Reproductive development and behavior of captive female chimpanzees

The reproductive histories and behavior of six chimpanzees were analyzed in order to evaluate the female reproductive pattern. This analysis indicated that captive females undergo an earlier menarche and an acceleration of subsequent reproductive phases as compared to wild chimpanzees. Menarche occurred between 7.7–9.3 years of age and first conceptions occurred between 8.2–10.7 years of age. Changes in the females' sex skins across menarche, pregnancy, and the menstrual cycle were also evaluated. The fluctuation in sex skin size was shown to have a marked effect on the occurrence of sexual behavior. In addition, the social context and relative familiarity of the chimpanzees affected the choice of sexual partners.     

Reproductive efficiency of Thoroughbred mares under Indian subtropical conditions: A retrospective survey over 7 years

Service records of 253 mares (1181 mare-years) spanning over 7 consecutive years, from nine organized Thoroughbred stud farms, situated in the subtropical northwestern India were retrospectively analyzed to assess their reproductive performance. The overall per cycle pregnancy rate at Day 16 and overall foaling rates were 50.30% and 68.95%, respectively, and were significantly higher in mares aged 3–7 years than ≥18 years old mares. The late embryonic losses (9.86%) that occurred between Days 16 and 39 post-ovulation contributed more than 50% of the overall detected pregnancy losses (19.11%). The overall percent detected pregnancy losses were lower in mares at ages 3–7 years compared to those at ages ≥18 years (14.78% vs. 46.43%, respectively; P < 0.0001). Chronic barren and habitual aborter mares tended to affect reproductive efficiency of mares. Fifty percent of the mares that experienced ≥2 consecutive abortions or barren years, again stayed aborted or barren in the next seasons, respectively. No effect of numbers of matings per oestrus was observed on overall fertility. Neither the induction of oestrus nor ovulation by exogenous hormonal treatment had any effect on most of the analyzed reproductive parameters. Regarding breeding month or years, the reproductive efficiency did not differ significantly. The incidence of multiple pregnancies was 5.40% and percent late embryonic loses were higher (P = 0.0016) in twin (21.98%) than singleton (8.64%) pregnancies. In conclusion, comparatively lower fertility rates were recorded in Thoroughbred mares bred under Indian subtropical climatic conditions than those reported from temperate regions that might be due to difference in breeding management rather than prevailing environment.     

Weaned horses,especially females,still prefer their dam after five months of separation

Under natural conditions, foals stop nursing from their dam at approximately-9 months old, but their bond persists until 1.5–2.5 years of age. In contrast, in horse breeding, foals are generally artificially weaned and totally separated from their dam at 5–7 months. However, it is not known whether the bond between the dam and her foal is maintained after artificial weaning. The aim of this study was (1) to assess whether foals still recognise and prefer their dam over other familiar mares several months after weaning and (2) to evaluate whether the preference for the dam is more pronounced in fillies or colts. Fifteen fillies and 19 colts were weaned at the age of 7 months old (complete separation from the mother). At the age of one year (i.e., 5 months after the separation), they underwent a test evaluating their preference for their dam or a familiar mare from their natal group. Significantly more foals first approached their dam; they also sniffed and tended to look more often at her. This finding indicates that artificially weaned horses remember and still exhibit a preference for their dam, suggesting that the bond persists even after 5 months of separation. Moreover, fillies exhibited a stronger preference for both mares than colts: they looked at them more frequently, sniffed them for a longer duration and spent more time in proximity to both mares than colts. This suggests that fillies generally have an even stronger attachment to their dam as well as to other mares from their natal group. This study calls into question the practice of artificial weaning at 5–7 months of age.     

Reproduction in the White Eared Opossum (Didelphis albiventris) under Temperate Conditions in Argentina

The reproductive cycle of the white eared opossum Didelphis albiventris Lund was studied in an area of temperate climate conditions in the region of Buenos Aires Province, Argentina. In 1987–1988 and 1988–1989 the reproductive season began in late August and continued until mid-March. Two distinct breeding periods per season were evident, the first in early September and a second in December. Litter size ranged from 4 to 9. The sex ratio in the pouch was 1:1. Only those females that gave birth early in the season could produce a second litter in December. Progressive pouch mortality was not found. Young opossums were weaned at 3.5 months of age and remained sexually inactive until the next reproductive season. Ages of captured opossums suggest that few animals survive for more than 20 months. Therefore, in the study area, most specimens breed only during one reproductive season. In South America, a latitudinal cline in the timing of the breeding season extends from tropical to temperate areas.     

Effects of long-term consumption of sewage solids on reproductive performance and serum progesterone and estradiol-17beta in mature fine-wool ewes

Eighty-four mature fine-wool ewes were randomly allotted to one of three dietary treatments to examine the influence of dietary sewage solids on reproductive performance. Treatments were a basal diet or basal plus either 3.5% cottonseed meal (CSM) or 7% sewage solids. Serum progesterone and estradiol-17beta measured in blood samples collected through an estrous cycle just before the breeding season of the first and second years were not adversely influenced by sewage feeding. Progesterone concentrations quantified in weekly blood samples collected during gestation and lactation of both years did not differ (P>0.10) among treatments. During the first lambing season, lambing data showed that ewes consuming CSM produced slightly more lambs per ewe exposed (P<0.10) than either basal- or sewage-fed ewes. During the second breeding season, midventral laparotomies revealed that ovaries from ewes receiving CSM contained more (P<0.10) corpora lutea than did those from either basal- or sewage-fed animals. However, at the end of the second lambing season, no differences (P>0.10) among treatments in the number of lambs born or weaned were observed. A third year's lambing data also revealed no difference (P>0.10) among treatments for lambing rates. These data indicate that prolonged consumption of a diet containing 7% sewage solids has little, if any, detrimental effect on reproductive performance of mature fine-wool ewes.     

Contraception in the chimpanzee: 12-year experience at the CIRMF Primate Centre, Gabon

The Primate Centre of the International Center for Medical Research in Franceville, Gabon (CIRMF) has had a chimpanzee (Pan troglodytes) colony for more than 20 years. A contraceptive policy was started in 1990, following a rapid increase in the number of individuals in the 1980s. Intrauterine devices were first used in 24 females; 10 failures occurred over a period of about 4 years. Depo-provera was then used in 28 chimpanzees for between 4 months and 4 years; 10 failures occurred, the animals gained weight, and cyclic swelling of the external genitalia was markedly modified. Starting in late 2000, 25 females received progestin implants; only one failure has so far occurred, and the main side effect is a complete abrogation of cyclic sexual swelling.     

Reproductive performance of the Y'ankasa sheep of Nigeria

The reproductive performance of the Y'ankasa sheep of Nigeria was studied to determine any seasonal effects on their breeding pattern. The study also aimed at documenting the reproductive performance levels of these animals as a basis for evaluating the results of any breeding improvement program in Nigeria using indigenous or exotic breeds of sheep. The study shows that this breed of sheep is a year-round breeder although some months favor lambing more than others. The age at first lambing and the lambing interval for these animals are 15.5 months and 9.7 months respectively. For 1975, the conception and lambing rates were 87.8% and 139% respectively. The rates for 1976 were 91% and 148%. Seventy-seven percent of the lambing were singles and the rest twins. The sex ratio, number of females per 100 lambs, was 49%. Lamb mortality rate up to 3 months of age was 12.0%. The lambs in our study grew very slowly, gaining only 15 kg in a 9-month period. By 10 months of age these lambs wieghed only 20 kg.     

Demography,female life history,and reproductive profiles among the chimpanzees of Mahale

Demography provides critical data to increase our understanding of the evolution, ecology, and conservation of primate populations. The chimpanzees of the Mahale Mountains National Park, Tanzania, have been studied for more than 34 yr on the basis of individual identification and standardized attendance records. From this long-term study, we derived the following demographic data: The major cause of death was disease (48%), followed by senescence (24%) and within-species aggression (16%). Fifty percent of Mahale chimpanzees died before weaning. The median ages of female life history variables were: first maximal swelling, 10.0 yr (n = 5); emigration, 11.0 yr (n = 11); and first birth, 13.1 yr (n = 5). The median period of adolescent infertility was 2.8 yr (n = 4) when calculated from the age at immigration to that at first birth. Female fecundity was highest between 20 and 35 yr of age, with an annual birth rate of 0.2. Twenty-six females that were observed from a young age (10-13 yr) to death at various ages (15-40 yr) gave birth to an average of 3.9 and weaned an average of 1.4 offspring. Twenty-five females that were observed from middle age (18-33 yr) to death in older age (31-48) gave birth to an average of 2.7 and weaned an average of 2.0 offspring. The post-reproductive lifespan for female chimpanzees was defined as the number of years that passed from the year when the last offspring was born to the year when the female died, minus 5. Twenty-five percent of old females had a post-reproductive lifespan. The interbirth interval after the birth of a son (x = 72 mo) tended to be longer than that after the birth of a daughter (x = 66 mo). The extent of female transfer, which is a rule in chimpanzees, is influenced by the size and composition of the unit group and size of the overall local community.     

The social structure and reproductive biology of the common mole-rat, Cryptomys h. hottentotus and remarks on the trends in reproduction and sociality in the family Bathyergidae

Cryptomys h. hottentotus occurs in mesic and semi-arid regions of South Africa. It lives in colonies ofup to 14 individuals, in which reproduction is limited to one ofthe largest males and the largest female in the colony.
Reproduction and details of colony size and the number of breeding animals in a colony are described for one complete and two incomplete field-captured colonies. One incomplete colony was kept in captivity for more than 3 years, whereas the complete colony was autopsied.
The social hierarchy of the common mole-rat is almost linear and has a value of 0.7 1 calculated from Landau's linearity index. Subordinate mole-rats weighed less than dominant mole-rats. The reproductive mole-rats are the dominant animals within the colony. The non-reproductive females rank lowest in the hierarchy.
Mating was not confined to a particular season, but occurred more frequently during the summer months. The alpha male initiated pre-copulatory behaviour. The gestation length is 55 to 66 days (n = 2). Two litters each of three pups were born to the colony during captivity. The pups were relatively altricial. They wandered out of the nest 5 days after birth, began to eat solids when 10 days old, and were fully weaned at 4 weeks. When 2 weeks old the pups began to spar with each other and with some of the adult colony members, but levels of aggression were never high. The pups did not disperse but were incorporated into the colony, this suggesting that C. h. hottentotus colonies are largely composed of related individuals.     

Selection for reproductive rate in Rambouillet sheep: Estimated genetic change in reproductive rate

Rambouillet sheep were selected for high or low reproductive rate based on an index of reproductive rate, reproductive INDEX = dam's total lifetime lambs born/(age in years-1). Selected ewes were first born in 1969. A random bred control line was established from the remaining foundation ewes with the first ewes born in 1973. Genetic variances were estimated with REML procedures for reproductive index and number of lambs born or weaned either per ewe exposed for breeding or per ewe lambing. Heritability of reproductive index was 0.26. Heritability for number of lambs born were 0.11 and 0.12 and for number of lambs weaned were 0.04 and 0.04 per ewe exposed for breeding and per ewe lambing, respectively. Breeding values for the reproductive index were estimated using the full animal model (BLUP) with the complete numerator relationship matrix on reproductive index, with fixed effect of year of birth of ewe and breeding values for number of lambs born or weaned either per ewe exposed for breeding or per ewe lambing, estimated with the fixed effects of year of birth of ewe and age of ewe and the random effect of permanent environmental effect among repeated lambing records. Genetic change was estimated as the regression of breeding value for reproductive index and number of lambs born or weaned either per ewe exposed for breeding or per ewe lambing on year of birth of the ewe. High and low lines both responded to selection for reproductive index as compared to the control line (P<0.01). The high line increased at a rate of b = 0.0134 ± 0.0006 reproductive index units and the low line decreased at a rate of b = − 0.0098 ± 0.0005 reproductive index units per year. Response to selection for reproductive index resulted in b = − 0.0074 ± 0.0007 and b = 0.0163 ± 0.0006 lambs per ewe exposed for breeding, and b = − 0.0041 ± 0.0002 and b = 0.0075 ± 0.0002 lambs weaned per ewe exposed for breeding in the low and high lines, respectively. There was a greater response to selection in the high line compared to the low line. These results indicated that the reproductive index did respond to selection.     

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.     

Comparison of Development and Life History in Pan and Cebus

We examined growth and development in capuchins and chimpanzees in relation to weaning, onset of reproduction, and reproductive life span. Striking differences are evident in neurobehavioral status at birth (more mature in capuchins), the relative duration of infancy (longer in chimpanzees), and the proportional weight of the infant at the time of weaning (greater in capuchins). Although capuchins and chimpanzees spend a similar proportion of life in a weaned but reproductively immature state, chimpanzees spend so much more of their lives as nursing infants that reproductive output per individual is much lower than in capuchins. Discussion centers around tolerated transfers of food (food-sharing) as a potential adaptation to limited foraging success by immature foragers. Perhaps food transfers from adult to infant, which is a more prominent feature of behavior in chimpanzees than in capuchins in natural environments, allow a very small weanling chimpanzee to survive.     

Spontaneous amyloidosis in twelve chimpanzees, Pan troglodytes

Spontaneous amyloidosis was diagnosed in 11 male and 1 female chimpanzees and confirmed histologically and immunohistochemically. The chimpanzees were > or = 15 years of age when first diagnosed and averaged 22.4 years of age. The average survival time after diagnosis of systemic amyloidosis was 1.86 years with a standard deviation of 4.06 years (n = 7). The chimpanzees with amyloidosis were asymptomatic except for hepatomegaly, which became more detectable with age. Significant increases in clinical chemistry values, as compared with referenced normals and established normals, of blood urea nitrogen (BUN), asparate aminotransferase (AST), gamma-glutamyltransferase (GGT), globulin, total protein, creatinine phosphokinase (CPK), sedimentation rate, and triglycerides were found in animals 7 years of age or older with amyloidosis. These serum chemistry values, while increased in chimpanzees with amyloidosis, were generally within normal limits. Immunohistochemistry for both amyloid A protein and amyloid P component-labeled extracellular amyloid in all chimpanzees with amyloidosis was determined. Amyloid was deposited primarily in the liver. Amyloidosis in the chimpanzee is a chronic, intractable, progressive, fatal disease, and appears to be similar to secondary amy loidosis in other species.     

Abnormal behaviors,with a special focus on rocking,and reproductive competence in a large sample of captive chimpanzees (Pan troglodytes)

Chimpanzees (Pan troglodytes) are endangered in the wild and may no longer be imported into the United States. Of those animals presently in captivity, candidates for breeding programs must be identified to insure a self-sustaining captive population. Some have suggested that poor reproductive performance might be linked to the performance of abnormal behaviors. In Study 1, three institutions housing breeding colonies of chimpanzees (86 males, 103 females) surveyed their animals for abnormal behaviors, copulatory performance, and, for females, maternal competence. In neither sex was there a positive association between absence of copulation and the presence of any of 18 forms of abnormal behavior. No one abnormal behavior was positively associated with inadequate maternal performance. Contrary to expectations, in both sexes (significantly for females), copulators exhibited more forms of abnormal behaviors than did noncopulators. In contrast, good mothers did show slightly fewer different forms of abnormal behaviors than did inadequate mothers. No specific combination of abnormal behaviors was associated with lack of copulatory performance in either sex or with inadequacy of maternal behavior. Significant sex differences occurred only in 2 of the 18 abnormal behaviors (coprophagy and self-clinging), both with females showing the higher prevalence. In Study 2, the rate of rocking in 5 male and 14 female chimpanzees at the Primate Foundation of Arizona was found to be relatively high among some reproductively competent subjects. Some rocking had forms and contexts indicative of aggressive displays. We conclude that chimpanzees should not be disregarded as potential breeders or as candidates for resocialization and breeding programs solely because they exhibit abnormal behaviors. Anyone involved in assessing well-being needs to be aware of individual differences among animals in the stimuli that may elicit rocking behavior. If they are strangers, they may elicit display rocking which does not indicate a lack of well-being. © 1992 Wiley-Liss, Inc.     

Comparison of Boer, Kiko, and Spanish meat goat does for stayability and cumulative reproductive output in the humid subtropical southeastern United States

ABSTRACT: BACKGROUND: Longevity is the amount of time breeding females stay active in a herd by avoiding death or culling because of illness or reproductive failure. This is a trait of economic relevance in commercial small ruminant breeding herds as it affects lifetime reproductive output. The purpose of this study was to determine if breed of meat goat influences breeding doe survival rates and cumulative reproductive performance under semi-intensive management. RESULTS: Boer (n[THIN SPACE]=[THIN SPACE]132), Kiko (n[THIN SPACE]=[THIN SPACE]92) and Spanish (n[THIN SPACE]=[THIN SPACE]79) does were evaluated for longevity trends and cumulative kid production. The herd was managed on humid subtropical pasture. Does had the chance to complete 2 to 6 production years. Survival curves were analyzed for 2 culling methods. The actual culling practice removed does after two failures to wean a kid. An alternative culling protocol removed doe records after the first failure to wean a kid. Kid production traits analyzed across herd life were the total number of kids weaned and cumulative kid weight weaned to the 2-, 3-, and 5-year stayability endpoints. Most (82 %) doe exits were illness-related under the actual culling method. Reproductive failure represented 51 % of doe exits under the alternative culling protocol. Boer does had greater survival declines (P[THIN SPACE]<[THIN SPACE]0.01) from 2 to 6 years of herd life compared to Kiko and Spanish under both culling protocols. Boer does had lower stayability rates (P[THIN SPACE]<[THIN SPACE]0.01) at each year endpoint for both culling protocols. Under the alternative protocol, over 50 % of Boer does failed to complete 2 years, whereas over 50 % of Kiko and Spanish does successfully completed 4 years. Boer does had lower (P[THIN SPACE]<[THIN SPACE]0.01) total number of kids weaned and cumulative weight weaned through each stayability endpoint compared to Kiko and Spanish. CONCLUSION: Boer does had low stayability and cumulative kid production rates compared to Kiko and Spanish does. Poor health was the primary driver of does exiting the herd. Kiko and Spanish does did not differ for longevity and lifetime performance indicators.     

Reproductive biology and ecology of female polar bears (Ursus maritimus)

Data on age-specific natality rates, litter size, interbirth interval, age of first reproduction, reproductive senescence, age of weaning and cub survival were determined for a free-ranging population of polar bears inhabiting Hudson Bay, Canada, near the southern limit of the species range. Serum progesterone levels were also determined for females at different stages of their reproductive cycle to provide corroborative support for the reproductive parameters described. Animals were live captured using immobilizing drugs and each animal uniquely marked for future identification. First parturition occurred at four or five years of age and the age-specific natality rate increased with age until approximately 20 years, after which it dropped markedly. At least 40% of adult females displayed two-year interbirth intervals and 55% of cubs in their second year were independent of their mother. Mean size of cub litters in spring was 1.9 and 13% of litters had three or more cubs. The natality rate for 5–20-year-old females was estimated as 0.9, higher than that reported for any more northerly polar bear populations where two-year interbirth intervals are rare, fewer than 5% of yearling cubs are weaned and triplet litters occur with less than 1% frequency. Cub mortality was initially high and declined with age. Although cubs in western Hudson Bay were weaned at a younger age and a lighter weight than their counterparts in more northern populations, cub mortality rates were similar. The reason for the marked differences in reproductive parameters in the western Hudson Bay population is not known. We speculate that sea-ice conditions may be sufficiently different to allow weaned bears at a lighter body weight to hunt seals more successfully there than further north.     

BREEDING BIOLOGY OF SOUTHERN ELEPHANT SEALS IN PATAGONIA

Abstract: Elephant seals breed in Patagonia (Península Valdés, Argentina) from late August to early November, reaching peak numbers during the first week in October. Observations of this population over the past ten years yielded similar results. Eighty percent of the pups were born by 2 October. Most (96%) of 663 females marked during three breeding seasons gave birth to a pup. Females stayed on land a mean of 28 d, gave birth 6 d after arrival, nursed their pups for 22 d, and copulated a mean of 2.5 times 20 d after parturition and 2 d before departure. Copulations peaked during the third week in October. Males spent 57–80 d on land fasting and defending harems of up to 134 females (median 11–13 females, depending on year). Most (96%) marked females that gave birth ( n = 636) also weaned their pups successfully. Pup sex ratio was unity. Harems were smaller and breeding occurred about three weeks earlier in Patagonia than in other colonies. Thermal conditions, day length and food availability may explain clines in the timing of breeding events between populations, Other parameters of the breeding season for the expanding Patagonia colony are similar to those for declining southern elephant seal populations elsewhere.     

Reproduction of the greater bushbaby (Galago crassicaudatus panganiensis) under laboratory conditions

A breeding colony of Galago crassicaudatus panganiensis was maintained under laboratory conditions for four years, August 1971 to July 1975. The estrous cycle and the gestation period averaged 39 and 132 days in length, respectively. Twinning occurred in 14.5% of the pregnancies. No postpartum estrus was observed, although 1.2 conceptions per female per year were recorded. Breeding occurred throughout the year. Puberty occurred between ten and 18 months of age, the first conception occurred between 15 and 18 months of age, and reproductive capability decreased rapidly between ten and 12 years of age.     

Reproduction in captive common marmosets (Callithrix jacchus)

Though sexual maturation may begin at around one year of age, first successful reproduction of the common marmoset (Callithrix jacchus) is likely to be later, and it is generally recommended that animals not be mated before 1.5 years of age. The average gestation period is estimated to be 143 to 144 days. A crown-rump length measurement taken by use of ultrasonography during the linear, rapid, prenatal growth phase (between approx. days 60 and 95) can be compared against standard growth curves to estimate delivery date to within 3 to 4 days, on average. Marmosets produce more young per delivery than does any other anthropoid primate, and have more variation in litter size. Many long-established colonies report that triplets are the most common litter size, and there is documented association between higher maternal body weight and higher ovulation numbers. Higher litter sizes generally do not generate higher numbers of viable young. Marmosets are unusual among primates in having a postpartum ovulation that typically results in conception and successful delivery; reported median inter-birth intervals range from 154 to 162 days. However, pregnancy losses are quite common; one study of a large breeding colony indicated 50 percent loss between conception and term delivery. The average life span for breeding females is around six years; the range of reported average lifetime number of litters for a breeding pair is 3.45 to 4.0. Our purpose is to provide an overview of reproduction in the common marmoset, including basic reproductive life history, lactation and weaning, social housing requirements, and common problems encountered in the captive breeding of this species. A brief comparison between marmoset and tamarin reproduction also will be provided.     

Analysis of chimpanzee history based on genome sequence alignments

Population geneticists often study small numbers of carefully chosen loci, but it has become possible to obtain orders of magnitude for more data from overlaps of genome sequences. Here, we generate tens of millions of base pairs of multiple sequence alignments from combinations of three western chimpanzees, three central chimpanzees, an eastern chimpanzee, a bonobo, a human, an orangutan, and a macaque. Analysis provides a more precise understanding of demographic history than was previously available. We show that bonobos and common chimpanzees were separated ~1,290,000 years ago, western and other common chimpanzees ~510,000 years ago, and eastern and central chimpanzees at least 50,000 years ago. We infer that the central chimpanzee population size increased by at least a factor of 4 since its separation from western chimpanzees, while the western chimpanzee effective population size decreased. Surprisingly, in about one percent of the genome, the genetic relationships between humans, chimpanzees, and bonobos appear to be different from the species relationships. We used PCR-based resequencing to confirm 11 regions where chimpanzees and bonobos are not most closely related. Study of such loci should provide information about the period of time 5–7 million years ago when the ancestors of humans separated from those of the chimpanzees.