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The AfriCat Predator Population Density Study in the Okonjima Nature Reserve

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okonjima location map

Leopard Collaring
Filmed by Wild Dog Productions Pty.
Mafana, a large male leopard in the Okonjima Game Reserve, was always eluding rangers who needed to put a collar on him.

 



 

Research Proposal: Submitted by Jenny Noack

Title of proposed project: The assessment of leopard (Panthera pardus) density and population size via a capture – recapture framework in an island bound conservation area in Namibia.

Principal Investigator: Jenny Noack

Co-Worker: Louis Heyns, Wayne Hanssen, Donna Hanssen, Diethardt Rodenwoldt

Contact Details:
AfriCat Foundation Research Department
Contact Person: Jenny Noack
Tel: +264-67-304566 / Mobile: +264-81-3987594
Fax: +264-67-687051
E-mail: research@africat.org
P.O. Box 1889, Otjiwarongo, Namibia

 

Background and Introduction

Project Need

In the last decades human activities have led to the devastating destruction of large parts of natural habitats (Gaston, 2008) leading to the dramatic decrease and threatened status of many wildlife species worldwide. The leopard (Panthera pardus) is classified as "Near Threatened" according to the IUCN Red List of Threatened Species (IUCN, 2014.2.). Leopards occur across wide ranges of sub-Saharan Africa as well as inhabit parts of Northern Africa and tropical Asia (Friedmann & Holzer, 2008). Their adaptability and tolerance towards a wide range of various habitats as well as their secretive and elusive nature had let them survive in marginal areas from which other felid have disappeared completely (IUCN, 2014.2). Despite their wide distribution throughout sub-Saharan Africa, the felids are declining dramatically in numbers and have disappeared from approximately 36.7% of their historical range (Ray et al. 2005) due to habitat fragmentation as well as intense persecution by humans. While sub-populations in North Africa and Asia are on the verge of extinction, Namibia’s population maintains stable numbers (Stein, Andreas & Aschenborn, 2011). With a total of only 17% of protected areas in forms of national parks, game reserves and recreational areas in Namibia (Turpi et al 2010), the majority of leopards occurs on commercial and communal farmland where the sporadic predation of livestock induces an inevitable conflict between man and carnivore. The necessity for the development and expansion of protected areas as well as the implementation and execution of improved livestock farming techniques on farmland are therefore of utmost importance to secure the survival of the Namibian leopard population.

 

Protected areas maintain a higher density of predators than un-protected areas (Stein et al. 2011). Carnivore abundance is often correlated by the level of inter- and intraspecific competition as well as the availability and accessibility of resources such as water and prey. Human-caused mortalities and habitat loss are additional factors playing a crucial role in determining felid densities (Khorozyan et al, 2008). Reliable large carnivore estimations are essential for the development of sustainable and long-term management and conservation strategies (Hayward, Brien, Kerley, 2007).

 

The majority of large felids are nocturnal and characterized by a secretive and elusive nature, consequently most techniques used to estimate species richness and abundance provide unreliable results. The application of camera traps proved to be the most dependable method when it comes to the creation of reliable quantitive estimations of large carnivore species.

 

Project Description

Goal: To assess the density and population size of leopards (Panthera pardus) in the Okonjima Nature Reserve using photographic capture-recapture sampling and provide scientific data on their demography as well as spatial and temporal distribution patterns.


Study Area

study area map

 

Fig. 2: The Okonjima Nature Reserve located in central Namibia compromises a total area of 22 000 ha. Okonjima is home to The AfriCat Foundation whose mission is the long-term survival of Namibia’s carnivores in their natural habitat.

 

The Okonjima Nature Reserve (Lat/Lon: 20º49’19.36’’S, 16º38’21.25’’E) is located in central Namibia approximately 50 km south of Otjiwarongo and compromises a total area of 22 000 ha. The study area is semi-arid and characterized by a marked seasonality. The annual precipitation averages approximately 450 mm. The Okonjima farm boundary traces a central plateau, at average an altitude of 1 600 meters, surrounded by the Omboroko Mountains. The vegetation can be mainly described as tree- and scrub savannah, interspersed with Yellow wood (Terminalia sericea) and several Acacia-species. Artificially constructed water reservoirs ensure the perennial supply of surface water.

Okonjima was used intensively for the purpose of cattle farming from 1920 until 1993. Since then the private nature reserve has been used for carnivore rehabilitation and tourism purposes exclusively.

The reserve is surrounded by a 96 km electrified perimeter fence, completed in 2010, and is bordered entirely by commercial farmland. An additional fence is erected within the reserve and creates a 20 000 ha reserve for carnivore rehabilitation and a 2 000 ha “lodge area” that includes lodges and campsites as well as the AfriCat headquarters and the Environmental Education Centre.

Leopards as well as brown hyenas (Hyena brunnae) occur naturally within the borders while cheetahs (Acinonyx jubatus), African wild dogs (Lycaon pictus) and spotted hyenas (Crocuta crocuta) are part of AfriCat’s rehabilitation program that have been released into the area.  Lions (Panthera leo) are absent from the study area. Thus, leopards belong to the apex predators in the reserve that are playing an important role in maintaining the health of the ecosystem.

 

Study Objectives

  1. To determine leopard density and population size via a capture-recapture framework using remote camera traps
  2. To determine the demography of leopards within the Okonjima Nature Reserve
  3. To develop a dataset that can be applied as a baseline for comparisons to similar areas
  4. To develop a long-term population monitoring program

 

Methodology and Data Analysis

Camera Trapping

Photography as a method of studying and observing wildlife goes way back in the nineteenth century and finds their early beginning in the 1860’s. George Shiras, lawyer, conservationist and pioneer of remote wildlife photography, was the first one to capture animals on photograph without human presence by using a trip wire that triggered his magnesium flash-gun camera. The use of camera traps by the hunting community promoted their increased application and advanced technical improvements (Sollmann et al. 2012). Since then the methodology of camera traps experienced major enhancements - not only in a technical but also in a visual way: Trip wires and step treadles that used to trigger old models have been replaced by infrared triggered systems placed in compact, camouflaged and waterproof housings. Today camera trapping is an essential and widely used tool that has revolutionized wildlife research and is applied for a variety of different research questions including the detection of species and their distribution and activity patterns as well as abundance and density estimations. 

The individual identification of animals via the use of wildlife camera traps has become a vital tool in animal ecology and has been applied to a wide range of individually recognizable species: tiger Panthera tigris (Karanth 1995), jaguar Panthera onca (Wallace et al. 2003), puma Puma concolor (Kelly et al. 2008), Iberian lynx Lynx pardinus (Gil-Sanchez et al. 2011), striped hyena Hyena hyena (Harihar et al. 2010), tapir Tapirus terrestris (Noss et al. 2003).

Individual recognition relies on capture - recapture surveys (Foster & Harmsen, 2011) and is based on the assumption that an animal can be identified by a natural or artificial marking and that this marking clearly distinguishes it from other individuals (Heilbrunn et al. 2003). Natural markings that allow for individual identification include stripe and spot pattern (e.g. tigers P. tigris, leopards P. pardus, cheetahs Acinonyx jubatus) and whisker patterns in lions Panthera leo.

The application of camera traps to study faunal populations offer a number of advantages and benefits when compared to other field methods and techniques: In contrast to other methods photographic captures provide undeniable evidence of an individual, animal and/or species being present at a certain site at a certain time and therefore, makes review by other researchers possible. 

leopard tree cameraleopard tree cameraleopard tree camera

Camera trap captures of (A) Jo Jo Farque and (B) Mafana (centre and right) at two different trap sites and while tracking him with radio telemetry. Distinctive spot patterns are visible. Besides individual patterns, additional features e.g. body size, colouring, sex, external marks (scars, ear cuts) can be used for identification purposes.

Camera trapping is a non-invasive method of observing wildlife with only a minimum of disturbance to the target species and therefore, provides ideal conditions to study the rare, cryptic and elusive. They are able to illustrate a 24h-cycle providing day and night time data whereas many other methods are restricted to diurnal use only (e.g. line transect surveys, VHF-telemetry) which makes them ideal to study and observe nocturnal species. Remote cameras can be left in the field unattended for several weeks without the supervision or the continuous presence of a researcher or field assistant. Depending on the environmental conditions as well as technical factors such as battery operating time, cameras can operate in the field for several weeks and months.

Since camera traps are mainly independent from field conditions such as weather, vegetation, habitat and terrain their use can be applied for a huge variety of ecosystems (Silveira et al. 2003) in which the use of alternative methods can be limited or restricted. Thus, camera trap studies have been conducted in different ecosystems and vegetation zones ranging from humid and dry forests (Karanth, 1998; Cuellar et al. 2006) over grassland habitats (Harris, 1996) to alpine ecosystems (Jackson et al. 2006; Xu et al. 2008).

camera set-up

Fig. 1: (A) Exemplary camera set – up: The camouflaged white flash camera trap is fastened to a trunk of a tree targeting a piece of bait on the opposite site. (B) Diagram of a Cuddeback Capture: Cameras are equipped with a heat-and-motion sensor that triggers the camera when a difference between the target- and the surrounding background temperature is detected.
 

The leopard population size will be sampled via the application of remote wildlife camera traps. Therefore a closed model capture – recapture sampling approach will be used that is estimating abundance based on the number of individuals captured and the frequency of their recaptures (Silver et al. 2004). Capture-recapture (C-R) models take into account that not all individuals in the population are necessarily detected (Rovero et al. 2013) and rely on the assumption that the target species can be individually identified via natural marker or other phenotypic features (Karanth & Nicholas 1998). C-R models are the commonly used strategy to estimate abundance of elusive terrestrial mammals with particular focus on felids (Foster & Harmsen, 2012). Because leopards are easily identifiable via their spot patterns, camera trapping provides a reliable and practical tool to estimate abundance and density.

Secondly, C-R assumes that the sampled population is geographically as well as demographically closed during the study period and that no births, deaths, immigrations or emigrations occur during the sampling period (Karanth & Nicholas 1998; Silver at al. 2004). It is crucial that the sampling period ensures demographic closure and therefore doesn’t exceed a certain threshold. Karanth et al. (1998, 2000) sampled a tiger population in India for 3 months while Silver at al. (2004) used a 2-months period to estimate abundance of a jaguar population in South America. The long intervals before a leopard returns to a defined starting point within its natural movement need to be considered additionally when determining the time frame of the study.

Finally, it is critical to optimise camera trap placement without leaving any gaps to ensure a complete coverage of the entire study area to maximize capture probability and the number of individuals caught on the camera (Karanth & Nicholas 1998). The determination of inter-camera trap distances is crucial so that no leopard has a zero capture probability during the sampling period. The smallest home range size of a female representative of the target species is commonly used to determine the area that should be covered by at least one trap site (Karanth & Nicholas 1998, 2000; Silver at al. 204; Soisalo & Cavalcanti, 2006, Marnewick et al. 2008). The set-up of more than one trap within the defined home range increases the chances that also individuals of the opposite sex as well as from different age classes will be exposed to the camera (Soisalo & Cavalcanti, 2006) and decreases the probability that individuals inhabit home ranges that fall between trap sites (Foster & Harmsen, 2012). Available VHF-telemetry data on the collared cats in the reserve as well as consulted literature will be used to determine the inter-camera trap distance based on average female leopard home range sizes.

 

Field Methods

Due to a limited number of available camera traps the study area will be divided into equal sized blocks that will be covered sequentially for the same amount of time (Karanth & Nicholas, 2002; O’Brien et al. 2003). A preliminary inspection will be implemented in order to identify suitable camera trap locations and Geographic Position System (GPS) points will be taken of every trap site and transferred onto a digital map. Camera trap studies aim to capture as many individuals as possible. To enhance accuracy of abundance estimates it is essential to chose trap sites that increase capture probability. Therefore camera traps should preferably be placed in areas that suggest an elevated frequency of leopard occurrence such as dry riverbeds, riverbanks and/or frequently used roads and pathways.

To increase leopard capture frequencies baited camera stations were used. Tall trees at each trap site were selected and baits positioned approximately 1.5 – 2.5 metres above the ground to prevent theft from other carnivores. A single camera trap was placed 2 - 3 metres perpendicular to the branch on which the leopard was expected to occur to access the bait. Cameras were housed in a protective case (CuddeSafe®) to prevent cameras from wildlife damage. Baits were fastened with double wound wire to ensure that leopards didn’t remove bait and feed out of sight.

Cameras were serviced every 6 – 7 days in order to verify battery status, change of SD cards, renew bait and ensure the correct functionality of each trap.

 

Identification

Photographs from each trap site will be evaluated. Leopards are identified via their unique spot patterns. Each individually identified leopard will be assigned with a unique identification number. If applicable, sex and age class are recorded. A capture history will be generated for each sub-sampled block and afterwards combined to form one data set for the entire study period (Soisalo & Calvalcanti 2006).

Therefore a standard X-matrix format is used where "1" indicates the presence of an individual during a sampling occasion and "0" the absence during that occasion (Karanth & Nicholas, 2004) (Tab. 1). Due to the fact that leopards are mainly nocturnal, a sampling occasion is defined as a 24-hour period starting and ending at midday (12h00) (Du Preez et al. 2014). After the effective sampling period, camera traps will be used as part of a long-term monitoring program of the sampled leopard population. 

Tab. 1: Example of a capture history data set showing the number of positively identified leopards (Pp) and the number of days per sampling period (sampling occasion). 0 = no visit; 1 = visit detected. The capture history of Pp1 (00011000) indicates its capture during sampling occasion 4 and 5 and absence during 1, 2, 3, 6, 7 and 8.

  Sampling Occasions (n=8)
ID 1 2 3 4 5 6 7 8 ...
Pp1 0 0 0 1 1 0 0 0  
Pp2 1 0 0 0 1 1 0 0  
Pp3 ...                
...                  

The number of captured and re-captured animals will be analysed by using a statistical model (e.g. CAPTURE) and abundance and population size (N) generated. The calculated abundance will be used to derive an estimation on the leopard density (D): D = N/A whereas A is defined as the actively sampled area.

Trap success between trap stations will be evaluated and compared in order to reflect spatial distribution patterns. Distances moved between traps during the sampling period will be used to calculate minimum home range estimates for each cat.

After the effective sampling period camera traps will be used as part of a long-term monitoring program for the sampled leopard population.

 

Carnivore Collaring

Besides their central objective to capture and identify leopards in the study area, camera traps are assisting to locate sites of high leopard occurrences or to identify hot spot area of certain individuals. In identified areas, remotely triggered steel mesh box traps are deployed in order to catch and collar specific individuals to study and monitor their ecology and behavior in an island-bound conservation area.

The box traps are specially designed to reduce the stress that live trapping often causes to these species and therefore reduce injury to the animals. The traps are larger than standard box traps (the smaller traps measure 2.8m x 1m x 0.8m and the larger 3m x 1.5m x 2.4m) to allow more space for the trapped animal. To attract carnivores, the traps will contain bait fastened in a back corner of the trap to prevent it from being removed. The box traps will be thatched with grass on the inside and outside and are additionally surrounded with thorn bushes to blend into the natural surroundings and ensure camouflage (McCarthy et al., 2013). A live camera will be positioned approximately 4 - 5 meters in front of the trap entrance, facing the bait inside the trap. An infrared light will be installed on top of the camera to ensure night vision. Electricity for the setup will be provided by a solar panel. A single antenna will be connected to a Wi-Fi system that enables the transmission of data to a Live ClientSystem (VIVOTEK, Vast©¬) and that allows for real-time remote monitoring. A 12 Volt battery will be connected to a car starter motor solenoid provided with a relay system on top of the trap. Once the solenoid is activated by an operatorvia the Live ClientSystem (VIVOTEK, VAST©), an output signal will be triggered which closes the gate of the trap with a 4 second delay. As soon an animal is captured the darting team including a veterinarian will be deployed to the site of the trap in order to immobilize the animal within one hour of being trapped.

The live camera system has the advantage that an operator can specifically distinguish between target and non-target animals, as well as previously collared individuals. It also prevents the need for a trigger plate in the middle of the box which often causes injury to trapped carnivores in traditional cage traps.

During each immobilization, animals will undergo a full veterinary health evaluation and will be thoroughly evaluated for any injuries. Morphometric data and biological samples will also be collected on each animal.

Photographs of both flanks and the face (both sides and a frontal view) will be taken, to catalogue individual spot patterns. Each carnivore will be fitted with a VHF radio collar. Sex will be recorded and age class will be estimated based on size, tooth abrasion (Stander, 1997) and general condition. Afterwards animals will be moved into a recovery crate for recovery. To prevent danger to the animals after immobilization they will be kept in a wooden crate until fully recovered before release into the reserve.

VHF radio telemetry can provide detailed information on species specific movement patterns, home range utilization, habitat use and inter- and intraspecific interactions and is particularly suitable for species that are nocturnal, secretive and occur at low densities. Information gathered over time will give indication on how much space an animal requires, which habitat is preferred and if space is shared with other individuals. The study of ecological parameters such as home range and territory sizes, prey preferences and spatial utilization patterns are important tools that help to develop long-term and sustainable conservation and management plans.

box trap set up

box trap capture

mafana capture

boxtrap for africat predator study 

In order to specifically catch leopards, the entrance to the box trap will be elevated and is, thus inaccessible for brown or spotted hyenas

 

leopard in box trap carrying caught leopard madiba transporting leopard
measuring leopard blood sample leopard collaring leopard
taking blood sample leopard measuring teeth leopard bwana 2014 collar
radio collars visitors with leopard jo jo farque leopard release

 

References:

Cuellar, E., Maffei, L., Arispe, R. & Noss, A. (2006) Geoffroy’s cat at the northern limit of their range: activity pattern and density estimates from camera trapping in Bolivian dry forests. Studies on Neoptropical Fauna and environment, 41, 169 - 177.

Foster, R. & Harmsen, B. J. (2012) A critique of density estimation from camera-trap data. The Journal of Wildlife Management, 9999, 1 - 13.

Gaston, Kevin J., et al. The ecological performance of protected areas. Annual Review of Ecology, Evolution, and Systematics 39 (2008): 93-113.

Gil-Sanchez, J. M., et al. (2011) The use of camera trapping for estimating Iberian lynx (Lynx pardinus) home ranges. European Journal of Wildlife Research, 57, 1203 - 1211.

Hayward, M. W., O'Brien, J., and Kerley, G. I. H. (2007) Carrying capacity of large African predators: predictions and tests. Biol. Conserv. 139: 219-229.

Harihar, A., Gosh, M., Fernandes, M., Pandav, B. & Goyal, S. P. (2010) Use of photographic capture-recapture sampling to estimate desity of striped hyena (Hyaena hyaena): implications for conservation. Mammalia, 74, 471 - 479.

Harris, R. B. (1996) Wild ungulate surveys in grassland habitats: Satisfying methodological assumptions. Chinese Journal of Zoology, 31, 16 - 21.

Heilbrunn, R. D., Silvy, N. J., Tewes, M. E. & Peterson, M. J. (2003) Using automatically triggered cameras to individually identify bobcats. Wildlife Society Bulletin, 748 - 755.

Henschel, P., Hunter, L., Breitenmoser, U., Purchase, N., Packer, C., Khorozyan, I., Bauer, H., Marker, L., Sogbohossou, E. & Breitenmoser-Wursten, C. 2008. Panthera pardus. The IUCN Red List of Threatened Species. Version 2014.2. (http://www.iucnredlist.org) Downloaded on 13 November 2014.

Jackson, R. M., Roe, J. D., Wangchuk, R., & Hunter, D. O. (2006). Estimating Snow Leopard Population Abundance Using Photography and Capture‐Recapture Techniques. Wildlife Society Bulletin, 34(3), 772-781.

Khorozyan, I. G., Alexander G. M., and Alexei V. A. (2008) Presence–absence surveys of prey and their use in predicting leopard (Panthera pardus) densities: a case study from Armenia." Integrative Zoology 3.4 322-332.

Karanth, K. U (1995): Estimating tiger (Panthera tigris) populations from camera trap data using capture-recapture models. Biological Conservations, 71, 333 - 338.

Kelly, M. J. (2008) Design, evaluate, refine: camera trap studies for elusive species. Animal Conservation, 11, 182 -184.

Marnewick, K., Funston, P. J., & Karanth, K. U. (2008). Evaluating camera trapping as a method for estimating cheetah abundance in ranching areas. South African Journal of Wildlife Research, 38(1), 59-65.

Noss, A. J., Cuellar, R. L., Barrientos, J., Maffei, L., Cuellar, E., Arispe, R., Rumiz, D. & Rivero, K. (2003) A camera trapping and radio telemetry study of lowland tapir (Tapirus terrestris) in Bolivian dry forests. Newsletter of the IUCN/SSC Tapir Specialist Group. Tapir Conservation, 12, 24 - 32.

O'Brien, T. G., & Kinnaird, M. F. (2008). A picture is worth a thousand words: the application of camera trapping to the study of birds. Bird Conservation International, 18, 144.

Ray, J. C., Hunter, L.T.B & Zigouris, J. (2005) Setting conservation and research priorities for larger African carnivores. Working paper 24. Wildlife Conservation Society, New York.

Rovero, F., Zimmermann, F., Berzi, D., & Meek, P. (2013). " Which camera trap type and how many do I need?" A review of camera features and study designs for a range of wildlife research applications. Hystrix, the Italian Journal of Mammalogy, 24(2), 148-156.

Silveira, L., Jacomo, A. T. A. Diniz-Filho, J. A. F. (2003) Camera trap, line transect census and track surveys: a comparative evaluation. Biological Conservation, 114, 351 - 355.

Silver, S. C., Ostro, L. E., Marsh, L. K., Maffei, L., Noss, A. J., Kelly, M. J., ... & Ayala, G. (2004). The use of camera traps for estimating jaguar Panthera onca abundance and density using capture/recapture analysis. Oryx, 38(02), 148-154.

Soisalo, M. K., & Cavalcanti, S. (2006). Estimating the density of a jaguar population in the Brazilian Pantanal using camera-traps and capture–recapture sampling in combination with GPS radio-telemetry. Biological Conservation, 129(4), 487-496.

Sollmann, R., Mohamed, A., Samejima, H. & Wilting, A. (2012) Risky business or simple solution - Relative abundance indices from camera trapping. Biological conservation, 159, 405 - 412.

Stein, A., Andreas, A. & Aschenborn, O (2011) Namibian National Leopard Survey – 2011. Final Report. Ministry of Environment and Tourism.

Turpi, J., Barns, J., De Longcamp, M. & Paxton, M. (2010) Sustainable Financing Plan for Namibia’s Protected Area System: Ministry of Environment and Tourism. Directorate of Parks and Wildlife Management.

Wallace, R. B., Gomez, H., Ayala, G. & Espinoza, F. (2003) Camera trapping for jaguar (Panthera onca) in theTuichi Valley, Bolivia. Journal of Neotropical Mammals, 10, 133 - 139.

Xu, A. et al. "Status and conservation of the snow leopard Panthera uncia in the Gouli Region, Kunlun Mountains, China." Oryx 42.03 (2008): 460-463.

 

Leopard Behaviour in the Okonjima Nature Reserve

1 female 2 cubs
1 female and 2 cubs.
1 female scent marking
Female leopard scent marking.
leopard hunting warthog
Leopard hunting warthog.
leopard cleaning prey before eating it
Leopard 'cleaning' prey before eating it.
female and male during mating
Mating leopards.
female and male during mating
Mating leopards.
female and male during mating
Mating leopards.
female and male during mating
Mating leopards.
female and male during mating
Mating leopards.

Mating
"Both leopards and lions have exactly the same mating rituals which, when averaged out, has them mating every 15 minutes for up to 5 days. This means that if they last a full 5 days, they can mate more than 250 times. This may seem a little excessive, but there is a good reason for this. In humans, females produce an egg every 28 days. If it is not fertilised, the egg and the lining of the uterus will be discarded. This does not happen with leopards though. The act of producing something that is not used is a waste of energy, so the female leopard requires a stimulus to start ovulation.

 

The female leopard will therefore go into oestrus. This is a state in which her hormones are at a level at which she is able to produce eggs. She will also leave a scent in her urine which will indicate to the male that she is ready to mate. As she enters oestrus the female will begin to mark her territory more often than usual and will call to attract the dominant male in the area. He will latch onto her scent using a gland on his palate called the organ of Jacobson, which is able to measure hormone levels and determine whether or not the female is ready to mate.

 

Once the mating begins it is a non-stop affair, filled with uneasiness and violence. In order to stimulate the female to ovulate, the male has barbs on his penis which dig into the female. When the penis is retracted it hurts the female causing her to lash out at the male. As the mating ritual continues she will produce eggs. Due to his weak sperm, a male leopard has to mate with a female often enough to ensure that fertilisation takes place.

 

It is interesting to note that current research suggests female leopards can tell whether or not a male is capable of being the territorial male and remain dominant for a long period of time. If she determines that he is not, she has the ability to make herself less fertile. This is because if she conceives and the male is ousted from his territory, a new male will kill her existing cubs. If this happens she would have completely wasted her energy.

 

It has also been established that females will mate with more than one male within a very short period, thus lowering the chances of infanticide. This is particularly important for female leopards whose territories overlap those of two dominant males. Infanticide in leopards accounts for 40% of cub deaths and is therefore one of the major considerations for females when choosing a mate.” Sabi Sabi


Updates:

Phase 1: AOPPDS Phase 1
Phase 2: AOPPDS Phase 2
Phase 3: AOPPDS Phase 3
Phase 4: AOPPDS Phase 4
Phase 5: AOPPDS Phase 5

 

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