Cheetah (Acinonyx jubatus)- Data, Pictures & Videos

[Acinonyx sp.]

Using GPS collars to investigate the frequency and behavioural outcomes of intraspecific interactions among carnivores: A case study of male cheetahs in the Maasai Mara, Kenya

Abstract
Intraspecific interactions between individuals or groups of individuals of the same species are an important component of population dynamics. Interactions can be static, such as spatial overlap, or dynamic based on the interactions of movements, and can be mediated through communication, such as the deployment of scent marks. Interactions and their behavioural outcomes can be difficult to determine, especially for species that live at low densities. With the use of GPS collars we quantify both static and dynamic interactions between male cheetahs (Acinonyx jubatus) and the behavioural outcomes. The 99% home-ranges of males overlapped significantly while there was little overlap of the 50% home-ranges. Despite this overlap, male cheetahs rarely came into close proximity of one another, possibly because presence was communicated through frequent visits to marking posts. The minimum distance between individuals in a dyad ranged from 89m to 196m but the average proximity between individuals ranged from 17,145 ± 6,865m to 26,367 ± 11,288m. Possible interactions took place more frequently at night than by day and occurred mostly in the 50% home-range of one individual of a dyad or where cores of both individuals overlapped. After a possible encounter male cheetahs stayed in close proximity to each other for up to 6 hours, which could be the result of a territory defence strategy or the presence of a receptive female. We believe that one of the encounters between a singleton and a 5-male coalition resulted in the death of the singleton. Our results give new insights into cheetah interactions, which could help our understanding of ecological processes such as disease transmission.

Introduction

Intraspecific interactions, or interactions between members of the same species, are an important component of population dynamics as they play a role in sociality [1], mating events [2], disease transmission [3, 4] and competition which could influence access to resources [5, 6], spatial organisation [7, 8] and mortality [9]. Interactions, which can be defined as “actions directed towards, or affecting, the behaviour of another animal” (sensu [10]), can be categorised into two groups; static or indirect interactions and dynamic or direct interactions [11]. Static interactions lack a temporal element and do not take into account the proximity between individuals. For example, individuals could use similar areas, but at different times. Static interactions can be determined by quantifying the amount of spatial overlap, which can give an indication as to the possibility of dynamic interactions occurring [12]. Unlike static interactions, dynamic interactions (also referred to as encounters or associations) include a temporal component and are based on the spatial proximity of simultaneous locations of individuals. The nature of dynamic interactions can differ and can include mating events or the sharing or defending of resources [2, 13]. Interactions can be mediated through communication, for example olfaction or vocalisation, as this can attract mates or allow conspecifics to assess potential threats which could minimise the occurrence of costly, potentially fatal, encounters [14–16]. When interactions occur they can elicit a change in movement and spacing behaviour, which can vary depending on the nature of the interaction [17, 18].

The likelihood, frequency and outcomes of interactions can be influenced by social structure which can differ significantly amongst species [19]. Felids, for example, are often considered to be predominantly solitary [20] yet the sociality of felids lies on a continuum with lions (Panthera leo), which live in social groups, on one end of this spectrum [21]. Even felid species that are often believed to be solitary can engage in social interactions (e.g. [13]). In some species these associations occur occasionally, such as at kills sites [13], whereas in other species these can be more enduring. In cheetahs (Acinonyx jubatus), for example, females are solitary, unless they are accompanied by dependent cubs, but male cheetahs can either be solitary or form stable, same-sex groups known as coalitions [22]. Coalitions generally consist of two to three related or unrelated individuals, but a rare five-male coalition has been seen in the Maasai Mara, Kenya (this study). The land tenure system of male cheetahs can broadly be categorised into two groups: floaters, who roam over vast areas that they do not defend, and resident males, who defend small territories possibly based on access to resources such as females [22, 23]. Territorial boundaries can however be fluid [24] and it is believed that cheetahs use a ‘time-share’ approach [25] where territories and home-ranges can overlap but where direct interactions between cheetahs are minimised through olfactory communication. Territorial males advertise their presence by scent marking (urinating and defecating) on marking posts which are usually prominent landscape features such as termite mounds, logs or trees [23, 26]. Despite cheetahs communicating their presence, males can encounter one another and encounters can range from passive [25] to acutely aggressive [27].

Until recently, research on the spatial organisation of male cheetahs has mostly been based on VHF telemetry or behavioural observations (e.g. [22, 27, 28]). These methods of data collection, while informative, make it difficult to continuously monitor several individuals at a time. In addition, while encounters between males have been observed, it is unknown how often they occur and what the behavioural outcomes are [25, 27]. This paucity of data is partly because cheetahs live at low densities and interactions are therefore difficult to observe. However, with the help of data loggers, such as GPS collars, it is possible to detect and quantify interactions when multiple individuals are tagged simultaneously (e.g. [17]).
Here we investigate interactions between male cheetahs using location data collected with GPS collars by investigating 1) static interactions by quantifying spatial overlap and visits to marking posts to determine the frequency of indirect interactions to try and understand the role that marking posts play in cheetah ecology, 2) dynamic interactions by quantifying the proximity between different individuals and 3) the outcomes of possible interactions in terms of movement behaviour and mortalities. Based on previous research we predict that males will overlap spatially but that there will be little overlap of the core areas [29]. We also predict that marking posts are frequently visited by both individuals in a dyad, i.e. pair of cheetahs, and that occasions where individuals of the dyad are in close proximity to each other are infrequent. Because encounters between males can be aggressive [27] we predict that the movement behaviour after a possible encounter would indicate avoidance behaviour (moving away from the encounter location, moving away from one another, increased distance travelled and decreased path tortuosity).


Methods

Study area
The study was conducted in the Maasai Mara, in the southwest of Kenya (centred at 1°S and 35°E), which is part of the larger Serengeti-Mara ecosystem. The study area (~2,600 km2) included the Maasai Mara National Reserve and the surrounding wildlife conservancies. The area experiences one wet season spanning from November to June and one dry season spanning from July to October [30]. After the wet season, the long grass attracts large numbers of migratory ungulates, including the white-bearded wildebeest (Connochaetes taurinus) and the common zebra (Equus quagga), from the Serengeti in Tanzania. Throughout the year there is an abundance of cheetah prey including resident white-bearded wildebeest, Thomson’s gazelle (Eudorcas thomsonii), Grant’s gazelle (Nanger granti) and impala (Aepyceros melampus) [31]. The habitat in the Maasai Mara varies, ranging from open grasslands and shrubland, to riverine forests found along the major rivers and their tributaries [32]. The open grassland plains, which are dominated by red oat grass (Themeda triandra), are mostly found toward the south and west of the study area, while the north and north-east consist mostly of Croton thickets (Croton dichogamous) and Vachellia woodlands (Vachellia drepanolobium and V. gerrardii).
Cheetah collaring
Global Positioning System (GPS) satellite collars (African Wildlife Tracking - www.awt.co.za) were fitted on four adult male cheetahs between 19th October 2016 and 9th February 2018. In compliance with Kenyan law, all immobilizations for deployment/removal of collars were performed by a Kenya Wildlife Service veterinarian. Cheetahs were free-darted and immobilized using a combination of ketamine (2–2.5 mg/kg) and medetomidine (0.07 mg/kg), remotely administered by a Dan-Inject CO2 rifle (Dan-Inject, Denmark), and reversed with atipamezole (0.3 mg/ml; following [33]). Sedation time was kept to a minimum, typically less than 1 hour. After immobilization all cheetahs recovered fully, showing no signs of distress and no apparent side effects were observed in the short- and long-term. Collars, which were only fitted on adults, weighed 400 grams which is the recommended weight for cheetah collars [34]. All collars were removed if they malfunctioned or if the batteries were low. The animal handling protocols used conformed to the standards of the American Society of Mammalogists [35] and permissions to deploy collars were provided by the Kenya Wildlife Service (Permit No.: KWS/BRM/5001) and the National Commission for Science, Technology and Innovation (Permit No.: NACOSTI/P/16/69633/10821).
The collared males were all singletons, except for one male (M03), who was part of a five-male coalition. Over an 18 month period, the five-male coalition were sighted on 73 occasions and only on one occasion did the coalition separate for a period of <24 hours. We therefore collected data on a total of eight individuals in four social groups. While this is a relatively small sample size, it is a quarter of the entire population as there are approximately 32 adults within the study area [36]. The collars collected GPS coordinates every three hours (00h, 03h, 06h, 09h, 12h, 15h, 18h and 21h) and when there was satellite communication, data were uploaded on a daily basis at 06h. On average, the collars were deployed for 285 days ranging from 115 to 349 days (Table 1). For each pair of cheetahs (hereafter referred to as a dyad), we only used simultaneously collected data for the analyses.

Data processing and analysis

Static interactions.
To determine the static interactions between male cheetahs we calculated their space use and the amount of overlap for each dyad to determine the possibility that individuals could encounter each other either directly or indirectly. Space use for each individual per dyad was based on their utilisation distributions, which is the distribution of an individual’s locations over time [37], using the adehabitat package [38] in R [39]. To calculate the utilisation distributions we used a fixed kernel density estimate using a bivariate normal kernel. We used the reference bandwidth parameter (href) as the smoothing factor unless href > 1000 then we used 80% of the href to minimise over-smoothing of the data. Using the resulting utilisation distribution, we determined both the 99% and 50% kernels which respectively represent an individual’s total space use and their core areas. For each dyad we then calculated the amount of overlap of the 99% kernels and the 50% kernels.
Marking posts are used by male cheetahs to communicate their presence to conspecifics. Using the methods described by [29], we located marking posts based on a cluster analysis using data from the GPS collars and opportunistically when conducting fieldwork. For each dyad, we determined how many marking posts were found within the 99% kernel overlap. We used the recurse package [40] to calculate 1) how many marking posts were visited by each individual and 2) how many marking posts were visited by both individuals in a dyad (hereafter referred to as mutual marking posts). We classified a visit when a cheetah came within 500m (half the average step-length, see Dynamic interaction for details) of a marking post. For the mutual marking posts we calculated 1) the time between individual visits and 2) the time spent within 500m of a mutual marking post by each individual and tested whether there were differences between individuals. In addition, we calculated the time between visits from two different individuals i.e. we calculated how long it took for individual x to visit a mutual marking post once individual y had visited and vice versa. We then tested whether the time it took for an individual to visit a marking post once the other individual had been there differed between the individuals. We tested the data for normality using the Shapiro–Wilk test and used a t-test if the data were normally distributed and a Wilcoxon test if they were not. If the data were not normally distributed then we provided the median ± median absolute deviation in addition to the mean ± standard deviation.

Dynamic interaction.
We first explored the movement of the different individuals using the moveVis package [41]. Then, to determine whether interactions between two individuals in a dyad were likely to occur, we calculated the proximity between simultaneous locations using the wildlifeDI package [42]. The 3hr resolution of the data is quite coarse so we used different proximity thresholds to group possible encounters based on the average 3-hour step-length which was 1,021m ± 1,487m (mean ± standard deviation). We used four proximity thresholds: <500m, <1000m, <1500m and <2000m which correspond to 0.5, 1, 1.5 and 2 times the average step-length. We then determined whether these possible encounters took place at night (fixes at 21hr, 00hr, 03hr or 06hr) or during the day (fixes at 09hr, 12hr, 15hr or 18hr), whether they occurred within 50% kernels and the distance to the nearest known marking post. The half-way points between the two individuals were used as the estimated location where a possible encounter occurred.
Encounter outcomes.
The GPS data were examined once every few days. If any unusual behaviour was detected, such as no movement after a possible encounter, the field team would investigate to establish whether any injuries or deaths occurred as a result. In addition, for each possible encounter we compared the behaviour before to the behaviour after at four different time lags; 3hrs, 6hrs, 12hrs and 24hrs. First, we calculated the distance between two individuals before and after a possible encounter to determine whether males moved away from one another after an encounter. Then, per dyad, we calculated 1) the distance to the encounter location for each individual, 2) the distance travelled per individual and 3) the path tortuosity, or straightness. Path tortuosity was calculated by dividing the net displacement by the total distance travelled. A value of around 1 would indicate that the individual travelled in a straight line whereas a value <1 would be indicative of a tortuous path. We compared the proximity of the two individuals within a dyad, distance to encounter location, distance travelled and tortuosity before and after a possible encounter. We tested the data for normality using the Shapiro–Wilk test and used a t-test if the data were normally distributed and a Wilcoxon test if they were not.

Results

In total, we secured simultaneous data on four male cheetah dyads ranging from 113 to 329 days per dyad. One individual (M01) was part of three of the four dyads, two individuals (M02 and M03) were part of two dyads and one individual (M04) was part of one dyad (Table 2).

Static interaction

Across the four dyads the 99% kernels ranged from 253 km2 to 1,903 km2 and the 50% kernels ranged from 11 km2 to 359 km2 (Table 2). For all the dyads, the 99% kernels overlapped but the amount of overlap ranged from 10% to 100% where in Dyad 2 the 99% kernel of individual M01 fell completely within the 99% kernel of individual M03 (Fig 1). Only the core areas (50% kernel) of Dyad 3 had an extensive area of overlap (28% and 48%), whereas the cores of the other dyads did not overlap or the area of overlap was minimal (< 4%).

We found 125 marking posts in the study area and the number of mutual marking posts per dyad ranged from 4 to 37 with the average number of visits to these marking posts ranging from 1 to 46.16 (Table 3). For the three dyads that had the most extensive spatial overlap and the largest number of mutual marking posts (Dyads 1, 2 and 3) the average time that an individual was within 500m of a mutual marking post did not vary significantly between individuals in the same dyad, ranging from 5.06 ± 4.56 hours to 7.95 ± 5.85 hours (mean ± standard deviation; Table 3). The data were not normally distributed and the median for the same three dyads ranged from 2.52 ± 2.35 hours to 7.02 ± 6.01 hours (median ± median absolute deviation). For all the dyads, the average time between visits of a mutual marking post varied significantly between the individuals (Table 3). In the case of Dyad 1 and 2, individual M01, who we classified as territorial, visited mutual marking posts more frequently compared to individuals M02 and M03, neither of which were strictly territorial (Table 1). The time between different individuals visiting the same mutual marking post ranged from 3.11 ± 4.01 days to 13.64 ± 8.51 days and did not differ significantly across the dyads (Table 3).

Dynamic interaction

For three of the four dyads (Dyads 1, 2 and 3) the possibility of individuals within each dyad encountering each other was high as they overlapped extensively in space and had a large number of mutual marking posts. For these three dyads we explored their simultaneous movements and calculated the proximity between each individual within a dyad. The individuals within the three dyads did on occasions come into close proximity to one another as can be seen in the animation provided in the S1 Movie. The minimum distance between individuals in a dyad ranged from 89 m to 196 m but the average proximity between individuals ranged from 17,145 ± 6,865 m to 26,367 ± 11,288 m (Fig 2). Possible encounters were classified according to four different thresholds and we detected four possible encounters with a proximity threshold of <500m, 11 with a proximity threshold of <1000m, 21 with a proximity threshold of <1500m and 25 with a proximity threshold of <2000m (Table 4). Possible encounters were more likely to occur at night than during the day (χ = 8, df = 1, p = 0.005) and occurred most frequently at 21hr and midnight. Of the 25 possible encounters, 64% (n = 16) occurred within the core area of one individual, 28% (n = 7) occurred where the 50% kernels overlapped and two possible encounters in Dyad 3 did not occur in any of the core areas. Eleven (44%) of the possible encounters occurred within 500m of a known marking post.\

Encounter outcomes

For possible encounters with proximity thresholds of <500m, <1000m and <1500m the distance between males was overall significantly less during the period 3 to 6 hours after a possible encounter, compared to the 3 and 6 hours before a possible encounter (Table 5). In other words, rather than moving away from each other, male cheetahs stayed in close proximity to each other for up to 6 hours after a possible encounter.For the distance to the encounter location, distance travelled and tortuosity we wanted to determine whether there was individual variation within each dyad. However, because of the paucity in the number of possible encounters that were detected per dyad we were only able to carry out the analysis for possible encounters with a proximity threshold <2000m. In general, cheetahs were closer to the encounter location after a possible encounter compared to before for all four time lags, apart from individual M03 in Dyad 3 where the opposite trend was observed, however none of the results were significant (S1 Table). Similarly, cheetahs travelled less after a potential encounter compared to before, apart from individual M03 in Dyad 3 where the opposite trend was observed. Some of the results, especially at the 12hr and 24hr lag were significant for Dyad 1 and 2 (S1 Table).

On the 11th February 2017 the collar on M04 stopped transmitting but when the team visited the last location sent by the collar, neither cheetah nor collar could be found and the individual has not been seen since. On the 1st October 2017 the collar on M01 stopped transmitting data after the collar data showed that individuals M01 and M03 had come within 89m of each other. The team went to the last GPS coordinate that was transmitted by the collar and found the remains of M01 70m from the last GPS fix sent by the collar. Upon inspecting the carcass, a puncture wound was found on the left side of the skull. Based on the circumstantial evidence, we believe that the death of M01 was either a direct or an indirect result of an aggressive interaction between him and the 5-male coalition (M03). Interestingly, three months prior to this encounter M03 and his coalition mates started establishing a territory approximately 25km southwest from M01’s territory. Two days before the encounter the coalition travelled 25 km to the encounter location, spent 12 hours within 150 m of M01 and then travelled 19km straight back to the core of their territory. M03 did not return to the vicinity of the encounter between the 1st October 2017, when the encounter took place, and 3rd February 2018, when M03’s collar was removed. The closest they came to M01’s territory during that time was approximately 10 km (Fig 3 and the animation in S2 Movie).


Discussion

Using GPS collar data we documented static and dynamic interactions between male cheetahs in Kenya’s Maasai Mara and investigated the outcomes of these interactions in terms of movement behaviour and mortalities. As we predicted, male cheetahs showed extensive spatial overlap of the 99% kernels. This high degree of overlap observed in the Maasai Mara could be related to the pattern of prey availability [43], although we do not have the data to test this. However, apart from one dyad, there was little overlap of core areas (50% kernels) and it could be that core areas are defended more intensively than the peripheral areas [29]. Similar to observations in other areas, marking posts were frequently visited by males [29, 44] and this could indicate the mechanism that results, despite the extensive spatial overlap, in the rarity of occasions when members of a dyad were in close proximity [27]. Interestingly, our results show that possible encounters were most likely to take place in the core area of one individual of a dyad or where cores of both individuals overlapped. We also found that, similar to African wild dogs (Lycaon pictus), possible encounters occurred more at night than during the day [17]. While cheetahs, like African wild dogs, are predominantly diurnal they can be active at night [45] and nocturnal activity for males has been found to be considerably higher than for females [46]. Data from camera traps set at marking posts found that visits occurred more at night than during the day ([44]; KK unpublished data) suggesting that male nocturnal activity is partly driven by patrolling behaviour which is probably why encounters predominantly took place at night.

In some species, including African wild dogs and white-faced capuchins (Cebus capucinus), avoidance behaviours, characterised by an increase in distance and speed travelled post-encounter, were observed as a result of interactions between different groups [17, 18]. However our results, in contrast to our predictions, did not show avoidance behaviour post-encounter as males stayed in close proximity to each other 3–6 hours after a potential encounter. It is possible that males stayed in close proximity to each other, as part of a territorial defense strategy, if a recent scent of a conspecific was detected. This behaviour has been observed in dwarf mongoose (Helogale parvula) groups, who moved slower and covered shorter distances in the hour following the encounter of rival faeces at a latrine site within their territory [16] and red fox (Vulpes vulpes) males who spent more time in scent-marked areas [47]. Alternatively, males could come into close proximity to one another if they are attracted to a resource, such as a female in oestrus [27, 48]. Cheetahs exhibit a high rate of multiple paternity [49] so it is possible that multiple males stay in the vicinity of a receptive female with the hope of getting a chance to mate. These encounters could however result in fatalities if the removal of competition increases future mating opportunities [50]. If encounters occur as a result of access to a receptive female rather than to a static, long-term resource such as a territory then this could explain why the five-male coalition did not take-over the territory of individual M01 after he died.

Aggressive interactions with fatal consequences are not uncommon in cheetahs. Caro [22] reported three cases in Serengeti where singletons were killed by coalitions (all three-male coalitions). Similarly, Mills and Mills [27] found that 50% of male-male encounters recorded in the Kgalagadi Transfrontier Park in Botswana/South Africa resulted in death. To our knowledge, fatal interactions have not been observed between female cheetahs. This could explain why male mortality is higher and life expectancy lower for males compared to females [51, 52] resulting in a female biased sex ratio [52]. For some species, such as voles (Microtus oeconomus), lions and grizzly bear (Ursus arctos), the removal of males, through either displacement or mortality, has a negative effect on population growth as a result of increased infanticide [53–55]. Infanticide has however not been observed amongst cheetahs [56] possibly because it rarely occurs in predominantly solitary species [57]. The removal of males could however have other population-level consequences [58] but the impact of male mortality on population dynamics in cheetahs is unclear.

Static and dynamic interactions can play a role in disease transmission [3, 4]. In the Mara-Serengeti ecosystem there is a relatively high prevalence of mange [59, 60] and in Southern Africa cheetahs have been positively tested for feline coronavirus (FCoV) and feline panleukopenia virus (FPV), which can be highly contagious and fatal [61, 62]. Pathogens such as these can easily spread through faeces and other bodily fluids, which are deposited and investigated by male cheetahs at marking posts. This could explain why in 2015 several males in the Maasai Mara, who overlapped spatially, died of a yet unknown disease within a short space of time [63]. We suggest that future epidemiological research should investigate the role of scent marking posts and movement in disease transmission [64].
Here we give a descriptive analysis of the static and dynamic interactions between male cheetahs and the outcomes of these encounters. Despite the clear patterns that were observed, there are several caveats that warrant discussion. Firstly, we were only able to use data from four collared males, one of which was part of a 5-male coalition. It is therefore possible that other uncollared individuals, including the other members of the 5-male coalition, could have influenced the results. Secondly, because of the resolution of the collar data we might have missed visits to marking posts and we inferred when interactions took place rather than being able to detect actual interaction (apart from one occasion). Our results are therefore likely to be on the conservative side and we suggest that future studies use higher resolution data and/or proximity loggers to investigate actual interactions between individuals (e.g. [17, 65]). However, even with a relatively coarse resolution of data and only a small number of individuals we managed to investigate interactions and subsequent outcomes between males giving a first detailed insight into intraspecific interactions in cheetah.

Table 1:

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Figure 1:

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Figure 2:

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Table 4:

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^Table 5

Figure 3

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[Acinonyx sp.]

Cheetah coalition hunts a wildebeest, one of the cheetahs takes the lead and grapples the wildebeest down the rest of the cheetahs kill it and eat it.

This is the first time I have seen a cheetah use solely grappling and not tripping+grappling to bring down prey larger than itself.

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[Acinonyx sp.]

Focus on the Cheetah (Acinonyx jubatus)

Cheetah

Acinonyx jubatus  (Schreber, 1775)

Afrikaans    Jagluiperd  

German    Gepard

French    Guépard

Swahili    Duma / Msongo

isiNdebele    Ihlosi

isiZulu     Ihlosi

isiXhosa    Ihlosi

seSotho    Lengau

seTswana    Lengau

Shona     Ihlosi

Shangaan    Ndloti

Venda     Didingwe

Nama/Damara  !Arub

Herero    Shitona
Ovambo    Shinga

IUCN Conservation Status:

Vulnerable (VU), C2a(i) = a continuing decline in numbers of mature individuals.
No  sub-population  was found  to  contain  more  than  1  000  mature  individuals.    In  2006 heetah numbers worldwide were estimated to be <15 000.





The cheetah’s speed, its hunting skills and its daylight activity led to its choice as a tame

hunting companion in earlier  times.   Evidence of this can  be  seen on a silver urn  found

recently in the Caucasian mountains.  The urn, dating back to 700-300 BC, is etched with

a tamed cheetah wearing a neck collar.  In South Africa, bushman from the Kalahari have
a tradition of tracking cheetah to its kill in order to snatch meat for their own use.

Taxonomy 
Kingdom:    ANIMALIA

     Phylum:    CORDATA

     Class:     MAMMALIA

     Supercohort:   LAURASIATHERIA

     Cohort:    FERUNGULATA

     Superorder:    FERAE

     Order:     CARNIVORA

     Suborder:    FELIFORMIA

     Family:    Felidae

     Subfamily:    Acinonychinae

     Genus:    Acinonyx
     Species:    jubatus

In 1775 J.  C. D. von Schreber  described the animal  after a specimen from the Cape of

Good Hope as a “purring” cat and classified it as Felis jubatus.  More recently the cheetah

was reclassified as Acinonyx jubatus as it differs from other cats in having claws that are

not fully  retractable.   The name  “cheetah” is  derived from  the  Hindu  word “chita”  which

means “spotted”.  The  cheetah  is not related to  other  spotted cats such as  the leopard,

jaguar or serval.  The genus is represented by a single species A. jubatus and is widely
distributed from Africa to the Middle East

the  legitimacy  of  their
sub-specie classification has been bought into doubt by genetic analysis which shows only
a minimal genetic variation between the populations.  This may be due to several recent bottlenecks in the genetic development of the species.

Distribution
In  the  past,  cheetah  were  widely  distributed  from  the  Cape  of  Good  Hope  to  the

Mediterranean,  excluding  the  rainforests  of  central  and  western  Africa,  to  the  Arabian

Peninsula, the Middle  East, Pakistan, India  and  the southern  parts  of Russia.   Cheetah

were extinct in India by 1952 and in Russia by 1989.  Only a reduced population of 40-50

cheetah  remains  in  the  Middle  East, most  being  found  in  Iran  and  a few  in  Pakistan.  

Cheetah  are  now  believed  to  be  extinct  in  the  western  Sahara,  Ghana,  Nigeria  and

Djibouti.  The two largest cheetah populations are in East Africa in the Serengeti and  in
southern Africa, in Namibia and Botswana.





Description

Cheetah have a slender build, relatively long legs and, in comparison to the stronger body

built of most other cat species, are light in weight in relation to size.  Its build resembles

that of  the dog  family rather  than that  of the cat  family.   The average  mass of  an adult

cheetah male is  64 kg and  of  a female, 45  kg.   Its aerodynamic  build  allows it to  reach

speeds of up to 104 km/hour during a chase, the highest speed of all land mammals.



Cheetah have partly retractable claws in poorly developed nail beds.  This distinguishes

them from  other cats  that have  fully retractable  claws and from the  dog family  that  has

non-retractable claws.



The spots of cheetah differ from  the rosette-like spots on the pelage  of the leopard and

jaguar  as  they  are  round  or  oval  and  fully  coloured.    The  typical  black  teardrop-stripe

commences from the inner eye and stretches down the face to the corner of the mouth.



The  colouring  and  spotted  pattern  of  the  pelage  varies  with  habitat  and  distribution.  

Cheetah from the Sahara desert are lighter in colour with hazelnut-coloured spots and dull

teardrop  stripes  and  tail  rings,  some  being  so  light  that  they  are  referred  to  as  white

cheetah.  Individuals inhabiting the black rocky mountain areas of the Sahara usually have

a brighter colouring.



A unique colour variation known as the king cheetah is found in southern Africa.  These

free roaming cheetahs were first recorded in 1928 with the most recent sighting being in

the Sabi  Sand Game  Reserve  in 1992.   Several  king cheetah  litters have  been born  in

captivity, 70 being bred in the De Wildt Cheetah Research Centre between 1980 and 2004.  

Research indicates that the king cheetah is not a genetic sub-species but merely a colour

variant of  A. jubatus,  with longer,  more  silk-like  hair.    The  spots along  the spine  of this

variant meld together to form stripes.  The end quarter of the tail is solid black compared

to the rings of the common cheetah.



In the 17th century, a cheetah with dark blue spots on a blue-tinted, white skin was found

in Jahaner,  India.    Sightings of  a  spotless  cheetah  in  Tanzania  in 1921  and  of  a  black
cheetah in Kenya in 1972 were also reported.

Spoor

The cheetah has the rounded print of a cat but shows definite, dog-like claw marks at the

front.    The  claw  marks  distinguish  it  from  other  cat  spoor  while  the  rounded  shape

distinguishes it from the lengthened, oval spoor of a dog or hyaena spoor.  The front print

of an adult cheetah spoor measures 110x85 mm and the hind 95x75 mm, inclusive of the
claw markings.

*This image is copyright of its original author

Trophy

The trophy measurement is the accumulative measure of the maximum length of the skull
from the nostril to the base of the cranium together with the maximum diameter of the skull.

*This image is copyright of its original author

Habitat requirement

Although  cheetah  fare  well  on  grass  plains  they  prefer  open  savannah  woodland  with

ample  visibility.    Hunting  success  is  increased  by  a  moderate  growth  of  vegetation  for

refuge but  dense  thicket limits  their  chasing  strategy.   Cheetah  are  mostly  restricted  to

sub-tropical  and  arid  habitats  with  an  annual  rainfall  of  100-600  mm.    Although  past

distribution  included  temperate  highveld  grassland,  this  habitat  is  marginal  as  cheetah

cannot  tolerate  snow  and extreme  wet,  cold conditions  with  a temperature  below -5°C.  

Mountainous terrain and riverine thickets are totally avoided.  The most determining habitat
parameter is the abundance of suitable prey.

Behaviour

Unlike most of the larger predator species that are nocturnal, the cheetah is predominantly

diurnal.    Most hunting  takes  place  in  early  mornings  after  dawn  and  in  late  afternoons

before  dusk,  as  they  need  good  visibility  to  outrun  their  prey.    This  strategy  reduces

competition with other large predators that kill cheetah such as the lion Panthera leo, the

leopard P. pardus and the spotted hyaena Crocuta crocuta.



Communication between individuals consists of a high pitched cry that can be heard up to

two kilometres away.  Cheetah purr like a domestic cat when happy and at rest.



Cheetah display a unique behaviour in their use of “play-trees”.  These act as a beacon

for  social  gatherings  of  homebound  groups  and  for  single,  adult,  nomad  cheetah  of  a

different birth origin.  Generally, a play-tree is visible from a distance and is a lone, tall tree

consisting  of  a  single  large  stem,  ample  canopy  shading  and  sparse  vegetation
underneath.  The stem is marked to a height of 1.7 m by numerous claw scratches.  Visiting cheetah  often  remain  at  a  play  tree  for  several  days.    Cheetah  from  different  groups

tolerate each other at play-trees and there is interchange of individuals between groups.  

It is a preferred site for alpha female/male bonding.



When trapping cheetah, the best results are achieved by placing traps at play-trees and,

if possible, by scenting them with cheetah urine rather than by setting them with dead bait.  

The trap should not be removed immediately after a successful capture as more than one

cheetah  may  be  trapped  at  the  same  site.    Translocated  cheetah  generally  attempt  to

return to their original home ground.  


Feeding & Nutrition

Free roaming, adult cheetah require an average of 2.5-3.5 kg fresh meat per day in order

to  maintain  their  health.    The  composition  of  cheetah  prey  depends  on  the  relative

abundance of prey species in the area.  Cheetah seldom take prey larger than 60 kg and

do not scavenge carcasses killed by other predators, preferring fresh meat from their own

kills.  Thus if bait is used it should be fresh meat.  In comparison to other large predators,

cheetah hunt young animals rather than old, injured or sick individuals.



A  cheetah  hunt  follows  one  of  three  basic  strategies  pouncing  on  unsuspecting  prey

searching for prey using a vantage point such as a high termite mound or a large, fallen

tree trunk ambushing, pushing or charging prey against a game or stock fence. Cheetah

are masters of this strategy



A kill usually starts with a short, high speed chase of 100-250 m.  The cheetah then trips

its prey with a  smack against the hind  legs,  jumps over it and  smothers  it by sinking its

fangs into the  throat.  The average  speed of cheetah  recorded  in the Serengeti was  87

km/h, although they can reach a top speed of 104 km/h.  The breathing rate of a cheetah

increases  tenfold  during  a  chase.    An  unnecessary  energy  loss  is  prevented  by

determining the potential of a successful outcome early in the chase and aborting it if it is

low.  The  average recorded chase  distance for successful  kills  in the southern  Kalahari

was 218 m and for unsuccessful chases, 122 m.



The kill prey is consumed rapidly in order to prevent its theft by other larger predators such

as lion and hyaena.  Even vultures can succeed in taking over a cheetah kill.  When more

than one cheetah feed together their bodies  lie in  a circle  around the carcass forming a

unique  star-like  pattern.    The  cheetah’s  water  requirements  are  met  by  the  blood  that

accumulates in the hollows of a kill.  Thus they are independent of surface water although

they will drink if water is available.  Wild fruit such as the tsamma and gemsbok cucumber

found  in  the  arid  Kalahari,  have  high  moisture  contents  and  are  frequently  chewed  by
cheetah in order to supplement their water intake.

Territory & Home range

Territorial male and brother groups scent-mark their ranges more intensely than nomadic,

solitary males.  As the health of nomads is generally weaker than that of territorial males,

they  seldom  remain  in  an  area  for  longer  than  a  few  days.    The  most  common  scent

marking is a backwards spray of urine against the stem of a tree or bush.  The tail is lifted

with the  hindquarters  facing the  tree.   The  urine  is then  sprayed  up the  tree  trunk to  a

maximum  height  by  stretching  the  hind  legs  and  lifting  the  hind  quarters  as  high  as

possible.  Dominance of the cheetah is established by the height of the marking; the higher

the marking, the  greater the dominance.   Play-trees  are priority sites for  scent marking.  

Scat deposited at fixed sites on the periphery of the range also act as a warning signal to

intruders.



The cheetah is unique among the large cats as the female has a larger home range than

the  male.    In  the  Serengeti,  adjacent  female  home  ranges  overlap  and  measure  an

average  of  833  km²  compared  to  the  37  km²  of  male  groups  and  777  km²  of  nomadic

males.  In Namibia, the recorded mean home range of females is 1 500 km² and for males,

800 km².  The home ranges of male groups do not overlap.  The size of these ranges is

limited by prey abundance and thus differs between regions.  The entire range is not used

equally and activities such as hunting, refuge and breeding are concentrated in different
areas.

Social structure

Lion and cheetah are the only large cats in the world that form social groups.  Groups of

cheetah consist of an adult alpha female and her sub-adult cubs, or of 2-4 sub-adult (chi)

or adult (beta or alpha) brothers or, less often, beta sisters.  Adult males of the same litter

either live  together as a  brother group  or  split to  become solitary nomads.   With  a  high

cheetah  density  and  a  good  supply  of  prey,  non-related  male  groups  may  form  large,

temporary groups  of up  to 20  individuals.   Related brothers generally group  together for

life  and  inhabit  the  same  home  range,  while  solitary  males  will  only  fight  to defend  a

territory or home range for a few months and then become nomadic.  Male groups do not
have a hierarchy of dominance and all members may mate with an available alpha female.

*This image is copyright of its original author

Reproduction

An  adult  alpha  female  has  little  contact with  the  alpha  males  and  even  courtship  and

mating  seldom  lasts  longer  than  24  hours.    Despite  this,  an  alpha female  is  extremely

particular in her choice of a male partner.  She becomes sexually mature at 21-24 months

and  generally  starts  mating at  about  three  years.   Mating  usually  occurs  at  night  and

gestation lasts  for 90-95  days.  Young  are born  at any  time of  the year as  females are

seasonally poly-oestrus and are not restricted to a breeding season.  When in pro-oestrus

she releases hormones in her urine that attracts alpha males.  During courtship oestrus is

further stimulated, with full oestrus being reached after 7-14 days.  In the interim the male

is rejected aggressively.  Ovulation is stimulated only during the final mating.  A high level
of major sperm abnormalities caused by  the  limited genetic variation of the species has negatively affected the population growth of cheetah.



The cubs are  hidden  in a den  until  an age of  5-8  weeks.  The den  is usually close  to  a

rocky outcrop used by the female as a vantage point and the surrounding area must have

ample prey and tall grass cover for  hunting.   The mother  translocates the litter to a new

den if prey moves away from the area or if predators endanger the site.  She constantly

remains in close vicinity to the den and restricts her hunting to this area.  The alpha male

has no role in raising the young and never accompanies the mother.



The cubs remain dependent for 12 to 20 months.  The weaning process starts at six weeks

and lasts until  an age of  3-4 months.   Cubs accompany the  hunt from an  age of seven
months and only leave the mother after 18 months.

Page 11 of 14



negatively affected the population growth of cheetah.



The cubs are  hidden  in a den  until  an age of  5-8  weeks.  The den  is usually close  to  a

rocky outcrop used by the female as a vantage point and the surrounding area must have

ample prey and tall grass cover for  hunting.   The mother  translocates the litter to a new

den if prey moves away from the area or if predators endanger the site.  She constantly

remains in close vicinity to the den and restricts her hunting to this area.  The alpha male

has no role in raising the young and never accompanies the mother.



The cubs remain dependent for 12 to 20 months.  The weaning process starts at six weeks

and lasts until  an age of  3-4 months.   Cubs accompany the  hunt from an  age of seven

months and only leave the mother after 18 months.





Production

The cheetah’s worst enemies are the degradation of suitable habitat, an inadequate supply

of prey and the low genetic diversity of the species.  As few cheetah in the Serengeti live

beyond four years, most females can produce only one litter.  Diseases commonly causing

mortalities are anthrax, tick fever, mange and catynteritis.  Cheetah are also susceptible

to internal parasites.



Cubs have a birth mass of 150-300 grams and a thick, greyish-blue coat on the back of

the neck that disappears after three months.  In the Serengeti 83% of the cubs die before

reaching an age of  14  months as they are prone  to  injury, disease and starvation.    The

density  of  both  cubs  and  adults  in  an  area  varies  in  relation  to  predation  and  prey

abundance.    A  large  number  of  females  will  concentrate  in  areas  with  a  good  prey

resource.  The average long-term cheetah density in the Serengeti is 0.8-1.0 cheetah/10

000 ha but can range up to 40 cheetah/10 000 ha.  The density of cheetah in savannah
woodland is lower than that of larger predators as a result of their predation.

*This image is copyright of its original author

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[Acinonyx sp.]

An assessment of the success of a cheetah re-introduction project in Matusadona National Park Reply

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[Acinonyx sp.]

Reassessment of an introduced cheetah Acinonyx jubatus population in Matusadona National Park, Zimbabwe

Abstract
Translocations are used to mitigate human–wildlife conflict, secure population viability of isolated populations and introduce or reintroduce populations in former or new range. With wild species increasingly confined to small patches of habitat embedded in human-dominated landscapes, the use of translocations is likely to increase. The cheetah Acinonyx jubatus is a large carnivore species with a long history of translocations. As for other species, evaluation of the success of cheetah translocations is complicated by a scarcity of published results, especially of failed attempts. Yet, such information is crucial to improve future translocations. A relatively well documented case is the translocation of alleged problem cheetahs into Matusadona National Park, Zimbabwe, in the early 1990s. In this study we used a combination of survey methods to reassess the status of Matusadona cheetah population and model current occupancy in relation to densities of competing carnivores and altitude. Our findings indicate this cheetah population has effectively been extirpated, highlighting the importance of thorough planning and standardized long-term monitoring of translocated populations for the understanding of the factors that affect translocation success.

Introduction

As a consequence of increasing human populations and the resulting demand for agricultural land and pressure on natural resources, the ranges of most large carnivores have contracted substantially (Di Minin et al., 2016). Maintaining viable populations of large carnivores requires large connected habitat networks (Di Minin et al., 2016). However, such networks are often inhabited by people, who degrade carnivore habitat and persecute large carnivores because of actual or perceived livestock losses, confining them to isolated patches of land (Di Minin et al., 2016).

Translocation is regularly used to mitigate conflict, secure viability of isolated populations and reintroduce populations in former or new range (Linnell et al., 1997; Fischer & Lindenmayer, 2000). Success depends on the aim of the translocation, the source of the translocated individuals, the number of individuals released and the cause of the original decline (Fischer & Lindenmayer, 2000). Translocations generally fail to reduce human–carnivore conflict and it is now widely accepted that such negative interactions are best mitigated by promoting coexistence (Linnell et al., 1997; Fischer & Lindenmayer, 2000; Boast et al., 2018a). However, translocations to establish or re-establish and augment populations of native, non-vulnerable, wildlife species seem more successful (Griffith et al., 1989). Translocation success increases when the source population is wild, the founder population is large (Griffith et al., 1989; Fischer & Lindenmayer, 2000) and the original cause of the population decline has been eliminated (Fischer & Lindenmayer, 2000). Although it is recognized that monitoring is crucial to improve the success of translocations (IUCN/SSC, 2013), there has been little follow-up of translocated animals to determine success (Linnell et al., 1997) and long-term evaluation of introduced or reintroduced populations is often lacking (Fischer & Lindenmayer, 2000).
The cheetah is a large carnivore species regularly translocated for conservation purposes (Boast et al., 2018a). A relatively well-documented translocation is the 1993–1994 translocation of alleged problem cheetahs from commercial farmland into Matusadona National Park, Zimbabwe, an unfenced protected area without resident cheetahs (Zank, 1995; Pitman, 2007). Although it was initially thought the introduced cheetah population would not be viable without intensive management (Zank, 1995), it persisted without intervention and, with the establishment of a breeding population, the introduction was cautiously deemed a success (Purchase & Vhurumuku, 2005). Whether the translocation reduced human–cheetah conflict on commercial farmland is unknown (Purchase, 1998). In this study, we used citizen science, questionnaire-based information, camera-trap and spoor surveys to reassess the status of Matusadona's cheetah population and model current cheetah occupancy in relation to densities of competing carnivores and terrain elevation. Our follow-up highlights the importance of thorough planning and standardized long-term monitoring of translocated populations to understand the factors affecting translocation success and to improve future translocation programmes.

Study area and species

The 1,370 km2 unfenced Matusadona National Park, utilized for photographic safaris, lies in northern Zimbabwe on the shore of Lake Kariba. Depending on the lake level, the Park comprises a valley of c. 400 km2 and an escarpment of c. 1,000 km2 (Fig. 1). During 1993–1994, 21 alleged problem cheetahs were opportunistically removed from commercial farmland in southern Zimbabwe; six died shortly after capture, and 15 were kept in a captive enclosure (boma) for 6 weeks and then soft released into the Park, which held no resident cheetahs at the time (Zank, 1995; Pitman, 2007). Although it was assumed cheetahs were once resident in Matusadona, there are no records to support this (Zank, 1995) and it remains unknown whether cheetahs had become extinct or had never been present at all (Purchase, 1998). The introduced cheetahs utilized only the valley floor (Purchase, 1998), which has a grassland foreshore but largely consists of woodland with Colophospermum mopane, Combretum spp. and Terminalia spp. The escarpment is dominated by miombo woodland and is characterized by steep valleys with limited water availability in the dry season (Zank, 1995; Purchase, 1998). The cheetah's competitors, lion Panthera leo, leopard Panthera pardus and spotted hyaena Crocuta crocuta, occur in the Park. For Matusadona's large carnivores, buffalo Syncerus caffer and impala Aepyceros melampus have historically been the most abundant prey species (Zank, 1995; Purchase, 2004). The Park is bordered by the Sanyati and Ume rivers and surrounded by communal land used for subsistence farming and trophy hunting under the Communal Areas Management Programme for Indigenous Resources (CAMPFIRE; Fig. 1).

Methods

Data collection
As part of a nationwide survey (van der Meer, 2018), we launched a citizen science programme (2012–present) to collect cheetah sightings and photographs from tourists, safari operators and others. In addition, we interviewed 49 respondents (August–September 2013) about cheetah presence in Matusadona and surrounding areas: 23 Zimbabwe Parks and Wildlife Management Authority rangers, seven CAMPFIRE rangers, six safari guides, five hunters and eight hunting scouts (van der Meer, 2018). If respondents had seen cheetahs, we recorded sighting date and location, number of individuals, age, sex and size. We only included sightings from respondents who could identify the cheetah from carnivore pictures and clearly recalled sighting details. Some respondents provided photographs with their sightings. We also asked respondents about human–cheetah conflict and anthropogenic cheetah mortality, and checked reports at the Zimbabwe Parks and Wildlife Management Authority and CAMPFIRE offices for records of conflict and mortality.
During June–August 2016 we carried out a camera-trap survey using a pre-planned grid of 49 traps set up 4–5 km apart on the valley floor and part of the escarpment (Fig. 1). Each trap station consisted of two Cuddeback (Cuddeback, Wisconsin, USA) or Panthera V2 (Panthera, New York, USA) cameras, c. 6 m apart, facing each other but slightly offset, to limit interference from the flash. The cameras were attached to trees or metal standards c. 60 cm above ground level. Following establishment, the survey was run for 40 days, after which traps were removed within the same time frame and order in which they were deployed.
In June 2013 we drove three road transects on the valley floor (total length 59.8 km), and in July–August 2016 we drove four road transects on the valley floor (total length 68.2 km), all of which were driven three times in the early morning, on different days, and three of which repeated routes surveyed in 2013. Transects were driven at 10–20 km/h, with an experienced tracker on a seat mounted at the front of the vehicle. When large mammal spoor was located the vehicle stopped and spoor was identified and date, location (with a GPS) and the number of animals recorded. To avoid double counting, transects were never surveyed on consecutive days and we only recorded tracks < 24 hours old.
We consulted expert authorities and literature to gain insight into historical cheetah population trends in Matusadona and possible underlying causes of changes in cheetah numbers; e.g. fluctuations in lake levels, which affect the area of valley floor foreshore exposed and therefore prey and predator densities. After 2005 there have been no prey surveys other than aerial counts. Therefore to facilitate a comparison between studies we used dry season aerial count data with comparable transects as an indication of impala and buffalo densities. Although aerial counts have limited detection ability, they provide an indication of population trends (Dunham et al., 2015). From 2005 lion and spotted hyaena densities were based on spoor surveys rather than on direct sightings or call-ups (playback recordings of vocalizations).


Analyses

From the photographs collected via sightings (n = 64, 38% with photographs) and the camera-trap survey (n = 9), we identified individual cheetahs based on coat markings (Zank, 1995). Sightings of known individuals (n = 33) were used in a presence–absence model and to determine space use based on minimum convex polygons (MCPs). Although MCPs create unpredictable bias when sampling is not systematic, as with opportunistic sightings, (Börger et al., 2006), we nevertheless decided to use this method, to facilitate comparison between studies.

Substrate and cheetah density in Matusadona were sufficient to use the calibration model of Winterbach et al. (2016), in which observed spoor density = 3.26 × carnivore density. In the absence of a species-specific equation based on a number of populations, this is the recommended calibration equation for the cheetah (Boast et al., 2018b). For each transect we calculated spoor density (individuals per 100 km). Mean spoor density, standard deviation and 95% confidence intervals were calculated across all transects driven. Based on these calculations we determined mean, minimum and maximum cheetah numbers for the valley floor.
For the camera-trap survey, the MCP encompassing all traps buffered by 5 km was gridded at 500 × 500 m resolution. This gridded habitat mask was used to extract the spatially explicit recapture modelled densities for sympatric carnivores and altitude (GLOBE Task Team, 1999). In addition, total numbers of cheetah sightings were calculated per grid cell by counting the presences (1) and absences (0) of identified individuals. Given the number of zeros in the data, a zero-inflated Poisson generalised additive model was applied to model the probability of occurrence of cheetahs in relation to altitude and densities of competing carnivore species using zipGAM in package mgcv in R 3.3.1 (Wood, 2011; R Core Team, 2017).

Results

Population estimate, movement and range
Based on photographic identification, the cheetah population of Matusadona comprised three adult cheetahs: a female (first sighting September 2010, last sighting May 2014) and a coalition of two males (first sighting December 2009, last sighting May 2018). We did not receive photographs of other individuals, nor did sightings without photographs indicate the existence of additional cheetahs or breeding. The analyses of spoor survey data indicated a mean population size of 2–3 cheetahs (Supplementary Table 1). We only encountered spoor of adult cheetahs, on their own or in a group of two individuals. Cheetahs predominantly utilized the north-eastern part of the valley floor; they were not sighted on the escarpment or in the neighbouring communal lands. Home ranges overlapped by 63–66% (c. 46.4 km2) and were 70.5 km2 for the cheetah female and 74.3 km2 for the male coalition (Fig. 2), which is larger than the 11.3–53.8 km2 home ranges in 1998 (Purchase, 1998). The cheetah female was last seen in May 2014 and is presumed dead.

Factors affecting current occurrence

The presence–absence model indicated that terrain is the strongest explanatory variable for cheetah occurrence. Cheetah presence was restricted to below 800 m altitude (Fig. 3). Cheetah presence was also related to the occurrence of competitors: although spotted hyaena densities did not significantly influence cheetah presence, both lion and leopard densities did. As lion and leopard (> 0.2 individuals/100 km2) densities increased, the likelihood of cheetah occurrence decreased (Fig. 3; Supplementary Table 2, Supplementary Fig. 1). The model explained 93% of the deviance in cheetah presence/absence and, based on altitude and densities of competing predators, predicted cheetah occurrence to be higher along the foreshore of the valley floor (Fig. 4).

Population trends and mortality

Our estimate of the cheetah population from spoor surveys corresponded well with the estimate from sightings, but in the study of Purchase & Vhurumuku (2005) the estimate based on spoor density was substantially higher than the estimate based on sightings. To facilitate comparison we therefore only present estimates based on sightings. During 1995–2005 Matusadona's cheetah population reproduced and remained relatively stable at a mean of 13.6 ± SE 1.2 individuals (Table 1). However, the proportion of subadult cheetahs increased, indicating lower juvenile mortality but possible higher adult mortality (Table 1). Data on survival rates of cubs and adults are scarce: in 1998 mean litter size was 2.8 cubs (n = 8) and cub survival at 3–24 months was 60% (Purchase, 1998). In this same study, it was confirmed that two cheetahs of the founder population were still alive. Matusadona is surrounded by communal land and opportunities for cheetahs to disperse are limited (Zank, 1995). Although human–cheetah conflict in the communal land surrounding Matusadona has historically been minimal (one report in 1998; Purchase & Vhurumuku, 2005), cheetahs ranging into this communal land have suffered from anthropogenic mortality (Zank, 1995). We did not record any cases of human–cheetah conflict or reports of anthropogenic mortality.

Ecological factors affecting population trends

Shortly after the cheetah introduction, the lake level increased, with a peak in 2000 and high levels until 2004 (Fig. 5), decreasing the area of foreshore exposed. Aerial surveys suggest this negatively affected the impala population, but this did not seem to impact the cheetah population. Within this period the cheetah population fared well, which was attributed to a decline in lion numbers (Purchase & Vhurumuku, 2005). However, despite lion numbers remaining low (Fig. 5), during 2005–2009 the cheetah population decreased dramatically (Table 1). Within this period the lake level decreased, reaching its lowest point in 2006 (Fig. 5), exposing a larger area of foreshore, which coincided with high impala numbers (Fig. 5). Disease-related mortality of wild cheetahs is minimal, but cheetahs are susceptible to anthrax (Terio et al., 2018). Since 2005 there have been no records of anthrax outbreaks (Department of Livestock and Veterinary Services, pers. comm.), nor did we come across records of sudden simultaneous die-offs of cheetahs or other carnivores in Matusadona.

Discussion

Based on our survey and recent citizen science sightings, the Matusadona cheetah population currently comprises a coalition of two male cheetahs. This population is considered isolated (see also Zank, 1995; Purchase, 1998) and, with only two males > 9 years old remaining, is functionally extirpated. The reasons for the drastic decline of this cheetah population remain unclear. Aerial surveys suggest that a reduction in productive habitat as a result of high lake levels negatively affected prey availability, which may have resulted in a decrease in cheetah numbers. However, prey estimates based on road transects and block counts do not support this hypothesis (Purchase & Vhurumuku, 2005). Post-release population viability analyses showed a viable cheetah population could be established if cub mortality was ≤ 60% (Zank, 1995). It was initially expected that cub mortality would be higher (95%) and that intensive metapopulation management would be necessary to maintain the population, but later studies showed cub mortality was 60% and the population would be viable at a carrying capacity of ≥ 25 cheetahs (Purchase, 1998).

Whether Matusadona could ever sustain this number of cheetahs is questionable. Cheetahs select home ranges based on prey availability and vegetation characteristics, and within these home ranges they preferably utilize open savannah habitat (Caro, 1994; Broomhall et al., 2003). Matusadona's cheetahs only utilized the flat valley floor, within which they made extensive use of a small core area of open habitat along the north-eastern lake shore. Regardless of lake level, this area has the largest foreshore (Fig. 1; Purchase & Vhurumuku, 2005), highest impala density (Zank, 1995; Dunham et al., 2006, 2015) and optimal habitat to hunt and rest (Purchase, 1998). Although cheetah females tolerate each other, males actively defend territories (Caro, 1994). Even though the territories of Matusadona's male cheetahs overlapped considerably (Zank, 1995; Purchase, 1998), territorial behaviour in combination with limited availability of preferred habitat is likely to restrict the number of cheetahs that can successfully utilize the 400 km2 valley floor (see also Purchase, 1998). This limits the carrying capacity of the Park which, based on historical sighting-based population sizes, is likely to be < 25 cheetahs.

Intraspecific competition with larger carnivores affects cheetah densities (Lindsey et al., 2011) and, apart from the availability of optimal habitat and prey, impacts cheetah occurrence on the valley floor. Our presence–absence model showed no significant effect of spotted hyaena densities but a negative relationship with lion and leopard densities. Despite signs of leopard predation of cheetahs (Purchase, 1998), previous studies in Matusadona focused on competition with lions and spotted hyaenas; the potential impact of leopards was not taken into account. Although lions and spotted hyaenas are considered the main competitors of cheetahs (Caro, 1994), leopards have considerable dietary overlap with cheetahs (Lindsey et al., 2011) and cause cheetah mortality (Caro, 1994; Broekhuis, 2015). Consequently, cheetahs actively avoid interactions with leopards (Vanak et al., 2013) and the presence of leopards increases the cheetah's space requirements (Lindsey et al., 2011).

For free-ranging mammal populations in which individuals mature late and have few offspring per year, adult and juvenile survival rates contribute more to population growth than fecundity (Heppell et al., 2000). Although Matusadona's 20.5% adult and subadult mortality rate was considered low (Purchase, 1998), the mortality rate of dispersers is likely to be considerably higher. Cheetahs that moved into neighbouring communal land were not resighted (Zank, 1995) and there are no indications that Matusadona's cheetahs colonized nearby wildlife areas or connected to other resident cheetah populations (van der Meer, 2018). Since the release, human population density (ZPWMA, 2015) and numbers of livestock in the region have increased, and numbers of wild prey decreased (Dunham et al., 2015), thereby further reducing dispersal potential for cheetahs (Winterbach et al., 2015). As suggested by Purchase (1998), the viability of Matusadona's cheetah population is likely to have been impaired by high mortality rates of subadults, which were forced to disperse into suboptimal habitat by already established cheetahs.

The Matusadona cheetah introduction was based on assumptions rather than a feasibility study to assess whether the cheetah's ecological needs could be fulfilled (Zank, 1995; Pitman, 2007). Twenty-one cheetahs were captured opportunistically without considering factors such as relatedness and ideal age and sex-ratio for introduction (IUCN/SSC, 1987). In addition, this number of removals and translocations was unlikely to significantly reduce human–cheetah conflict at the capture site (Purchase, 1998) nor was it sufficient to establish a viable population at the release site (Zank, 1995). If international guidelines had been followed (IUCN/SSC, 1987), the translocation should not have proceeded, especially as without any attempt to identify and mitigate factors causing low cheetah densities inside protected areas and declining cheetah numbers outside protected areas, the translocation was unlikely to significantly contribute to cheetah conservation in Zimbabwe (Zank, 1995).
Long-term protocols in which key parameters are monitored at previously specified time intervals are crucial to understand the factors affecting translocation success and improve future translocations (Fischer & Lindenmayer, 2000; IUCN/SSC, 2013). Compared to other translocated populations (Boast et al., 2018a), the Matusadona cheetah population is relatively well studied but, nevertheless, a lack of standardized long-term monitoring prevents us from determining the exact causes of the population's extirpation. Translocation success largely depends on habitat quality and quantity at the release site and, in the case of translocations in unfenced environments, the surrounding areas (Griffith et al., 1989; Fischer & Lindenmayer, 2000; Boast et al., 2018a). Although the Matusadona cheetah population persisted for over a decade, the limited availability of optimal cheetah habitat in combination with interspecific competition and minimal dispersal abilities appear to make the area unsuitable to harbour a viable cheetah population. Unless the exact causes for the extirpation of Matusadona's cheetah population can be clearly understood and properly addressed, further reintroductions (IUCN/SSC, 2013) of cheetahs into the area are not advisable.

https://www.cambridge.org/core/journals/oryx/article/reassessment-of-an-introduced-cheetah-acinonyx-jubatus-population-in-matusadona-national-park-zimbabwe/163848EED4B2F49F81C65E306E6735B0 Reply

[Acinonyx sp.]

A Brief History of Cheetah Conservation

Abstract

The cheetah, Acinonyx jubatus, is a charismatic, and iconic species that, due to its uniqueness and extreme popularity over the epochs, has been exploited to near extinction. Once found on three continents, this streamline carnivore is disappearing rapidly from its former range and faces a bleak future due to numerous factors, including, popularity by nobility as pets and hunting companions, human population growth, human-wildlife conflict, illegal trafficking, loss of genetic diversity, and many others. The conservation crisis this species is facing is overwhelming, yet a core of hard-working conservationists has been working diligently for decades to slow, and in some areas try to even reverse, their downward spiral. This narrative will give the reader an insight into the history of the conservation efforts for this species, what has been done to date, and what still needs to be developed to save this unique species.

Introduction
The cheetah (Acinonyx jubatus), the most unique of the 41 species of felids, (Kitchener et al., 2016) is at the crossroads of its survival. With an estimated population of 7100 adult and adolescent cheetahs in their natural habitat (Durant et al., 2017), long-term conservation research programs are collectively working on strategies to ensure their survival. Over the past 4 decades, a small but prolific group of international researchers and conservation biologists has emerged, all dedicated to solving the problems that threaten cheetah survival. Their collective research is presented in the chapters of this book and brings together what we currently know about the cheetah, the challenges it is facing, and the solutions that have been developed. Many of the cheetah conservation strategies that currently are being undertaken have a unique and interesting history of how they began. Here we endeavor to offer a historical overview and timeline that ties together the information presented in the chapters of this book.

Historical context

Ancient History—Challenges for the Cheetah Populations
The cheetah is a survivor; its challenging evolutionary history has shaped a unique physiology, optimized for speed (Chapter 7). The first fossil records of cheetah (Acinonyx) date from approximately 4 million years ago and evidence of related species was retrieved in America, Europe, Asia, and Africa (Chapter 3). Following a founder effect approximately 100,000 years ago, the cheetah escaped extinction in the Pleistocene, which left the species with both reduced numbers and diminished genetic diversity (Chapter 6).
Modern History—Human Pressure on the Cheetah Populations
For the past 5000 years, humans throughout Asia, Europe, and Africa have revered the cheetah (Chapter 2); however, humans’ fascination with the cheetah has manifested in ways that have contributed to the species’ near demise (Wrogemann, 1975; Chapters 2, 14, and 2221422).
The first recognized use of the word cheetah was in 1610, but it wasn’t until 1775 that a German naturalist, Johann Christian Daniel von Schreber, published the first description of the species, which at that time was commonly found throughout Asia and Africa. Cheetahs were appreciated as hunting companions in India, and in Hindu, cheetahs were known as “Chita” or the “Spotted One,” and were often referred to as “hunting leopard.” The pressure on the wild cheetah populations due to the harvest of cheetahs for Maharajahs’ hunting parties, and later for safari hunting, contributed to the decline and eventual extinction of cheetahs from the wild in India (Chapters 4, 5, and 224522).
In Africa, the first nationally protected area, Kruger National Park, was established in the 1890s. Measures to protect wildlife were in part motivated by plummeting wildlife populations, as over exploitation, habitat loss, and human persecution (Schreber, 1775) arose as a result of a shift from traditional lifestyle to agriculture under colonial influences. However, large carnivores were viewed as vermin, and until the 1970s, cheetahs and other predators were killed in many African national parks to protect game (in addition to their persecution on farmland) (Linnell et al., 2001, Woodroffe and Ginsberg, 1997).
In addition, during the 1960s, cheetahs, along with other exotic wildlife species, were in high demand to stock the world’s zoos. Due to poor captive breeding success, most zoo-bound cheetahs were captured from wild populations, putting big pressure on populations in East Africa and Namibia (Chapter 22). As a consequence, game dealers showed farmers how to catch cheetahs with cage traps. From the late 1960s through the 1970s several thousand cheetahs were caught for the world’s zoos (Marker-Kraus et al., 1996). Cage traps typically were set at cheetah marking trees, visited primarily by males; but zoos preferred females. For every female cheetah caught, up to 20 males were captured, many of which were killed by the farmers (Marker-Kraus et al., 1996).
As a result of human development, cheetah numbers are estimated to have dropped from 100,000 in 1900 to 7,100 in 2016 (Durant et al., 2017, Myers, 1975; Chapters 4 and 29429).

1960s—The beginning of knowledge

In the 1960s, the first reports of concern over declining populations of cheetahs emerged when the East African Wildlife Society began an investigation into the species’ status (Graham, 1966). A couple of years later the results of the first study of cheetahs in the wild was published, where George Schaller shared his findings from Tanzania’s Serengeti National Park (Schaller, 1968). His work described the unique hunting style of the cheetah, illuminating traits and strategies (Chapter 9). In 1969, Joy Adamson wrote about raising an orphaned cheetah, which she reintroduced into the wild in Kenya’s Meru National Park (Adamson, 1969). Adamson’s reports included the first close observations of birthing, cub behavior, and development (Adamson, 1972).

1970s—The need for conservation action is recognized

Learning About the Cheetah—Early Research Studies in Africa
In the 1970s, Randal Eaton shared additional insight into cheetah behavior, describing breeding and hunting behaviors, social structure, and prey preferences he observed in Kenya (Eaton, 1974; Chapters 8 and 989). In the Serengeti, George and Lory Frame studied behavioral ecology to evaluate the status and survival of cheetahs. They recorded hunting methods, social interactions, maternal behavior, family structure, mother–cub interactions, coexistence with other predators, and developed a methodology for identifying cheetahs by spot patterns on the face (Chapter 32). Their method of weighing carcasses after meals and noting the portion consumed became the norm for many cheetah feeding ecology studies (Frame and Frame, 1981).
In 1975, Norman Myers was the first to publish about the range wide decline in cheetah due to habitat loss and human-wildlife conflict (Myers, 1975). While there were perhaps 40,000 wild cheetahs in 1960, there were reportedly fewer than 20,000 by 1975, and of those, fewer than 3,000 in Africa’s protected areas (Myers, 1975; Chapter 19). The reduced numbers were attributed to conflict with larger predators inside protected areas and conflict with the growing human populations outside protected areas (Myers, 1975). It was recognized that the mere existence of protected areas was insufficient to guarantee the long-term survival of this wide-ranging carnivore. Myers reported that cheetahs occurred at low densities with a limited distribution in only a small portion of sub-Saharan Africa and was the first to call for action “in the near future to reverse this decline.”
Myers voiced his concerns over a growing African human population and the human disturbances impacting Africa’s wildlife. In particular, carnivores like the cheetah were under pressure due to the potential threat they posed to livestock. In 1975, Africa’s population was 450 million people and growing by 3.5%–4.5% per year, exerting unsustainable pressures on wild lands and wildlife. He reported that livestock farmers in Kenya, Namibia, and Zimbabwe were motivated to engage in cheetah persecution as they were receiving “compensation revenue” through the sales of skins (Myers, 1975). The countries Myers considered having the greatest potential for cheetah conservation initiatives were Botswana, Kenya, Namibia, Tanzania, and Zimbabwe, and sustainable land management was encouraged to balance the needs of wildlife, people, livestock, and the land (Myers, 1975).

Conflict with African Farmers Acknowledged

When CITES (1975) put an end to the export of wild cheetahs, Namibian farmers no longer had a market for trapped cheetahs (Marker-Kraus et al., 1996). As a result, trapped cheetahs generally were killed, but a few were translocated to national parks and game reserves (Marker-Kraus et al., 1996; Chapter 20). To provide care for cheetahs captured in conflict with farmers, the Pretoria Zoo, in partnership with Anne Van Dyk, developed the De Wildt Cheetah and Wildlife Centre in South Africa in 1971 (renamed Anne Van Dyk Cheetah Centre in 2014). In the years to follow, they also became the most successful breeding center in the world, providing captive bred cheetahs to the world’s zoos (Marker-Kraus, 1990; Chapter 22).
Conflict between the farmers and cheetahs continued (Chapter 13). During the 1980s, Namibian game and livestock farmers reported killing over 800 cheetahs per year (CITES, 1992). The problems facing wild cheetahs were brought forward to the US conservation community in 1977, when Marker conducted research in Namibia on cheetah rehabilitation, and learned firsthand about the cheetah and farmer conflict (Marker-Kraus and Kraus, 1990). Cheetahs in South Africa and Zimbabwe were confronted with similar issues (Marker-Kraus and Kraus, 1990), while those in Kenya and Tanzania faced habitat loss, illegal snaring, and poaching (Myers, 1975).

Conflict with African Farmers Acknowledged

When CITES (1975) put an end to the export of wild cheetahs, Namibian farmers no longer had a market for trapped cheetahs (Marker-Kraus et al., 1996). As a result, trapped cheetahs generally were killed, but a few were translocated to national parks and game reserves (Marker-Kraus et al., 1996; Chapter 20). To provide care for cheetahs captured in conflict with farmers, the Pretoria Zoo, in partnership with Anne Van Dyk, developed the De Wildt Cheetah and Wildlife Centre in South Africa in 1971 (renamed Anne Van Dyk Cheetah Centre in 2014). In the years to follow, they also became the most successful breeding center in the world, providing captive bred cheetahs to the world’s zoos (Marker-Kraus, 1990; Chapter 22).
Conflict between the farmers and cheetahs continued (Chapter 13). During the 1980s, Namibian game and livestock farmers reported killing over 800 cheetahs per year (CITES, 1992). The problems facing wild cheetahs were brought forward to the US conservation community in 1977, when Marker conducted research in Namibia on cheetah rehabilitation, and learned firsthand about the cheetah and farmer conflict (Marker-Kraus and Kraus, 1990). Cheetahs in South Africa and Zimbabwe were confronted with similar issues (Marker-Kraus and Kraus, 1990), while those in Kenya and Tanzania faced habitat loss, illegal snaring, and poaching (Myers, 1975).


Cooperative Captive Cheetah Programs Begin
In response to the world’s declining biodiversity, the United States (US) Endangered Species Act (ESA, 1973) and the World Conservation Union’s (IUCN) Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES, 1975) were passed (Chapter 21). As a result, sourcing wild cheetahs for zoo exhibits stopped, and zoos began to collaborate through managed breeding programs to maximize genetic diversity, promote conservation, and educate the public (Chapters 22 and 232223).
By the early 1970s, after returning from Kenya, Eaton began developing safari parks in the United States. His recommendation was for large habitats to foster better environments for breeding cheetahs. With this plan, he helped develop several safari parks in the United States, including the Wildlife Safari in Oregon, where Laurie Marker began working with cheetahs in 1974. Wildlife Safari became one of the first successful breeding facilities in the United States. During the same period, the San Diego Wild Animal Park in California and Whipsnade Safari Park in the United Kingdom also were developed. These facilities were some of the few facilities in the world to begin breeding cheetahs successfully.

Captive Cheetah Management and the Cheetah Species Survival Plan® (SSP)

In 1982, Wildlife Safari hosted the first US national cheetah meeting, bringing together zoos willing to cooperate on breeding. The foundation for managing demographic structure in captive cheetahs began with the first regional cheetah studbook for North America (Marker, 1983), followed in 1988 with the first International Cheetah Studbook (Marker-Kraus, 1990). The studbook is a registry that lists all known animals belonging to zoos and private facilities, thus creating the preconditions for selecting breeding animals. Marker has maintained the International Cheetah Studbook since its inception (Chapter 22).

In 1984, the North American Cheetah Species Survival Plan® (SSP) of the American Association of Zoos and Aquariums (AZA) was developed (Chapter 23). The SSP brought collaborative research and management to the forefront and helped develop comprehensive plans to conserve captive and wild cheetahs.
In 1987, under the leadership of the Cheetah SSP Species Coordinator, Jack Grisham, and Dr. Ulysses Seal, founding chair of IUCN’s Conservation Breeding Specialist Group (CBSG), a national meeting on cheetahs convened. Here, the Cheetah SSP designated the US captive cheetah population as a research population (Grisham and Lindburg, 1989), and the first systematic research plan was designed and implemented. The initial 3-year, multidisciplinary research project provided a basic understanding of cheetah biology and was a critical step in forming conservation strategies for ex situ populations. Its results appeared in a special edition of Zoo Biology (Wildt and Grisham, 1993). Fig. 1.1A shows some of the research collaborators involved in designing and implementing the research plan.

Cheetah Studies in Africa
In 1980, photographer and tour guide David Drummond reintroduced 3 orphaned cheetah cubs in the Maasai Mara National Reserve (Drummond, 2005). The female cub started a long lineage of cheetahs that were resident at the Governor’s Camp area of the Maasai Mara. Drummond was one of the first to present the issues of poaching, wildlife interactions, and challenges of pastoral communities in terms of predators.

Tanzania—The Serengeti Research Project
In 1980, Dr. Tim Caro and his students continued the longitudinal study of cheetah behavior initiated by the Frames in the Serengeti National Park. Today, the research program is overseen by the Zoological Society of London (ZSL) through Dr. Sarah Durant, making it the longest-running cheetah research project. Caro’s team also was interested in understanding wild versus captive behavior and used the Serengeti population to provide baseline data on breeding behavior and mother cub interactions (Chapter 9). Cheetahs of the Serengeti Plains: Group Living in an Asocial Species (Caro, 1994) has been the primary reference book on cheetah behavioral ecology since its publication.
During the same time frame, Hamilton studied the ecology of cheetahs in sub-Saharan Africa. His findings showed that, albeit in low densities, cheetahs were persisting even in areas where they were predicted to be extinct due to rising human populations. He reported cheetahs to be remarkably successful predators, well adapted to coexistence with nomadic pastoralists in arid and semiarid lands over large areas (Hamilton, 1986).



1990s—Population research studies, African conservation programs, health analyses of wild cheetahs, and population viability analyses

Range-Wide Population Studies
At the end of the 1980s there was little understanding of the population distribution or the ecology and biology of healthy, free-ranging cheetahs outside of protected areas. In the 1990s, Paula Gros undertook cheetah population surveys in East Africa (Gros, 1996, Gros, 1998, Gros, 2002, Gros and Rejmanek, 1999) and provided a rich insight into population distribution. During the same period, the cheetah’s global status was summarized using country specialist information gathered in the 1980s from all range countries (Marker, 1998).

The Beginning of African Conservation Programs

Namibia—The Cheetah Conservation Fund (CCF)
In response to accumulating evidence about the cheetah’s habitat and population decline and to gain knowledge on cheetahs in nonprotected areas, Laurie Marker founded the Cheetah Conservation Fund (CCF) in 1990 and set up an international field research and education center along with a model farm in Namibia. Marker began working with Dieter Morsbach, research scientist from the Namibian Ministry of the Environment (MET), and the livestock farming communities who were trapping and killing high numbers of cheetahs each year (Chapter 13). In 1991 an extensive survey of Namibian rural farming communities provided a better understanding of the threats to the cheetah and the techniques employed to prevent livestock depredation by cheetahs (Marker-Kraus et al., 1996, Marker et al., 2005). This community survey—along with health, disease, reproduction, genetic, and ecological surveys on Namibian cheetahs—provided the groundwork for many cheetah-range country programs (Marker et al. 2010).
CCF also launched national and international outreach and education programs to raise awareness of the cheetah’s vulnerable status (Marker and Boast, 2015; Chapter 18). In 1994, CCF launched the first of many African livestock guarding dog programs in cooperation with Dr. Ray Coppinger and the Livestock Guarding Dog Association of America (Chapter 15).

South Africa—Cheetah Outreach
In 1997, Cheetah Outreach (CO), near Cape Town, was founded by Annie Beckheling and Mandy Schumann. Launched as an educational cheetah encounter facility, its focus is conserving South Africa’s wild cheetahs. Adapting CCF’s educational outreach programs, Cheetah Outreach began to educate school children and the public and developed a cheetah ambassador program (Chapter 28) working closely to develop nutritional guidelines for captive cheetahs (Chapter 26). In addition, they developed a livestock guarding-dog program and work closely with livestock farmers to help reduce conflict.

Systematic Biobanking and Reproductive Research
In 1994, an international research collaboration between the Namibian MET, a SSP research team, and CCF initiated the investigation of links between reproductive traits, nutrition, and diet. CCF’s Genome Resource Bank, which included the first “field banked” sperm samples, was initiated (Crosier et al., 2007; Chapter 27). At the same time, reproductive research on cheetah males was being conducted at a few zoos in the United States (Wildt and Grisham, 1993) and in South Africa at the De Wildt Cheetah and Wildlife Center (Bertschinger et al., 2008; Chapter 27). Data on the basic biology of the female cheetah took longer to complete, with the first results in 2011 bringing insight into ovarian development and reproductive cycling (Crosier et al., 2011, Wachter et al., 2011; Chapter 27).
In 1996, Dr. Linda Munson, cheetah SSP veterinarian pathologist, trained Namibian veterinarians and field biologists in systematic sample collection and assisted in developing sample collection research protocols that led to long term, collaborative global disease studies (Chapter 25). Further training brought together pathologists from the United States, Europe, and South Africa.
Population, Habitat, and Viability Analysis (PHVA)
In 1996, the cheetah SSP sponsored a Population, Habitat and Viability Assessment (PHVA; Chapter 38) for the Namibian Cheetah and Lion. It was hosted by CCF and gathered the IUCN CBSG, the IUCN Cat Specialist Group, the MET, and members of the Cheetah SSP. The workshop provided a platform for Namibian farmers, wildlife officials, international scientists, and other stakeholders and set forth a strategy for managing cheetahs in Namibia while addressing issues affecting neighboring cheetah-range countries (Berry et al., 1997). The first comprehensive conservation plan for managing the wild Namibian cheetah (Berry et al., 1997), and the first National Cheetah Plan followed (Nowell, 1996).

2000s—Cheetah conservation programs, range-wide workshop, and programs, off the beaten track

Expansion of Cheetah-Specific Conservation Programs Across the Cheetah’s Range
The new millennium saw the development and expansion of cheetah conservation programs and new areas of research.

Kenya—CCF, Kenya/Action for Cheetah, Kenya (ACK)
In 2001, the Kenyan government and its citizens voiced concern about the decline of their cheetah population. The decline was attributed to a reduction in wild prey caused by poaching and the transition from large, collective ranches to smaller farms (Chapter 11), and also to habitat fragmentation (Chapter 10). Although cheetahs had been photographed extensively in the Maasai Mara (Ammann and Ammann, 1984, Scott and Scott, 1998), there was no cheetah conservation work being undertaken in the country. To address the void, Mary Wykstra and Laurie Marker developed CCF Kenya in 2001, which later became Action for Cheetahs in Kenya (ACK).

Iran—Iranian Cheetah Society (ICS)
In 2001, Mohammad Farhadinia, Kaveh Hobeali and Morteza Eslami founded the Iranian Cheetah Society (ICS), an NGO at the forefront of cheetah conservation in Iran (Chapter 5).

Zimbabwe—Cheetah Conservation Program and Cheetah Conservation Project
In 2002, following long-term cheetah studies from Vivian Wilson, in Zimbabwe, a cheetah conservation program was developed under the leadership of Netty Purchase, the carnivore project coordinator from the Marwell Zimbabwe Trust, and Verity Bowman from the Dambari Trust. In 2012, Dr. Esther van der Meer launched the Cheetah Conservation Project Zimbabwe.

Namibia—Cheetah Research Project
In 2002, the Leibniz Institute of Zoo and Wildlife Research (IZW) from Germany established the Cheetah Research Project in the southeast of Namibia under the direction of Dr. Bettina Wachter.

Botswana—Cheetah Conservation Botswana (CCB)
In 2004, Rebecca Klein, Anne-Marie Houser, and Dr. Kyle Good established Cheetah Conservation Botswana (CCB) with the assistance of the Mokolodi Wildlife Foundation and CCF. Their first research camp, at Jwaneng Diamond Mine’s game reserve, monitored the reserve’s cheetah population, and conducted outreach, education, and research on nearby farmland. CCB developed an administration and education base at Mokolodi Nature Reserve near Gaborone, followed by a field station, model farm, and education center on farmland near Ghanzi.
Carnivore Projects
In addition to the afore-mentioned cheetah projects, several carnivore projects have put a strong emphasis on cheetah conservation. These programs include: Endangered Wildlife Trust (EWT, South Africa), The Africat Foundation and N/a’an ku sê (Namibia), Botswana Predator Conservation Trust, Tanzanian Wildlife Research Institute, and Ruaha carnivore project (Tanzania).

ange-Wide Workshops and Collaboration

In 2001, the Cheetah SSP joined with in situ cheetah conservation efforts and sponsored the first Global Cheetah Action Plan Workshop in cooperation with the CBSG, CBSG South Africa, and CCF. Over 50 people from 11 countries attended the workshop in South Africa presenting on in situ and ex situ cheetah research (Fig. 1.1B). Working groups convened to discuss census research, protection of cheetahs outside protected areas, education and communication, and health and viability of the ex situ population. The findings provided the basis for the first Global Cheetah Action Plan that helped link research initiatives and enhance collaborations (Bartels et al., 2002a). This group called itself the “Global Cheetah Forum” (GCF).

Keeping the momentum, a second GCF took place in 2002. Participants included collaborators working on the Asiatic cheetah and members of the IUCN Cat Specialist Group. The highest priority was completing a census of free-ranging cheetahs to determine how and where range-wide conservation efforts could be implemented (Bartels et al., 2002b). In addition, the Forum members determined that conservation education and training programs should continue to be a top priority in range countries (Chapter 18).

Following this meeting in 2002, cheetah conservation organizations and representatives of the South African farming community met formally for the first time and developed the National Cheetah Conservation Forum (NCCF). Led by a team from the De Wildt Cheetah and Wildlife Centre, EWT, several universities, the National Research Foundation, the Agricultural Research Council and other governmental institutions, new research and conservation initiatives were launched, including a census of the South African cheetah populations.

Part of South Africa’s approach to reducing conflict with livestock and game farmers was to begin a large-scale reintroduction program into private game reserves (Chapter 20). The cheetah populations on these game reserves needed to be artificially connected through animal movement. To achieve this, the cheetah metapopulation strategy was launched in 2009, and is managed by the EWT. In addition, these private reserves allowed for additional ecological studies on cheetah in these protected areas (Chapter 8).

The next GCF meeting took place in 2004, in the Serengeti in Tanzania with over 30 delegates from 7 countries (Fig. 1.1C). The SSP-sponsored “Cheetah Census Technique Development Workshop” was hosted by Dr. Sarah Durant from the ZSL and Wildlife Conservation Society (WCS). Census techniques for acquiring reliable, quantitative information on cheetah distribution and their numbers across Africa were discussed, and a cheetah census technique manual was developed to standardize “best practice” guidelines (Bashir et al., 2004).

Building on this collaboration, in 2005 a southern African Cheetah Regional Workshop brought over 30 people from 6 countries to CCF in Namibia (Dickman et al., 2006). This workshop was facilitated by IUCN CBSG southern Africa and moderated by cochair of IUCN’s Cat Specialist Group, Dr. Christine Breitenmoser. The aims were to assess and evaluate accomplishments in the southern African region and to set new objectives. Key determinations appeared in a special issue of Cat News (Breitenmoser and Breitenmoser, 2007).
Following the 2005 workshop, the Cat Specialist Group in Switzerland undertook the Cheetah Compendium (http://www.catsg.org/cheetah/20_cc-compendium/index.htm). It is a web-based communication tool that houses a library of information, data, documents, maps, and other material relevant to the conservation of the cheetah.

Range Wide Cheetah Program

The Range Wide Conservation Program for Cheetah and African Wild Dogs (RWCP) was established in 2007 through a collaboration of Canid and Cat Specialist Groups of IUCN, and led by Drs. Sarah Durant and Rosie Woodruff. Under this program, regional cheetah workshops for southern and eastern Africa took place in 2007 (IUCN/SSC, 2007a, IUCN/SSC, 2007b; Figs. 1.1D and 1.2A ), with initial meetings for central, northern, and western Africa in 2012 (IUCN/SSC, 2012; Fig. 1.2B) and a follow up meeting for southern Africa in 2015 (RWCP and IUCN/SSC, 2015; Fig. 1.2C). Range-wide priority conservation plans were developed with government officials for both the cheetah and African wild dog, drawing on the similarities of these species’ conservation requirements (Chapter 39).

Drawing on regional plans, national workshops developed country-specific plans in many cheetah-range states (Chapter 39). These workshops led to increased government awareness and support throughout the cheetah’s range—as well as ongoing census research allowing for mapping of cheetah populations by national and regional experts—and an understanding of the threats the species was facing and will likely face in the future (Durant et al., 2017).
The lack of local capacity was a key finding of regional plans. To address this challenge, CCF, in cooperation with the Howard G. Buffett Foundation and the RWCP, trained more than 300 government wildlife officials, university professors, scientists, conservation managers, conservation NGO officers, and community extension officers from 15 cheetah-range countries between 2008 and 2011. The aim of the courses was to teach research techniques (Chapters 29–3829303132333435363738) and to promote a unified and systematic approach to cheetah research and conservation (Marker and Boast, 2015).

Off the Beaten Track—Other Cheetah Populations


Iran
Cheetahs were known to still be present in Iran after the Iranian Revolution ended in 1979 (Joslin, 1984), although little information was available on their population size or distribution before 2000. The current population is estimated at less than 50 adult and adolescent individuals (Durant et al., 2017; Chapter 5).
In 2001 the Iranian government arranged two separate meeting one headed by Dr. George Schaller from the Wildlife Conservation Society (WCS) and the other headed by Dr. Laurie Marker from the CCF to discuss options to save Iran’s small population of Asiatic cheetah (Acinonyx jubatus venaticus). Attendees included government officials from the Department of Environment (DOE), the United Nations Development Program (UNDP) and various Iranian researchers involved with the Iranian cheetah. The outcome of the meetings provided support for Iran to begin working under a multiyear UNDP grant aimed at saving the critically endangered Asiatic cheetah population.
Two important meetings followed. In 2004, the Iranian Centre for Sustainable Development (CENESTA) hosted an International Workshop on the Conservation of Asiatic Cheetah, with participation by Asiatic cheetah conservation partners and local communities to discuss conservation strategies with stakeholders throughout the cheetah’s Iranian range (Fig. 1.2D); and in 2010, an Iranian Cheetah Strategic Planning meeting reviewed the previous decade of work and planned Iranian’s cheetah survival strategies for the following 5 years (Breitenmoser et al., 2010). An overview of the Iranian cheetah situation is found in Chapter 5.


NW Africa—Algeria
The first survey of cheetahs in Algeria was undertaken in 2005 in the Ahaggar National Park, Central Sahara (Busby et al., 2009, Wacher et al., 2005). Interviews with nomadic herders helped assess the nature of interactions between nomads, cheetahs, and other wildlife. A year later, the North African Region Cheetah Organization (Observatoire du Guépard en Régions d’Afrique du Nord (OGRAN) met in Tamanrasset, Algeria, for a 3-day conference to discuss conservation strategies in Algeria highlighting data collection, census techniques, training and education needed to conserve this critically endangered cheetah population. As a result, PhD fieldwork began in the Ahaggar National Park, collecting the region’s first camera trap evidence of cheetahs (Belbachir et al., 2015).

India
The cheetah disappeared from India in 1956 (Divyabhanusinh, 1999). For some time, there have been discussions on reintroducing cheetahs to India (Ranjitsinh and Jhala, 2010). In 2009, the Wildlife Trust of India (WTI), headed by Chairman Dr. M.K. Ranjitsinh, hosted a team of experts including representatives from the IUCN (including its Cat Specialist Group, Reintroduction Specialist Group, and Veterinary Specialist Group), Oxford University’s WildCRU, Cheetah Outreach, and CCF, along with Indian authorities and forestry directors from various regions.
They concluded the following: The original cause of the extinction of the cheetah in India had been adequately addressed; a network of protected areas had been established; and effective wildlife legislation, change in the conservation ethos, and nationwide awareness could lead to a successful cheetah reintroduction (Ranjitsinh and Jhala, 2010). However, the reintroduction project has been stalled indefinitely due to political issues concerning what subspecies could or should be used (Laing and Nelson, 2012, O’Brien, 2013).

International Attention to Illegal Wildlife Trafficking of Cheetahs—UAE/North Africa

By 2006, it was evident that there was need for a long-term plan for combatting illegal wildlife trafficking and for cheetah conservation awareness in Ethiopia. The Ethiopian Wildlife Conservation Authority assigned a task force to develop guidelines and recommendations for a sanctuary for wild orphan animals, and in 2010, the Born Free Foundation in Ethiopia built a sanctuary to hold cheetahs, lions, and other confiscated animals.

The first solicited report on illegal trade was compiled by Nowell to CITES in 2015 (Chapter 14). In 2016, at the CITES Convention of the Parties (CoP17), several resolutions were accepted to work toward the reduction of supply and demand for illegally trafficked cheetah cubs (CITES, 2016; Chapter 14). Work continues between the Horn of Africa and the Gulf States.
Go to:
2015 Onwards—the recent years
In 2015, the AZA launched their Saving Animals From Extinction (SAFE) program to focus on conservation of the cheetah along with nine other species. The goal of SAFE is to restore healthy populations in the wild by connecting scientists with stakeholders and to identify threats, launch action plans, find new resources and engage the public (Chapter 23).
In mid 2015, the RWCP met again to update the southern African regional cheetah plan and population maps (RWCP and IUCN/SSC, 2015). This time the meeting was supported by SAFE with representatives from the AZA community, IUCN SSC Cat Specialist Group, seven national governments, and a multitude of cheetah research and conservation NGOs.
In December 2016, a 54 coauthored paper was published presenting the current estimate on cheetah numbers and distribution (Durant et al., 2017; Chapter 39). It highlighted that the cheetah populations continue to decline in range and number in addition to the need for further investment into their conservation. With 77% of the remaining 7100 adult and adolescent free-ranging population found outside protected areas, it is being recommended the species be uplisted to endangered status on the IUCN red list (Durant et al., 2017; Chapter 39).

Conclusions
The survival of the cheetah needs to be the responsibility of everyone, not just governments and conservationists. In light of largely human-caused global changes in the environment (Chapter 12), people have the responsibility to help ensure the availability of wide-ranging spaces for cheetah conservation (Chapter 17). This is only possible if the livelihoods (Chapter 16) of local populations living in the same habitat as the cheetah are also taken into consideration in a holistic conservation approach. We hope that the passion and cooperative efforts of active cheetah conservationists will be augmented by the world at large—everyday people who care about wildlife and cheetahs—to help the cheetah reverse indefinitely its fragile march toward extinction. The last chapters of this book Chapters, 39 and 403940) will help define the way forward to help secure a future for the cheetah.

https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7150087/ Reply

[Acinonyx sp.]

Cincinnati’s cheetah brothers turn 17 – perhaps oldest cheetahs in the world

CINCINNATI —

The Cincinnati Zoo celebrated the birthdays of two of its cheetahs Monday.

Cheetahs Chance and Bravo celebrated their 17th birthdays and the Cincinnati Zoo believe this makes them the oldest cheetahs in the United States and quite possibly the world. According to the zoo, cheetah live an average lifespan of 10 to 12 years, so Chance and Bravo are now well above that average.

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Zoo officials say the two cheetah's longevities can be attributed to the special geriatric enrichment, diet and tender, loving care they receive at the Cincinnati Zoo.

Born at the DeWildt Breeding Center in South Africa in 2004, Bravo and Chance came to Cincinnati when they were six months old. They remain a coalition here, as brother cheetahs often stick together in the wild, and are the zoo's only cheetahs housed together. They spend more time in the zoo's Africa exhibit yard than the other cheetahs.
Cheetahs are endangered and their populations continue to decline worldwide. The Cincinnati Zoo is proud to be called "The Cheetah Capital of the World" because of its conservation efforts and education for cheetahs in its care.

https://www.wlwt.com/article/cincinnatis-cheetah-brothers-turn-17-perhaps-oldest-cheetahs-in-the-world/36323551# Reply

[Acinonyx sp.]

Optimal hunting conditions drive circalunar behavior of a diurnal carnivore

Foraging requirements and predation risk shape activity patterns and temporal behavior patterns widely across taxa. Although this has been extensively studied in small mammals, the influence of predation and prey acquisition on the activity and behavior of large carnivores has received little attention. The diurnal activity described as typical for cheetahs (Acinonyx jubatus) has been explained in terms of their avoidance of antagonistic interactions with other larger predators. However, a recent study revealed that cheetahs are frequently active at night, especially during periods of full moon. Being both predator and “prey” in an environment with comparatively high densities of larger and competitively dominant nocturnal predator species, we investigated whether cheetah nocturnal behavior could be explained by favorable conditions for 1) predator avoidance or 2) prey acquisition. We used a data set of continuously recorded behavior created using machine-learning techniques on behavioral data collected in the field to transform recorded 2D activity values from radio-collars into 3 distinct behavioral states (feeding, moving, and resting). We found that 32.5% of cheetah feeding behavior occurred at night and that, in the dry season, nocturnal feeding behavior was positively correlated with moonlight intensity. Our results suggest that nocturnal and circalunar behavior of cheetahs is driven by optimal hunting conditions, outweighing the risks of encountering other predators. Using novel methodology, the results provide new insights into the temporal distribution of behavior, contributing to our understanding of the importance of moonlight and season on the behavior patterns of diurnal species.

https://www.researchgate.net/publication/264554420_Optimal_hunting_conditions_drive_circalunar_behavior_of_a_diurnal_carnivore Reply

[Acinonyx sp.]

Timed running speed of a cheetah (Acinonyx jubatus) 

Introduction 
The cheetah is generally believed to be the fastest running animal, but its maximum speed has been uncertain. A widely-quoted speed of 71 miles per hour (32ms-’) has been discredited: a tame animal was claimed to have run 80 yards (73 m) in 2.25 s, but the enclosure in which the run was made was later shown to be only 65 yards (59 m) long (Hildebrand, 1959, who also remarks that the time is very imprecise, and that there is an error of arithmetic). Hildebrand’s own (1961) estimate of 56 miles per hour (25ms-’) does not come from a timed run over a measured distance, but was obtained by analysing film, taking the scale of distance from an estimated length of the animal’s body. The measurements reported in this paper were made in 1965, but are published only now because the author was not aware that previously published records were unreliable.

Method 
The subject was an adult female cheetah weighing approximately 35 kg, which had been orphaned as a cub, and reared on Kenplains Farm, Athi River, Kenya, by Dr David Hopcraft. She was returned to the wild, but visited Kenplains occasionally. The measurements were made on one of these visits, between 10:00 h and 11 :30 h, on firm level ground with a slight covering of murram dust. There was insufficient wind to move wool hanging vertically from a post. A 220 yard (201.2 m) course was measured, with a surveyor’s tape, and marked with 2 posts at each end. The posts at the start had a taut length of white wool tied between them, approximately 0.45 m from the ground. A clear line was drawn between the posts at the finish. The author, who undertook the timing, was then a current athletics coach well used to hand-timing. He used an analogue stopwatch (Omega), reading to 0.1 s, which had been calibrated against 2 stopwatches in The Voice of Kenya broadcasting house, and 2 stopwatches of the Kenya Amateur Athletics Association. Each trial was started with the cheetah held 18m behind the start line. A running start was decided upon as hunting cheetah launch into their major sprint from a moving start, and sustained maximum running speed over a reasonable distance was the desired parameter. An open Landrover stood, with engine running, about 75 m down the course. The stopwatch operator stood in the back of the Land Rover holding a piece of meat which had been shown to the cheetah. On a command, the cheetah was released, and when the timer saw her break the wool at the start he activated the stopwatch. The Land Rover accelerated down the course and stopped after the finish line, when the meat was thrown down. The watch was stopped at the instant when the cheetah’s anterior thorax was deemed to have crossed the finish line. Three runs were made, with successive runs in opposite directions. A 30 min rest was allowed between trials. 

Results and discussion 
The recorded times were 7.0, 6.9 and 7.2 s. The mean time of 7.0 s gives a speed of 29 m s-' (64 miles per hour). The only other large animals for which accurate, reliable speeds have been published are greyhounds and racehorses. Data from the sporting pages of newspapers show that most greyhound races are won at 15-16ms-', and most horse races of less than one mile (1.6km) at 16-17m s-'. The highest horse speed recorded by Matthews (1994) is 19.2ms-' over 402m. The times used to calculate these speeds were all measured from a standing start, so are not strictly comparable with the speed of 29 m s-' for the cheetah. Nevertheless, it is clear that the cheetah is very much faster than horses or dogs. Garland (1983) quotes a speed of 20 m s-' or more for 17 species of mammal. Apart from the cheetah (the discredited record discussed in the Introduction), these are Vulpes fulva, two species of Lepus and 13 ungulate species. For the great majority of these, there is no information about the method of measuring speed. Indeed, the speeds may be mere subjective estimates. In six cases the speeds are recorded as being speedometer readings, but these cannot be accepted as reliable unless the animal is running close to the vehicle, parallel to it in a straight line, and the speedometer calibration has been checked. There is no assurance that these conditions were ever met. Elliot, Cowan & Holling (1977) filmed lion attacks on various prey. Triangulation from two cameras, or a grid marked on the ground, enabled them to measure speeds. They measured each animal's speed at intervals as it accelerated and estimated maximum speed by fitting a curve to the data. They obtained a value of 26.5 m s-' for Gazella thomsoni (and much lower speeds for other animals). Unfortunately, they show no points on their graphs of speed against time, so we cannot tell whether this speed was actually observed or is merely an extrapolation from the earlier stages of acceleration. Thus the speed of 29 ni s-', recorded for the cheetah in this paper, is by far the highest reliablyrecorded running speed for any animal. Reply

[Acinonyx sp.]

Spatial ecology of free-ranging cheetah (Acinonyx jubatus) and its applications to mitigate the farmer-cheetah conflict in Namibia

Abstract:
The global distribution of the cheetah (Acinonyx jubatus) has decreased dramatically during the past decades. Cheetahs are currently confined to only 9% of their former range. Approximately 77% of the cheetah range lies outside protected areas, frequently exposing cheetahs to conflict with people. Southern Africa represents with approximately 4,000 individuals the stronghold of the global cheetah population which comprises approximately 7,100 individuals. Namibia hosts approximately 1,500 individuals, which together with the cheetahs in Botswana form the largest connected population worldwide. This population is threatened because most of these cheetahs roam on livestock farms and are persecuted by farmers. From a conservation point of view it is therefore of utmost importance to develop non-lethal mitigation strategies to reduce this long-lasting farmer-cheetah conflict. This dissertation thesis was conducted to use long-term data of the Cheetah Research Project (CRP) of the Leibniz Institute for Zoo and Wildlife Research (IZW) in Berlin, Germany, to develop and test such a mitigation strategy in Namibia. In chapter 5 (publication one), I investigated the socio-spatial organization of free-ranging cheetahs on commercial farmland in central Namibia. Although cheetahs have been studied in several areas in the world for decades, their socio-spatial organization had not yet been clarified. The most comprehensive study was conducted in the 1990s in the Serengeti National Park (NP) in Tanzania and described a unique social organisation in mammals. Adult males either defended small territories separated from each other by some distance or roamed in large home ranges that encompassed several territories. The latter males are termed “floaters” and regularly visited the territories within their home ranges. Females also roamed in large home ranges which encompassed several territories but stayed mainly in the area between territories. Both territory holders and floaters can be solitary or occur in coalitions of two to three males. Subsequent studies elsewhere did not recognise such socio-spatial organization because of the method by which they categorized the individuals for data analysis. In this chapter I analysed the movement data of 133 males and 31 females to demonstrate that the socio-spatial organization of cheetahs described in the Serengeti NP also exists in Namibia. Several predictions were derived from the social organisation described for the Tanzanian cheetahs and tested with the data of the CRP. Consecutively I re-analysed published data of previous studies and could confirm the two tactics also in these datasets. Territory holder have preferred access to females, floaters heavily fight for territories, and the pattern can be found in all studies populations. Therefore I conclude, that this behavior may be a general trait of the species. In chapter 6 (publication two) we investigated the consequences of this spatial system for camera trap studies and capture-recapture models that estimate abundance and density of animal populations. Such models require high capture and recapture rates and a homogeneous detection probability of all individuals at camera trap stations. Capture and recapture rates were highest at the marking trees of territories, where territory holders frequently marked and which were regularly visited by floaters and irregularly visited by females. This meant that the detection probability at marking trees differed strongly between territory holders, floaters and females. Thus, the assumptions of most capture-recaptures models were violated in previous studies that provided abundance and density results of cheetahs by applying such techniques and are therefore likely to be biased. Chapter 6 tested the performance of four types of capture-recapture models whose assumptions permitted heterogeneity in the detection probability and compared the estimated abundance with the true abundance of cheetahs in the study area. This revealed two best suited models for the socio-spatial organization of cheetahs with one being favorable if spatial tactic is not known a priori. The results matched with the known abundances. On the basis of this information, I looked in chapter 7 (manuscript) at the local density and activity of cheetahs within the landscape and in relation to human-wildlife conflict and farm management. Some farms overlapped with a cheetah territory, others did not. The marking trees in the core area of cheetah territories had a substantially higher cheetah activity than other areas. We termed these core areas of the territories “cheetah communication hubs”, because they play an important role in the social system of cheetahs. From the perspective of the farmers, these hubs are local hotspots of predation risk for the cattle. We used an experimental approach to demonstrate that farmers who stationed their suckler cows and calves in such a hub suffered substantially higher losses of calves than farmers who had stationed their breeding herds far away from such a hub. This discovery was used to develop a livestock management plan for farmers overlapping with a cheetah hub. When their breeding herds were shifted away from the hub, their losses decreased substantially. This is because the hub did not shift, nor did the cheetahs conduct excursions to pursue and hunt the calves. Instead, they preyed on the local, naturally occurring wildlife prey. The mitigation strategy presented and tested here is therefore a highly effective and sustainable solution to reduce the farmer-cheetah conflict in Namibia and potentially also elsewhere.

https://refubium.fu-berlin.de/handle/fub188/27339 Reply

[Acinonyx sp.]

Cheetahs: Biology and Conservation: Biodiversity of the World: Conservation from Genes to Landscapes (Chapters 1-9)

https://books.google.ca/books?id=H3rXDgAAQBAJ&pg=PA451&dqhl=en&sa=X&ved=2ahUKEwjtgNHI287tAhVGGs0KHSb7Ak8Q6AEwAXoECAUQAg#v=onepage&q&f=false

All other chapters can be found by searching in google scholar: google scholar
 
It contains amazing information on cheetahs. Reply

[Acinonyx sp.]

Wild cheetahs in Sir Bani Yas Island, UAE!!!!

http://www.cnn.com/2010/WORLD/meast/04/14/cheetah.cubs.uae/index.html


Tracking cheetahs on Sir Bani Yas Island 

Cheetah cubs

*This image is copyright of its original author

Reply

[Acinonyx sp.]

Amazing cheetah research paper

Understanding the Role of Semiochemicals on the Reproductive Behaviour of Cheetahs (Acinonyx jubatus)—A Review

Simple Summary
This review aims to provide an in-depth overview of the reproductive physiology and behaviour of cheetahs (Acinonyx jubatus). Specifically, it focuses on the role that pheromones (a class of semiochemicals) play by directly affecting the reproductive (e.g., precopulatory and copulatory) behaviour. Furthermore, it aims to critically analyze current research and provide new insights on study areas needing further investigation. It is clear, for instance, that further research is necessary to investigate the role of semiochemicals in the reproductive behaviour of cheetahs in order to rectify the current behavioural difficulties experienced when breeding younger females. This, in turn, would aid in improving captive breeding and the prevention of asymmetric reproductive aging.

Abstract
The cheetah species (Acinonyx jubatus) is currently listed as vulnerable according to the International Union for Conservation of Nature (IUCN). Captive breeding has long since been used as a method of conservation of the species, with the aim to produce a healthy, strong population of cheetahs with an increased genetic variety when compared to their wild counterparts. This would then increase the likelihood of survivability once released into protected areas. Unfortunately, breeding females have been reported to be difficult due to the age of these animals. Older females are less fertile, have more difficult parturition, and are susceptible to asymmetric reproductive aging whereas younger females tend to show a significantly lower frequency of mating behaviour than that of older females, which negatively affects breeding introductions, and therefore mating. Nonetheless, the experience from breeding methods used in some breeding centres in South Africa and the Netherlands, which also rely on the role that semiochemicals play in breeding, proves that cheetahs can be bred successfully in captivity. This review aims to give the reader an in-depth overview of cheetahs’ reproductive physiology and behaviour, focusing on the role that pheromones play in this species. Furthermore, it aims to provide new insight into the use of semiochemicals to improve conservation strategies through captive breeding.

1. Introduction
With less than 7100 cheetahs left in the world [1] and an overall classification as “vulnerable with a decreasing population trend” [2], the cheetah is one of many species in need of conservation efforts [1]. During the last ice age, the cheetah species almost went extinct [3] and experienced a “population bottleneck” that led to reduced genetic diversity, followed by high levels of inbreeding [4]. As reported by Frankham [5], the International Union for Conservation of Nature (IUCN) considers genetic diversity as “one of the three levels of biodiversity requiring conservation”. Therefore, captive breeding aims to create a captive population that would act as a vital genetic reservoir in the case of another sudden, unforeseen loss of wild populations [6]. Besides the fact that reproduction is a crucial part in ensuring the conservation and survival of a species [7], the main aim of captive breeding is to produce a population of cheetahs with a higher genetic variety when compared to their wild counterparts. This has become a necessity in creating a sustainable, healthy population of cheetahs that would survive after possible reintroduction into protected wild areas [8,9]. Therefore, studies focusing on increasing natural reproduction in supervised populations are crucial to obtaining a self-sustainable cheetah population.
Research advancement in animal behaviour related to reproduction, raising of offspring, breeding, and communication, including that of chemical communication (the predominant form of interaction between many organisms) [10], is crucial for the conservation of numerous endangered animals [11].
Cheetahs rely on their advanced olfactory systems for chemical communication [12] which contributes largely to regulating animal behaviour [13], including sexual receptivity [10]. Chemical communication occurs through volatile chemical signals and/or non-volatile chemical signals. The former is generally fixed by lipids, released into the air, and detected as an odour, whereas non-volatile chemical signals are transferred to the vomeronasal organ of an animal’s olfactory system [14,15]. The odours that are released bind to corresponding receptors on the sensory neurons to stimulate a response; thus, the shape and structure of the chemical compounds are important in the discrimination of different odours [12]. The release and reception of chemical signals (semiochemicals) from one animal to another is involved in a very intricate system from initial detection to final response [16] and contributes largely to regulating animal behaviour [13], including sexual receptivity [10].
Therefore, this review aims to provide the reader with an overview of the cheetah’s characteristics and, in particular, on its reproductive behaviour focusing on the role that semiochemicals play in this species. Furthermore, it aims to provide new insight into the use of semiochemicals to improve conservation strategies through captive breeding.
2. The Cheetah
Well-known for its speed, the cheetah (Acinonyx jubatus) is a medium-sized feline and the only member of its genus [17]. In the past, the cheetah could be found throughout Asia and Africa. Today, however, they can only be found in Iran and sub-Saharan Africa in grassland and savannah habitats, with larger populations in areas with an abundance of prey and a large stretch of land [12].
In the 1900s, the cheetah population was composed of over 100,000 individuals worldwide. Today, less than 7100 cheetahs are left [1], with a 76% loss in Africa and a reduction of the Asian population to very small numbers [6]. The Asian population is therefore listed as critically endangered [6] and the overall cheetah population is listed as vulnerable with a decreasing population trend [2]. The decreasing trend emphasizes the need for more effort in conserving the species [6].

2.1. Threats

Some of the leading causes relating to the endangerment of many wild cat species, including tigers and leopards, are the disturbance humans have on the environment [18,19,20], along with harmful human activities such as poaching and climate change [11,17]. With regards to cheetahs, the increasing human population and, therefore, the utilization of more land for activities such as farming [21] has led to the fragmentation or total loss of habitat [17] and has resulted in a total loss of 91% of historic range since the 1900s [1]. Competition, therefore, arises between farmers and cheetahs, where cheetahs are killed because they are perceived as pests and a threat to livestock [12,22].

As an intermediate predator, a cheetah faces a great deal of competition from larger predators [23]. Unfortunately, with cheetahs being built solely for speed there is the disadvantage of insufficient physical strength to protect its offspring, territories, and prey from stronger predators, predominantly lions and hyenas [24,25,26,27]. Therefore, a decline in habitat would cause an increase in the abundance of these predators in certain areas, thus causing cheetahs to avoid specially protected areas that serve as a refuge for most large predators [12].

2.2. Possible Reasons for Population Decline

Generally, felids have small litter sizes (1–5 with an average of 3) because females usually have no aid from males or other females in hunting, providing and caring for their young, and later teaching their offspring how to hunt [25]; in contrast, cheetahs have large litter sizes (1–8 with an average of 4) [22,27], with low birth weights (approximately 380–700 g) [27] and high growth rates. These traits, along with the swift return to oestrus after the loss of an unweaned litter of cubs, have been speculated to have evolved as a response to the high mortality rate of cheetah cubs in order to increase survival rate [23].

Studies evaluating cub mortality and possible causes, showed that the most common causes of death were predation [23,28], abandonment [23] starvation and injury [28]. Unfortunately, cheetah cub survival in predator-free, unprotected areas, is not much greater than in protected areas containing large predators [28] because cheetahs inhabiting unprotected areas are more prone to human conflict, including the illegal pet trade [22].

In captivity, the highest amount of cub deaths is attributed to bad mothering ability [29]. In the wild, the ability to find and obtain prey plays a primary role in the abandonment of cubs by their mothers [23]. In captivity, however, it could be speculated that a contributing factor most likely responsible for the high cub mortality could be related to the mother’s genetics.

Several studies have proved that mothering ability is a genetically linked trait, and it is essentially related to the rate of offspring survival before weaning [30,31,32,33,34]. While it is possible to select for good mothering ability in a captive breeding program, selective breeding is difficult for species with a low genetic diversity [35].

Cheetahs population is said to have undergone a genetic bottleneck that severely reduced genetic diversity [36], followed by constant inbreeding [4]. It has been known for a long time that inbreeding reduces survival and reproduction in a species, including mothering ability, sperm production, and adult and juvenile survival [5].

2.3. Reproductive Characteristics

Female cheetahs reach sexual maturity at 20–24 months of age [1] and have a considerably short oestrous cycle if compared to those of other large felids, which lasts 20–30 days [24]; in fact, the duration of the oestrous cycle is between 7–21 days, with oestrus lasting between 2–6 days on average and inter-oestrous periods lasting for 13.9 ± 0.7 days [17,37].

Cheetahs are induced ovulators [9], meaning that frequent copulation contributes in maintaining high levels of oestrogen, and, in combination with the low levels of progesterone, stimulates the anterior pituitary to release luteinizing hormone (LH) and trigger ovulation [38,39,40,41,42].

Female cheetahs are also polyoestrous; therefore, oestrus and breeding occur throughout the year [9,12,24]. In the wild, however, it has been seen that females “decide” when it is the right time to come on oestrus [12] and are therefore thought of as being seasonally polyoestrous [22]. In captive breeding facilities, with no environmental pressure, reproduction and oestrous onset can occur at any time throughout the year [6]. At 6 years of age, females reach maximum reproductive capability, which continues until they are 8 years old and declines thereafter [22].

Unfortunately, female cheetahs are susceptible to asymmetric reproductive aging (ARP); therefore, the older a female is when she has her first litter, the shorter her reproductive lifespan will be. This occurs due to continuous oestrous cycling, where the fluctuating levels of oestrogen cause the reproductive organs to age at a faster rate [8]. Failure to reproduce after a long period of not reproducing is observed in a wide variety of species [43]. In female cheetahs it has been observed that although older cheetahs produced less recoverable oocytes and ovarian follicles, the quality of the gametes and function of the ovaries were unexpectedly normal. The main factor being the cause of infertility in older cheetahs was seen to be uterine integrity, with endometrial hyperplasia seen in 50% of the females considered to be in their “prime” years (e.g. best years fo reproduction; 6–8 years) and in more than 85% of the older females (9–15 years) [44]. These results are similar to that of another study with 23 species of felids that had been exposed to progestins for at least 6 months, which is believed to have stimulated the endometrial epithelial cells to start differentiating [45].

This rapid aging process can be prevented by breeding females when they are still young adults [8], which would also increase their chances of being able to reproduce for the rest of their reproductive lifespan [43].

2.4. Reproductive Behaviour

Although cheetah females display characteristic feline oestrus behaviour, there is a large amount of variability between individuals [9]. The periods in which a female cheetah will accept a male to mate with her is also very short, difficult to identify or lacking [46]. A few authors have hypothesized this as “silent” oestrus [47,48,49]. In captive breeding programs, silent oestrus is seen in younger, more inexperienced females. Therefore, even though an active male can still detect a female on oestrus by her scent, if she does not display oestrus behaviour, he will not be able to find her [50].

According to Wielebnowski and Brown [9], in cheetahs, the increased frequency of certain behaviours is correlated with increasing concentrations of faecal oestradiol and, therefore, to oestrus. In this study, they classified seven females as breeders, four as non-breeders, and three as undetermined. The breeders had all previously given birth and were older than the age of 6 years. The non-breeders were younger than 4 years and had never bred before, despite being introduced to two different males more than 15 times. The undetermined group consisted of one 2-year-old female and two 5-year-old females who had never been introduced to a male. Although this study had few limitations (females were usually housed together; they were under different management systems; the undetermined group was classified as non-breeders during data analyses), it did have interesting results. Based on their observations, in fact, the behaviours “rub”, “object sniff”, “roll”, “urine-spray” and “meow-chirp” (refer to Wielebnowski and Brown [9] for definitions) significantly correlated in a positive manner with increases in oestradiol concentrations; all females of varying ages showed typical feline oestrous behaviour; there was a significant difference in age and breeding status with regards to the frequency of oestrous behaviour displayed, where non-breeders and younger females showed significantly less characteristic oestrous behaviour. Based on their results, the researchers concluded (i) that oestrus could not be regarded as silent since all ages displayed oestrous behaviour; (ii) the frequency of oestrous behaviour displayed increased with age; and (iii) age and experience are the two main factors that influence the frequency of oestrous behaviour.

Based on the results and conclusions of this study, the following assumption can be made: Since all non-breeders were under 4 years of age (breeders were over 6 years of age) and had all been previously introduced to males (>15 introductions) with no success, it can be assumed that even though mating behaviour is displayed by younger females, if the frequency is not enough, mating will not occur.

Whereas the above study proved that female cheetahs do show characteristic feline oestrous behaviour, it also proved the variation between individuals regarding the frequency and type of behaviour displayed by each female. This is different from other felids, where the sequence of behavioural changes is displayed in a foreseeable manner during oestrus [6]. In a captive breeding setting, the sequence displayed by each individual would need to be analysed in order to determine the oestrous state, which is difficult and time-consuming [6].

3. The Physiology of Olfaction

3.1. The Main Versus the Vomeronasal (Accessory) Olfactory System

All mammals possess a main and an accessory (vomeronasal) olfactory system besides bats, marine mammals, and humans [51,52]. These systems contain sensory neurons that are necessary to detect semiochemicals and, therefore, scent-marks [11]. Both systems are astonishingly consistent in all species of mammals, and although there are structural differences, there are also similarities in how each system functions [12,53]. The olfactory and vomeronasal receptors are very different from each other with regards to their primary structures and location [11,16]. The receptors from each system are only able to detect certain categories of compounds [51]. For example, volatile compounds are generally detected by the olfactory system via the nasal epithelium, whereas non-volatile compounds are said to be detected by the vomeronasal organ, which is located in the nasal septum [54,55]. Although the neural pathways of each system do run parallel to each other [55], the olfactory receptors transmit signals to the main olfactory bulb, and the vomeronasal receptors transmit signals to the accessory olfactory bulb [14,53]. Both neural pathways from each system, however, are involved in managing behavioural and endocrine responses [56].

In terms of detection and response to pheromones, previous misconceptions included the fact that the vomeronasal system detected only pheromones, whereas the main olfactory system could not [53]. This is not true as some pheromones utilize the vomeronasal system while others utilize the main olfactory system [16]. Furthermore, some semiochemicals can be detected by both olfactory systems [53].

Another misconception was that the main olfactory system initiated general reproductive behaviour and the vomeronasal system moderated specific behavioural signals relevant to each sex [57]. Both olfactory systems do coincide with each other [16]. In fact, although reduced, a neuroendocrine reaction in response to certain pheromones is still seen after the removal of the vomeronasal organ [55]. For example, the mating stance, which is a behavioural response to the steroidal pheromone, androstenone, was not affected after blocking the vomeronasal duct. Only in immature mammals that do not possess a functional vomeronasal organ, reproductive behaviour is affected [16].

3.2. The Main Olfactory System

The arrangement of the olfactory system is similar across species with regards to how the olfactory receptors function, the physiological activity thereafter, as well as the arrangement of the olfactory central nervous system [58]. Although the main olfactory epithelium generally detects volatile compounds [54], detection of non-volatile compounds occurs when an animal expresses behaviour that consists of direct physical contact [56].

The two kinds of receptors in the main olfactory epithelium are known as the olfactory receptors and the trace amine-associated receptors [11]. In rats, the suckling pheromone triggers a particular segment of the main olfactory bulb [57]. This pathway is also involved in the identification of the opposite sex [16]. The fact that both pheromones and common odours can influence the neuroendocrine system [57] is most likely due to the fact that the olfactory epithelium also contains a group of receptors dedicated to detecting pheromones [59]. These receptors are otherwise known as V1R-positive and are, in fact, a type of vomeronasal receptor [56], which means that the detection of pheromones is not a specific function of the vomeronasal system [53].

3.3. The Vomeronasal (Accessory) Olfactory System

The vomeronasal system is generally believed to be the main detector of semiochemicals, especially pheromones [56], and can detect chemical compounds of both high and low volatility [51]. Detection of pheromones via the vomeronasal system plays a large role in reproduction through the influence of sexual behaviour, as well as hormone levels involved with reproduction [55]. The vomeronasal organ, or otherwise known as the organ of Jacobson [12], is where the detection of chemical compounds occurs [60]. Cats transport non-volatile semiochemicals, such as steroid conjugates and proteins, to the vomeronasal organ by using the flehmen grimace [61]. Flehmen is a behavioural response where the lower jaw opens halfway while the cat pauses and breathes steadily [62]. The flehmen response to a scent mark seems to be universal amongst all felids, including the cheetah, domestic cat, tiger, and lion [61,63,64,65]. It is used to analyse urine [51], although males tend to show this reaction at a higher rate than females [64].

The vomeronasal organ is found in most terrestrial mammals [16], apart from bats and some species of primates. It is a tubular structure that is found in between the oral and nasal cavities [51], consisting of two sacs filled with fluid that forms an opening at only one end [12]. The vomeronasal receptors are the V1r receptors [53] and V2r receptors [66].

3.4. The Link to Olfaction and Hormone Release

The connection between the gonadotropin-releasing hormone (GnRH) and olfactory systems, as well as between mediation of sexual behavioural/hormonal responses is very intricate [16]. The GnRH neurons are ultimately the main drivers of reproductive status. They have been identified to possess bidirectional connections with neurons involved in odour and pheromone processing, such as in the olfactory cortex, medial amygdala, and posterocortical amygdalar nucleus, thus proving that signals are received from both the vomeronasal and olfactory system [55]. This connection of neurons is generally referred to as the “hypothamalmic-pituitary-gonadal axis” [16]. Noradrenaline and serotonin levels also act as an indication of the psychological state of the animal and thus affect the reproductive endocrinology and behaviour of the animal [57]. GnRH neurons, therefore, influence the endocrinology of the animal in correlation with the psychological state of the animal [55]. Since the GnRH neurons are involved in bidirectional connections with neurons from the olfactory systems [55], semiochemicals and scent-marking would, therefore, play a role in reproduction and the endocrinology of the animal as well.

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[Acinonyx sp.]

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4. How Is Olfaction Linked to Reproduction?

4.1. The Effect of Pheromones on Behaviour

Although it is true that non-chemical signals, such as hearing and visualization, the physiological state, as well as the experience the animal has had in the past, are a necessity for eliciting behavioural responses [54], it has already been stated many times that semiochemicals directly affects an animal’s behaviour [12]. This is especially behaviour related to locomotion, interactions with other animals and reproduction [11,67]. The observed effects that pheromones have on an animal’s behaviour are correlated with the animal’s endocrine status [14]. The stereotypical behaviours caused by pheromones may be an unconscious response to this particular group of odours [16].

In predator-prey relationships, prey animals will exhibit defensive behaviour and try to escape once detecting the odour of a predator [68]. The semiochemicals in the urine of a dominant animal act to suppress the physiology, as well as the reproductive behaviour (such as a decrease in the rate of marking), of the subordinate animal [14].

Pheromones involved with reproductive behaviour are used as a way of attracting the opposite sex [11] and acts as an aphrodisiac in causing the appropriate precopulatory and copulatory responses [14]. The odour of a boar increases the receptivity of a female to sexual interactions, as well as promotes lordosis in 50–60% of females in oestrus [10]. Aggressive behaviour is another known response to reproductive pheromones, although it is not always just the reproductive behaviour of an animal that is affected, but the reproductive physiology as well. Pheromones have been known to accelerate puberty [12] and stimulate the onset of oestrus [69].

4.2. Captive Breeding

Captive breeding of cheetahs occurred in response to the rapid decrease in cheetah population numbers over the past years; therefore, it initially aimed at increasing numbers, with the management of the last free-roaming cheetahs regarded as important for the species’ overall survival [37]. Unfortunately, cheetahs were found to be difficult to breed in captivity, even when there was no difference in nutrition, health, genetics, the function of the hypophysis, seminal quality in males, and anatomy of the reproductive tract in females [6,9,12,17,24,70,71].

Various behavioural methods were attempted by zoological facilities to rectify this and increase breeding success for cheetahs. Whereas some of these attempts have resulted in success, not one method was proven to work at all the various facilities. The key to success was said to be based on the knowledge of the reproductive behaviour of free-roaming cheetahs [21], as well as th cheetah’s natural social conditions, such as adhering to their solitary nature [9] and keeping breeding females separate [17]. With these management methods in place, females are observed for any behavioural signs that are indicative of oestrus [70], although this has proven to be very difficult in determining when females are actually on oestrus [6] and negatively affects the occurrence of breeding introductions [17].

Allowing each individual animal to choose their own mate is a significant part of successful breeding [72]. By using the correct housing and breeding management methods, the Ann van Dyk Cheetah Centre in South Africa (Brits, South Africa), previously known as de Wildt, has been very successful in breeding with cheetahs. It was found that the presence of male cheetahs around the females is the best method to detect when a female is on oestrus by observing her for any changes in behaviour towards the males [22]. Males are released into a walkway known as “lover’s lane” located between the female enclosures in order to smell which females are on oestrus [50]. The presence of the males has been noted to trigger the oestrous cycle in females, with interactions between the sexes resulting in a display of oestrous behaviour after the female walks towards the males in the walkway [22].

Another successful breeding method used at Wassenaar Wildlife Breeding Centre in the Netherlands (Wassenaar, The Netherlands) includes the utilization of one enclosure by the male and the female in turns. They are not allowed any direct visual or physical contact at first and are each observed according to their reaction to each other’s scent. Only after a lot of interest has been shown by both to the other’s scent, and the male starts stutter-calling or “yipping”, they are allowed visual exposure to one another. Females are observed once again for any aggression towards the male before physical introductions are allowed. If the female does not roll for the male or display oestrous behaviour then they are once again separated [73].

From the above methods, it can be speculated that semiochemicals play an important role in the initial process of breeding at Wassenaar, whereas they only seem to come into play during the last stage of the breeding process at the Ann van Dyk centre. Nevertheless, the breeding results achieved from both facilities prove that cheetahs can be bred successfully in captivity without the need for assisted reproductive techniques, as well as emphasizes the role that semiochemicals play in cheetah courtship.

5. Semiochemicals
Semiochemicals are defined as chemical substances released by a life form that instigates a physiological or behavioural reaction from another living organism [74,75]. The earliest and most studies on semiochemicals have been on insects [14]. Semiochemicals have since been classified according to the relationship between the individual that released the chemical signal and the individual that received the signal [14], thus resulting in changes to the behaviour and/or physiology of the receiver. Semiochemicals that act as chemical signals between individuals of the same species (intraspecific) are classified as pheromones, whereas chemical signals acting between members of two different species (interspecific) are classified as allelochemicals [12].
Pheromones can be both odorous, thus originating from volatile chemical compounds, or non-odorous, which are from non-volatile chemical compounds [11]. Pheromones are divided into two categories based on the response to the chemical signal, namely releasers and primers. Releasers are pheromones that result in an immediate response in the behaviour of the receiver [11] and are otherwise known as signalling pheromones involved in communication [11,12]. Primers act along with the nervous system of an animal and generate a physiological response after a longer period of time [11]; therefore, primers affect mainly the endocrine system [14].

5.1. Complex vs. Single Compounds

There has been very little evidence to support the theory that a chemical signal is reliant on a number of chemical compounds [52] and it has been shown that a single chemical compound is as efficient as a mixture of compounds in evoking a behavioural response [76]. The fact that multiple chemical compounds, with each coding for the same or similar chemical signal/behavioural response, are released in scent markings is because the interaction of the chemical compounds increases detection of the signal, memorization, and learning of different chemical signals released by different species, such as those released by predators, as well as discrimination between different chemical signals [77].

5.2. Semiochemicals in Communication

Semiochemicals that are involved in communication act between animals/organisms that have more advanced olfactory systems, with which they rely on more than they do with hearing or vision [11,12]. Scent-marking releases a large amount of volatile and non-volatile chemical compounds that are usually present at low concentrations. The different concentrations of chemical compounds could be important with reference to the message that is sent [78]; and could release specific characteristic odours, although exactly what compounds are inhaled and how they are analysed by the animal has not yet been fully discovered [11].

Territorial marking, identification of neighbours, detection of larger predators and food/prey, signalling of alarm and attraction of members of the opposite sex are all functions of scent-marking [12,14,19,54]. Scent marks, in fact, also provide information on the animal’s sex, age and the reproductive state of females [14,19], which is also an indication of whether or not she is sexually receptive [79]. Genetic relatedness and compatibility of future mates can also be assessed from scent marks [54].

5.3. Identification of Individuals through Semiochemicals

Determining whether individuals have their own specific body odour, or ‘biochemical fingerprint’ or not, has become a topic of increasing interest since the identification of individuals through semiochemicals is of great significance [54]. Soso, [11] stated that semiochemicals in exocrine secretions (scent marks), which can be complex or simple mixtures, code for individuality based on the amount and the presence of particular chemical compounds. This has also been confirmed in humans where volatile compounds, identified from the sweat, urine and saliva, were shown to be both individual and gender-specific [80]. It has also been demonstrated that dogs could discriminate between the body odours of several different humans. This study showed that the dogs were able to differentiate the body odours of not only individuals of the same family, but between a set of twins as well [81]. In previous studies done with lions, over 50 volatile chemical compounds were identified in the marking fluid, with each individual having a different chemical composition [82]. Seven chemical compounds were found in the marking fluid of almost all the lions, and therefore, it is possible that some, or all of these compounds, are involved in species identification, either individually or in a combination [78].

5.4. Semiochemicals in Cheetahs

Although many studies have proved the importance of semiochemicals in domestic and wild felids [11,14,54,62,63,64,78,83], very little is known about their role in cheetahs.

Unfortunately, cheetah urine itself has not acquired much interest from scientists, probably due to the very little amount of odour it emits [26]. As reported by Visser [12], who analysed the urine of three males and two females, and Burger et al. [26], who analysed the urine of six males and one female, there was a much wider variety of volatile constituents identified in female urine than male urine. The constituents that were common in the males from both studies included that of 2-butanone, phenol, hexanal, octanal, nonanal, benzaldehyde, 2-pentanone, 2-hexanone, 2-heptanone, 3-pentanone, 4-heptanone, 3-hexanone, acetophenone, cyclohexanone, 3-methylcyclopentanone, ethyl propyl ether, butyl ethyl ether, butyl propyl ether, dimethyl disulfide, dimethyl sulfone, sulfur S6, sulfur S8 and urea. Of these compounds, only a few are known to be involved in certain behaviours in various species such as 2-butanone, phenol, octanal, nonanal, benzaldehyde, 2-pentanone, 4-heptanone, acetophenone, cyclohexanone, 3-methylcyclopentanone and dimethyl disulfide [10,11,51,67]. Many of the abovementioned compounds have been identified in the marking fluid/urine of various other feline species as well. These compounds, together with the animals in which they have been found and the behaviour they elicit are listed in Table 1.

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5.5. Current Uses of Synthetic Scent to Influence Behaviour and Its Potential Use in Cheetahs

Providing enrichment for captive animals aims mainly to improve the welfare of the animals [89] by decreasing stress and improving natural and, therefore, reproductive behaviour as well [90]. Pheromones are used quite frequently in captive facilities as a form of olfactory enrichment by introducing new scents or objects that have been scented to an animal’s enclosure [91]. Felids display cheek rubbing behaviour as a result of the use of novel scents (e.g., perfume for behavioural enrichment in zoological parks) [92]; lions respond to olfactory enrichment by behaving more socially and displaying an increase in the activity and frequency of various behaviours [76]. Besides enrichment, semiochemicals have also been specifically used for reproductive purposes in some species. For instance, synthetic boar pheromone is used to identify when a sow is on oestrus. These pheromones are already produced and sold on a commercial level in aerosol cans [51]. A synthetic scent known as Feliway® has proven useful in lowering the level of corticosteroid metabolites in the faeces of tigers after artificial insemination operation procedures [67].

With regards to cheetahs, since the presence/odour of a male cheetah has been noted to trigger the oestrous cycle in the females [22,73] and the scent is known to play an important role in reproduction with cheetahs as well as other felids [21,39,62,63,70,73,88,93], there is an opportunity to further investigate the role of semiochemicals in cheetahs and their possible use to improve oestrus behaviours for successful mating. For instance, it could be possible to use a synthetic or natural scent in cheetahs as well in order to increase the frequency of oestrus behaviours displayed in younger females to ensure successful mating when the females are introduced to the males.

6. Conclusions
The overall cheetah population is currently listed as vulnerable, with a decreasing population trend [2]. The management of the last free-roaming cheetahs is therefore regarded as important for the species’ overall survival [37]. Captive breeding of cheetahs occurred in response to the rapid decrease in numbers in the past and therefore is aimed at increasing cheetah population numbers with a higher genetic variety when compared to their wild counterparts. This would lead to the creation of a sustainable, healthy population of cheetahs that would be able to survive after possible reintroduction [8,9,37]. Despite the fact that reproduction in wild cheetahs is not an issue, cheetahs were found to be difficult to breed in captivity [6,7,9,12,17,23,24,71,94].
The key to success is based on the knowledge of the reproductive behaviour of free-roaming cheetahs [21] and, therefore, managing individuals according to their natural social conditions [71]. This would include keeping males and females in separate enclosures and only introducing pairs during mating [9] and keeping breeding females separate since even small amounts of social aggression acts to suppress the oestrous cycle [17]. Using male cheetahs to detect when a female is on oestrus [22] and allowing each individual animal to choose their own mate also played a significant part in ensuring successful breeding [72].
Since female cheetahs are susceptible to the effects of asymmetric reproductive aging (ARP), it would be necessary for them to breed when they are still young adults in order to prevent this aging process [8], such as after reaching sexual maturity at 20–24 months of age [1]. Unfortunately, females younger than the age of four years display a significantly lower frequency of characteristic oestrous behaviour than females older than the age of six years [9], this means that even though a male is still able to detect a female on oestrus by her scent, if she does not display oestrus behaviour, he will not be able to find her [50] and the chance of mating is severely reduced.
Intraspecific semiochemicals [12] that are involved with reproductive behaviour are used as a way of attracting the opposite sex [11] and acts as an “aphrodisiac” in causing the appropriate precopulatory and copulatory responses [14]. These semiochemicals are synthesized by an animal [54] and released as exocrine secretions to create a ‘scent-mark’ [11], such as urine and marking fluid [67].
Since the presence/odour of a male cheetah has been noted to trigger the oestrous cycle in the females [22,73] and the scent is known to play an important role in reproduction with cheetahs as well as other felids [7,21,39,62,63,73,88,93], if one were to analyse the chemical compounds in the marking fluid of male cheetahs (with a specific focus on high impact odorants), it could lead to the identification of a single compound (or a limited number of compounds in a mixture) that would be able to increase the frequency of reproductive behaviour displayed by the females in the hopes that intercourse and pregnancy would occur at a higher rate.

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[Acinonyx sp.]

Categorising cheetah behaviour using tri-axial accelerometer data loggers: a comparison of model resolution and data logger performance


Abstract
Background
Extinction is one of the greatest threats to the living world, endangering organisms globally, advancing conservation to the forefront of species research. To maximise the efficacy of conservation efforts, understanding the ecological, physiological, and behavioural requirements of vulnerable species is vital. Technological advances, particularly in remote sensing, enable researchers to continuously monitor movement and behaviours of multiple individuals simultaneously with minimal human intervention. Cheetahs, Acinonyx jubatus, constitute a “vulnerable” species for which only coarse behaviours have been elucidated. The aims of this study were to use animal-attached accelerometers to (1) determine fine-scale behaviours in cheetahs, (2) compare the performances of different devices in behaviour categorisation, and (3) provide a behavioural categorisation framework.
Methods
Two different accelerometer devices (CEFAS, frequency: 30 Hz, maximum capacity: ~ 2 g; GCDC, frequency: 50 Hz, maximum capacity: ~ 8 g) were mounted onto collars, fitted to five individual captive cheetahs. The cheetahs chased a lure around a track, during which time their behaviours were videoed. Accelerometer data were temporally aligned with corresponding video footage and labelled with one of 17 behaviours. Six separate random forest models were run (three per device type) to determine the categorisation accuracy for behaviours at a fine, medium, and coarse resolution.
Results
Fine- and medium-scale models had an overall categorisation accuracy of 83–86% and 84–88% respectively. Non-locomotory behaviours were best categorised on both loggers with GCDC outperforming CEFAS devices overall. On a coarse scale, both devices performed well when categorising activity (86.9% (CEFAS) vs. 89.3% (GCDC) accuracy) and inactivity (95.5% (CEFAS) vs. 95.0% (GCDC) accuracy). This study defined cheetah behaviour beyond three categories and accurately determined stalking behaviours by remote sensing. We also show that device specification and configuration may affect categorisation accuracy, so we recommend deploying several different loggers simultaneously on the same individual.
Conclusion
The results of this study will be useful in determining wild cheetah behaviour. The methods used here allowed broad-scale (active/inactive) as well as fine-scale (e.g. stalking) behaviours to be categorised remotely. These findings and methodological approaches will be useful in monitoring the behaviour of wild cheetahs and other species of conservation interest.

Background
Global biodiversity loss is one of the biggest crises currently threatening the natural world [1,2,3]. Approximately 40% of plant species, 23% of invertebrates, and 18% of vertebrates assessed are considered to be threatened [4]. For mammals, 22% of described species [4] and 26% of assessed carnivores are considered to be threatened [5]. Some of the primary threats to carnivores include reductions in prey [6] and habitat [7], human-wildlife conflict (primarily in terms of livestock losses) [8,9,10,11], and illegal trade in animals or animal parts [11,12,13,14,15]. Conservation is therefore at the forefront of policy-making decisions worldwide [16,17,18,19].
To implement effective conservation and species management strategies, an understanding of target species populations, ecology, and behaviour is important to provide an insight into the status of the species, its needs, and putative causes of decline. When monitored over time, population data can indicate trends in particular groups of animals or in species as a whole [20, 21] or the efficacy of conservation efforts in comparison to control areas [22]. Remote sensing technologies such as camera traps have aided in species population assessments [23,24,25,26], as well as in our understanding of species ecology, and behaviour [23]. Advances in Global Positioning System (GPS) devices have further contributed towards understanding species ecology by providing insights into movements and habitat use. Knowledge of behaviour in space and time can provide insights into the importance of particular habitats and microhabitats for a species. For example, Wege et al. [27] identified novel foraging sites used by fur seals, contributing towards conservation policy-making as these sites were heavily utilised during the winter and were not previously considered when making assessments for potential marine protected areas (MPAs), where summer use is considered to be more important.
Accelerometer data loggers have been used independent of [28, 29] and in combination with [30,31,32] other remote sensing technologies such as GPS devices, magnetometers, and gyroscopes. Tri-axial accelerometers measure acceleration in three orthogonal axes (heave, surge, and sway), providing information on omnidirectional dynamic movement of an animal, as well as its posture (via static acceleration) [33, 34]. When accelerometers are used alongside devices such as GPS loggers, detailed behaviour patterns in space and time can be elucidated (e.g. [35, 36]). Unlike other remote sensing technology such as camera traps, loggers are fitted to the animals of interest (either directly or via collars or harnesses), providing data on the individual for the entire deployment period, not simply when activated. This feature is particularly useful for assessing the behaviours of cryptic species with large home ranges or that utilise difficult-to-monitor habitats (e.g. dense forests/bush, burrows, or expansive deserts). Although, the relative affordability and ease with which loggers can be deployed has led to their widespread use, less consideration appears to be given to device selection and subsequent downstream data processing. Most applications of animal-borne accelerometers have been to examine behaviours (e.g. [31, 37,38,39,40]), with several resulting in the categorisation of coarse-scale descriptions (i.e. three or four different behaviours) [28, 33, 40, 41]. While several studies have categorised behaviours manually by coarsely examining the acceleration traces generated (e.g. [33, 42]), others have implemented machine-learning techniques (many described in [43]), including random forests (RFs) [29, 31, 37,38,39,40, 43, 44], to classify behaviours to datasets using training and test data. Other approaches, such as the use of magnetometers, have proven successful in the determination of specific behaviours (e.g. biting and chewing in grazing herbivores) [45].
Cheetahs (Acinonyx jubatus) are medium-large felids inhabiting Africa and Iran [46,47,48]. They are classified as ‘Vulnerable’ by the IUCN [46] with the most recent population assessment (2014) suggesting just under 7100 adolescent and adult cheetah remain in the wild [47]. Population strongholds exist in southern and eastern Africa [46, 47]. Whilst conservation measures such as confiscation of traded animals and parts and reducing conflict with humans have been put in place, cheetah populations continue to decrease, with habitat loss, persecution, and illegal hunting and trade comprising major threats [7, 11, 47]. As such, detailed monitoring of cheetah movements, habitat use, and behaviour can assist with conservation efforts to ensure stringent monitoring of frequently used areas to reduce poaching and the adequate provision of resources to meet the needs of the species. To date, only coarse behaviours (active, inactive, and feeding) have been defined for cheetahs using remote sensing technology (accelerometers) [33, 41]. However, other ecological information such as different hunting strategies they may adopt and the associated costs of chasing prey [30, 32, 49, 50] (using GPS and accelerometers) have also been elucidated. However, while fine-scale behaviours, such as stalks (which may not result in a hunt), different movement gaits (e.g. walking vs. sprinting), and resting, have yet to be described for cheetahs, such data are available in other species (e.g. [31, 37,38,39, 43, 44]). Cheetahs are considered to be “extreme” movers, potentially reaching top speeds of 64mph (103kph) in a matter of seconds [51]. Therefore, the ability to distinguish between fine-scale behaviours may help to define the ecological needs of cheetahs, including hunting success rate, and, thus, contribute to conservation efforts. However, due to the high power and accelerations attained by cheetahs, monitoring their behaviour remotely may be limited by the capacity of individual devices.
The overall aim of the current study was to ground-truth behaviours performed by cheetahs against data collected using tri-axial accelerometers. Specifically, we wanted to (1) determine the accuracy with which a suite of behaviours in a cheetah’s repertoire could be defined; (2) determine whether this could be affected by the technical specifications of two different accelerometer devices, and; (3) provide a framework in the form of a vignette containing “R” code to develop behaviour categorisation models for other species of policy or conservation interest.
Methods
Study animals and collar preparation
This study was carried out in October 2012 at the Cheetah Conservation Fund (CCF) research centre near Otjiwarongo, Namibia (− 20.447763° N, 16.677918° E). Five resident adult cheetahs (three males and two females) were fitted with their own neck collars (nylon dog collars with plastic clip buckle: mass = 75 g, length = 570 mm, width = 20 mm) equipped with two tri-axial accelerometer data loggers: 1. G6, CEFAS Technology Limited, Lowestoft, UK (maximum = 2.3 g, size = 40 × 28 × 15 mm (L × B × D), mass = 18 g including urethane encasement, recording frequency = 30 Hz); 2. X8M-3, Gulf Coast Data Concepts (GCDC), LLC, Waveland, MS, USA (maximum = 8.6 g, resolution = 0.001 g, size = 50 × 30 × 12 mm (L × B × D), mass = 21.6 g including epoxy encasement, recording frequency = 50 Hz). To ensure the collar remained centred on the ventral side of the neck, an additional weight comprising four steel nuts (120 g) was added. The total weight of the fully equipped collars was approximately 235 g (see Additional file 3: Figure S1a for constructed collar design). Prior to being fitted to cheetahs, collars were hung on a metal rail with the accelerometers located at the bottom of the collar to allow for the devices to be calibrated (see “Data processing—accelerometers” below).
Exercise arena and video capture
Cheetahs were exercised by chasing a lure (cloth rag) attached to ~ 285 m of cord around a pre-determined track. The lure machine, powered by an electric motor, was remotely controlled by a keeper, such that the speed and direction of the lure could be altered at will. The keeper changed the direction of the lure strategically to attempt to outwit the chasing animals and prevent capture of the lure. The chasing animals were thereby encouraged to employ different strategies to try to catch the lure, including stalking behaviour and high-speed pursuits. Each cheetah was exercised individually and behaviour was recorded using a video camera (Canon PowerShot SX230 HS; Canon, Japan). Typically exercise bouts lasted 10–15 min and consisted of three or four active chases (e.g. running, stalking) punctuated by two or three lower intensity rest periods (e.g. lying down, walking, standing). Collars were retrieved when the animal had finished exercising. As exercise bouts comprised periods of activity and inactivity, data associated with both hunting and resting were collected and ground-truthed against video footage.
Data processing—accelerometers
Following exercise bouts, data loggers were removed from collars and data were downloaded. The data collected for both devices were calibrated to correct for non-centred mounting of the devices on the collars using the region of the dataset where the collars had been attached to the metal rail (see Additional file 1: Study details, collar calibration, and calculations). The data corresponding to the times of captured video footage were selected and the rest of the data were removed. Static acceleration (acceleration due to gravity; Additional file 5: Figure S2, static acceleration diagram) was derived for each axis from the corrected heave (acceleration in vertical axis), surge (acceleration in longitudinal axis), and sway (acceleration in transverse axis) data by calculating a rolling mean over a two-second window [52]. Dynamic acceleration was then calculated for each axis as the absolute result of subtracting static acceleration for a particular axis from its raw acceleration. Vectorial Dynamic Body Acceleration (VeDBA), Vectorial Static Body Acceleration (VeSBA), animal static acceleration (Anim.stat), pitch, and roll were also determined (Additional file 1: Study details, collar calibration, and calculations).
Data processing—video footage
All video footage (approximately 58 min; 103,869 CEFAS logging events; 174,185 GCDC logging events) was synchronised with its complementary accelerometer datasets. Video footage was assessed frame-by-frame (Avidemux software; Developer: Mean) and cheetah behaviour was matched with the accelerometer data. Initially, 22 behaviours and behaviour combinations were identified (Table 1). Any other behaviour was recorded as ‘other’ and instances where behaviour could not be assigned (e.g. if an object obstructed a clear view of the animal) were removed from the dataset as we could not be certain of categorisation, resulting in a loss of approximately six minutes’ worth of data. Each labelled dataset was amalgamated to give two master spreadsheets of labelled accelerometer data; one for each model of accelerometer device (CEFAS and GCDC).

Data analysis

Data analysis was carried out in ‘R’ version 3.4.3 [54] using the ‘h2o’ package version 3.16.0.2 [53]. RF analysis (Additional file 2: Code) was conducted on the datasets labelled with behaviours. The datasets were split into three, such that 60% of cases were selected at random to entrain models (training dataset), 20% of cases were selected at random to validate the model (validation dataset), and the remaining 20% were used to test model performance (test dataset). The training data were used to entrain the RF model to categorise specific behaviours (see Table 2 for behaviour list). The validation dataset was then used to assess the performance of the model via model accuracy, (root) mean square error (RMSE and MSE), and r2. The validation data were also used to refine the model by altering model parameters and comparing the metrics listed above. The test data were only used once at the end of the process to compare model accuracy after validation to the outputs of the training dataset.


Model structure
Initially, models were entrained to categorise 17 behaviours (Table 2, fine-scale). The predictor variables were: heave, surge, sway, static heave, static surge, static sway, dynamic heave, dynamic surge, dynamic sway, VeDBA, VeSBA, Anim.stat, pitch, and roll. A stopping criterion (stopping-rounds = 2) was implemented to optimise the duration for which models were run. A stopping criterion of two stops fitting the model when the two-tree average is within 0.1% accuracy of the previous two-tree average. If this criterion is increased, the average is taken over the specified number of trees. Models were refined by changing their depth and comparing their overall accuracy (percentage of correctly categorised behaviours divided by percentage of incorrectly categorised behaviours). The model with the highest accuracy was retained. Models were re-run using coarser behavioural categories (Table 2). For each model, cross-validation was performed using five folds and comparing mean accuracy, RMSE, MSE, and r2 to the training dataset. ‘R’ code for RF model constructs and additional model information are provided in the supplement (Additional file 2: Code). Logger performances were compared for the categorisation of each individual behaviour using chi-squared tests.



Results

There was no indication of significant overfitting when cross-validation of models was carried out (Table 3).


CEFAS loggers

In the first model, behaviours were categorised on a fine scale. The behaviours sought to be categorised were: crouch, lie, sit, stand, head movement, crouching stalk, lying stalk, sitting stalk, standing stalk, walking stalk, trotting stalk, walk, trot, canter, gallop, pounce, and other. The overall accuracy of the model was 83.3% (MSE = 0.18, RMSE = 0.42, r2 = 0.99). However, as ‘other’ was an uninformative category, which didn’t require correct positive categorisation in the training dataset as it comprised a ‘rag bag’ of various movements, its categorisation could be disregarded (but the variable still remained in the model). Once disregarded, the accuracy of the model increased to 84.2%. Sitting stalk, lying stalk, lying, and standing were categorised with over 90% accuracy. Behaviours with < 50% categorisation accuracy included pouncing, crouching, trotting, and trotting stalk (see Table 4 ; Fig. 1 for full description of classification accuracy). Crouching behaviour was most often confused with lying (14.5%), other (27.6%), and standing (47.4%). Trotting and trotting stalk were most often confused with cantering (trotting: 21.1%; trotting stalk: 19.4%), galloping (trotting: 6.1%; trotting stalk: 13.2%), other (trotting: 42.2%; trotting stalk: 45.8%), and walking (trotting: 19.7%; trotting stalk: 9.7%). In addition, trotting stalk was confused with walking stalk (6.3%) (Fig. 2A). In terms of predictor variables, static acceleration in all three axes was most important in categorising behaviours (heave: scaled importance (improvement of MSE relative to maximum improvement across all predictors) = 100%, explanatory power = 14.4%; sway: scaled importance = 80.0%, explanatory power = 11.5%; surge: scaled importance = 71.1%, explanatory power = 10.2%), followed by VeDBA (scaled importance = 53.3%, explanatory power = 7.7%), roll (scaled importance = 52.0%, explanatory power = 7.5%), and heave acceleration (scaled importance = 51.6%, explanatory power = 7.4%). In all, these six predictors explained 58.7% of the RF model variance.

In the second, coarser model, several behaviours from the previous model were combined in an attempt to reduce the error rate. ‘Pounce’ was entered as ‘other’ as it could not be categorised reliably and was often confused with several other behaviours. Behaviours in this model included: Sedentary (‘crouch’, ‘lie’, ‘sit’, ‘stand’), head movement, moving stalk (‘trotting stalk’, ‘walking stalk’), crouching stalk, sitting stalk, lying stalk, standing stalk, gallop, canter, trot, and walk. The overall accuracy of this model was 84.7% (MSE = 0.16, RMSE = 0.40, r2 = 0.98), which increased to 86.6% when ‘other’ behaviours were removed. Sedentary, sitting stalk and lying stalk were the only behaviours where the prediction accuracy surpassed 90%. The prediction accuracy for two behaviours was lower than 50%; cantering and trotting (see Table 4 and Fig. 1 for full description of classification accuracy). Cantering was most often confused with galloping (43.8%), other (34.4%), and trotting (8.0%), while trotting was most often confused with cantering (20.1%), other (40.9%), and walking (20.8%) (Fig. 2B). Once again, static accelerations (heave: scaled importance = 100%, explanatory power = 13.0%; surge: scaled importance = 87.8%, explanatory power = 11.5%; sway: scaled importance = 76.2%, explanatory power = 9.9%), VeDBA (scaled importance = 73.7%, explanatory power = 9.6%), and VeSBA (scaled importance = 50.8%, explanatory power = 6.6%) featured among the important predictor variables. Overall, the top five predictors explained 50.7% of the RF model variance.

The final, coarsest model comprised a simplified RF where behaviours were either deemed to be active (‘walk’, ‘trot’, ‘canter’, ‘gallop’, ‘walking stalk’, ‘trotting stalk’, ‘pounce’), inactive (‘crouch’, ‘lie’, ‘sit’, ‘stand’, ‘crouching stalk’, ‘lying stalk’, ‘sitting stalk’, ‘standing stalk’), head movement, or other. The accuracy of this final model was 88.2% (MSE = 0.11, RMSE = 0.32, r2 = 0.89). When ‘other’ was removed, model accuracy increased to 92.7%. Inactivity was predicted to the highest degree of accuracy and head movement with the lowest. Activity was predicted with 86.9% accuracy (see Table 4; Fig. 1 for full description of classification accuracy). Whilst head movement was most often confused with all three behaviour categories; inactivity (47.3%), activity (28.8%), and other (24.0%), activity was most often confused with inactivity (58.9%) and ‘other’ behaviours (40.2%) (Fig. 2C). The most important predictors in this model were VeDBA (scaled importance = 100%, explanatory power = 15.1%), static acceleration in the heave axis (scaled importance = 84.1%, explanatory power = 12.7%), and dynamic acceleration in the sway (scaled importance = 63.0%, explanatory power = 9.5%), heave (scaled importance = 56.8%, explanatory power = 8.6%), and surge (scaled importance = 52.5%, explanatory power = 7.9%) axes as well as static acceleration in the surge axis (scaled importance = 50.4%, explanatory power = 7.6%). In total, the top six variables explained 61.6% of the model variance.

GCDC loggers
The models outlined above were repeated for the GCDC data loggers. The model containing the finest-scale behaviours was 85.5% accurate (MSE = 0.17, RMSE = 0.41, r2 = 0.99). Accuracy increased to 85.8% when the category ‘other’ was omitted. The sedentary behaviours of lying, lying stalk, and sitting stalk were categorised with > 90% accuracy. Crouching was the only behaviour that was categorised with < 50% accuracy (see Table 4 and Fig. 1 for full description of classification accuracy); it was most often confused with standing (41.1%) and other behaviours (41.1%) (Fig. 3A). Static acceleration in all three axes was the most important predictor for behaviour (heave: scaled importance = 100%, explanatory power = 16.0%; sway: scaled importance = 80.1%, explanatory power = 12.8%; surge: scaled importance = 72.8%, explanatory power = 11.6%). In total, static acceleration variables explained 40.4% of the model variance.

When the second model outlined above was reproduced for the GCDC logger, it had an accuracy of 86.2% (MSE = 0.15, RMSE = 0.39, r2 = 0.98), which increased to 87.4% when the ‘other’ category was removed. Behaviours categorised with > 90% accuracy were sedentary, lying stalk, and sitting stalk. No behaviour categorisation was < 50% accurate; those behaviours that were most difficult to categorise were trotting, cantering, and head movement (see Table 4, Fig. 1 for full description of classification accuracy, and Fig. 3B for confusion matrix). In terms of predictor variables, static acceleration in all axes (heave: scaled importance = 100%, explanatory power = 14.7%; surge: scaled importance = 78.6%, explanatory power = 11.6%; sway: scaled importance = 78.2%, explanatory power = 11.5%), VeSBA (scaled importance = 51.8%, explanatory power = 7.6%), and Anim.stat (scaled importance = 51.5%, explanatory power = 7.6%) were most important in determining behaviours in this model. The top five predictor variables explained 53.1% of the model variance.

In the final RF model for the GCDC data loggers, activity, inactivity, head movement, and other behaviours were categorised. This model performed with a categorisation accuracy of 90.2% (MSE = 0.09, RMSE = 0.31, r2 = 0.90), which increased to 92.9% when the ‘other’ behaviour category was omitted. Inactivity was most easily classified and no behaviour had a classification accuracy below 61.3% (see Table 4, Fig. 1 for full description of classification accuracy, and Fig. 3C for confusion matrix). In this model, static acceleration in the heave (scaled importance = 100%, explanatory power = 12.7%) and surge (scaled importance = 84.8%, explanatory power = 10.8%) axes were most important for categorisation, followed by VeDBA (scaled importance = 83.2%, explanatory power = 10.6%), static acceleration in the sway axis (scaled importance = 75.8%, explanatory power = 9.6%), Anim.stat (scaled importance = 61.1%, explanatory power = 7.8%), VeSBA (scaled importance = 57.4%, explanatory power = 7.3%), and dynamic acceleration in the heave (scaled importance = 52.0%, explanatory power = 6.6%) and sway (scaled importance = 52.0%, explanatory power = 6.6%) axes. Together these eight variables explain 79.1% of the model variance.

Logger comparison
Generally, the higher capacity loggers, with higher recoding frequency (GCDC) outperformed their lower capacity (CEFAS) counterparts when determining cheetah behaviour (Table 5; Fig. 1). This was particularly evident during medium- and fine-scale behaviour categorisation. Of the 30 different behaviour-model combinations run, the CEFAS loggers significantly outperformed the GCDC loggers only four times: standing (χ2 = 11.63, df = 1, p < 0.001) and galloping (χ2 = 19.46, df = 1, p < 0.001) in the fine-scale model, galloping (χ2 = 23.15, df = 1, p < 0.001) in the medium-scale model, and inactivity (χ2 = 4.42, df = 1, p = 0.035) in the coarse model (Table 5; Fig. 1). Conversely, the GCDC loggers were significantly better than the CEFAS loggers at behaviour categorisation on 11 occasions (Table 5; Fig. 1), notably when defining trotting, moving stalks, head movement, pouncing, and, in the fine-scale model, most sedentary behaviours. Whilst the GCDC loggers were better overall at defining behaviours on a fine- and medium-scale, there was no significant difference between the two devices when categorising behaviours on a coarse-scale (Table 5).

*This image is copyright of its original author

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