Reproductive aging in captive and wild common chimpanzees: factors influencing the rate of follicular depletion

We examine and discuss evidence of contrasting differences in fertility patterns between captive and wild female chimpanzees, Pan troglodytes, as they age; in the wild females reproduce in their 40s, but captive studies suggest that menopause occurs around that time. Thus, despite the increased longevity generally observed in captive populations reproductive life span is shortened. We outline a hypothesis to explain the apparent differential pace of reproductive decline observed between wild and captive populations. The breeding schedules of captive primates may contribute to accelerated reproductive senescence because continuous cycling in captive animals results in early depletion of the ovarian stock and premature senescence. Available evidence supports the hypothesis that women with patterns of high oocyte loss experience earlier menopause. Chimpanzees in captivity live longer, and thus, similar to humans, they may experience follicular depletion that precedes death by many years. In captivity, chimpanzees typically have an early age at menarche and first birth, shorter interbirth intervals associated with short lactational periods as young mature faster, and nursery rearing, which allows mothers to begin cycling earlier. Variables typical of wild chimpanzee populations, including late age at menarche and first birth, long interbirth intervals associated with prolonged lactational periods, and a long period of female infertility after immigration, spare ovulations and may be responsible for the later age at reproductive termination. Finally, we describe and discuss the timing of specific reproductive landmarks that occur as female chimpanzees age, distinguishing between functional menopause (age at last birth) and operational menopause (end of cycling). Am. J. Primatol. 71:271–282, 2009. © 2008 Wiley‐Liss, Inc.     

Reproductive Parameters of Female<Emphasis Type="Italic">Pan paniscus</Emphasis> and <Emphasis Type="Italic">P. troglodytes</Emphasis>: Quality versus Quantity

We investigated intra- and interspecific differences in life history and reproductive parameters in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes). We compare the parameters of wild and captive females in order to shed light on the influence of habitat or specific differences or both on reproduction. We present new and additional information on reproductive parameters from captive bonobos and chimpanzees. Captive chimpanzees birth more live offspring and have a shorter interbirth interval, but experience higher infant mortality than captive bonobos. Although captive bonobo females tend to start reproduction at a younger age than chimpanzees, this is effectively only so for wild-born females of both species. Ultimately both species reach the same rate of production of offspring surviving to 5 yr. These results contrast with data from the wild. Wild bonobos tend to have higher reproductive success, a higher fertility rate and a shorter interbirth interval than wild chimpanzees. Reproduction is similar for wild and captive bonobos, which suggests that they are producing at their maximum under both conditions. Overall captive chimpanzees perform better than their wild conspecifics, probably because of lower feeding competition. Infant survival is the only specific difference not affected by captivity. Bonobo infants survive better, which suggests that chimpanzee infants are more at risk. We argue that the interspecific variation in reproductive parameters in captivity is related to the different influence of captivity on reproduction and different pressures of external sources of infant and juvenile mortality.     

Reproduction of the quokka, Setonix brachyurus, in captivity

This study records estimations of the chronology of certain events in the calendar of development of the quokka. One gestation period of between 25 and 26 days was observed and 22 joeys (young) had an average birth weight of 0.3852±0.0436 g. Vacation of the pouch occurred between 185 and 195 days of age. One thousand joeys were found to have a masculinity of 0–453 which is statistically smaller than parity. An estimation was made of the age at which the separate teeth erupt and a method of approximating the ages of juveniles and yearlings proposed by using dental eruption stages. The youngest recorded ages at which males and females became reproductively mature were 389 and 252 days respectively. After attaining reproductive maturity all animals born in captivity bred throughout the year. Animals taken from Bald Island, although showing seasonal breeding in the wild, will breed without periodic anoestrus interruption when taken into captivity. Captive animals may live seven years or longer. Using the chronology of some of the above events a comparison was made of the reproductive potential of the wild and domesticated populations.     

The effect of long non-reproductive periods on the genital health in captive female white rhinoceroses (Ceratotherium simum simum, C.s. cottoni)

White rhinoceroses suffer from a low reproductive rate in captivity. Intensive efforts to propagate specifically the northern white rhinoceros have been very limited. The dismal outlook for this subspecies in the wild makes successful ex situ breeding programs paramount. In this context, this study examined 48 southern and 6 northern white rhinoceroses using ultrasound and faecal hormone analysis to elucidate causes for female reproductive failure and to determine whether long non-reproductive periods have a detrimental impact on genital health. Results showed that 76% of the nulliparous females had intact hymenal membrane indicating these females had never been bred, at an age when their wild counterparts have delivered multiple offspring. Fifty-six percent of the studied population had various reproductive pathology. Cystic endometrial hyperplasia; leiomyomas of the cervix, uterus and ovary, adenoma; para-ovarian cysts and hydromucometra represent the scope of lesions identified. The stages of the lesions in nulliparous females correlated with age (r = 0.4, P < 0.05). Due to the severity of the lesions, 28% of the study population was considered post-reproductive. Therefore, the reproductive life span in some individuals was 10-20 years shorter than expected. However, in parous females the incidence of pathological lesions was significantly lower (P < 0.0001). Seventy-eight percent females studied had erratic or absent luteal activity. The hormone data corresponded with two ultrasonographic levels of ovarian activity, active and inactive, occurring within an age range of 3-19 years and 15-38 years, respectively. This suggests the lack of ovarian activity by reproductive mid-life in non-reproducing females. The accuracy of the ultrasound findings was validated by necropsy in nine animals showing a strong positive correlation (r2 = 0.9, p < 0.001). Our data suggests that the development of reproductive pathology and ovarian inactivity in white rhinoceros is an age-related consequence of long non-reproductive periods. This asymmetric ageing process of the genital organs can be prevented with the achievement of at least one pregnancy.     

Use of RAPD-PCR-analysis of cellular DNA for the evaluation of genetic polymorphism and subspecies diagnostics of the Far Eastern leopard Panthera pardus orientalis]

Genetic polymorphism in the Far-Eastern leopard subspecies Panthera pardus orientalis was examined by RAPD-PCR analysis of total cellular DNA. Primers diagnostic for the leopard subspecies were detected. These primers can be used for differentiating wild-living animals from those kept in captivity and distinguishing different breeding lines. Genetic diversity was shown to be lower in animals from the zoo (D = 0.058) and higher in a natural population (D = 0.137). Two lines of the Persian subspecies were genetically closest (D = 0.042), and the Persian and Far-Eastern subspecies represented by wild animals, genetically most remote (D = 0.274). The level of genetic divergence of the Persian subspecies and Far-Eastern subspecies (zoo animals) was lower (D = 0.108) than between representatives of the Persian subspecies from the captive and natural population (D = 0.220). On the basis of analysis of relatedness, performed using NTSYS and PHYLIP software packages, the possibility cannot be excluded that some P. p. orientalis animals from the zoo have a hybrid origin and may be assigned to another Far-Eastern subspecies.     

Negligible senescence in the longest living rodent,the naked mole-rat: insights from a successfully aging species

Aging refers to a gradual deterioration in function that, over time, leads to increased mortality risk, and declining fertility. This pervasive process occurs in almost all organisms, although some long-lived trees and cold water inhabitants reportedly show insignificant aging. Negligible senescence is characterized by attenuated age-related change in reproductive and physiological functions, as well as no observable age-related gradual increase in mortality rate. It was questioned whether the longest living rodent, the naked mole-rat, met these three strict criteria. Naked mole-rats live in captivity for more than 28.3 years, ∼9 times longer than similar-sized mice. They maintain body composition from 2 to 24 years, and show only slight age-related changes in all physiological and morphological characteristics studied to date. Surprisingly breeding females show no decline in fertility even when well into their third decade of life. Moreover, these animals have never been observed to develop any spontaneous neoplasm. As such they do not show the typical age-associated acceleration in mortality risk that characterizes every other known mammalian species and may therefore be the first reported mammal showing negligible senescence over the majority of their long lifespan. Clearly physiological and biochemical processes in this species have evolved to dramatically extend healthy lifespan. The challenge that lies ahead is to understand what these mechanisms are.     

Evaluation of the role of the national parks for Persian leopard (<Emphasis Type="Italic">Panther pardus saxicolor</Emphasis>, Pocock 1927) habitat conservation (case study: Tandooreh National Park,Iran)

Having knowledge of the habitat requirements of top predators is important for their conservation and also for the stability of wildlife communities since they are keystone species of ecosystems. This study aims to predict suitable habitats for the Persian leopard (Panther pardus saxicolor, Pocock 1927) in Tandooreh National Park, Iran. This species is the largest leopard subspecies in the Middle East and is at high risk of extinction. Presence points for the species have been obtained from field data, literature review, atlas data, and IUCN records. Variables related to human disturbance, terrain, land use/land cover, and the leopards’ prey were used to predict the habitat selection of the Persian leopard by employing binary logistic regression. Our model predicted that the presence of Persian leopards is restricted to the southern, north-western, and western areas of the park which are under severe fragmentation threat. Moreover, leopards avoided areas near human development such as roads. We have also found that the availability of prey such as Capra aegagrus, Ovis vignei, and Ovis orientalis is one of the important parameters to predict habitat suitability for leopards. Our model provides a suitable tool for national park managers to identify biodiversity hotspots since leopards are often associated with high biodiversity.     

Reproductive parameters of the common genet <Emphasis Type="Italic">Genetta genetta</Emphasis> (Linnaeus, 1758) in Southwest Europe

We used information from 148 litters belonging to 119 different wild and captive female common genets (Genetta genetta) in SW Europe over a period of almost 50 years to study certain reproductive parameters. Data were obtained from a combination of sources including carcasses, live captures, telemetry-based studies, captive genets, and reports in the literature. The circannual pattern of breeding was fairly similar in both wild and captive genets, with births occurring in all months except December, albeit with a large peak in spring and a secondary peak in autumn. The sex ratio at birth was near to 1:1, and mean litter size was 2.16 ± 0.76 cubs/litter with a range of 1–5 cubs. Litters of two cubs were the most frequent both in wild and captive settings. In captivity, we have observed the births of replacement litters and double litters; however, we did not find evidence of wild females that having two litters in the same year. We conclude that the possibility of breeding at any time of the year is facilitated by the continuous spermatogenesis of the males and by the ability of females to enter into estrus after losing a litter prematurely and to breed twice in the same year.     

Unterartenfrage Persischer Leoparden geklärt

The Studbook for the Persian Leopard, Panthera pardus saxicolor, was analyzed. The whole population derives from a few founder animals, imported in the midth fifties from Iran and in the late sixties from Afghanistan. To avoid inbreeding later on the Iranian and the Afghan lines were mixed. A female imported in 1968 from Kabul to Cologne is represented in each of the more than 100 animals living today.This study deals with the question of subspecies of leopards in Afghanistan. Out of the 27 subspecies described four are believed to exist in Afghanistan. However, according to a molecular-biological revision of the species there is only one subspecies in Afghanistan, Panthera pardus saxicolor. To clarify the subspecies question various measures of furs have been taken in the bazars. The results revealed that the leopards in Afghanistan are the biggest of its species. However a further differentiation according to the area of origin within the country was not possible. Also the traditional differentiation on the basis of colours and patterns on the furs was not possible.In contrast to the molecular-biological investigations published, not only samples of zoo animals were available in this study but also samples from the wild. The results confirm that almost all leopards from Afghanistan and Iran belong to one and the same subspecies. Only in the most eastern part of Afghanistan, the Indian leopard, Panthera pardus fusca, can be found.Mixing the two lines subsequently is justified by the results of this study. Recently acquired animals from the Caucasus, however, should be tested genetically before integrating them into the zoo population.     

Applicability of age-based hunting regulations for African leopards

In species in which juvenile survival depends strongly on male tenure, excessive trophy hunting can artificially elevate male turnover and increase infanticide, potentially to unsustainable levels. Simulation models show that the likelihood of safe harvests can be improved by restricting offtakes to males old enough to have reared their first cohort of offspring to independence; in the case of African leopards, males were ≥7 years old. Here, we explore the applicability of an age-based approach for regulating trophy hunting of leopards. We conducted a structured survey comprising photographs of known-age leopards to assess the ability of wildlife practitioners to sex and age leopards. We also evaluated the utility of four phenotypic traits for use by trophy hunters to age male leopards in the field. Our logistic regression models showed that male leopard age affected the likelihood of survey respondents identifying the correct sex; notably, males <2 years were typically misidentified as females, while mature males (≥4 years) were sexed correctly. Mature male leopards were also more likely to be aged correctly, as were portrait photographs. Aging proficiency was also influenced by the profession of respondents, with hunters recording the lowest scores. A discriminant model including dewlap size, the condition of the ears, and the extent of facial scarring accurately discriminated among male leopard age classes. Model classification rates were considerably higher than the respective scores attained by survey respondents, implying that the aging ability of hunters could theoretically improve with appropriate training. Dewlap size was a particularly reliable indicator of males ≥7 years and a review of online trophy galleries suggested its wider utility as an aging criterion. Our study demonstrated that an age-based hunting approach is practically applicable for leopards. However, implementation would require major reform within the regulatory framework and the hunting industry.     

雪豹保护遗传学和基因组学研究进展

雪豹(Panthera uncia)隶属于食肉目猫科豹属,是生活在青藏高原及其周边地区的旗舰物种。随着分子生物学和高通量测序技术的发展,雪豹保护遗传学和保护基因组学研究得到快速的发展,其中非损伤性遗传取样法显著推动了雪豹保护遗传学研究。本文综述了非损伤性遗传取样法在雪豹物种鉴定、个体识别和性别鉴定等研究中的应用,雪豹的系统发生地位、系统地理格局和种群遗传结构及其亚种争议、演化历史、适应性演化和基因组特征等保护遗传学和基因组学方面的研究现状和进展,并对雪豹保护遗传学和基因组学未来发展趋势进行了展望,以期促进雪豹保护生物学研究和保护对策的科学制定。     

Female reproductive cycles of wild female felids

Many felid species are endangered because of destructive human activities. As a result, zoos are being tasked with sustaining genetically healthy populations in case of catastrophic extinctions. Unfortunately, with the exception of a few species, most felids do not reproduce well in captivity. The ability to track reproductive activity via hormones is key to developing successful ex situ breeding programs. Through the development of noninvasive fecal hormone monitoring techniques, a high degree of variability in estrous cycle characteristics has been found to exist across the taxon, including the type of ovulation. For example, although all felids have induced ovulations, the occurrence of spontaneous ovulations varies across species, and even between individuals within a species. Clouded leopards, fishing cats and margays frequently have spontaneous ovulations, whereas these are rarely observed in the cheetah, tigrina and ocelot. There are marked species differences in the impact of season on reproductive function, with some being exquisitely sensitive to photoperiod (e.g., Pallas' cat), some moderately affected (tiger, clouded leopard, snow leopard), and others that are not influenced at all (e.g., ocelot, tigrina, margay, lion, leopard, fishing cat). One of the greatest challenges remaining is overcoming the problems associated with highly variable ovarian responses to ovulation induction therapies used with assisted reproductive procedures, like artificial insemination (AI). Success is relatively high in the cheetah and ocelot, but few pregnancies have resulted after AI in clouded leopard, fishing cat and tiger. Current knowledge of the reproductive physiology of nondomestic felids, including aspects of the anatomy, behavior and ovarian cycles will be presented, and how the rapidly growing endocrine database is aiding ex situ management efforts.     

Genetic management of captive prosimian populations,a report of experience withLoris tardigradus

Extinction of small, closed populations in captivity as well as in the wild is believed to be nearly inevitable, because inbreeding will adversely effect reproductive success, mortality, sex ratio and also the susceptibility to epidemic diseases and environmental stress. An ever increasing number of primate species exist only in small isolated populations, which contain only a part of the original genetic variability. In captive breeding programs research about genetic management strategies is, therefore, of essential importance. In 1980 we imported 9Loris tardigrdus nordicus (4 females, 5 males) from NE-Sri Lanka. The founders came from one natural breeding population. All sexual mature females are breeding. Up to now the colony contains 36 living individuals. The main goal of our long-term genetic management plan was to minimize inbreeding and to preserve the genetic diversity. Therefore, we try to pass the founder bottleneck rapidly by enlarging the population to a desired minimum population size of 25 pairs and to equalize the founder representation within any generation. The need to control the spread of sublethal genes, introduced by one of the founders, conflicts directly with the aim of equalizing the founder representation. A solution of this problem is discussed. To produce a sufficiently large population we intend to give animals to other institutions and to build up an exchange-system for offspring individuals, which should lead to an international studbook.     

Factors influencing reproductive success and litter size in captive island foxes

A severe decline of island foxes (Urocyon littoralis) on the northern Channel Islands in the 1990s prompted the National Park Service to begin a captive breeding program to increase their numbers. Using detailed records of all the fox pairs (N = 267) that were part of the program on San Miguel, Santa Rosa, and Santa Cruz Islands from its inception in 2000 through 2007, we identified factors influencing the breeding success of pairs in captivity in the interest of formulating strategies that could increase captive productivity. We compiled a database of variables including litter size, reproductive success, distance to nearest occupied pen during the breeding season, subspecies, exposure, female age, male age, age difference, female and male origin (wild vs. captive born), same versus different origin, years paired, previous reproductive success by the pair, previous reproductive success by the female, mate aggression-related injuries, male previous involvement in a pair with mate aggression, and female previous involvement in a pair with mate aggression. We used multiple linear regression to identify factors predictive of litter size, and logistic regression to predict the probability of reproductive success. A larger inter-pen distance, older male age, less exposure, and a smaller intra-pair age difference positively affected litter size. The probabilities of reproductive success increased with fewer years paired and less exposure. Comparatively, pairs with wild born females (vs. captive born females), and previously successful pairs (vs. previously unsuccessful and new pairs) were most likely to be successful. These results indicate that the optimal situation was to pair wild-caught females with older males in sheltered pens that were as far from other pens as possible, to maintain successful pairs and repair unsuccessful ones. Published 2013. This article is a U.S. Government work and is in the public domain in the USA.     

Differential infant carrying in captive and wild common marmosets (Callithrix jacchus)

Callitrichids are communal breeders that lack sexual dimorphism, and only a few studies have examined behavioral gender differences among them. The purpose of this study was to investigate gender differences in infant carrying in 16 captive and seven wild common marmoset groups. Our results showed that female–female twin pairs were carried significantly more often by fathers than were male–male and male–female pairs both in the wild and in captivity. We suggest these differences may be related to different reproductive potentials of male and female Callithrix jacchus and possibly to future breeding competition among females.     

Impact of leopards on a working ranch in Laikipia,Kenya

This study of leopard predation on livestock was carried out on the Lolldaiga Hills ranch (200 km²) in the Laikipia District in Kenya between September 1989 and August 1995. The aims of the study were to find out whether leopards depend on livestock as a food resource on the ranch, and to quantify the economic impact of leopards on the livestock production systems. The density of leopards was estimated in three ways, and the biomass density of potential prey for leopards, both wild and domestic. Diet analysis of leopards revealed that leopards are not relying on livestock as an important food resource. To quantify the effects of leopards, an approach used in the economic assessment of agricultural development schemes was adopted ( Gittinger, 1982 ) and models were derived by following those criteria. It was found that leopards, not being persecuted, live at a relatively high density on the ranch but have less of an adverse impact on livestock than might be expected. Wildlife as a food resource for leopards and a buffer against the killing of livestock could be of benefit on a working ranch to reduce the economic impact of predators on the production system.     

太行山华北豹袭击家畜的时空特点与管理建议: 以山西省和顺县为例

人兽冲突是全球野生动物保护面临的共同挑战。袭击家畜是大型食肉动物引发人兽冲突的主要原因之一, 如果管理不当还会导致针对大型食肉动物的报复性猎杀等严重威胁。冲突的缓解需要深入了解冲突发生的规律与原因, 评估当地社区与居民的态度与看法, 以采取因地制宜的管理措施。太行山是华北豹(Panthera pardus japonensis)的核心分布区, 华北豹袭击家牛的问题是该区域野生华北豹保护中面临的一大挑战。本研究以山西省和顺县为研究区域, 在多方联合发起的“和顺县华北豹袭击家牛肇事补偿项目”实施过程中, 系统收集了2015-2019年期间华北豹袭击家畜事件的记录, 结合同期的红外相机监测数据, 探究了袭击事件的时空分布格局, 并从生境和猎物的角度分析了背后的原因。本文还评估了豹肇事造成的经济损失, 以及受损农户对华北豹的态度和对补偿的满意度。2015-2019年, 项目区域内共记录到华北豹袭击家牛事件195起, 向116户受损农户提供补偿资金共27.05万元。分析结果显示, 华北豹袭击家牛事件在夏季最多, 时间上与家牛上山散放的季节重叠; 袭击风险随狍(Capreolus pygargus)相对多度的上升而下降, 随家牛相对多度和距村庄距离的增加而上升, 但保护区内外和华北豹相对多度对袭击风险的影响不显著。对107名受损农户的访谈结果显示, 华北豹袭击家牛事件虽然对当地畜牧业所造成的整体损失较小, 但对受损农户而言其家庭收入损失严重, 因此导致受损农户对华北豹持负面态度, 且认为现有的补偿金额偏低。基于本研究的结果, 我们提出如下管理建议以期缓解未来该地区华北豹带来的人兽冲突: (1)加强放牧管理、改进管理方式, 特别是限制牛群离开村庄的距离, 以及在家牛散放期间将牛群每晚赶回牛圈以降低家畜被袭击的风险; (2)开展禁牧试点, 严格划分允许放牧与禁止放牧的区域; (3)继续开展华北豹袭击家牛的定损与补偿工作, 降低当地农户的经济损失, 同时开展社区走访、自然教育, 提升当地居民对华北豹的容忍度; (4)加强野生动物种群的监测与保护, 促进人与野生动物的和谐共存。     

Successful ovulation induction and laparoscopic intrauterine artificial insemination in the clouded leopard (Neofelis nebulosa)

Exogenous gonadotropins and a laparoscopic intrauterine artificial insemination (AI) technique were assessed for effectiveness in the clouded leopard (Neofelis nebulosa), a species difficult to breed in captivity due to severe mate incompatibility. Fourteen hormone trials using 10 female clouded leopards were performed to evaluate the ability of 50, 100, or 200 i.u. pregnant mares' serum gonadotropin (PMSG) and 75 or 100 i.u. human chorionic gonadotropin (hCG) to induce folliculogenesis and ovulation, respectively. Laparoscopic evaluation of ovarian activity was conducted at 29–48 hr after hCG administration. Time of ovulation in PMSG/hCG-treated clouded leopards was approximately 38–39 hr after hCG. Excessive follicular development was observed using the high hormone dosages (200 i.u. PMSG/100 i.u. hCG), whereas the lower dosages avoided ovarian hyperstimulation. Previous ovulation sites and mature corpora lutea were detected upon laparoscopic examinations in two of the 10 females housed alone, indicating that this species occasionally spontaneously ovulates. Five females were inseminated by depositing electroejaculated, washed sperm transabdominally into the proximal aspect of each uterine horn. One postovulatory female, previously treated with 100 i.u. PMSG and 75 i.u. hCG and inseminated in utero with 88 × 10⁶ motile sperm at 45 hr post-hCG, produced a pregnancy and two live cubs after an 89 day gestation. These results demonstrate: (1) an exquisite ovarian sensitivity to exogenous gonadotropins in clouded leopards; and (2) that artificial insemination has the potential of resulting in offspring in this species. (This article is a US Government work and, as such, is in the public domain in the United States of America.) © 1996 Wiley-Liss, Inc.     

Life expectancy,maximum longevity and lifetime reproductive success in female Thornicroft's giraffe in Zambia

Gestation and longevity scale with body mass across taxa, yet within size dimorphic taxa, males tend to have reduced lifespans compared with females. Testing life history models, and accounting for sex differences in longevity, requires obtaining accurate longitudinal data from wild populations. We provide the first report describing key life history parameters from a long‐term study of giraffes in Africa. We followed a population of Thornicroft's giraffe (Giraffa camelopardalis thornicrofti) in Zambia for over 40 years. Maximum longevity among females was approximately 28 years, with lifespan accounting for 81% of the variance in lifetime reproductive success. Average adult female life expectancy was no different than average adult male life expectancy. However, the breeding lifespan of males was about half that of females, while maximum lifespan of males was 75% that of females. Our findings support the suggestion that sex differences in maximum lifespan arise from stronger selection for lengthy lives in females than in males. Among females, longer lives are associated with greater reproductive output.     

RFLP-based assay of Sorghum bicolor (L.) Moench genetic diversity

Sixty-two single-copy sorghum DNA clones were used to compare restriction fragment patterns of 53 sorghum accessions from Africa, Asia and the United States. Included were accessions from five morphological races of the cultivated subspecies bicolor, and four races of the wild subspecies verticilliflorum. From two to twelve alleles were detected with each probe. There was greater nuclear diversity in the wild subspecies (255 alleles in ten accessions) than in the domestic accessions (236 alleles in 37 accessions). Overall, 204 of the 340 alleles (60%) that were detected occurred in both subspecies. Phylogenetic analysis using parsimony separated the subspecies into separate clusters, with one group of intermediate accessions. Though exceptions were common, especially for the race bicolor, accessions classified as the same morphological race tended to group together on the basis of RFLP similarities. Selection for traits such as forage quality may have led to accessions genetically more similar to other races being classified as bicolors, which have a loose, small-grained panicle similar to wild races. Population statistics, calculated using four nuclear and four cytoplasmic probes that detect two alleles each, revealed a low but significant amount of heterozygosity, and showed little differentiation in alleles in the wild and cultivated subspecies. Outcrossing with foreign pollen appears to have been more important than migration via seed dispersal as a mechanism for gene flow between the wild and domestic accessions included in this study.