The morphological basis of the underbite trait in langurs (P. melalophus and T. cristatus) with an analysis of adaptive and evolutionary implications

Underbites or mandibular incisor protrusions are widely prevalent among Coloboid monkeys. The morphological basis of the trait is explored in two langur species differing with respect to its incidence. P. melalophus, characterized by a high occurrence of underbites, has mandibular incisors which are significantly wider than those of T. cristatus, the more “normative” occlusal group. No other statistically significant incisor metric differences are noted. Comparisons of convexgent traits in the Colobinae, Alouattinae, and Ruminantia support the contention that underbites are a herbivorous adaptation. The zoogeographic distribution of the trait in langurs is discussed in the context of evolutionary dynamics.     

Females of the lekking great snipe do not prefer males with whiter tails

A previous experimental study of great snipe, Gallinago media, has reported an effect on male mating success of the amount of white in their tails. That result is one of a very limited set of existing experimental results supporting a female mate preference for a morphological trait in animals. However, a later observational study did not find any correlation between amount of white and male mating success. If females sample a limited number of males, their preferences need not result in strong relationships between mating success and trait values in males, possibly explaining the failure to find the predicted correlation. Yet, females of lekking species are thought to have ample opportunities for mate sampling. To resolve these contrasting results, we present in this paper (1) a larger correlational study (several leks during 10 years) showing no relationship between male mating success and whiteness of tails (measured in several ways), and most importantly (2) evidence that individual females do not mate predominantly with males with very white tails among those males that each female samples. These results show that females do not prefer males with whiter tails as mates, within the contemporary natural variation in the trait. They also show that there is no sexual selection of the trait at present. This does not necessarily imply that white tails are not a sexually selected adaptation in males, but the mechanisms are likely to have been different from direct mate choice of whiter tails per se. Copyright 2000 The Association for the Study of Animal Behaviour.     

A time series of evolution in action: a latitudinal cline in wing size in South American Drosophila subobscura

Drosophila subobscura is geographically widespread in the Old World. Around the late 1970s, it was accidentally introduced into both South and North America, where it spread rapidly over broad latitudinal ranges. This invading species offers opportunities to study the speed and predictability of trait evolution on a geographic scale. One trait of special interest is body size, which shows a strong and positive latitudinal cline in many Drosophila species, including Old World D. subobscura. Surveys made about a decade after the invasion found no evidence of a size cline in either North or South America. However, a survey made in North America about two decades after the invasion showed that a conspicuous size cline had evolved and (for females) was coincident with that for Old World flies. We have now conducted parallel studies on 10 populations (13 degrees of latitude) of flies, collected in Chile in spring 1999. After rearing flies in the laboratory for several generations, we measured wing sizes and compared geographic patterns (versus latitude or temperature) for flies on all three continents. South American females have now evolved a significant latitudinal size cline that is similar in slope to that of Old World and of North American flies. Rates of evolution (haldanes) for females are among the highest ever measured for quantitative traits. In contrast, the size cline is positive but not significant for South or North American males. At any given latitude, South American flies of both sexes are relatively large; this in part reflects the relatively cool climate of coastal Chile. Interestingly, the sections of the wing that generate the size cline for females differ among all three continents. Thus, although the evolution of overall wing size is predictable on a geographic scale (at least for females), the evolution of size of particular wing components is decidedly not.     

D4 dopamine-receptor (DRD4) alleles and novelty seeking in substance-dependent, personality-disorder, and control subjects.

Two reports have been published suggesting an association between the personality trait of novelty seeking and the DRD4*7R allele at the D4 dopamine-receptor locus (with heterozygotes or homozygotes for DRD4*7R having higher novelty seeking). We studied novelty seeking and four coding-sequence polymorphisms affecting protein structure in the D4 dopamine-receptor gene (DRD4) in a sample of 341 American subjects, of whom 224 are of primarily European ancestry and 117 are of primarily African ancestry. These subjects had diagnoses of substance dependence or personality disorder (PD) or were screened to exclude major psychiatric diagnosis. We found that, although the substance-dependent subjects had significantly higher novelty seeking than the control and PD subjects, they did not differ in DRD4*7R allele frequency. There was no association between any DRD4 polymorphism and novelty seeking in any population or diagnostic group, except for a significant association between the DRD4*7R allele and lower novelty seeking among European American females and African American substance abusers. The novelty seeking of subjects heterozygous for a null mutation did not differ from that of subjects with two functional alleles. We conclude that the most likely explanation of these results is that the DRD4 VNTR does not influence directly the trait of novelty seeking, in these samples.     

Reproductive strategies under multiparasitism in natural populations of the parasitoid wasp Nasonia (Hymenoptera)

Parasitoid Nasonia wasps adjust their progeny sex ratio to the presence of conspecifics to optimize their fitness. Another trait under female control is the induction of offspring diapause. We analysed progeny sex ratios and the proportion of diapausing offspring of individual Nasonia females in host patches parasitized by two species, Nasonia vitripennis and Nasonia giraulti, in North American field populations using microsatellite fingerprinting. Both Nasonia species produced similar sex ratios on hosts that were co‐parasitized by their own species as by the other species, indicating that females do not distinguish between con‐ and heterospecific clutches. The sex ratios of the diapause and adult fractions of mixed broods from single females were not correlated. We found further indications that N. vitripennis females take the emergence time of the offspring into account in their sex allocation. The reproductive strategies of Nasonia under multiparasitism are largely adaptive, but also partially constrained by information.     

Mate‐choice copying: A fitness‐enhancing behavior that evolves by indirect selection

A spatially explicit, individual‐based simulation model is used to study the spread of an allele for mate‐choice copying (MCC) through horizontal cultural transmission when female innate preferences do or do not coevolve with a male viability‐increasing trait. Evolution of MCC is unlikely when innate female preferences coevolve with the trait, as copier females cannot express a higher preference than noncopier females for high‐fitness males. However, if a genetic polymorphism for innate preference persists in the population, MCC can evolve by indirect selection through hitchhiking: the copying allele hitchhikes on the male trait. MCC can be an adaptive behavior—that is, a behavior that increases a population's average fitness relative to populations without MCC—even though the copying allele itself may be neutral or mildly deleterious.     

Fitness cost of pheromone production in signaling female moths

A secondary sexual character may act as an honest signal of the quality of the individual if the trait bears a cost and if its expression is phenotypically condition dependent. The cost of increasing the trait should be tolerable for individuals in good condition but not for those in a poor condition. The trait thus provides an honest signal of quality that enables the receiver to choose higher quality mates. Evidence for sex pheromones, which play a major role in shaping sexual evolution, inflicting a signaling cost is scarce. Here, we demonstrate that the amount of the major component of the pheromone in glands of Lobesia botrana (Lepidoptera) females at signaling time was significantly greater in large than in small females, that male moths preferred larger females as mates when responding to volatile signals, and small virgin females, but not large ones, exposed to conspecific pheromone, produced, when mated, significantly fewer eggs than nonexposed females. The latter indicates a condition-dependent cost of signaling. These results are in accordance with the predictions of condition-dependent honest signals. We therefore suggest that female signaling for males using sex pheromones bears a cost and thus calling may serve as honest advertisement for female quality.     

An analysis of cercopithecoid odontometrics. I. The scaling of the maxillary dentition.

The relations between two tooth indices, post-canine area and incisor width in the upper jaw, and three variables, diet, body weight and body weight dimorphism, were examined separately for the males and females of 29 cercopithecoid species. Each species was assigned to one of three diet classes (folivore, frugivore, omnivore). Data on the other variable consisted of species means (log-transformed) obtained from published sources. The analytic techniques used were bivariate and multiple regression, the tooth indices being the dependent variables. All tooth indices scaled isometrically within diet classes, and all except female incisor width scaled with positive allometry across diet classes. In both sexes, the body weight adjusted mean incisor width of folivores was significantly smaller than that of either frugivores or omnivores. In the females, the body weight adjusted mean post-canine areas did not differ significantly across diet classes, while in the males the omnivores had a larger body weight adjusted mean post-canine area than either the folivores or frugivores. Female post-canine area was the only tooth index for which body weight dimorphism was a significant predictor. Extrapolations of these findings to other extant and to fossil primate species are discussed.     

Rethinking incisor size and diet in anthropoids: diet, incisor wear and incisor breadth in the African apes

In a seminal study Hylander (1975) concluded that the length of the incisor row in catarrhines considered frugivores is longer relative to body mass than in those classified as folivores. Assuming that large fruits require greater incisal processing than do leaves, stems, berries, and seeds, he argued that the larger incisors of frugivores increased their resistance to wear. The present analysis examines diet, incisor wear, and incisor crown breadth in cranial samples of western lowland gorillas and chimpanzees. Incisor wear rate was assessed on the basis of the extent of incisor crown reduction observed at sequential stages of first molar wear. Incisor metrics were obtained from the unworn teeth of juveniles. Results suggest that incisor wear is greater in the more folivorous western lowland gorillas than in more frugivorous chimpanzees. Moreover, incisor crown dimensions do not differ appreciably among African apes. These findings fail to support the hypothesis that slower wear rates are associated with broader incisor crowns, and raise new questions regarding the significance of incisor row length in anthropoids.     

Relationship between canine dimorphism and mandibular morphology in the hamadryas baboon and the Japanese monkey

To examine morphological interrelationships between canine size and mandibular corpus shape, inter-sex comparisons were made in the hamadryas baboon and the Japanese monkey, known to display extreme and moderate canine dimorphism, respectively. Results of adult comparisons showed that all mandibular dimensions were significantly larger in the males than in the females in both species. In the hamadryas baboon, the males also exhibited a higher ratio of anterior to posterior corpus height than the females. This sex difference in corpus shape was not significant in the Japanese monkey, indicating lack of involvement of canine dimorphism. Analysis of mandibular growth patterns in the hamadryas baboon demonstrated that significant sexual size difference did not occur before incisor eruption, and that the anteriorly high corpus of the adult male mandible was associated with a rapidly increasing symphyseal height after incisor eruption. It was also shown that the female canine started to erupt shortly after incisor eruption, while the forming male canine continued to stay near the mandibular base and developed further in size until eruption. The relative positions of the incisors kept shifting upwards even after eruption in the males, while they hardly changed in the females. It is therefore suggested that the prolonged development and size increase of the male canine is accompanied by further enlargement of the symphysis, resulting in the higher anterior corpus of the adult males compared to the adult females. The results thus indicate the importance of understanding the spatial relationships of the developing teeth in interpreting mandibular morphology.     

Community functional trait composition at the continental scale: the effects of non‐ecological processes

Ecological communities and their response to environmental gradients are increasingly being described by measures of trait composition at the community level – the trait‐based approach. Whether ecological or non‐ecological processes influence trait composition between communities has been debated. Understanding the processes that influence trait composition is important for reconstructing paleoenvironmental conditions from fossil deposits and for understanding changes in community functionality through time. Here, we assess the influence of ecological and non‐ecological processes on the distribution of traits within North American mammals. We found that non‐ecological processes including historical contingency, spatial autocorrelation, and evolutionary history do not influence trait composition; however, the variance in trait composition is highly explained by climate gradients. Our results suggest that habitat breadth, terrestriality, diet breadth, and reproductive traits are strong candidates as proxies for measuring functional aspects of environments in the past and present.     

Defining Vertical Bars in Relation to Female Preference in the Swordtail Fish Xiphophorus cortezi (Cyprinodontiformes, Poeciliidae)

Defining a male trait in relation to female preference is an important step towards determining the role that female preference has played in the evolution of a male trait. The pigment pattern 'vertical bars' is a complex male trait that functions as a sexual signal in swordtail fishes. A previous study suggested that X. cortezi females had a preference for males with more vertical bars. In that study, however, bar number had been reduced on the stimulus males by removing the most posterior bars, which changed the distance over which the bars spanned, in addition to changing bar number. To determine whether X. cortezi females have a preference for greater bar span and/or number of bars, we tested for female preference in two experiments. First, we gave females a choice between males with a greater bar span or a smaller bar span, holding bar number and total pigmented area constant. Females spent significantly more time with the males with the smaller bar span. This result, together with results from the previous study, suggest that females do not assess bar span alone, but possibly a composite component of the bars, such as bar frequency (number of bars/bar span). Secondly, we gave females a choice between six thinner bars or one wider bar of equal pigmented area. The six-bar treatment had a greater bar span in addition to more bars. We detected no preference for either treatment. These results suggest that the previously detected preference for more bars may actually reflect a preference for greater pigmented area. Finally, we present data on natural variation in the bars for X. cortezi and use principal components analysis to provide a composite definition of this trait.     

Lingual incisor traits in modern hominoids and an assessment of their utility for fossil hominoid taxonomy

The morphology of the anterior dentition has received scant attention for purposes of taxonomic discrimination. Recently, however, lingual incisor morphology was used in differentiating several Miocene ape species and genera. This paper assesses the utility of this morphology for taxonomic discrimination by examining the nature and patterns of variation in lingual incisor morphology in extensive samples of modern chimpanzees, gorillas, orangutans, and gibbons. This paper documents discrete morphological traits on the lingual side of incisors. Trait frequencies are used in univariate and multivariate analyses to examine the apportionment of variation in species, subspecies, and populations. A correlation between lingual incisor traits, tooth dimensions, and sex attempts to determine if such factors affect the manifestation of traits. Finally, the findings are applied to understanding patterns of variation in the Miocene hominids. The study demonstrates that: 1) lingual incisor morphology differs substantially between the hylobatids and great apes; 2) variation in incisor traits is high within species, and most of it is found within local populations; and 3) incisor traits do not correlate significantly with incisor dimensions or sex. Species and to some extent subspecies of extant hominoids can be differentiated statistically using lingual incisor traits, but the frequency of traits such as continuous or discontinuous cingulum, or the presence or absence of pillars, differentiates them. Given this pattern of variation, I argue that it is necessary to assume and document similar patterns of variation in Miocene apes before incisor morphology is used for differentiating taxa.     

Is hissing behaviour of incubating great tits related to reproductive investment in the wild?

Anti-predator behaviour of breeding animals is a complex trait that depends on current reproductive investment as well as individual differences in risk-taking propensities. In response to nest predation, many bird species produce specific sounds, such as the hissing calls in incubating great tits (Parus major), that may provoke an acoustic startle response in the predator. However, it is still unclear whether the propensity of incubating females to produce hissing calls towards nest predators depends on the reproductive investment. With our 3-year study, we show that response type (females that do not hiss versus females giving hissing calls) to a potential nest predator, the woodpecker, is a repeatable trait. We found no differences in the studied reproductive traits between hissing and non-hissing birds. Interestingly, among hissing birds, fast-responding females started egg-laying earlier than slow-responding ones. Among non-hissing birds, heavier birds initiated clutches earlier. We also revealed that hissing birds breed in areas with decreased nest-box occupancy, suggesting either that they potentially select different areas to breed or that territory size is larger as a result of hissing birds being more aggressive. These findings demonstrate that response type is not related to the early reproductive value of the brood across distinct behavioural groups. However, our results do suggest that non-hissing and hissing females differ in terms of individual quality or dominance or personality related aspects.     

Sexual selection when the female directly benefits

Why do females of many species mate with males on the basis of traits apparently detrimental to male survival? The answer may lie in the fact that these male traits are correlated with male condition. We consider the argument that high male condition directly benefits female fecundity and/or viability (e.g. through lower transmission of parasites, improved control of resources, or better paternal care). Using a quantitative genetic model we show how female preferences for male traits that indicate condition can evolve, even if the male traits themselves have deleterious effects on both the male and the female's fecundity. So-called ‘arbitrary preferences’ can spread in this way because male traits subject to sexual selection are often under additional selection to become correlated with condition. At equilibrium the positive effects of male condition on a female's fecundity and the negative effects of the male trait on her fecundity are balanced and the female preference is under stabilizing selection. The male trait will often be correlated with viability, but not with fecundity, even though the preference evolved as a result of differences in male fecundity. The mean fecundity of females is not maximized, and can steadily decline as the male trait and female preference evolve. If the male trait has no direct deleterious effects on female fecundity, as may happen in species with no paternal care, female preferences are under continuous directional selection to increase.     

Demography can favour female-advantageous alleles

When female fecundity is relatively independent of male abundance, while male reproduction is proportional to female abundance, females have a larger effect on population dynamics than males (i.e. female demographic dominance). This population dynamic phenomenon might not appear to influence evolution, because male and female genomes still contribute equally much to the next generation. However, here we examine two evolutionary scenarios to provide a proof of principle that spatial structure can make female demographic dominance matter. Our two simulation models combine dispersal evolution with local adaptation subjected to intralocus sexual conflict and environmentally driven sex ratio biases, respectively. Both models have equilibria where one environment (without being intrinsically poorer) has so few reproductive females that trait evolution becomes disproportionately determined by those environments where females survive better (intralocus sexual conflict model), or where daughters are overproduced (environmental sex determination model). Surprisingly, however, the two facts that selection favours alleles that benefit females, and population growth is improved when female fitness is high, together do not imply that all measures of population performance are improved. The sex-specificity of the source–sink dynamics predicts that populations can evolve to fail to persist in habitats where alleles do poorly when expressed in females.     

THE VARIABILITY IS IN THE SEX CHROMOSOMES

Sex differences in the mean trait expression are well documented, not only for traits that are directly associated with reproduction. Less is known about how the variability of traits differs between males and females. In species with sex chromosomes and dosage compensation, the heterogametic sex is expected to show larger trait variability (“sex‐chromosome hypothesis”), yet this central prediction, based on fundamental genetic principles, has never been evaluated in detail. Here we show that in species with heterogametic males, male variability in body size is significantly larger than in females, whereas the opposite can be shown for species with heterogametic females. These results support the prediction of the sex‐chromosome hypothesis that individuals of the heterogametic sex should be more variable. We argue that the pattern demonstrated here for sex‐specific body size variability is likely to apply to any trait and needs to be considered when testing predictions about sex‐specific variability and sexual selection.     

Sexual and temporal variations in floral scent in the subdioecious shrub Eurya japonica Thunb

In many flowering plants, floral scents are a significant trait for visitors, playing an important role in attracting pollinators and/or detracting herbivores. The evolution of flowering plants from hermaphroditism to dioecy is often accompanied by sexual dimorphism in floral scent. In this study, floral scents emitted by different sexual morphs of the subdioecious shrub Eurya japonica Thunb. were collected using a dynamic headspace method, and sexual and temporal variations were evaluated by gas chromatography–mass spectrometry (GC–MS). Two volatiles, α‐pinene and linalool, were identified as the major components of floral scents in females, hermaphrodites, and males. The males emit higher amounts of floral scents, particularly α‐pinene, compared to females or hermaphrodites. Floral scents emitted by males generally decrease as flowers enter senescence, whereas those from females or hermaphrodites do not significantly differ. Intraspecific variations in floral scents of subdioecious species provided by this study would contribute to better understanding of sexual dimorphism in floral scent.     

Is the Expression of Male Traits in Female Lesser Kestrels Related to Sexual Selection?

Some lesser kestrel females (Falco naumanni) show male plumage traits, i.e. grey rumps and tails. This phenomenon has seldom been analyzed in birds, and two hypotheses have been suggested to explain it. The first proposes that, when sexual selection acts favouring the expression of a trait in males, females could show the analogous character by genetic correlation (indirect sexual selection). Alternatively, the expression of these traits in females could be favoured by intra-sexual competition or even by male mate choice selecting ornamented females (direct sexual selection). We have tested if females with male traits are favoured by direct sexual selection, through a 3-yr observational study of 239 female lesser kestrels. Our results cannot support the predictions, as females with grey plumages do not achieve access to better breeding opportunities or fitness benefits. These traits do not seem to be honest signals of phenotypic quality, since physical condition and survival did not differ between females which showed male traits and those which did not. The expression of male traits in these females increased with their ages, but showing a high individual variability. Finally, since the genetic correlation hypothesis is unlikely in this species because all males have grey rumps and tails, we propose a new age-related hormonal explanation.     

Sexual differences in response to larval food stress in two nuptial feeding orthopterans – implications for sexual selection

The extent to which a trait deteriorates in response to stress can indicate its fitness importance. Food limitation is a naturally-occurring stress in the katydid Kawanaphila nartee (Orthoptera: Tettigoniidae) and should cue larvae as to the level of sexual competition expected; hunger drives adult females to compete for copulations and thus acquire spermatophore meals. This increases sexual selection on females relative to males for increased body mass. As predicted, under experimental food-stress larval female K. nartee showed little loss of mass whereas male mass decreased. In contrast, the sizes of body parts less critical to mating success showed similar decreases in males and females. For katydids such as Conocephalus nigropleurum , where reversals in mating roles do not occur, larval food stress should result in a greater preservation of male body mass, an important trait in male mating success. This prediction was supported; male mass decreased less than that of females.