Eaton RL. 1970. Notes on the reproductive biology of the cheetah. International Zoo Yearbook 10, 86-90

This paper summarizes the information on cheetah reproduction biology in the literature and data from a field study of cheetah ecology in Nairobi National Park and Masai Amboseli Game Reserve, Kenya. Data on the reproductive biology of endangered species such as the cheetah is important as an aid to breeding them in captivity.
Births appear to be seasonal relating to game concentrations and rainy seasons. Factors affecting hunting success that are related to seasonal changes, other than game concentrations, are vegetation growth and colour. Breeding success would be highest in those females that gave birth during the wet season so that food would be most easily available when the cubs required it.

Eaton_1970_Notes_on_the_reproductive_biology_of_the_cheetah.pdf

Fitch-Snyder H. 1988. Environmental factors influencing captive reproduction of cheetahs.
Supplement to the AAZPA Cheetah SSP Husbandry Manual, 8 pp.

Five North American zoos, each having produced at least six litters at the time the survey was made, were examined and compared. A husbandry questionnaire was used which included questions concerning enclosure types, exposure to other animal species, diet, social groupings, oestrus behaviour, parturition and maternal care.

Fitch-Snyder_1988_Environmental_Factors_Influencing_Captive_Reproduction_of_Cheetahs.pdf

Frank J, Saffoe C. 2005. Breeding management strategy for cheetahs (Acinonyx jubatus) at the Smithsonian's National Zoological Park. Animal Keeper's Forum 7/8, 393-397.

It is no shocking revelation to any animal keeper that cheetahs (Acinonyx jubatus) are very difficult to breed in captivity. Many cat species reproduce poorly in zoos, but cheetahs seem to have more challenges, ranging from poor genetic variability and gamete malformation to behavior nuances that complicate husbandry practices. At the Smithsonian's National Zoological Park (SNZP) we have been trying hard to overcome these obstacles and breed cheetahs. Although we cannot control their genetic situation we can manipulate our management to increase the potential for creating optimum breeding conditions.

Frank_&_Saffoe_2005_Breeding_management_strategy_at_Smithonian.pdf

Kelly MJ, Laurenson MK, FitzGibbon CD, Collins DA, Durant SM, Frame GW, Bertram BCR, Caro TM. 1998. Demography of the Serengeti cheetah (Acinonyx jubatus) population: the first 25 years. J Zool , Lond 244:473-88.

Data are presented on the demography and reproductive success of cheetahs (Acinonyx jubatus) living on the Serengeti Plains, Tanzania over a 25-year period. Average age at independence was 17.1 months, females gave birth to their first litter at approximately 2.4 years old, interbirth interval was 20.1 months, and average litter size at independence was 2.1 cubs. Females who survived to independence lived on average 6.2 years while minimum male average longevity was 2.8 years for those born in the study area and 5.3 years for immigrants, with a large proportion of males dispersing out of the Plains population. Females produced on average only 1.7 cubs to independence in their entire lifetime and their average reproductive rates were 0.36 cubs per year or 0.17 litters per year to independence. Variance in lifetime reproductive success in the cheetah is similar to that of other mammals. No significant negative correlations were found between adult cheetah population size and numbers of cubs reaching independence, implying that the Plains population had not reached carrying capacity. Annual numbers of adult female cheetahs only were correlated with rainfall. Adult female cheetah numbers were not correlated with adult female lion numbers on the Plains, however, reproductive rates of cheetahs were negatively correlated with the presence of lions while cheetahs had cubs. Moreover, cheetah reproductive success was lower during the period of high lion abundance (1980±1994) than during the earlier period of relatively few lions (1969±1979). Litter size at independence dropped from 2.5 to 2.0, lifetime reproductive success declined from 2.1 to 1.6 cubs reared to independence, and the reproductive rate (cubs/year) decreased from 0.42 to 0.36 from the earlier to the later period. Cheetah reproductive success showed little association with the presence of Thomson's gazelle at sightings except for a negative correlation between large numbers of gazelle (200±500) and reproductive success possibly because hunting success decreases with increasing prey herd size, or because cheetahs always lose in direct competition with other predators which are attracted to large congregations of prey. In addition, cheetah reproductive success was negatively correlated with the presence of Grant's gazelles (11 or more) perhaps because Grant's gazelles were more likely to occur consistently in dry areas.

Kelly_et_al_1998_Long-term_cheetah_demography.pdf

Kelly MJ, Durant SM. 2000. Viability of the Serengeti cheetah population. Conservation Biology 14(3):786-97.

Most recent population viability analyses, especially those of long-litled species, rely on only a few years of data or data from a closely related species, combined with educated guesswork, to estimate model parameters and the variability surrounding those measures. This makes their conclusions or predictions difficult to evaluate. In our study, we used 20 years of demographic data on Serengeti cheetahs (Acinonyx jubatus) to conduct a population viability analysis. First we constructed a model of the deterministic growth rate and found that the cheetah population is nearly serf-replacing (A = 0.997). Our model showed that population growth was most strongly influenced by adult survival followed by juvenile survival, which is typical of long-lived, iteroparous species. We then examined extinction risk and long-term projections of cheetah population size with our stochastic model, Popgen. We compared the projections with over 20 years of field data and found that demographic stochasticity trials produced a stable population size, whereas environmental stochasticity trials were slightly more pessimistic. Extinction risk was highly sensitive to both adult survival and juvenile survival (from 0-1 years). Decreasing the variance in survival rates also decreased extinction risk. Because lions are the major predator on cheetah cubs, we used our demographic records to simulate the effect of different lion numbers on juvenile survival. High lion abundance and average lion abundance resulted in extinction of nearly all cheetah populations by 50 years, whereas with low lion abundance most cheetah populations remained extant. Conservation of cheetahs may not rely solely on their protection inside national parks, but may also rely on their protection in natural areas outside national parks where other large predators are absent.

Kelly_&_Durant_2000_Serengeti_cheetah_population_viability.pdf

Laurenson MK, Caro TM, Borner M. 1992. Female cheetah reproduction. National Geographic Research & Exploration 8(1):64-75.

To provide baseline information for the mounting effort to conserve cheetahs, female reproduction and offspring mortality were studied in a free-ranging population in the Serengeti National Park, Tanzania. Most adult females were capable of reproduction; conceptions were more frequent in wet season months, possibly as a result of increased food availability in the form of newborn Thomson's gazelles. Cub mortality was extremely high, and cheetahs had only a 6% chance of reaching independence at 18 months of age. Predation by lions was the principal source of mortality, although some litters were abandoned by their mothers when nearby prey was scarce. Mothers produced new litters rapidly following the loss of an unweaned litter. These findings suggest that cheetahs may not be able to maintain high densities in the presence of other large carnivores, and that many of the problems zoological institutions experience in breeding cheetahs are specific to the captive context.

Laurenson_et_al_1992_Cheetah_reproduction.pdf

Laurenson MK. 1995. Implications of high offspring mortality for cheetah population dynamics. In:Sinclas ARE, Arase P, editors. Serengeti II: Research, Conservation and Management of an Ecosystem. Chicago: University of Chicago Press.

In this chapter recent findings on the causes of juvenile mortality in cheetahs are discussed and preliminary data presented suggesting that predation on cheetah cubs is an important factor affecting the Serengeti cheetah population. First, the relative importance of factors affecting cheetah fecundity and mortality is considered and second, the effect of variation in fecundity and mortality factors on female lifetime reproduction and cub recruitment rates is simulated. In addition, the implications for cheetahs of recent changes in carnivores numbers in the Serengeti-Mara ecosystem are discussed. Finally, the relevance of these findings for cheetah populations elsewhere in Africa and their application to future conservation is considered. Although there is little information on the factors affecting adult mortality in females, these findings provide tentative evidence that offspring mortality, in particular from lion predation, may have a critical effect on the size of the Serengeti cheetah population. Interaction between cheetahs and other predators are potentially important for the population dynamics of cheetahs in the Serengeti-Mara ecosystem and deserve further scrutiny.The suggestion that other large predators have a detrimental effect on cheetah population size, is important from conservation perspective. Some pastoralists and ranchers tolerate cheetahs to a greater extent than lions or hyenas and cheetahs seem to prosper in these areas. As cheetahs may have difficulty in reaching large numbers in isolated protected areas, it is perhaps in these multiple land use areas that conservation efforts should be concentrated to find ways in which continuing conflicts between cheetahs and man can be minimized.

Laurenson_-_Implications_of_high_offspring_mortality_for_cheetah_population_dynamics.pdf

Laurenson MK. 1995. Behavioural costs and constraints of lactation in free-living cheetahs. Animal Behaviour 50:815-26.

Increased energetic expenditure during lactation must be met either from metabolic stores or by increasing food intake. Additional behavioural costs and constraints may be imposed on those species that conceal their young in a fixed place. This study examines how wild cheetahs, Acinonyx jubatus, alter their behaviour during lactation and whether denning involved extra costs or constraints. Females almost doubled their food intake when lactating, although only mothers with cubs in the lair increased the time they spent drinking. Higher levels of food intake were achieved by catching a greater proportion of larger prey items, hunting these at a higher rate and by increasing the success rate of hunts. Lactating females with emerged cubs increased the proportion of time they spent observing and hunting relative to time spent resting and moving. When lactating females had cubs in the lair they were on the move for longer each day, travelled further, had restricted ranging patterns and tended to make more kills in the heat of the day than when accompanied by their cubs. These latter results suggest that the need to find a safe place, near water, in which to conceal immobile and vulnerable cubs may impose additional behavioural constraints and costs and lactating females.

Laurenson_1995_Behaviour_of_cheetahs_during_lactation.pdf

Laurenson MK, Wielebnowski N, Caro TM. 1995. Extrinsic factors and juvenile mortality in cheetahs. Conservation Biology 9(5):1329-31.

Cheetahs are thought to exhibit low genetic variability and, as a consequence, may suffer high juvenile mortality, impaired reproduction, and susceptibility to disease. Recently, however, independent criticisms have been levelled at this body of work questioning both the analysis and the standards of evidence for lack of genetic diversity as well as the evidence that cheetahs are suffering as a consequence. O'Brien (1994a) combines a number of points in his first rejoinder to these criticisms, including novel interpretations of ecological data that we obtained and concern about their collection and analysis. Here, we first show that this interpretation is seriously flawed and that our conclusion that predation is the key source of mortality in the wild is valid. Then, extending previous criticisms and continuing to focus on this one consequence of lack of genetic variability, we discuss O'Brien et al.'s (1985) interspecific comparison of juvenile mortality in captivity by highlighting analytical problems and presenting new data on captivity.

Laurenson_et_al_1995_Cheetah_mortality.pdf

Laurenson MK. 1995. Cub growth and maternal care in cheetahs. Behavioral Ecology 6(4):405-9.

Using cub growth as an index, the influence of maternal nutrition, litter size, and cub sex on maternal care in cheetahs (Acinonyx jubatus) were examined and cub and litter growth rates compared with those of larger felids. Seventy-nine free-living cheetah cubs in 21 litters from 15 mothers were weighed at least once between 6 and 48 days of age. Eleven litters were weighed at the begging and end of a 5-day observation of their mothers. The mean cub growth rate varied significantly between litters, due primarily to differences in maternal food intake. Growth declined sharply when maternal foods intake was less than 1.5 kg/day, but did not increase with greater levels of food intake. Lower limits of growth rates may therefore have been set by the mother's food intake, whereas upper limits may be set by the intrinsic physiological ability of cubs to grow. Although male cubs were heavier than female cubs in the same litter when first weighed, major differences in growth rate between the sexes were not apparent at this stage. Both cheetah cubs and litters grow fast relative to other large felids, and it is argued that this may be an adaptation to the high rate of cheetah juvenile mortality from predation.

Laurenson_1995_Cheetah_cub_growth.pdf

Lindburg DG, Durrant BS, Millard SE, Oosterhuis JE. 1993. Fertility assessment of cheetah males with poor quality semen. Zoo Biology 12(1):97-104.

Reports on semen quality of the cheetah (Acinonyx jubatus) indicate that high percentages of abnormal morphs and sperm concentrations, 10 times lower than in domestic cats, are found in all populations. These characteristics are believed to result from unusual genetic homozygosity, hypothesized to have been caused by passage of the species through one or more population bottlenecks during its recent history. In a sample of 12 captive living males, we found semen characteristics to be equal or inferior to those previously reported for all males living in other captive facilities. Ten of these males (83.3%) nevertheless produced pregnancies. Seventeen of 19 pregnancies, resulted from matings during a single oestrus. This examination of the reproductive potential of males having comparatively inferior ejaculate quality supports the suggestion that husbandry programs may be more significant than physiological impairment in causing the low birth rates in captive cheetahs. These results also have implications for ascertaining fertility thresholds in mammalian populations undergoing increased levels of inbreeding as a consequence of habit deterioration.

Lindburg_et_al_1993_Fertility_assessment_of_cheetah_males.pdf

Marker LL, Dickman AJ, Leo RM, Mills MGL, MacDonald DW. 2003. Demography of the Namibian cheetah, Acinonyx jubatus jubatus. Biol Conserv 114:413-25.

Namibian cheetahs have suffered, and continue to suffer, high levels of removal due to conflict with local farmers, and it is important to understand the demography of this population in order to determine its likely persistence. Examination of cheetahs reported live-trapped or killed by local farmers, combined with subsequent information from radio-telemetry, allowed demographic parameters such as sex ratios, age and social structure, litter size, interbirth intervals and survivorship to be estimated for cheetahs on Namibian farmlands. Cub mortality was relatively low, but adult mortality was high, particularly for males, and peaked at 5-6 years of age. Neither marking nor relocating cheetahs seemed to affect survivorship, and there was no difference in survivorship between the sexes. Time spent in captivity did not appear to affect survival after release. These findings will be useful in formulating recommendations regarding the conservation and sustainable utilization of cheetah populations outside protected areas.

Marker_et_al_2003_Demography_of_the_Namibian_cheetah.pdf

Osthoff G, Hugo A, Wit M. 2006. The composition of cheetah (Acinonyx jubatus) milk. Comparative Biochemistry and Physiology 145, 265-269.

Milk was obtained from two captive bred cheetahs. The nutrient content was 99.6 g protein; 64.8 g fat; and 40.21 g lactose per kg milk. Small amounts of oligosaccharides, glucose, galactose and fucose were noted. The protein fraction respectively consisted of 34.2 g caseins per kg milk and of 65.3 g whey proteins per kg milk. Very little variation in milk composition among the individual cheetahs was noted. Electrophoresis and identification of protein bands showed a similar migrating sequence of proteins as seen in lion's and cat's milk, with small differences in the â-caseins. The lipid fraction contains 290.4 g saturated and 337.3 g mono-unsaturated fatty acids per kg milk fat respectively. The high content of 279.5 g kg-1 milk fat of polyunsaturated fatty acids is due to a high content in á-linolenic acid. No short chain fatty acids, but substantial levels of uneven carbon chain fatty acids were observed.

Osthoff_et_al_2006_Milk_composition_in_cheetah.pdf

Tennant MB, Craig SJ. 1977. Breeding cheetahs at the Lion Country Safari Parks: a summary. Int Zoo Yb 17(167):169.

A six-year study aimed at determining the conditions under which the cheetah is most likely to reproduce. Comprehensive behavioural notes were taken and still and cine cameras and tape recorders were used in the collection of data. This paper summarizes the conditions and events leading to births and attempts to evaluate the factors involved. Main factors that have contributed to the success of the cheetah breeding programme include a quality commercially prepared diet, a large compound with wide field of vision and which excludes sensory awareness of other large carnivores, females of at least three years of age, predetermined sexual activity levels for males and the fighting, females was apparently courted and impregnated by one male alone.

Tennant_&_Craig_1977_Breeding_Cheetahs_at_the_Lion_Country_Safari_Parks.pdf

Re-introduction programs rely on reproductively sound individuals and high offspring survival once captively bred species are released into their natural habitat. Some species involved in captive breeding programs reproduce poorly: one prominent example is the cheetah (Acinonyx jubatus). Here we use the cheetah as a case study to test predictions from all current hypotheses explaining this phenomenon by monitoring postemergence cub survival and assessing the reproductive status of free-ranging and captive female cheetahs in Namibia with ultrasonography. We show that reproductive activity and health of cheetah females is determined by reproductive history and age rather than innate rhythms, captive stress, or lack of genetic diversity, and that postemergence cub survival under natural conditions in a predator-free habitat is high. Our results suggest that management practices of captive breeding and re-introduction programs should encourage early  reproduction in females to induce long-lasting and healthy reproductive performance. With this practice, re-introduction projects might increase their chances of success.

Wachter_et_al_2011_The_cheetah_controversy_revisited.pdf

Wildt DE, Bush M, Howard JG, Meltzer DG, Van Dyk A, Ebedes H, Brand DJ.  Reproductive traits in the male south African cheetah. Ref Type: Unpublished Work

The purpose of the study has been to evaluate and compare reproductive traits in established populations of male Transvaal, South West or hybrid (Transvaal X South West) cheetahs maintained under two captive managements. We have determined that a great proportion of cheetah spermatozoa collected by electroejaculation are morphological abnormal. Furthermore we observed no major differences in reproductive traits based on a comparison of the Transvaal, South West or factor alone. However, a greater proportion of male cheetahs allowed free range with females in a large territorial enclosure produce ejaculates which tend to be of greater quality compared to cheetahs continuously supported in small camps with only occasional female exposure.

Wildt_et_al_-_Reproductive_traits_in_the_male_south_African_cheetah.pdf

Wildt DE, Brown JL, Bush ME, Barone MA, Cooper KA, Grisham J, Howard JG. 1993. Reproductive status of cheetahs (Acinonyx jubatus) in North American zoos: the benefits for physiological surveys for strategic planning. Zoo Biology 12(1):45-80.

Under the mandate of a Species Survival Plan (SSP), reproductive status was assessed in 128 cheetahs maintained in 18 different institutions in North America. A mobile laboratory research team evaluated cheetahs using anaesthesia, serial blood sampling, electroejaculation (males), and laparoscopy (females). Biomaterials were also collected for parallel studies of genetics, nutrition, and health. There was no mortality, and cheetahs were capable of reproducing naturally after these intense manipulatory examinations. No marked differences were observed in reproductive or endocrine characteristics between proven and unproven breeders. However, males consistently produced teratospermic ejaculates, and cheetah sperm were compromised in conspecific or heterologous in vitro fertilization systems. Structurally abnormal sperm were found to be filtered by the oocyte's zona pellucida. More than 80% of the females were anatomically sound, but morphological and endocrine evidence suggested that ~50% or more of the population may have had inactive ovaries at the time of the examination. Males ranging in age from 15 to 182 months produced spermic ejaculates, but motile sperm numbers/ejaculate and circulating testosterone concentrations were highest in males 60 to 120 months old. Parovarian cysts were observed in 51.5% of female cheetahs, but comparisons between proven and unproven subpopulations revealed that this abnormality likely had no influence on fertility. Fresh luteal tissue not observed in any nonpregnant or nonlactating female, strongly suggesting that the cheetah is an induced ovulator. Overall survey results were discussed in the context of the etiology of reproductive inefficiency, especially with respect to the potential importance of biological versus management factors. Four high priority research areas in cheetah reproductive biology were identified:1) continuous monitoring of ejaculate quality in the extant population, while studying the impact of pleiomorphisms on fertility; 2) determining the potential relationship between libido and androgen production (excretion) in males; 3) confirming the extent of cyclic, or acyclic, ovarian activity in females; and 4) continued development of assisted reproductive techniques for enhancing management. In summary, a multidisciplinary, multi-institutional survey coordinated through the SSP is both possible and useful for generating a physiological and health database beneficial to driving further research and management initiatives.

Wildt_et_al_1993_Reproductive_status_of_cheetahs_in_North_American_zoos.pdf