Size-dependent mating strategies and the risk of cannibalism

The role of sexual selection in determining the nature and direction of sexual size dimorphism may depend upon the timing of sexual selection, and this may also influence the variation in male size. For example, selection through sperm competition favours smaller males in the highly sexually size dimorphic orb-weaving spider Nephila edulis , whereas larger males are better able to exclude their smaller rivals from the central hub of the web where mating takes place. We investigate experimentally the role of body size and hub tenure in determining male fertilization success when males of different sizes compete for a single female over a 24-h period that includes a period of darkness. Our results confirm that small and large males obtain similar paternity share but that, in contrast with previous studies, hub tenure does not translate into greater paternity share. Unexpectedly, smaller males are at greater risk of postmating sexual cannibalism than larger males, suggesting that natural selection through sexual cannibalism may place a lower limit on male size.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 355–363.     

Forceps Size Does Not Determine Fighting Success in European Earwigs

Male European earwigs (Forficula auricularia) possess substantially larger forceps than females and use these forceps to batter rivals in intrasexual contests to determine dominance. Although previous investigations have shown that male fighting and mating success increases with forceps size, it is not clear that sexual selection acts directly on forceps size per se; increased forceps size may be a correlated response to selection for some other trait. We experimentally reduced forceps length of males and paired them with unmanipulated males in staged encounters. Although apparent (postmanipulation) forceps length did not affect contest outcomes, original (premanipulation) forceps length did: males with longer original forceps won more contests. These results suggest that weapon size itself does not determine success in contests between male European earwigs. Thus, sexual selection may operate on forceps size in some other context or may act on some other trait that covaries with forceps size.     

Male Reproductive Success in a Promiscuous Armoured Catfish Corydoras Aeneus (Callichthyidae)

Among a variety of fish mating systems, promiscuity with random-mating seems to be most prevalent. However, detailed studies of promiscuity have been rare due partly to the peculiar difficulty in examination of male mating and reproductive success in the random mating. Females of the armoured catfish Corydoras aeneus (no sexual dimorphism other than size of males > females) spawn 10–20 egg-clutches with multiple males at a time, but an entire egg clutch is inseminated by sperm of a single male. We studied mating system of this fish in aquarium. Males had neither mating territories nor monopolized females, never being aggressive against rival males. Evidence of female preference for certain male traits including size was not detected. Females mated a male in proportion to his relative courtship frequency among males. Courtship frequency was not related to male size, and male mating success was not different between small and large males. Clutch size and insemination rate were different neither between small and large males nor between frequently and less frequently courting males. Thus, the male reproductive success will not be related to the male size, but directly to courtship frequency, indicating the random mating in this fish. There seemed to be fecundity advantage with size in female, and the consequent sexual difference in energy allocation will be responsible to the sexual dimorphism. We also discuss the low male-GSI in this promiscuous fish in which sperm competition hardly occurred.     

Effects of Sexual Selection Upon Sperm Morphology and Sexual Skin Morphology in Primates

We review possible effects of sexual selection upon sperm morphology, and sexual skin morphology, in primates. Comparative morphometric studies, involving 31 species representing 21 primate genera, revealed a positive relationship between volume of the sperm midpiece, occurrences of multiple partner matings by females, and large relative testes sizes, which indicate sperm competition. The midpiece houses the mitochondria required to power sperm motility. Hence, sperm competition may have influenced the evolution of increased mitochondrial loading in species where females mate with multiple partners during the fertile period. Females of some Old World monkey species and female chimpanzees exhibit large estrogen-dependent sexual skin swellings during the follicular phase of the menstrual cycle. Studies of mandrills support the conclusion that swellings act primarily as sexually attractive, graded signals and that swelling size may indicate current reproductive quality. Measurements of the genitalia in chimpanzees indicate a secondary function for female swellings. The swelling increases the operating depth of the female's vagina by 50% during the fertile phase of her cycle. Males have evolved long, filiform penes capable of placing sperm close to the os cervix during competitive multipartner matings. This may exemplify how morphologic specializations in females can influence the coevolution of advantageous genitalic specializations in males: the phenomenon that Eberhard (1985) dubbed cryptic female choice.     

Mating Behavior, Sexual Selection, and Copulatory Courtship in a Promiscuous Beetle

The function of male movements during copulation is unclear. These movements may be a result of the necessary mechanics of insemination, or they may also have further function, for instance, stimulating or courting a female during mating, perhaps influencing female mate choice. We present data from three experiments exploring the mating behavior and copulatory movements of the highly promiscuous beetle Psilothrix viridicoeruleus. Male mating success in the struggle over mating was not related to male or female size (measured by weight) but successful males were more vigorous in terms of copulatory movements. These males took longer to mount females but copulated longer and remained mounted longer. We discuss these results in terms of the mating system of Psilothrix and also in terms of observations of the timing of insemination during copulation. We suggest that copulatory movements in this species are best understood as copulatory courtship.     

Morphological Variation in Populations of <Emphasis Type="Italic">Eulemur albocollaris</Emphasis> and <Emphasis Type="Italic">E. fulvus rufus</Emphasis>

Sexual dimorphism in body size and canine weaponry is commonly associated with high levels of male-male competition. When group living species do not rely heavily on male-male competition for access to females, sperm competition may represent a viable alternative strategy. Unlike most haplorhine primates, lemurs are typically monomorphic in body weight and canine height. We assessed variability of body mass dimorphism and canine size dimorphism in brown lemurs using morphometric data from 3 populations in southeastern Madagascar: Eulemur fulvus rufus, E. albocollaris, and hybrids of the species. We found significant male-biased canine dimorphism in E. albocollaris in conjunction with body-size monomorphism. We observed similar patterns in the hybrids, but E. fulvus rufus exhibited significant female-biased size dimorphism and canine monomorphism. Testes volume was relatively high across study populations. Thus, sperm competition appears to be strong in brown lemurs. E. albocollaris males combine sperm competition with large canines, but not higher body mass, indicating a difference in sexual strategy from most lemurs. Patterns of body mass and canine size dimorphism are not uniform across brown lemur populations, indicating that future work on these populations can explicitly test models that predict relationships between size dimorphism and various types of competition.     

Sexual selection research on spiders: progress and biases

The renaissance of interest in sexual selection during the last decades has fuelled an extraordinary increase of scientific papers on the subject in spiders. Research has focused both on the process of sexual selection itself, for example on the signals and various modalities involved, and on the patterns, that is the outcome of mate choice and competition depending on certain parameters. Sexual selection has most clearly been demonstrated in cases involving visual and acoustical signals but most spiders are myopic and mute, relying rather on vibrations, chemical and tactile stimuli. This review argues that research has been biased towards modalities that are relatively easily accessible to the human observer. Circumstantial and comparative evidence indicates that sexual selection working via substrate-borne vibrations and tactile as well as chemical stimuli may be common and widespread in spiders. Pattern-oriented research has focused on several phenomena for which spiders offer excellent model objects, like sexual size dimorphism, nuptial feeding, sexual cannibalism, and sperm competition. The accumulating evidence argues for a highly complex set of explanations for seemingly uniform patterns like size dimorphism and sexual cannibalism. Sexual selection appears involved as well as natural selection and mechanisms that are adaptive in other contexts only. Sperm competition has resulted in a plethora of morphological and behavioural adaptations, and simplistic models like those linking reproductive morphology with behaviour and sperm priority patterns in a straightforward way are being replaced by complex models involving an array of parameters. Male mating costs are increasingly being documented in spiders, and sexual selection by male mate choice is discussed as a potential result. Research on sexual selection in spiders has come a long way since Darwin, whose spider examples are reanalysed in the context of contemporary knowledge, but the same biases and methodological constraints have persisted almost unchanged through the current boom of research.     

Contrasting selection pressure on body and weapon size in a polygynous megaherbivore

Sexual size dimorphism (SSD) is a common morphological trait in ungulates, with polygyny considered the leading driver of larger male body mass and weapon size. However, not all polygynous species exhibit SSD, while molecular evidence has revealed a more complex relationship between paternity and mating system than originally predicted. SSD is, therefore, likely to be shaped by a range of social, ecological and physiological factors. We present the first definitive analysis of SSD in the common hippopotamus (Hippopotamus amphibius) using a unique morphological dataset collected from 2994 aged individuals. The results confirm that hippos exhibit SSD, but the mean body mass differed by only 5% between the sexes, which is rather limited compared with many other polygynous ungulates. However, jaw and canine mass are significantly greater in males than females (44% and 81% heavier, respectively), highlighting the considerable selection pressure for acquiring larger weapons. A predominantly aquatic lifestyle coupled with the physiological limitations of their foregut fermenting morphology likely restricts body size differences between the sexes. Indeed, hippos appear to be a rare example among ungulates whereby sexual selection favours increased weapon size over body mass, underlining the important role that species-specific ecology and physiology have in shaping SSD.     

Pre- and Post-Copulatory Mate Selection Mechanisms in an African Dung Beetle, <Emphasis Type="Italic">Circellium bacchus</Emphasis> (Coleoptera: Scarabaeidae)

Females of most taxa mate selectively. Mate selection may be: (1) pre-copulatory, involving active female choice and male-male competition, and (2) post-copulatory, with cryptic female choice and sperm competition. Because female dung beetles (Circellium bacchus) invest heavily in parental care by ball-rolling and remaining with developing larva they are, therefore, expected to be highly selective when mating. Mate choice in this species was investigated via behavioral observations and investigations of genital allometry of both sexes, leading to conclusions about the mechanisms of, and male characteristics important in, female choice. Male–male competition seems to be crucial in mate selection of C. bacchus, although the females appeared to show no active mate choice. There is a negative allometric relationship between genital size and body size of males as predicted by the ‘one size fits all’ hypothesis (where males have genitalia that fit average-sized females). For the females, no relationship was found between genital size and body size. This might be as a result of the non-sclerotized nature of female genitalia, which may allow for greater morphological plasticity.     

Sexual behaviour and evolution of sexual dimorphism in body size in Jaera (Isopoda Asellota)

Individuals of the genus Jaera do not mate at random. In the species from the Mediterranean group, J. italica and. J. nordmanni, large males and medium sized females are at an advantage and their sizes are positively assorted. These effects are attributable to sexual competition between males. In the Ponlo-caspian species J. istri, no advantage of large males exists, but sexual selection could be the cause for a long passive phase prior to copulation and for normalizing selection upon female size at pairing. In the Atlantic species, J. albifrons, no selection can be ascertained.
Differential mating success in males appears as one of the causes of the evolution of sexual dimorphism in body size, which makes males larger, of equal size, or smaller than females according to the species. The reason for this reversal in dimorphism seems to differ in the two sexes. Sexual selection provides an explanation for the evolution of male size, while the interspecific changes in female length are more likely due to ecological factors.     

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.     

Size Dependent Sexual Selection in Drosophila ananassae

Mate choice based on body size is widespread and can have numerous consequences. We present data, which show the effect of male and female body size on sexual selection in Drosophila ananassae. The relationships between wing size, locomotor activity, mating latency, courtship pattern, fertility and mating success were studied. Mating latency was negatively correlated with wing length and with locomotor activity, while wing length and locomotor activity was positively correlated in males as well as in females. In female- and male-choice, we found that mate choice influenced size-assortative mating by: (1) large and small males preferring to mate with large females, (2) large males successfully competing for large females, leaving small males to mate with small females. Males increased their reproductive success by mating with large and more fecund females. In addition, in pairs of long/short winged flies, long winged flies courted and mated more successfully than short winged flies and they also have longer duration of copulation and more progeny than short winged flies. We found sterile mating in pairs of small winged males and females.     

Male strategies and Plio-Pleistocene archaeology

Archaeological data are frequently cited in support of the idea that big game hunting drove the evolution of early Homo, mainly through its role in offspring provisioning. This argument has been disputed on two grounds: (1) ethnographic observations on modern foragers show that although hunting may contribute a large fraction of the overall diet, it is an unreliable day-to-day food source, pursued more for status than subsistence; (2) archaeological evidence from the Plio-Pleistocene, coincident with the emergence of Homo can be read to reflect low-yield scavenging, not hunting. Our review of the archaeology yields results consistent with these critiques: (1) early humans acquired large-bodied ungulates primarily by aggressive scavenging, not hunting; (2) meat was consumed at or near the point of acquisition, not at home bases, as the hunting hypothesis requires; (3) carcasses were taken at highly variable rates and in varying degrees of completeness, making meat from big game an even less reliable food source than it is among modern foragers. Collectively, Plio-Pleistocene site location and assemblage composition are consistent with the hypothesis that large carcasses were taken not for purposes of provisioning, but in the context of competitive male displays. Even if meat were acquired more reliably than the archaeology indicates, its consumption cannot account for the significant changes in life history now seen to distinguish early humans from ancestral australopiths. The coincidence between the earliest dates for Homo ergaster and an increase in the archaeological visibility of meat eating that many find so provocative instead reflects: (1) changes in the structure of the environment that concentrated scavenging opportunities in space, making evidence of their pursuit more obvious to archaeologists; (2) H. ergaster's larger body size (itself a consequence of other factors), which improved its ability at interference competition.     

Sexual Selection on male cuticular hydrocarbons via male–male competition and female choice

Traditional views of sexual selection assumed that male–male competition and female mate choice work in harmony, selecting upon the same traits in the same direction. However, we now know that this is not always the case and that these two mechanisms often impose conflicting selection on male sexual traits. Cuticular hydrocarbons (CHCs) have been shown to be linked to both social dominance and male attractiveness in several insect species. However, although several studies have estimated the strength and form of sexual selection imposed on male CHCs by female mate choice, none have established whether these chemical traits are also subject to sexual selection via male–male competition. Using a multivariate selection analysis, we estimate and compare sexual selection exerted by male–male competition and female mate choice on male CHC composition in the broad‐horned flour beetle Gnatocerus cornutus. We show that male–male competition exerts strong linear selection on both overall CHC abundance and body size in males, while female mate choice exerts a mixture of linear and nonlinear selection, targeting not just the overall amount of CHCs expressed but the relative abundance of specific hydrocarbons as well. We discuss the potential implications of this antagonistic selection with regard to male reproductive success.     

Sexual cannibalism, competition, and size dimorphism in the orb-weaving spider Nephila plumipes Latreille (Araneae: Araneoidea)

The degree and direction of sexual dimorphism varies widely,but in several taxa of orb-weaving spiders, including Nephila,males may be less than one-tenth the size of females. This differenceis commonly attributed to selection through precopulation sexualcannibalism: females may either fail to detect very small males,or ignore them as potential prey items. However, there is oftenthe potential for male-male competition in these species becauseseveral males can be found on the web of a single female. Weinvestigated experimentally the effects of sexual cannibalismand male-male competition on male body size and hence sexualdimorphism in the Australian golden orb-weaver (Nephila plumipes).Small males were less likely to be detected and cannibalizedthan larger males. However, larger males excluded small malesfrom the central hub of the web, where mating takes place. Theconflicting effects of sexual cannibalism and male-male competitionmay be responsible for the relatively large variation in malebody size in this species.     

端黑萤和边褐端黑萤两性异形和雌性个体生育力

检测了端黑萤(Abscondita chinensis)和边褐端黑萤(Abscondita terminalis)繁殖期个体大小和形态特征的两性异形以及雌性繁殖输出。两因素方差分析表明,端黑萤的全长显著小于边褐端黑萤的全长,雌性个体显著大于雄性个体。以全长为协变量的两因素协方差分析显示,端黑萤的前胸背板宽、鞘翅长、复眼宽、胸长、腹宽、发光器面积及体重均显著小于边褐端黑萤,而腹长显著大于边褐端黑萤,触角长物种间差异不显著;雌性的前胸背板宽、腹长、腹宽及体重均显著大于雄性,而鞘翅长、复眼宽、胸长、触角长、发光器面积均显著小于雄性;物种和性别的交互作用对前胸背板宽、复眼宽、腹长、触角长及发光器面积影响显著,对其余的鞘翅长、胸长、腹宽及体重影响不显著。端黑萤和边褐端黑萤的两性异形指数分别为0.144和0.091。9个形态特征变量的主成分分析(Eigenvalue ≥ 1)发现,前2个主成分共解释56.8%的变异;复眼宽和发光器面积在第一主成分有较高的负负载系数,腹长有较高的正负载系数(解释31.3%变异);前胸背板宽和鞘翅长在第二主成分有较高的负负载系数(解释25.5%变异)。端黑萤和边褐端黑萤的平均窝卵数分别为28.9粒和18粒。线性回归显示,端黑萤和边褐端黑萤的窝卵数均与母体的全长及体重呈显著的正相关。单因素方差分析表明,特定全长的端黑萤的窝卵数显著大于边褐端黑萤。端黑萤和边褐端黑萤均属于雌性偏大的两性异形,是生育力选择、能量分配和运动综合影响的结果。雌萤腹腔容量等关键局部特征的增大是对生育力选择的适应,且生育力大小与两性异形差异程度成正相关。     

Impact of intrasexual selection on sexual dimorphism and testes size in the Mexican howler monkeys Alouatta palliata and A. pigra

One of the goals of physical anthropology and primatology is to understand how primate social systems influence the evolution of sexually selected traits. Howler monkeys provide a good model for studying sexual selection due to differences in social systems between related species. Here, we examine data from the sister howler monkey species Alouatta palliata and A. pigra inhabiting southeastern Mexico and northern Guatemala. We use a resampling approach to analyze differences in sexual dimorphism of body and canine size. In addition, we compare testes size as a way of gauging the intensity of sperm competition in both species. Morphometric data were collected from wild-caught individuals, including body mass and length, and dental data were obtained from casts from wild individuals and from museum specimens. Although A. pigra individuals are larger than their A. palliata counterparts, we find that both species exhibit similar levels of sexual dimorphism for all of the variables considered. Testicular volume results indicate that A. palliata male testes are on average twice as large as those of A. pigra males, suggesting more intense sperm competition in the former species. Our study shows that A. pigra is not highly sexually dimorphic as was once thought, and testes size differences suggest the need for a clearer understanding of howler monkey social systems.