the meat is also considered a delicacy. In sourcing the animals, communities would either actively hunt pangolins or opportunistically make use of those that are found as road kills. In some cases, large sums of money are offered to source pangolins specifically for traditional medicine. Knowledge about the species was more prevalent among the elderly and generally the species was less well known amongst younger rural people. Nevertheless, the species is still regarded as an important and sought-after organism for traditional medicine in South Africa and, as such, traditional medicine can pose a significant threat to the species population status throughout its distributional range.

Ground pangolins Smutsia temminckii are inconspicuous, mainly nocturnal mammals that occur at low population densities. As a result, there is scant information available on the ecology and physiology of this species. To date the handful of studies focussing on this species were centred in the mesic eastern regions of its range, with no attention being given to ecological and biological traits in arid environments. To address these data shortfalls, a study was undertaken in the Kalahari Desert in north-western South Africa. Very High Frequency (VHF) transmitters or Global Positioning System (GPS) loggers were fitted to 16 ground pangolins and their ecology and physiology studied over a period of three years. Throughout its range the ground pangolin is increasingly endangered, predominantly due to anthropogenic threats. Previous studies have identified some threats facing this species, but very few of these studies included quantitative data as to the extent of these threats. The present study suggests that the main threats to ground pangolins in southern Africa are electrocution on electrified fences, the traditional medicine trade, habitat loss, road mortalities and capture in gin traps. Although accidental poisoning has previously been viewed as a threat, a review of the available literature suggests that this is not the case and that these views stemmed from two isolated incidents of captive animals. Electrocutions pose the greatest threat to ground pangolins, with mortality rates of one pangolin per 11 km of electrified fence per year recorded during this study. Home range sizes of this arid-zone population are comparable to the home range sizes recorded in mesic populations. Study animals in the Kalahari had Minimum Convex Polygon (MCP) home range values of 10.0 ± 8.9 km² SD for adults and 7.1 ± 1.1 km² SD for juveniles, in comparison to the 0.17 – 23.38 km² MCP in Zimbabwe and 1.3 –7.9 km2 MCP in mesic north-eastern South Africa. These results are surprising as arid-zone animals usually have larger home ranges than do their mesic counterparts. It is hypothesised that this discrepancy can be explained by ground pangolins in the Kalahari being on average 25 – 30 % smaller than their mesic counterparts. This smaller body size means that individuals require a smaller home range to meet their dietary requirements, which could be counteracting the trend of arid-zone individuals requiring larger home ranges.This research also found ground pangolins to be more active during the day in winter and nearly entirely nocturnal during summer. These changes in photoperiodicity are likely driven by adaptations to the extreme climatic conditions present at the study site rather than predation pressure or changes in food resources, although the latter may be playing a role as well. By being diurnal in winter, individuals avoid the extremely cold night-time temperatures, thus saving energy that would otherwise be spent on maintaining their body temperature. By contrast, becoming nocturnal in summer allows them to avoid the extreme daytime temperatures, thus conserving water. The results of this study reiterate previous findings that ground pangolins are entirely myrmecophagous and highly selective of the species that they eat. Study animals were recorded feeding on only four ant and two termite species, representing 7.5 % and 50 % of the total species richness of each of these families. This is similar to previous research in mesic savannahs where five ant one termite species constituted 97 % of the ground pangolin’s diet. No seasonal differences in prey selection were observed, although the species consumed did depend on the habitats that were available to individual ground pangolins. This study is the first to investigate the core body temperature (Tb) of a free-ranging pangolin. These data indicated that Tb fluctuated cyclically between 32 – 35 °C and reflected the activity periods of the animal, peaking when the animal was active. When the individual was inactive, Tb steadily decreased, suggesting that it used daily heterothermy to cope with the low food availability in this unpredictable environment.In summary, ecological parameters between arid and mesic ground pangolin populations are similar in some respects, but divergent in others. Arid-zone populations are smaller overall and show a higher degree of diurnal activity. Despite this smaller body size, home range values appear to be similar between arid and mesic populations. Although the prey species eaten by ground pangolins in the Kalahari differ from prey species in eastern South Africa and Zimbabwe, they belonged to the same genera and were of similar morphology and ecology. The core body temperature results confirm findings from previous captive studies that pangolin body temperatures are lower than those of other eutherian mammals of a similar size. Furthermore, these data indicate that core body temperature is a lot more variable than previously believed, and thus warrants further study. Overall ground pangolin densities in the Kalahari appear to be double those in eastern South Africa and Zimbabwe, although it is unclear whether this is genuine or whether it is an artefact of individuals being easier to locate in the Kalahari. This study also confirmed that electrocutions are probably the greatest threat to this species, and new threats such as accidental capture in gin traps and road mortalities have also come to light.