The phylogeny of diving beetles (Coleoptera: Dytiscidae) and the evolution of sexual conflict

A model of evolution based on conflicts of interest between the sexes over mating decisions is examined in relation to diving beetles (Dytiscidae). The model predicts the following evolutionary sequence: (1) cost to females of mating increases, (2) females evolve behavioural resistance to male mating attempts, (3) males evolve devices to overcome female resistance, and (4) females evolve morphological counter-adaptations to the male devices. This model is tested using species of Dytiscidae, in which (1) some species have a very long mating duration while others mate quickly, (2) females of some species resist male mating attempts by swift and erratic swimming when seized by a male, (3) males of some species possess a grasping device in the form of sucker-shaped setae on the legs used to adhere to the pronota or elytra of females prior to mating, and (4) females of some species have a modified dorsal cuticle with irregular sculpturing which appears to interfere with the male adhesive setae. The predicted order of evolution of some of these features was tested in a cladistic analysis of 52 taxa in Dytiscidae and Hygrobiidae using characters from adult and larval morphology and a portion of the gene wingless . The combined analysis resulted in nine most parsimonious cladograms. The consensus cladogram of these indicates that male sucker setae arose a single time in a clade of Dytiscinae. Nested within this clade are five groups with an independently derived, modified dorsal cuticle in females. This pattern of characters in Dytiscinae is consistent with the prediction implied by the model of sexual selection. The utility of wingless as a marker for phylogenetic analysis of diving beetles is discussed, and the resulting phylogeny is compared with previous analyses and current classification.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 359–388.     

Head shape allometry and proximate causes of head sexual dimorphism in Podarcis lizards: joining linear and geometric morphometrics

Podarcis bocagei and Podarcis carbonelli are two lacertid species endemic to the western Iberian Peninsula, and both show head size and shape sexual dimorphism. We studied immature and adult head sexual dimorphism and analysed ontogenetic trajectories of head traits with body and head size, aiming to shed light on the proximate mechanisms involved. Immatures were much less dimorphic than adults, but geometric morphometric techniques revealed that head shape sexual differences are already present at this stage. Males and females differed in allometry of all head characters with body size, with males showing a disproportionate increase of head size and dimensions. On the other hand, head dimensions and head shape changed with increasing head size following similar trends in both sexes, possibly indicating developmental restrictions. Consequently, adult sexual dimorphism for head characters in these species is the result of both shape differences in the immature stage and hypermetric growth of the head in relation to body size in males.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 111–124.     

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.     

Functional basis for sexual differences in bite force in the lizard Anolis carolinensis

In many species of lizards, males attain greater body size and have larger heads than female lizards of the same size. Often, the dimorphism in head size is paralleled by a dimorphism in bite force. However, the underlying functional morphological basis for the dimorphism in bite force remains unclear. Here, we test whether males are larger, and have larger heads and bite forces than females for a given body size in a large sample of Anolis carolinensis . Next, we test if overall head shape differs between the sexes, or if instead specific aspects of skull shape can explain differences in bite force. Our results show that A. carolinensis is indeed dimorphic in body and head size and that males bite harder than females. Geometric morphometric analyses show distinct differences in skull shape between males and females, principally reflecting an enlargement of the jaw adductor muscle chamber. Jaw adductor muscle mass data confirm this result and show that males have larger jaw adductors (but not jaw openers) for a given body and head size. Thus, the observed dimorphism in bite force in A. carolinensis is not merely the result of an increase in head size, but involves distinct morphological changes in skull structure and the associated jaw adductor musculature.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 111–119.     

Quantification of sexual dimorphism in Asellus aquaticus (Crustacea: Isopoda) using outline approaches

A marked sexual dimorphism is often observed in arthropods species in which males perform precopulatory mate guarding. It is generally thought to reflect the influence of sexual selection. Until now, sexual dimorphisms associated with mate guarding have mainly been qualitatively described. However, assessing the effects of sexual selection on sexual dimorphims requires a preliminary quantitative assessment of differences in morphology between sexes. Using Fourier analyses, we tested if morphological dimorphisms could be quantitatively assessed in the isopod Asellus aquaticus . In addition, we checked whether sexual dimorphism in shape was exclusively related to mate guarding through considering characters that are not, a priori , implicated in mating behaviour. To assess the potential role of sexual selection in shaping morphology, we then examined how dimorphic characters could influence males' pairing success. Three characters (pleotelson, paraeopods 4 and 5) differed significantly in shape between males and females. In addition, two characters (pleotelson and paraeopods 4) differed in shape between guarding males and non-guarding males, with the latter being closer in shape to females. This suggests that sexual selection may be partly responsible for the observed morphological divergence between sexes in A. aquaticus .  © 2002 The Linnean Society of London, Biological Society of the Linnean Society , 2002, 77 , 523–533.     

Sexual dimorphism and the differential mortality model: is behaviour related to survival?

There are numerous hypotheses to explain the evolution of sexual dimorphism in spiders. One of the most controversial is the differential mortality model (DMM) which proposes that differing rates of (adult) male and female mortality can result in a skewed operational sex ratio and lead to the evolution of small males. This hypothesis has been examined using a comparative approach which assumes that the behaviour of males and females could be used as a surrogate measure of mortality. We tested this assumption using two model species, Hogna helluo and Pardosa milvina (Araneae: Lycosidae) that differ in the degree of sexual dimorphism both in terms of body size and level of activity. Our data demonstrate that differences in male and female behaviour are not predictive of differences in mortality. Rather, as in other organisms, mortality is a complex phenomenon dependent on activity as well as size. These data call into question the methods previously used to test the DMM and suggest that understanding sexual size dimorphism (SSD) in spiders will require evaluation of historical constraints as well as how size currently influences fitness in each sex.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 97−103.     

Female-biased sexual size dimorphism in tinamous: a comparative test fails to support Rensch's rule

Rensch's rule states that degree of sexual dimorphism increases with body size in species with larger males, and decreases with body size in those with larger females. To test this rule, we assessed the pattern of sexual size dimorphism in tinamous using a comparative analysis of independent contrasts. Tinamous are a monophyletic group of primitive birds comprising at least 47 ground dwelling species with prominent or exclusive paternal care of eggs and offspring. Although the size of females exceeded that of males in most considered species, we found an isometric relationship between males and females, instead of the negative allometric one predicted by Rensch's rule. Previous studies in Strigiformes and Falconiformes found positive allometric and isometric relationships respectively, and, considering these findings with our results, we conclude that Rensch's rule is not supported by birds with exclusively female-biased sexual dimorphism in size.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 519–527     

Size dimorphism in Rankinia [Tympanocryptis] diemensis (Family Agamidae): sex-specific patterns and geographic variation

Sexual dimorphism has implications for a range of biological and ecological factors, and intersexual morphological differences within a species provide an ideal opportunity for investigating evolutionary influences on phenotypic variation. We investigated sexual size dimorphism (SSD) in an agamid species, Rankinia [Tympanocryptis] diemensis , to determine whether overall size and/or relative morphological trait size differences exist and whether geographic variation in size dimorphism occurs in this species. Relative morphological trait proportions included a range of head, limb, and inter-limb measurements. We found significant overall intersexual adult size differences; females were the larger sex across all sites but the degree of dimorphism between the sexes did not differ between sites. This female-biased size difference is atypical for agamid lizards, which are usually characterized by large male body size. In this species, large female-biased SSD appears to have evolved as a result of fecundity advantages. The size of relative morphological trait also differed significantly between the sexes, but in the opposite direction: relative head, tail, and limb sizes were significantly larger in males than females. This corresponds to patterns in trait size usually found in this taxonomic group, where male head and limb size is important in contest success such as male–male rivalry. There were site-specific morphological differences in hatchlings, including overall body size, tail, inter-limb, thigh, and hindlimb lengths; however, there were no sex-specific differences indicating the body size differences present in the adult form occur during ontogeny.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 699–709.     

Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)

Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons ( Morelia spilota ) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 113–125.     

Sexual dimorphism and sexual conflict in the diving beetle Agabus uliginosus (L.) (Coleoptera: Dytiscidae)

Sexual conflict can drive intersexual arms races, with female resistance and male persistence traits coevolving antagonistically. Such arms races are well documented in some diving beetles, although the extent of sexual conflict in this family remains unclear. The European dytiscid Agabus uliginosus has a strikingly dimorphic female; individuals from most regions are smooth and male‐like, whereas those from some populations have a strongly roughened dorsum, a trait that has attracted the name dispar. We demonstrate that rough and smooth females differ consistently in the development of dorsal surface microreticulation, and that these females are associated with males that differ in the development of their persistence traits. These findings extend the occurrence of pre‐insemination sexual conflict and associated intrasexual dimorphism in Dytiscidae, and suggest that such mating systems are relatively widespread in these beetles.     

Ontogenetically stable dimorphism in a lacertid lizard (Acanthodactylus boskianus) with tests of methodology and comments on life-history

Recent arguments in the literature prompted us to compare methods for assessing sexual dimorphism in body proportions of lacertid lizards, using Acanthodactylus boskianus . Although expressing body-part measurements as proportional to head length was the most effective method, we recommend using trunk length for the baseline as a general method for lizards. We also argue that, when aiming to assess sexual dimorphism in body proportions of lizards, if the context is ecological, all available adults should be included. However, for morphology and taxonomy, small sub-samples of the largest individuals that maximally express their genetic morphological potential should be used. In A. boskianus , the sexual dimorphism of mensural characters in adults was typical: males were larger, with relatively larger head and appendages. However, the ontogeny of this dimorphism was unusual in that the differences existed already in youth and thereafter persisted isometrically. The sexual dimorphism of meristic characters was male-biased in numbers of femoral pores and of caudal vertebrae, and female-biased in numbers of ventral plates along the trunk and of precaudal vertebrae. Size dimorphism may conceivably play a role in sex recognition because two potential visual cues (i.e. size dimorphism and dichromatism) appear to complement each other.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 97 , 275–288.     

Bite-force performance and head shape in a sexually dimorphic crevice-dwelling lizard, the common chuckwalla [Sauromalus ater (= obesus)]

The Common Chuckwalla [ Sauromalus ater (=  obesus )] is a large, sexually dimorphic lizard with a flattened head that takes refuge from predators in rock crevices. Males use their relatively large heads to bite competing males during territorial fights and to restrain females during copulation. Flattened heads with an antipredator function (i.e. seeking refuge in crevices) and enlarged heads with intrasexual competition and reproductive functions suggest possible antagonism between selective pressures on head morphology in males. To examine this hypothesis, we performed a morphometric analysis and measured the bite-force performance of 49 adult chuckwallas. Males had disproportionately wider heads than females, but did not have deeper heads. Males bit with nearly four times the force of females, consistent with the notion of sexual selection for high bite force in males. Although constrained by crevice-wedging behaviour, head depth was a good predictor of bite force in both sexes. In males, however, osteological head width also was a good predictor of bite force. These results are consistent with the hypothesis that head shape in males is under antagonistic selective pressures, which may partly explain the pattern of head shape dimorphism. The disproportionately wide head of males may reflect anatomical modifications to enhance bite force in response to sexual selection in spite of presumed constraints on head shape for crevice-wedging behaviour  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 215–222.     

The fitness advantage of a high-performance weapon

Weapons used in combat between males are usually attributed to sexual selection, which operates via a fitness advantage for males with weapons of better 'quality'. Because the performance capacity of morphological traits is typically considered the direct target of selection, Darwin's intrasexual selection hypothesis can be modified to predict that variation in reproductive success should be explained by variation in performance traits relevant to combat. Despite such a straightforward prediction, tests of this hypothesis are conspicuously lacking. We show that territorial male collared lizards with greater bite-force capacity sire more offspring than weaker biting rivals but exhibit no survival advantage. We did not detect stabilizing or disruptive selection on bite-force capacity. Taken together, these results support the hypothesis that superior weapon performance provides a fitness advantage through increased success in male contests. Sexual selection on weapon performance therefore appears to be a force driving the evolution and maintenance of sexual dimorphism in head shape.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 840–845.     

Sex‐specific evolution of bite performance in Liolaemus lizards (Iguania: Liolaemidae): the battle of the sexes

Although differential selective pressures on males and females of the same species may result in sex‐specific evolutionary trajectories, comparative studies of adaptive radiations have largely neglected within‐species variation. In this study, we explore the potential effects of natural selection, sexual selection, or a combination of both, on bite performance in males and females of 19 species of Liolaemus lizards. More specifically, we study the evolution of bite performance, and compare evolutionary relationships between the variation in head morphology, bite performance, ecological variation and sexual dimorphism between males and females. Our results suggest that in male Liolaemus, the variation in bite force is at least partly explained by the variation in the degree of sexual dimorphism in head width (i.e. our estimate of the intensity of sexual selection), and neither bite force nor the morphological variables were correlated with diet (i.e. our proxy for natural selection). On the contrary, in females, the variation in bite force and head size can, to a certain extent, be explained by variation in diet. These results suggest that whereas in males, sexual selection seems to be operating on bite performance, in the case of females, natural selection seems to be the most likely and most important selective pressure driving the variation in head size. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 461–475.     

Sexual size dimorphism and sex ratios in dragonflies (Odonata)

Sexual size dimorphism and biased sex ratios are common in animals. Rensch's rule states that sexual size dimorphism (SSD) would increase with body size in taxa where males are larger than females and decrease with body size in taxa where females are larger. We tested this trend in dragonflies (Odonata) by analysing body size of 21 species and found support for Rensch's rule. The increase in SSD with increasing size among species can be explained by sexual selection favouring large males. We also estimated the slope of the relationship between sex ratio and size ratio in populations of the 21 species. A negative slope would suggest that the larger sex suffers from high mortality in the larval stage, consistent with riskier foraging. The slope of this relationship was negative, but after correcting for phylogentic non-independence with independent contrasts the relationship was no longer statistically significant, perhaps because of phylogenic inertia or low sample size.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 507–513.     

Sexual dimorphism in the attachment ability of the Colorado potato beetle Leptinotarsa decemlineata (Coleoptera: Chrysomelidae) to rough substrates

Many representatives of the beetle family Chrysomelidae exhibit a distinctive sexual dimorphism in the structure of adhesive tarsal setae. The present study demonstrates the influence of surface roughness on the friction force of Leptinotarsa decemlineata males and females. The maximum friction force of individual beetles was measured on epoxy resin surfaces (smooth and with asperities ranging from 0.3 to 12.0 microm) using a centrifugal force tester. On the smooth surface, no considerable differences between males and females were found, whereas on rough surfaces, females attached significantly (up to two times) stronger than males. Clawless beetles generated lower forces than intact ones, but demonstrated similar differences between males and females. The results indicate that the female adhesive system has its main functional trait in a stronger specialisation to rough plant surfaces whereas the adhesive system of males possess a certain trade-off between attachment to rough plant surfaces during locomotion on vegetation and to the smooth surface of the female elytra, while mating.     

Morphology and somatometric growth of long-tailed macaques (Macaca fascicularis fascicularis) in Singapore

Crab-eating, or long-tailed, macaques [ Macaca fascicularis (Raffles, 1821)] have been studied extensively throughout their distribution in South and South-east Asia. Despite this extensive body of research, the island population of long-tailed macaques from Singapore remains virtually undescribed. In the present study, we compare the morphometric variability and patterns of growth observed in a population sample from Singapore with a composite sample from Thailand, north of the Isthmus of Kra. The results of our analyses indicate that there are statistically significant differences between the two populations in adult size and shape. For both males and females, the Singapore population is smaller than the Thai population. Relative to body length, the Singapore macaques exhibit significantly longer tails, and, relative to cranial length, they exhibit significantly more narrow faces than the Thai macaques. Although levels of sexual dimorphism for most morphometric traits are very similar, indicating similar levels of male–male competition for females, the Singapore males exhibit a significantly larger testicular volume relative to body weight, suggestive of an alternative male reproductive strategy. In addition to adult somatometric size and shape, comparisons of growth patterns relative to age and body size reveal significant differences between the two population samples. Combined, these results suggest either that statistically significant differences in adult morphology and patterns of growth can occur in presumably reproductively cohesive subspecies, or the Singapore macaques may be taxonomically distinct.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 675–694.     

Direct and indirect selection in moth pheromone evolution: population genetical simulations of asymmetric sexual interactions

Female moths generally use pheromones to attract males. Normally, all females in a population produce a specific chemical blend with only a limited variance, and the local males are highly attracted to this blend. To better understand the direct and indirect selective forces acting on this communication system, where, unusually, it is the reproductively limited sex that signals for matings, a population genetical model has been constructed and numerically analysed. Basic to the model is the assumption that the pheromone attraction system functions asymmetrically, leading to strong sexual selection between males but no direct sexual selection between females. Evolutionary simulations using the model show that sexual selection in males causes an indirect stabilizing selection on the pheromone blends produced by females. Thus, a more narrow range of pheromone variation is selected for, even in the absence of female sexual selection. The strength of the selection is analysed, and it is suggested that this indirect stabilizing selection becomes particularly important in situations where geographically adjacent populations have evolved different pheromone blends.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 117–123.     

Extreme ultraviolet sexual dimorphism in jumping spiders (Araneae: Salticidae)

Jumping spiders (Salticidae) have acute vision with some cells in the retina that are sensitive to ultraviolet (UV) spectra (< 400 nm). However, no study has documented the use of UV signals in salticids. To appreciate the function of UV vision, it is necessary to characterize the UV colours of salticids. In the present study, the UV and human-visible wavelengths of a tropical ornate salticid spider, Cosmophasis umbratica , were analysed using reflectance spectrometry to obtain evidence of sex-specific UV colours. An absolute sexual dimorphism in the UV colours of this salticid species was found. All of the body parts of adult males that are displayed to conspecifics during intra-specific interactions reflected UV (300–400 nm) light, whereas the adult females and juveniles did not reflect UV light from any body part. A great deal of variation was also found in the UV wavebands among males. This is the first full UV characterization of a salticid spider and the first study to demonstrate an extreme sexual UV dimorphism in jumping spiders. The findings obtained provide evidence that UV reflectance may comprise important sexual signals in jumping spiders.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 397–406.     

Male-biased size dimorphism in ichneumonine wasps (Hymenoptera: Ichneumonidae) – the role of sexual selection for large male size

Abstract.  1. Sexual differences in body size are expected to evolve when selection on female and male sizes favours different optima.
2. Insects have typically female-biased size dimorphism that is usually explained by the strong fecundity advantage of larger size in females. However, numerous exceptions to this general pattern have led to the search for selective pressures favouring larger size in males.
3. In this study, the benefits of large size were investigated in males of four species of ichneumonine wasps, a species-rich group of parasitoids, many representatives of which exhibit male-biased size dimorphism.
4. Mating behaviour of all ichneumonine wasps are characterised by pre-copulatory struggles, in the course of which males attempt to override female reluctance to mate. A series of laboratory trials was conducted to study the determinants of male mating success.
5. A tendency was found for larger males as well as those in better condition to be more successful in achieving copulations. Size dimorphism of the species studied, mostly male-biased in hind tibia length but female-biased in body weight, indicates that sexual selection in males favours longer bodies and appendages rather than larger weight.
6. The qualitative similarity of the mating patterns suggests that sexual selection cannot completely explain the considerable among-species differences in sexual size dimorphism.
7. The present study cautions against using various size indices as equivalents for calculating sexual size dimorphism.
8. It is suggested that female reluctance in ichneumonine wasps functions as a mechanism of female mate assessment.