Foraging behaviour of the slender loris (Loris lydekkerianus lydekkerianus): implications for theories of primate origins

Members of the Order Primates are characterised by a wide overlap of visual fields or optic convergence. It has been proposed that exploitation of either insects or angiosperm products in the terminal branches of trees, and the corresponding complex, three-dimensional environment associated with these foraging strategies, account for visual convergence. Although slender lorises (Loris sp.) are the most visually convergent of all the primates, very little is known about their feeding ecology. This study, carried out over 10 (1/2) months in South India, examines the feeding behaviour of L. lydekkerianus lydekkerianus in relation to hypotheses regarding visual predation of insects. Of 1238 feeding observations, 96% were of animal prey. Lorises showed an equal and overwhelming preference for terminal and middle branch feeding, using the undergrowth and trunk rarely. The type of prey caught on terminal branches (Lepidoptera, Odonata, Homoptera) differed significantly from those caught on middle branches (Hymenoptera, Coleoptera). A two-handed catch accompanied by bipedal postures was used almost exclusively on terminal branches where mobile prey was caught, whereas the more common capture technique of one-handed grab was used more often on sturdy middle branches to obtain slow moving prey. Although prey was detected with senses other than vision, vision was the key sense used upon the final strike. This study strongly supports the notion that hunting for animal prey was a key ecological determinant in selecting for visual convergence early on in primate evolution. The extreme specialisations of slender lorises, however, suggest that early primates were not dedicated faunivores and lend further support to the emerging view that both insects and fruits were probably important components of the diet of basal primates, and that exploitation of fruits may account for other key primate traits.     

Morphometry and allometry of the primate humerus

The primate distal humerus has been used both in phylogenetic reconstruction and in assessing locomotor and postural adaptations. This study uses an allometric approach to predict locomotor patterns of extant primates regardless of phylogenetic position. By showing the relationship between form and function in living primate taxa it will be possible to use this data set to predict locomotor behavior of extinct primates. Several linear measurements were taken from the distal humerus of 71 extant primate species (anthropoids and prosimians). Allometric regressions of each measurement were performed with mandibular M₂ area as a surrogate for body size. These measurements were used to determine if significant differences in distal humerus morphology exist among locomotor groups. The results were then used to test several hypotheses about the relationship between humeral form and function. For example, the hypothesis that suspensory primates have a large medial epicondyle is confirmed; the hypothesis that terrestrial quadrupeds have a deep olecranon fossa could not be confirmed with quantitative data. In addition to this hypothesis testing, the residuals from the allometric regressions of the humeral measurements were used in a discriminant functions analysis to estimate locomotor behavior from distal humerus morphology. The discriminant functions analysis correctly reclassified 64/71 (90%) species.     

Female reproductive strategies in orangutans,evidence for female choice and counterstrategies to infanticide in a species with frequent sexual coercion

Intersexual conflicts over mating can engender antagonistic coevolution of strategies, such as coercion by males and selective resistance by females. Orangutans are exceptional among mammals for their high levels of forced copulation. This has typically been viewed as an alternative mating tactic used by the competitively disadvantaged unflanged male morph, with little understanding of how female strategies may have shaped and responded to this behaviour. Here, we show that male morph is not by itself a good predictor of mating dynamics in wild Bornean orangutans but that female conception risk mediated the occurrence and quality of male–female interactions. Near ovulation, females mated cooperatively only with prime flanged males who they encountered at higher rates. When conception risk was low, willingness to associate and mate with non-prime males increased. Our results support the hypothesis that, together with concealed ovulation, facultative association is a mechanism of female choice in a species in which females can rarely avoid coercive mating attempts. Female resistance, which reduced copulation time, may provide an additional mechanism for mate selection. However, coercive factors were also important as prime males were frequently aggressive to females and females used mating strategies consistent with infanticide avoidance.     

Repeatability: Its role in evolutionary studies of mating behavior

Summary Repeatability, a concept derived from quantitative genetics theory, is a statistic that describes the degree to which variation within individuals contributes to total variation in a population. Its usual application has been to set an upper limit on heritability but it may also be useful for studies of stereotypy of behavior. The repeatability of the production of male mating signals gives information both about whether males differ sufficiently for selection to act and whether the differences could be appreciably heritable. Measures of the repeatability of female mating preferences will provide data that can describe the preference functions used in mathematical models of the evolution of sexually selected traits, as well as putting an upper bound on the heritability of preferences. A survey of the few measures in the literature shows that the repeatability of male signal production varies substantially (range 0.21–0.85) and does not necessarily reflect heritability. The repeatabilities of female preferences have not been published previously: for the response to conspecific pheromones by female flour beetles (Tribolium castaneum), my best estimate is zero. Measuring the repeatability of other traits such as parental care and foraging behavior may also lead to insights about selection on and the evolution of these traits.     

Allometry of primate hair density and the evolution of human hairlessness

Allometric analyses of hair densities in 23 anthropoid primate taxa reveal that increasingly massive primates have systematically fewer hairs per equal unit of body surface. Considering the absence of effective sweating in monkeys and apes, the negative allometry of relative hair density may represent an architectural adaptation to thermal constraints imposed by the decreasing ratios of surface area to volume in progressively massive primates. Judging by estimates of body volume, denudation of the earliest hominids should have progressed to a considerable extent prior to their shift from a forest to a grassland habitat during the Pliocene. We propose that, lacking a reflective coat of hair, the exploitation of eccrine sweating emerged as the primary mechanism for adaptation to the increased heat loads of man's new environment and permitted further reduction of the remnant coat to its present vestigial condition.     

Co-evolution of male and female reproductive characters across the Scathophagidae (Diptera)

Sperm morphometry is extremely variable across species, but a general adaptive explanation for this diversity is lacking. As sperm must function within the female, variation in sperm form may be associated with variation in female reproductive tract morphology. We investigated this and other potential evolutionary associations between male and female reproductive characters across the Scathophagidae. Sperm length was positively associated with the length of the spermathecal (sperm store) ducts, indicating correlated evolution between the two. No association was found between sperm length and spermathecal size. However, the size of the spermathecae was positively associated with testis size indicating co-evolution between male investment in sperm production and female sperm storage capacity. Furthermore, species with a higher degree of polyandry (larger testes) had longer spermathecal ducts. However, no associations between sperm length or length variation and testis size were found which suggests greater sperm competition sensu stricto does not select for longer sperm.     

The timing of mating influences reproductive success in Drosophila melanogaster: implications for sexual conflict

Despite its potential importance, the role of the timing of mating(s) as a source of variation in female lifetime reproductive success has been largely overlooked. Here, using a laboratory‐adapted population of the model species Drosophila melanogaster, we explore how temporal variation in the patterns of single and multiple matings influences female fecundity. We find that the boost to fecundity known to occur after a virgin female’s initial mating also extends to subsequent matings as nonvirgins, but only for a short duration. This fecundity boost at least partially offsets the direct costs of multiple matings to females in this population of D. melanogaster. The implications of these results for our understanding of the evolution and maintenance of polyandry in this species are discussed in the context of sexual conflict.     

Cephalic modifications in dimorphic dwarf spiders of the genus oedothorax (Erigoninae,Linyphiidae, Araneae) and their evolutionary implications

Reproductive competition among males selects for a broad variety of strategies and traits from mate guarding to nuptial food gifts. Males of many dwarf spider species possess conspicuous secondary cephalic modifications, and the few studies available suggest that these cephalic structures are connected to extensive glandular tissue. Because females were observed to contact the male head structures during mating, these traits may have evolved in the context of sexual selection. We investigated the structure, glandular equipment, and sensory equipment of the cephalic regions of several species of the dwarf spider genus Oedothorax with varying degrees of sexual dimorphism using light and electron microscopy. In one Oedothorax species, there are two male morphs that exhibit a cephalic modification (O. gibbosus gibbosus) or not (O. gibbosus tuberosus). Our study demonstrates that all males investigated produce cephalic secretions, irrespective of the morphology of their cephalic region, however, they may differ in amount of secretion and in cellular organization. In males of O. apicatus, O. gibbosus gibbosus and O. retusus the gland cells are very abundant in the area of a cephalic hump, whereas in the less conspicuous O. agrestis, and O. gibbosus tuberosus the gland cells are restricted to a small area behind the ocular region or include the ocular region as in O. fuscus. The glandular tissue consists of two gland types in O. agrestis, O. fuscus, O. gibbosus tuberosus and O. retusus and of only one type in O. apicatus and O. gibbosus gibbosus. The setae present on the head structure of all species seem to function as mechano‐ and/or chemoreceptors. The implications of our findings for the evolution of secretory head structures are discussed along with their potential for driving speciation. J. Morphol. 2011. © 2011Wiley‐Liss, Inc.     

A role for female ornamentation in the facultatively polygynous mating system of collared flycatchers

In a polygynous mating system, females settling with alreadymated males often experience low mating success due to the reducedparental contribution of the male. However, there are numerousfactors that may still make it advantageous for some femalesto choose this mating status. Facultative polygyny is believedto be dominated by male advertisement and female choice. Althoughquality differences and competition among females are increasinglyrecognized as important determinants of polygynous settlementpatterns, the importance of signals of female quality in thismating system is largely unknown. Here we examined the relationshipof the white wing patch size (WPS) of female collared flycatchers,a phenotypically plastic and age-dependent ornament, with socialmating status, while controlling for settlement date and age.At the population level, monogamous, primary, and secondaryfemales did not differ in WPS. However, the primary female ofindividual males was more ornamented than the secondary female,and this difference declined with increasing distance betweenprimary and secondary nests. Secondary female ornamentationincreased, whereas that of the primary female did not changewith nest distance. These results suggest a subtle role forfemale ornamentation at polygynous mating. Future studies shouldtherefore take into account mating status when assessing thecosts and benefits of female signals. Moreover, patterns inquality-indicating female traits may contribute to the explanationof differences in reproductive success among females of differentmating status.     

Population differences in female resource abundance, adult sex ratio, and male mating success in Dendrobates pumilio

In this study I examined the relationship among abundance ofreproductive resources, population density, and adult sex ratioin the strawberry dart-poison frog, Dendrobates pumilio, andhow these variables in turn influence the mating system, malereproductive success, and sexual selection. I studied the matingbehavior in two populations of D. pumilio living in a primaryand secondary rainforest on the Caribbean slope of Costa Rica.The abundance of tadpole-rearing sites (reproductive resourcesfor females) was approximately 10-fold higher in the secondary forest. Accordingly, the population density was higher and theadult sex ratio was strongly female biased in the secondaryforest, whereas the adult sex ratio was even in the primaryforest. The female-biased sex ratio was associated with a higherlevel of polygyny and higher male mating and reproductive successin the secondary forest. In contrast, the level of polyandrydid not differ between habitats. As expected, the opportunityfor sexual selection on male mating success was lower in thesecondary forest, the habitat with high female density. Inconclusion, my results suggest that ecological variables suchas resource availability have a great impact on the matingsystem and sexual selection through their effect on population structure. Moreover, the results of this study give furtherevidence that the opportunity for sexual selection is influencedby the adult sex ratio and hence by the operational sex ratioin a population.     

The evolution of primate communities and societies in Madagascar

During their 120 to 165 million years of isolation, the flora and fauna of Madagascar evolved, to a large extent, independently of the African mainland.¹ In contrast to other oceanic islands, Madagascar is large enough to house the major components of tropical ecosystems, allowing tests of evolutionary hypotheses on the level of complete communities. Taking lemurs, the primates of Madagascar, as an example, evolutionary hypotheses correctly predict the organization of their community structure with respect to ecological correlates. Lemur social systems and their morphological correlates, on the other hand, deviate in some respects from those of other primates. Apparently, lemur social systems are influenced by several selection pressures that are weak or rare in other primates. These include variable activity patterns and avoidance of infanticide. The interspecific variation in lemur social systems therefore offers a unique opportunity for a comprehensive study of the determinants of primate social systems.     

Relation between the dominance rank of female rhesus monkeys and their access to males

Systematic observations were made on 12 measures of the sexual, aggressive, and social interactions of 24 male–female pairs of rhesus monkeys in six social groups, each consisting of one male and four ovariectomized females tested in a large room. Each female in a group was treated in turn first with estradiol alone and then with estradiol and progesterone in combination. When hormone-treated, the female was also observed during pair tests with the male in the same large observation room (four males, eight females, 240 group tests, 240 pair tests). The dominance ranks of females during group tests were determined post hoc by means of the dominance index [Zumpe & Michael, American Journal of Primatology 10:291–300, 1986]. In all six groups, the most dominant female virtually monopolized the male, and the subordinate females' interactions with the male, assessed during pair tests, were almost completely suppressed during group tests. This “dominant female effect” was a robust phenomenon that depended solely on female dominance rank. It was independent of the identity and hormonal status of females and of the social preferences of males as expressed in pair tests. These findings demonstrate the existence of female mate competition in an Old World primate.     

Natural selection in the <Emphasis Type="Italic">TLR-</Emphasis>related genes in the course of primate evolution

The innate immune system constitutes the front line of host defense against pathogens. Toll-like receptors (TLRs) recognize molecules derived from pathogens and play crucial roles in the innate immune system. Here, we provide evidence that the TLR-related genes have come under natural selection pressure in the course of primate evolution. We compared the nucleotide sequences of 16 TLR-related genes, including TLRs (TLR1–10), MYD88, TILAP, TICAM1, TICAM2, MD2, and CD14, among seven primate species. Analysis of the non-synonymous/synonymous substitution ratio revealed the presence of both strictly conserved and rapidly evolving regions in the TLR-related genes. The genomic segments encoding the intracellular Toll/interleukin 1 receptor domains, which exhibited lower rates of non-synonymous substitution, have undergone purifying selection. In contrast, TLR4, which carried a high proportion of non-synonymous substitutions in the part of extracellular domain spanning 200 amino acids, was found to have been the suggestive target of positive Darwinian selection in primate evolution. However, sequence analyses from 25 primate species, including eight hominoids, six Old World monkeys, eight New World monkeys, and three prosimians, showed no evidence that the pressure of positive Darwinian selection has shaped the pattern of sequence variations in TLR4 among New World monkeys and prosimians. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Coevolution of male and female reproductive traits in a simultaneously hermaphroditic land snail

Inter- and intraspecific studies in gonochoristic animals reveal a covariation between sperm characteristics and the size of the female reproductive tract, indicating a rapid evolutionary divergence, which is consistent with the theory of post-copulatory sexual selection. Simultaneous hermaphrodites differ from species with separate sexes (gonochorists) in that they possess both functional male and female reproductive organs at the same time. We investigated whether in hermaphroditic animals intraspecific variation in reproductive traits results from divergent coevolution, by quantifying the variation in male and female traits among six natural populations of the snail Arianta arbustorum and examining the covariation in interacting traits. There was a significant among-population variation in spermatophore volume, number of sperm transferred and sperm length, as well as in volume of the sperm storage organ (spermatheca) and number of tubules, but not in spermatheca length. We found a positive association between sperm number transferred and spermatheca volume. This result suggests that the same post-copulatory mechanisms as in gonochorists drive the correlated evolution of reproductive characters in hermaphrodites.     

Immunolocalization of aquaporin 1, 5, and 9 in the female pig reproductive system

Thirteen mammalian aquaporin (AQP) isoforms have been identified, and they have a unique tissue-specific pattern of expression. AQPs have been documented in the reproductive system of both male and female humans, rats, and mice. However, tissue expression and cellular and subcellular localization of AQPs are unknown in the female reproductive system of pigs. In this study, AQP1 immunoreactivity was detected in the capillary endothelium of the ovary. Distinct immunolabeling of capillary endothelium was also observed in the oviduct and uterus. AQP5 was expressed in flattened follicle cells of primordial follicles, granulosa cells of developing ovarian follicles, and muscle cells of the oviduct and uterus. Staining of AQP5 was also observed in the epithelial cells of the oviduct and uterine epithelium. AQP9 immunoreactivity was observed in granulosa cells of developing follicles. AQP9 was also localized in the luminal epithelial cells of the oviduct and uterine epithelia cells. This is, to our knowledge, the first study that shows tissue expression and cellular and subcellular localization of AQPs in the reproductive system of the female pig. Moreover, these results suggest that several subtypes of the AQPs (AQP1, 5, and 9) are involved in regulation of water homeostasis in the reproductive system of gilts.     

Testosterone levels and stress in women: the role of stress coping strategies, anxiety and sex role identification

This study evaluated the relation between testosterone changes in response to anticipatory stress and several psychological variables that contribute to the stress reaction. Salivary testosterone was determined in 76 female students under stress-free conditions and before an important examination. Individual stress reactions were highly variable in direction and extent: both significant increases and decreases were found. Thus the data did not confirm previous findings of general increases in testosterone levels under stress in women. Depending on the women's level of trait anxiety (assessed via STAI) and the general use of positive or negative cognitive coping strategies (assessed via SVF), we found significant differences in their baseline testosterone levels. Individual endocrine changes under stress were correlated with baseline testosterone levels: High testosterone concentrations at rest were more likely to drop under anticipatory stress than low concentrations. These contrasting effects can be explained by the significant interaction of trait anxiety and the sex role dimension of masculinity (assessed via BSRI) with testosterone production in females.     

Relative reproductive success of female moorhens using conditional strategies of brood parasitism and parental care

In a population of moorhens (Gallinula chloropus), at least27% of netting females laid one or more eggs in a neighbor'snest Females laid parasitically under three conditions: 56%of parasitic eggs were from nesting females that preceded layinga dutch in their own nest by a parasitic laying bout, 19% werefrom females whose nests were depredated before clutch completionand that laid the following egg parasiticaDy, and 25% were froma small number of females without territories, "non-nesting"parasites, that each laid a series of parasitic eggs. Clutchsizes varied greatly between females, but nesting females eachlaid a consistent clutch size both within and between seasonsfor a given mate and territory. Nesting females that employeda dual strategy of brood parasitism and parental care producedextra eggs that they laid in the nests of neighbors before layinga dutch in their own nests. Two out of ten females whose dutchesI experimentally removed during the laying period were successfullyinduced to lay their next egg in the nest of a neighbor. Nestingfemales that laid parasitically selected their hosts opportunisticallyfrom among the nests dosest to their territories. An experimentin which parasitic eggs were removed and hosts left to rearonly their own young showed that parasites did not choose hoststhat were better parents than pairs with contemporary neststhat were not parasitized. Females that only laid parasiticaDywithin a given season timed their parasitic laying bouts poorlyand achieved no reproductive success. Parasitic young rarelyfledged, and the mean seasonal reproductive success of nestingbrood parasites did not differ from that of nonparasitic females.However, the variance in reproductive success of nesting broodparasites was significantly higher than that of nonparasiticfemales.