Parameters of Mating Effort and Success in Male European Ground Squirrels,Spermophilus citellus

Behavioural and physiological changes during the reproductive period were documented in male European ground squirrels including: initiated conflict, scent marking, locomotion and range size, as well as testis size, scrotal pigmentation, testosterone levels and changes in body mass. Spring emergence dates in adult males varied but preceded those of females. Gonadal maturation was complete in adult males at emergence. Most yearlings did not mature and emerged with females. After a quiescent premating phase, male range, range overlap, conflict rates and locomotion increased at female emergence. The synchrony of females, their small ranges and short latencies to ovulation produced a highly male-biased operational sex ratio (OSR) and potential for scramble competition polygyny. In contrast to other ground squirrel species, each female apparently mated with only a single male. Scramble competition did appear to occur, in that males tried to contact as many females as often as possible before copulation. Females placed high temporal prerequisites on males. Costs of scrambling were documented by body mass loss during mating, which covaried with the number of ‘acquired’ females. Testicular regression occurred earlier, and fattening, moult and onset of next hibernation were delayed in successful males. The determinant factor in these costs was found to be mass loss intensity during the main mating period. Surprisingly, this factor was best related to feeding changes and not to locomotion or aggression. Time budget constraints perhaps imposed by the female, with compounding influences of exercise and stress effects of conflict, then negatively affected ensuing seasonal activities. The individual changes may adversely affect individual survivability and/or future reproductive success.     

Seasonality and interindividual variation in mandrill feeding ecology revealed by stable isotope analyses of hair and blood

Mandrills are large-bodied terrestrial forest primates living in particularly large social groups of several hundred individuals. Following these groups in the wild to assess differences in diet over time as well as among individuals is demanding. We here use isotope analyses in blood and hair obtained during repeated captures of 43 identified free-ranging mandrills (Mandrillus sphinx) from Southern Gabon, to test how dietary variation relates to the season as well as an individual's age and sex. We measured the stable carbon (δ¹³C‰) and nitrogen (δ¹⁵N‰) isotope ratios in 46 blood and 214 hair section samples as well as from a small selection of mandrill foods (n = 24). We found some seasonal isotopic effects, with lower δ¹³C values but higher δ¹⁵N values observed during the highly competitive long dry season compared to the fruit-rich long rainy season. Variation in δ¹³C was further predicted by individual age, with higher δ¹³C values generally found in younger individuals suggesting that they may consume more high canopy fruit than older individuals, or that older individuals consume more low canopy foliage. The best predictor for δ¹⁵N values was the interaction between age and sex, with mature and reproductively active males revealing the highest δ¹⁵N values, despite the observation that males consume substantially less animal food items than females. We interpret high δ¹⁵N values in these mature male mandrill blood and hair sections to be the result of nutritional stress associated with intense male–male competition, particularly during mating season. This is the first study showing isotopic evidence for nutritional stress in a free-ranging primate species and may spark further investigations into male mandrill diet and energy balance.     

Individual Facial Coloration in Male <Emphasis Type="Italic">Eulemur fulvus rufus</Emphasis>: A Condition-dependent Ornament?

Researchers studying individual variation in conspicuous skin coloration in primates have suggested that color indicates male quality. Although primate fur color can also be flamboyant, the potential condition dependence and thus signaling function of fur remains poorly studied. We studied sources of variation in sexually dichromatic facial hair coloration in red-fronted lemurs (Eulemur fulvus rufus). We collected data on 13 adult males in Kirindy Forest, Madagascar, during two study periods in 2006 and 2007, to determine whether variation in facial hair coloration correlates with male age, rank, androgen status, and reproductive success. We quantified facial hair coloration via standardized digital photographs of each male, assessed androgen status using fecal hormone measurements, and obtained data on reproductive success through genetic paternity analyses. Male facial hair coloration showed high individual variation, and baseline coloration was related to individual androgen status but not to any other parameter tested. Color did not reflect rapid androgen changes during the mating season. However, pronounced long-term changes in androgen levels between years were accompanied by changes in facial hair coloration. Our data suggest that facial hair coloration in red-fronted lemur males is under proximate control of androgens and may provide some information about male quality, but it does not correlate with dominance rank or male reproductive success.     

Prominent ornaments and rapid color change: use of horns as a social and reproductive signal in unicornfish (Acanthuridae: <Emphasis Type="Italic">Naso</Emphasis>)

The behavior of two unicornfishes, Naso unicornis with a horn-shaped protuberance on the forehead and Naso vlamingii with a round protuberance, was observed in social and reproductive contexts in an aquarium. Males of both species performed displays that were associated with quick changes in the colors of the protuberance and other body parts, highlighting the protuberance by color contrast. The displays with color changes of the protuberance took place when a male courted a female in the evening or in the night. The same displays were occasionally performed by males toward females throughout the daylight hours. In N. vlamingii, dominant males displayed the protuberance toward subordinate smaller males. Although the sizes and shapes of the protuberance were sexually monomorphic, females of both species rarely made displays of the protuberance. A hypothesis about the function of the protuberance in unicornfishes is proposed: that males use the protuberance as a conspicuous signal in courtship and contests among males by emphasizing it with quick changes of its color contrasts. We argue that the combination of morphologically distinct protuberances and quick changes of their color is a sexually selected trait among unicornfishes, because these characters play important roles in intersexual and intrasexual interactions.     

Context and Development of Sexual Behavior of Wild Bonobos (Pan paniscus) at Wamba, Zaire

I studied sexual behavior of immature bonobos (Pan paniscus) in a wild group living at Wamba, Zaire, with special reference to its development. Even immature individuals under 1 year old performed sexual behavior. Sexual behavior occurred in almost all age–sex combinations, except between immature and mature females. Based on analyses of behavioral pattern and context, I classified sexual behavior involving immature individuals into three categories. (1) Genital contact between immature individuals was observed during play, and was performed by males more frequently than by females. This sexual behavior shared many traits with that of other great apes. (2) Copulation-like genital contact was observed between immature males and mature females. Its frequency increased with the immature male's age; it developed into copulation in adulthood. (3) Genital contact used to regulate interindividual relationships. This behavior, which is unique to bonobos, was absent among infants. It developed between late juvenile and early adolescent periods in association with changes in social circumstances.     

LOSS OF NUPTIAL COLOR IN THREESPINE STICKLEBACKS (GASTEROSTEUS ACULEATUS)

Loss of conspicuous nuptial color in Gasterosteus aculeatus (threespine stickleback) has been reported from several localities in western North America and is ascribed to increased rates of predation or to convergent threat displays. I have examined 66 populations of G. aculeatus from the Queen Charlotte Archipelago and found extensive variation in the expression of red nuptial color. Males in 31 of the populations lack red throats, while males in five populations have major expression of red nuptial color. I test two hypotheses for nuptial-color loss: that the loss results from increased predation rates and that it involves differences in water spectra (relative transmission at 400 nm). Results, which are consistent with the second hypothesis, show that the greatest expression of red pigment occurs in habitats with the highest water clarity, while loss of red nuptial color is generally found in heavily stained waters. There is no correlation between nuptial-color loss and presence or absence of vertebrate predators. Two new hypotheses for these associations are proposed: signal-masking in spectrally restricted habitats and carotenoid deficiencies in the diet. Previous studies of red nuptial color and its loss in Gasterosteus merit additional attention, given the associations with underwater spectra.     

九寨沟自然保护区川金丝猴的分布及种群数量

2014年2月至5月,对四川省九寨沟自然保护区野生川金丝猴的种群分布、数量及其社会结构进行调查,初步探讨该种群与其他种群在外部形态和社会结构方面的差异性。采用"V"型路线法调查,发现九寨沟保护区共有川金丝猴4群。然后结合长时间的持续跟踪观察,证实家庭单元19个,全雄群3个,能区分不同个体296只。OMU大小平均为12.7只,其中OMU内的成年个体占38%,青少年个体占53%,幼仔占9%。成年与青少年比例为1∶1.39,成年雌雄的比例为3.84∶1。与秦岭种群的外部形态相比,该种群的成年雄性更为壮大,毛发弯曲程度明显;成年雌性毛发略显灰黑色,金黄色暗淡,大部分个体嘴角长有很小的肉瘤;青少年个体毛色泛白。亦发现九寨沟种群倾向于选择海拔偏高(2400—3550 m)的针叶林带活动。     

Scrotal circumference does not accurately predict degree of germinal epithelial loss or semen quality in yearling Hereford and Simmental bulls

The accuracy of scrotal circumference as a predictor of specific pathologic changes within testicular parenchyma was assessed by scoring 121 Hereford and Simmental bulls for scrotal circumference, degree of germinal epithelial loss and semen quality. Scrotal circumference was not linearly related to the degree of germinal epithelial loss or to the percentage of tubules (Grade 4+ or higher, no germinal epithelium) for either breed (P > 0.1). Thirty-two centimeters was the minimal acceptable scrotal circumference to ensure both a low prevalence of tubules with irreversible loss of germinal epithelium and acceptable semen quality. Scrotal circumference had a positive predictive value (the proportion of test-positive animals truly diseased) of 0.07 (95% confidence interval [CI] of 0 to 0.29) and a specificity (the proportion of test postive individuals that are diseased) of 0.89 (95% CI of 0.83 to 0.95) for 25% Grade 4 + tubules. For acceptable semen morphology (>/= 75% morphologically normal spermatozoa), scrotal circumference had a positive predictive value of 0.08 (95% CI of 0 to 0.34), a sensitivity (the proportion of diseased individuals testing positively) of 0.06 (95% CI of 0 to 0.24), and a specificity of 0.88 (95% CI of 0.80 to 0.95). For acceptable sperm motility (>/= 30% progressively motile spermatozoa) scrotal circumference had a positive predictive value of 0.0 (95% CI of 0 to 0.22), a sensitivity of 0.0 (95% CI of 0 to 0.15), and a specificity of 0.87 (95% CI of 0.80 to 0.94). Although scrotal circumference is widely accepted as a useful test, its accuracy should be recognized as low when 32 cm is used as the minimal acceptable scrotal circumference in predicting unacceptable testicular histology or unacceptable semen quality in a healthy population of yearling bulls.     

The Evolutionary Decoupling of Behavioral and Color Cues in a Multicomponent Signal in Two Sceloporus Lizards

The study of multicomponent signals in the context of social systems has generated interesting results demonstrating that complex signals are used in many communication systems. The multicomponent signal in the majority of Sceloporus lizards consists of a color signal (blue abdominal coloration) and the behavioral display of the color signal (fullshow behavior). In a small number of species, males have lost the color signal. We staged outdoor trials between conspecific males in two closely related Sceloporus lizards that differ in the presence of blue abdominal coloration. In the species with the evolutionary loss of blue abdominal coloration (Sceloporus virgatus), fullshow behavior is present but reduced compared with that of the species with male abdominal coloration (Sceloporus undulatus consobrinus). In comparison, the mean rates of other behaviors that do not display abdominal skin (push‐up, head‐bob) did not differ between these species. We also found that S. virgatus males were more likely to show a neutral response following the first fullshow during the 60‐min trial. While, S. u. consobrinus males were more likely to respond to the first fullshow with an aggressive response. Thus, in this case, the color signal and the behavioral signal are evolutionarily decoupled because in S. virgatus the loss of the color signal is not coincident with the loss of the behavioral signal.     

Developmental changes in the facial morphology of the Borneo orangutan (Pongo pygmaeus): possible signals in visual communication

Orangutans display remarkable developmental changes and sexual differences in facial morphology, such as the flanges or cheek-pads that develop only on the face of dominant adult males. These changes suggest that facial morphology is an important factor in visual communication. However, developmental changes in facial morphology have not been examined in detail. We studied developmental changes in the facial morphology of the Borneo orangutan (Pongo pygmaeus) by observing 79 individuals of various ages living in the Sepilok Orangutan Rehabilitation Centre (SORC) in Malaysia and in Japanese zoos. We also analyzed photographs of one captive male that were taken over a period of more than 16 years. There were clear morphological changes that occurred with growth, and we identified previously unreported sexual and developmental differences in facial morphology. Light-colored skin around the eyes and mouth is most prominent in animals younger than 3 years, and rapidly decreases in area through the age of approximately 7 years. At the same time, the scattered, erect hairs on the head (infant hair) become thick, dense hairs lying on the head (adult hair) in both sexes. The results suggest that these features are infant signals, and that adult signals may include darkened face color, adult hair, whiskers, and a beard, which begin to develop after the age of approximately 7 years in both sexes. In females, the eyelids remain white even after 10 years, and turn black at around the age of 20; in males, the eyelids turn black before the age of 10. The whiskers and beards of adults are thicker in males than in females, and are fully developed before the age of 10 in males, while they begin to develop in females only after approximately 20 years. White eyelids and undeveloped whiskers and beards may be visual signals that are indicative of young adult females. Our results also show that the facial morphology of the unflanged male is similar to that of the adult female, although it has also been pointed out that unflanged males resemble younger individuals.     

Thermal tolerance of Frankliniella occidentalis: Effects of temperature, exposure time, and gender

Western flower thrips (WFT), Frankliniella occidentalis, is an invasive worldwide pest that causes great economic loss. Temperature plays an important role in shaping insect distributions. In this study, we report the survival rates of 3-day-old WFT exposed to rapid changes in extreme high and low temperatures. Temperature, exposure time, and gender all significantly affected the survival of adult WFT, with the lowest survival associated with more extreme temperatures and/or longer exposures. The temperature required to kill 50% of exposed WFT individuals (LT₅₀) decreased with extended exposure time, but females were more tolerant to extreme temperature than males. Investigation of rapid cold or heat hardening suggested that a short prior exposure to a sub-lethal low or high temperature increased WFT survival during a subsequent exposure to a lethal temperature. Tolerance of extreme temperatures and an ability to undergo rapid hardening are of great ecological relevance in determining the geographic distribution of WFT, allowing it to survive better in temporary bouts of extreme temperature stress. Our findings provide useful information on the environmental limits on the distribution of WFT, which have implications for control of this pest.     

Sexual Selection of Zorion guttigerum Westwood (Coleoptera: Cerambycidae: Cerambycinae) in Relation to Body Size and Color

Zorion guttigerum is a flower-visiting longhorned beetle endemic to New Zealand. Sexual selection of this species in relation to the body size and color form of different sexes was investigated in the field. The population sex ratio, based on censuses of feeding and mating sites (flowers), is male-biased. Females are significantly larger than males. Both sexes have antennae of similar length but the antennal length relative to the elytral length is greater in males than in females, and the antennal length of males increases more with an increase in body size than that of females. Both sexes have dark blue (DB) and yellowish-brown (YB) individuals. Both pair-bonded and solitary males are similar in elytral and antennal length. In pair-bonded males, DB individuals are significantly more numerous than YB ones, but in solitary males, the number of both color forms is similar. Males tend to have territory protection behavior, fighting with and chasing away rival males from feeding and mating sites. Larger males usually win the fight but the size-dependent fighting advantage does not translate into mating success. Male color plays an important role in mating success, with DB males having a significantly better chance to mate than YB males. Furthermore, male body size and color also have interactions in mating success: males of DB color morph obtain a greater mating advantage according to body size. Pair-bonded females are significantly larger and have longer antennae than solitary females, suggesting that males prefer larger females for mating. In addition, females of DB color morph with longer antennae are also preferred by males for mating. The significance of these findings is discussed.     

Sex and genetic differences in hair color changes during early childhood.

Hair color was assessed routinely from three months to six years for children participating in a longitudinal study of twins: 169 female twin pairs, 161 male pairs, and 60 opposite-sex pairs. Age trends, established by sampling only one number of every pair, showed marked changes in hair color for both sexes, but there was a consistent excess of light-haired males and dark-haired females. Within-pair concordance rates were calculated for same-sex pairs whose zygosity had been determined independently through bloodtyping. A high rate of concordance was found for MZ twins at every age in spite of the general change in hair color, indicating a strong genetic influence in the timing of color changes. The results are discussed in terms of accelerated maturation of females, and the need for genetic models of the inheritance of hair color which are age- and sex-specific.     

Social Dispersion and Genetic Variation in Two Species of Emballonurid Bats

Saccopteryx leptura and S. bilineata are closely related, sympacric species of neotropical bats that exhibit marked differences in social organization and dispersion patterns. S. leptura is monogamous and roosts in small groups of 1 to 5 (Y = 2.6) individuals that are finely dispersed. S. bilineata is harem-polygamous and roosts in larger groups of 1 to 42 (Y = 8.1) individuals that are more coarsely dispersed. Allozyme genetic studies of individuals from Trinidad, W I. demonstrate that both species carry similar, high levels of polymorphism and heterozygosity. Although significant genetic heterogeneity was observed among geographical populations of S. leptura, there is no evidence for either species that social structuring leads to inbreeding and the loss of heterozygosity, or that it promotes the development of genetic heterogeneity among social units that could accelerate the evolutionary diversification of these taxa. These results do not support a recently proposed hypothesis that social structuring in mammals has these effects. Observed intercolony genetic heterogeneity among adult males resident in multimale groups of S. bilineata is greater than intercolony heterogeneity of adult females. This is an apparent result of female dispersal and the recruitment of males into their parental groups. These results suggest that the adult males within large colonies may be kin-related.     

Chest Color and Social Status in Male Geladas (<Emphasis Type="Italic">Theropithecus gelada</Emphasis>)

Conspicuous colored patches on animals often serve as sexually selected signals that advertise male quality. Such colored traits facilitate assessment of risks associated with a specific contest or benefits associated with a specific mate choice. Here, we investigate whether a colored patch of skin on the chests of male geladas (Theropithecus gelada) is a sexually selected signal. Specifically, we examine the relationship between color (redness), social status (a proxy for reproductive success), and age. We use observational data from known individuals from a population of wild geladas living in Ethiopia. We digitally quantified chest color using a previously-validated method for measuring color under field conditions. Results from this study are consistent with the hypothesis that redness is a quality signal in males. Baseline color correlates with status even when controlling for age. Indeed, males with redder chests were members of “better” groups: 1) leader males—the only males with reproductive access to females—had the reddest chests, and 2) within leader males, males with large units (>6 females) had redder chests than males with small units. At present, we are unable to address whether male chest color is directed at potential rivals or mates. Nevertheless, our data support the hypothesis that quality signals should prevail in large, fluid groups, where it is unlikely that individuals recognize all other group members. If individual recognition is limited in gelada society, this would favor the evolution of alternative means of assessment for making reproductive decisions.     

The sexual function of wallowing in male wild boar (Sus scrofa)

The wild boar (Sus scrofa) is a medium-sized ungulate that produces a large variety of marks. Traditionally, wallowing is an activity that has fundamentally been related to several processes such as thermoregulation, reduction in the number of ectoparasites or disinfection of wounds in the skin of males resulting from fights to reach females. In this study, I analysed another possible function of wallowing in free-ranging wild boar populations when temperatures are low and parasitic numbers are not significant. The results indicate that during the sampling time, from October to February, the number of wild boars with a mud layer on their skin was 47, out of a total of 558 wild boar analysed. Of these 47 wild boars, 44 were males and within this group, 40 individuals were adult males. The wallowed males were bigger and older than non-wallowed males hunted at the same time. In addition, I found that males wallowed mainly during autumn (90.9% of all cases). In the total number of wild boars analysed, I did not notice individuals with wounds resulting from fights other than from the hunting day. On the other hand, the analysis of the females reproductive systems indicated that the rut period took place during the last days of October and the first days of November. From these results, the coincidence of the rut period with the maximum of wallowing in adult males could be indicative of a sexual function of this activity.     

Social control of sex change in the Red Sea razorfish Xyrichtys pentadactylus (Teleostei,Labridae)

Synopsis The Red Sea razorfish, Xyrichtys pentadactylus, a territorial haremic labrid with dominance hierarchies within the harems. Theory predicts that primary males (fish developing initially as males) should be rare or nonexistent in haremic territorial species because the larger secondary males (males which have undergone sex and/or color change) limit access to females. Histological examination of gonads of 95 specimens showed that all males are derived from females by sex change (i.e. they are secondary males). During five months of field studies 100% of more than 200 observed matings were pair spawnings — the usual mating practice for monandric (having one type of male) species. Sex change in females was induced by male removal in nature. Isolation of four groups of females in aquaria showed that the largest female in the social group changes sex in the absence of a male, demonstrating that sex change is socially-controlled in this species.     

Do Female Mandrills Prefer Brightly Colored Males?

Although secondary sexual adornments are widespread in male primates, few studies have examined female choice for these characters. Mandrills (Mandrillus sphinx) present an extreme example of sexual dimorphism, with males exhibiting an array of striking adornments. The most dominant adult male in a group exhibits the brightest and most extensive red coloration, while the other males are less brightly colored. I examined whether female mandrills prefer brightly colored males using data on periovulatory sexual behavior during the 1996 mating season for all males 8 years old (n = 5) and all parous females (n = 9) in a semifree-ranging colony at CIRMF, Gabon. Brightness of male coloration is significantly positively correlated with time spent within 2 m of females, female responsibility for proximity, number of sexual presentations received, % approaches accepted by females, and % inspections with which females cooperated. Females also groomed only the brightest male. Behaviors indicating female preference are not correlated significantly with male dominance rank, and partial correlations confirm that the influence of male color on female behavior is stronger than that of male rank. With the influence of male dominance rank controlled, correlation coefficients between female behaviors and male mating success are high and positive. In further support of the hypothesis that females show mate choice for brightly colored males, independent of dominance rank, I report an unusual case wherein the alpha male fell in rank without loss of coloration. He experienced no significant change in female responsibility for proximity, sexual presentations received, or female reaction to approaches or inspections, though he was no longer observed to mate. Accordingly, female mandrills attend to differences in male secondary sexual characters and favor brightly colored males. As brightly colored males are also dominant this reinforces the influence of male-male competition on male reproductive success and may explain the very high reproductive skew in mandrill males and their extraordinary appearance.     

The Association of Intergroup Encounters,Dominance Status,and Fecal Androgen and Glucocorticoid Profiles in Wild Male White‐Faced Capuchins (Cebus capucinus)

Androgens play a role in male reproductive competition, frequently via aggression, while glucocorticoids are associated with the stress response. However, the relationships of these hormones with different sources of competition (intra‐ vs. intergroup) and dominance status are highly variable. Here, we consider the fecal androgen (fA) and glucocorticoid (fGC) profiles of alpha and subordinate male Cebus capucinus in the context of intergroup competition during a rare period of low intragroup competition (i.e. all females were either pregnant or lactating). Intergroup encounters (IGEs) are a long‐term reproductive strategy in male white‐faced capuchins, enabling them to assess the composition of neighboring groups. IGEs pose a threat to resident males as these can result in injury or death, loss of dominance rank, group eviction, and group takeovers that are frequently associated with infanticide. From February to July 2007, fecal samples were collected from eight males in three groups of white‐faced capuchins in the Santa Rosa Sector of the Área de Conservación Guanacaste, Costa Rica. IGE rate was positively associated with both fA and fGC levels, indicating that IGEs are perceived as reproductive challenges by resident males, and may be associated with elevated metabolic costs. Alpha males sire the majority of group offspring and, accordingly, the threat of IGEs to both future (via rank loss or eviction) and current (via infanticide) reproductive success is greater than for subordinate males. Consistent with this observation, alpha males had higher fA and fGC levels than subordinate males. Given that all females were either pregnant or lactating and pronounced overt intragroup competition was absent, we interpret the difference in hormone profiles of alpha and subordinate males as being primarily associated with variation in the perceived threats of IGEs according to dominance status. Future studies should focus on the interaction of intra‐ and intergroup competition by examining hormone levels in the presence of periovulatory females. Am. J. Primatol. 75:107‐115, 2013. © 2012 Wiley Periodicals, Inc.