Sexual dimorphisms in the dermal structure of the lesser‐spotted catshark,Scyliorhinus canicula (Linnaeus, 1758)

Sexual dimorphisms in the dermal structures of two elasmobranch species have previously been reported and it has been linked to the use of the mouth by males during copulation. Until relatively recently, the fact, that male Scyliorhinus canicula use their mouths for grasping and biting females during copulation was unknown. This study reveals that not only do adult (M ≥ 525 mm, F ≥ 550 mm) S. canicula show a sexual dimorphism in the epidermis and dermis, but that hatchling S. canicula are born with a sexually dimorphic epidermal layer and this persists into the juvenile stage (M < 525 mm, F < 550 mm). A sexual dimorphism was found in all size classes with both hatchling and juvenile female S. canicula having significantly thicker epidermal layers than hatchling and juvenile male S. canicula. Adult female S. canicula were found to possess both a significantly thicker epidermal and dermal layer than adult male S. canicula. The presence of a sexual dimorphism in the epidermal and dermal layers of adult S. canicula could be directly related to reproductive behaviour in response to the male biting the female prior to copulation.     

Sex‐specific repeatabilities and effects of relatedness and mating status on copulation duration in an acridid grasshopper

In species with direct sperm transfer, copulation duration is a crucial trait that may affect male and female reproductive success and that may vary with the quality of the mating partner. Furthermore, traits such as copulation duration represent the outcome of behavioral interactions between the sexes, for which it is important—but often difficult—to determine which sex is in phenotypic control. Using a double‐mating protocol, we compared copulation durations between (1) virgin and nonvirgin and (2) sibling and nonsibling mating pairs in rufous grasshoppers Gomphocerippus rufus. Nonvirgin copulations took on average approximately 30% longer than virgin copulations, whereas relatedness of mating partners was not a significant predictor of copulation duration. Longer nonvirgin copulations may represent a male adaptation to sperm competition if longer copulations allow more sperm to be transferred or function as postinsemination mate guarding. The absence of differences between pairs with different degrees of relatedness suggests no precopulatory or preinsemination inbreeding avoidance mechanism has evolved in this species, perhaps because there is no inbreeding depression in this species, or because inbreeding avoidance occurs after copulation. Controlling for the effects of male and female mating status (virgin vs. nonvirgin) and relatedness (sibling vs. nonsibling), we found significant repeatabilities (R) in copulation duration for males (R = 0.33; 95% CI: 0.09–0.55) but not for females (R = 0.09; 95% CI: 0.00–0.30). Thus, copulation durations of males more strongly represent a nontransient trait expressed in a consistent manner with different mating partners, suggesting that some aspect of the male phenotype may determine copulation duration in this species. However, overlapping confidence intervals for our sex‐specific repeatability estimates indicate that higher sampling effort is required for conclusive evidence.     

Exaggerated male genitalia intensify interspecific reproductive interference by damaging heterospecific female genitalia

Male–male competition over fertilization can select for harmful male genital structures that reduce the fitness of their mates, if the structures increase the male's fertilization success. During secondary contact between two allopatrically formed, closely related species, harmful male genitalia may also reduce the fitness of heterospecific females given interspecific copulation. We performed a laboratory experiment to determine whether the extent of genital spine exaggeration in Callosobruchus chinensis males affects the fitness of C. maculatus females by injuring their reproductive organs. We found that males with more exaggerated genital spines were more likely to injure the females via interspecific copulation and that the genital injury translated into fecundity loss. Thus, as predicted, reproductive interference by C. chinensis males on C. maculatus females is mediated by exaggeration of the genital spine, which is the evolutionary consequence of intraspecific male–male competition. Harmful male traits, such as genital spines, might generally affect the extent of interaction between closely related species.     

Male mate choice in the chameleon grasshopper (Kosciuscola tristis)

In many species, males can increase their fitness by mating with the highest quality females. Female quality can be indicated by cues, such as body size, age and mating status. In the alpine grasshopper Kosciuscola tristis, males can be found riding on subadult females early in the season, and as the season progresses, males engage in fights over ovipositing females. These observations suggest that males may be competing for females that are either unmated (early season) or sperm‐depleted (late season). We thus hypothesised that male K. tristis may be choosy in relation to female mating status, and specifically, we predicted that males prefer females that are unmated. We conducted behavioural experiments in which males were given the choice of two females, one mated and one unmated. Contrary to our prediction, males did not mate preferentially with unmated females. However, copulation duration with unmated females was, on average, 24 times the length of copulation with mated females. While female K. tristis can reject mates, we did not observe any evidence of overt female choice during our trials. Females may gain additional benefits from mating multiply and may therefore not readily reject males. While our experiment cannot definitively disentangle female from male control over copulation duration, we suggest that males choose to invest more time in copula with unmated females, perhaps for paternity assurance, and that male mate assessment occurs during copulation rather than beforehand.     

Mating Compatibility between Geographic Populations of the Seed Beetle <Emphasis Type="Italic">Callosobruchus maculatus</Emphasis>

Allopatric populations sometimes display reproductive barriers in incipient form, and may thus reveal potential mechanisms of speciation. We conducted mating trials to estimate the degree of precopulatory (behavioral) and postcopulatory (gametic) prezygotic isolation between African and Asian populations of the seed beetle Callosobruchus maculatus. The populations differ in several fitness-related traits, and have been associated with different legume hosts. In single-pair trials, the probability of copulation within 10 min was lower when Asian females were paired with African males than it was in the other three pairing combinations. This pairing also provided evidence for gametic isolation, as Asian females mated once to African males laid fewer eggs than did females in the other treatments. Variation in fecundity could not be explained by the duration of copulation, which did not differ among pairing combinations. There was also no effect of pairing combination on the number of copulation attempts, latency to copulation, or rate of egg hatch. When females were simultaneously presented a male from each population, Asian males obtained a disproportionate share of the matings, and were more likely to disturb a pair already in copulo. Surprisingly, male size did not influence the probability of copulation in the single-male trials, the outcome of male–male competition, or the fecundity of once-mated females. Although the African and Asian populations showed some evidence of mating incompatibility, the absence of symmetrical isolating factors suggests minimal barriers to gene flow.     

Ultrasonic courtship song of the yellow peach moth, <Emphasis Type="Italic">Conogethes punctiferalis</Emphasis> (Lepidoptera: Crambidae)

Although generation of ultrasound during courtship has been reported for an increasing number of moth species, the effect of the ultrasound on mating remains uncertain in many cases because of a lack of proper verification. Here we report that males of the yellow peach moth Conogethes punctiferalis (Crambidae) sexually communicate with females by emitting loud ultrasound (103 dB peak equivalent sound pressure level at 1 cm; dominant frequency 82 kHz) before attempting copulation. The male ultrasound consists of consecutive clicks (pulses) in the early phase of the sound train and consecutive pulses (burst) in the late phase. When females were deafened by puncturing the abdominal tympanic membranes, copulation never occurred. We found that deafened females did not assume the wing-raising posture, which, for normal pairs, always precedes successful copulation. Our findings indicate that male courtship ultrasound evokes wing-raising as an acceptance behavior from females, which in turn evokes a copulation attempt by a male.     

Mating-induced reduction in accessory reproductive organ size in the stalk-eyed fly <Emphasis Type="Italic">Cyrtodiopsis dalmanni</Emphasis>

Background  

Internal reproductive organ size is an important determinant of male reproductive success. While the response of testis length to variation in the intensity of sperm competition is well documented across many taxa, few studies address the importance of testis size in determining other components of male reproductive success (such as mating frequency) or the significance of size variation in accessory reproductive organs. Accessory gland length, but not testis length, is both phenotypically and genetically correlated with male mating frequency in the stalk-eyed fly Cyrtodiopsis dalmanni. Here we directly manipulate male mating status to investigate the effect of copulation on the size of both the testes and the accessory glands of C. dalmanni.     

Microscale Laser Surgery Demonstrates the Grasping Function of the Male Sex Combs in Drosophila melanogaster and Drosophila bipectinata

Male secondary sexual traits of animals are richly diversified in form and complexity, yet there are many species in which their precise function remains unknown. Within the genus Drosophila, species belonging to the melanogaster and obscura species groups have evolved a remarkable variety of sex combs, male‐limited secondary sexual traits located on the tarsi of both front legs. Information concerning sex comb function is minimal or absent, except for D. melanogaster, where previous studies indicate that the sex combs are used for grasping the female prior to copulation. These studies, however, do not unambiguously demonstrate comb function, because it has not been possible to ascribe observed behavioral outcomes of the various comb manipulations to changes in the combs per se. We used microscale laser surgery to manipulate comb size in D. melanogaster and D. bipectinata, and tested the hypothesis that the sex combs function as grasping devices in courtship, making them essential for copulation to ensue. Results of high‐resolution behavioral analysis in small observation arenas demonstrated that in both species in which sex combs were surgically eliminated, males were unable to grasp, mount or copulate. The combless foretarsi of these altered males slipped off the end (D. melanogaster) and sides (D. bipectinata) of the female abdomen when courting males attempted to grasp. In most cases, males whose sex combs were reduced but not completely removed exhibited similar copulation probabilities as surgical control males, a result we demonstrated in observation chambers as well as under more ecologically realistic conditions inside population cages where males and females interacted on the surface of fruit substrates. Thus, the sex combs in D. melanogaster and D. bipectinata are grasping devices, essential for mounting and copulation.     

Larval aggregations of the blister beetle Stenoria analis (Schaum) (Coleoptera: Meloidae) sexually deceive patrolling males of their host,the solitary bee Colletes hederae Schmidt & Westrich (Hymenoptera: Colletidae)

We here report on observations on the parasitism of the solitary bee Colletes hederae Schmidt & Westrich (Hymenoptera: Colletidae) by triungulins (first instar larvae) of the European beetle Stenoria analis (Schaum) (Coleoptera: Meloidae). Our observations carried out in western France provide behavioural evidence that patrolling males of C. hederae hover in front of, and are strongly drawn to, larval aggregations of S. analis in a manner reminiscent of approaching flights to emerging, conspecific females. Not only were the male bees strongly attracted to clusters of newly hatched triungulins, but they also attempted copulation (i.e., pseudocopulation) with the latter, which regularly resulted in small groups of triungulins being transferred onto the thorax of the male bees. Decent-sized groups of triungulins were exclusively found on the thorax of male bees, which suggests that triungulins of S. analis lure only males of the targeted host species, presumably by mimicking the female pheromonal cues of C. hederae. This is to our knowledge the first record of this kind of interaction for the whole West-Palaearctic, and only the second account on sexual deception between blister beetles and wild bees described to date.     

Geometric characters of the radius and tibia in <Emphasis Type="Italic">Macaca mulatta</Emphasis> and <Emphasis Type="Italic">Macaca fascicularis</Emphasis>

We performed comparative analyses of four cross-sections of the distal radius and tibia in two species of macaque to clarify the relationships between bone morphology and locomotor type. The lengths of bones and five bone geometric properties in each section were examined and compared separately in both female and male Macaca mulatta and Macaca fascicularis. In M. mulatta, there were no significant gender-specific differences in either the radius or the tibia. In contrast, the radius and tibia of male M. fascicularis had greater geometric parameters in the 20% and 40% positions relative to the 5% and 10% positions from the distal end than those of their female counterparts. The radius and tibia of M. mulatta were relatively longer than those of M. fascicularis, and the sectional parameters of the tibia of M. mulatta were relatively larger than those of M. fascicularis. Standardization of the log-transformed bone length between the species revealed larger radial cortical bone areas in M. fascicularis. In contrast, there were minimal differences in the tibial cortical bone areas between the two species. This study suggests that the observed distinctions in bone geometry in female and male M. fascicularis may be due to gender-specific differences in the muscle weights of the forearm and calf, which may underlie the divergence in the leaping abilities of females and males of this species. Taken together, these results of interspecies comparisons may be related to the fact that arboreal primates such as M. fascicularis undergo compressive mechanical stress due to the forelimb lead that occurs as the animal descends a sloping trunk or bridges a tree gap downward, while terrestrial primates such as M. mulatta move on nearly flat substrates. Differences in fore- and hind-limb bone properties between the two species are discussed with regard to functional morphology and locomotor type.     

Patterns of reproductive isolation within and between two Lygaeus species characterized by sexual conflicts over mating

Theory suggests that, under some circumstances, sexual conflict over mating can lead to divergent sexually antagonistic coevolution among populations for traits associated with mating, and that this can promote reproductive isolation and hence speciation. However, sexual conflict over mating may also select for traits (e.g. male willingness to mate) that enhance gene flow between populations, limiting population divergence. In the present study, we compare pre‐ and post‐mating isolation within and between two species characterized by male–female conflict over mating rate. We quantify sexual isolation among five populations of the seed bug Lygaeus equestris collected from Italy and Sweden, and two replicates of a population of the sister‐species Lygaeus simulans, also collected from Italy. We find no evidence of reproductive isolation amongst populations of L. equestris, suggesting that sexual conflict over mating has not led to population divergence in relevant mating traits in L. equestris. However, there was strong asymmetric pre‐mating isolation between L. equestris and L. simulans: male L. simulans were able to mate successfully with female L. equestris, whereas male L. equestris were largely unable to mate with female L. simulans. We found little evidence for strong post‐mating isolation between the two species, however, with hybrid F₂ offspring being produced. Our results suggest that sexual conflict over mating has not led to population divergence, and indeed perhaps supports the contrary theoretical prediction that male willingness to mate may retard speciation by promoting gene flow.     

Origin of the white shark Carcharodon (Lamniformes: Lamnidae) based on recalibration of the Upper Neogene Pisco Formation of Peru

Abstract: The taxonomic origin of the white shark, Carcharodon, is a highly debated subject. New fossil evidence presented in this study suggests that the genus is derived from the broad‐toothed ‘mako’, Carcharodon (Cosmopolitodus) hastalis, and includes the new species C. hubbelli sp. nov. – a taxon that demonstrates a transition between C. hastalis and Carcharodon carcharias. Specimens from the Pisco Formation clearly demonstrate an evolutionary mosaic of characters of both recent C. carcharias and fossil C. hastalis. Characters diagnostic to C. carcharias include the presence tooth serrations and a symmetrical first upper anterior tooth that is the largest in the tooth row, while those indicative of C. hastalis include a mesially slanted third anterior (intermediate) tooth. We also provide a recalibration of critical fossil horizons within the Pisco Formation, Peru using zircon U‐Pb dating and strontium‐ratio isotopic analysis. The recalibration of the absolute dates suggests that Carcharodon hubbelli sp. nov. is Late Miocene (6–8 Ma) in age. This research revises and elucidates lamnid shark evolution based on the calibration of the Neogene Pisco Formation.     

The role of functional constraints in nonrandom mating patterns for a dance fly with female ornaments

Most hypotheses to explain nonrandom mating patterns invoke mate choice, particularly in species that display elaborate ornaments. However, conflicting selection pressures on traits can result in functional constraints that can also cause nonrandom mating patterns. We tested for functional load‐lifting constraints during aerial copulation in Rhamphomyia longicauda, a species of dance fly that displays multiple extravagant female‐specific ornaments that are unusual among sexual traits because they are under stabilizing selection. R. longicauda males provide females with a nuptial gift before engaging in aerial mating, and the male bears the entire weight of the female and nuptial gift for the duration of copulation. In theory, a male's ability to carry females and nuptial gifts could constrain pairing opportunities for the heaviest females, as reported for nonornamented dance flies. In concert with directional preferences for large females with mature eggs, such a load‐lifting constraint could produce the stabilizing selection on female size previously observed in this species. We therefore tested whether wild‐caught male R. longicauda collected during copulation were experiencing load‐lift limitations by comparing the mass carried by males during copulation with the male's wing loading traits. We also performed permutation tests to determine whether the loads carried by males during copulation were lighter than expected. We found that heavier males are more often found mating with heavier females suggesting that whereas R. longicauda males do not experience a load‐lift constraint, there is a strong relationship of assortative mating by mass. We suggest that active male mate choice for intermediately adorned females is more likely to be causing the nonrandom mating patterns observed in R. longicauda.     

Structural evidence why males of Panorpa liui offer prey rather than salivary mass as their nuptial gift

Ma, N. and Hua, B. 2010. Structural evidence why males of Panorpa liui offer prey rather than salivary mass as their nuptial gift. —Acta Zoologica (Stockholm) 92 : 398–403. The scorpionflies are considered as ideal model animals for the study of mating systems in insects. The males generally offer both prey and salivary mass as nuptial gifts to the females during copulation. Our field observations show that Panorpa liui is peculiar because the males offer only prey rather than salivary secretions as nuptial gift. Through anatomical and histological examinations, the salivary glands of P. liui were found to be devoid of sexual dimorphism in Panorpa for the first time. Both the male and female P. liui bear simple salivary glands, which are only composed of a common duct and two short sac‐like glands. This is the first attempt to explore the relationship between the salivary glands and the mating tactics from the structural aspect in Panorpa, speculating that the simple structure of the male salivary glands in P. liui might be responsible for its failing to produce salivary mass as a nuptial gift during copulation. Compared with Boreidae, Meropidae, Bittacidae and Panorpidae, we presume that the absence of sexual dimorphism of the salivary glands might represent a plesiomorphy in P. liui. The origin and evolutionary process of the nuptial gift behaviour are tentatively speculated in Panorpa.     

Non-parental infanticide in a dense population of the Black-billed Magpie (<Emphasis Type="Italic">Pica pica</Emphasis>)

We report the first case of non-parental infanticide in the Black-billed Magpie Pica pica. Using a video camera installed in the victims’ nest, we recorded repeated visits (over 4 days) of an adult (each time one bird) who attacked six nestlings at each visit until they died or were evicted. The nest was one of 58 nests filmed over four breeding seasons. Collected evidence suggests that the perpetrator(s) might have been the female breeder of the neighbouring nest, possibly also her male partner. The parental female aggressively attacked the perpetrator. Post-infanticide expansion of breeding territory by the suspected perpetrator is the hypothetical ultimate explanation of the observed infanticide. Movie clips from inside the nest are shown at: , , , and .     

A six-year study of fire-related flowering cues and coexistence of two perennial grasses in a wet longleaf pine (<Emphasis Type="Italic">Pinus palustris</Emphasis>) savanna

Integrating population and community ecology can improve our understanding of the impacts of natural disturbances. Fire-stimulated flowering occurs in many long-lived herbaceous species of fire-maintained grasslands and savannas. Coexistence of these long-lived species may be affected in part by interspecific differences in the effect of fire-stimulated flowering on resource conservation, clonal growth, and longevity. This study comprised two parts. The first part investigated the effectiveness of different fire-related cues on fire-stimulated flowering in two cooccurring dominant grass species in a wet longleaf pine (Pinus palustris) savanna in southeastern Mississippi, USA. The second part investigated the immediate effects of the most important of these cues (damage and removal of aboveground vegetation and surface litter in May) along with nutrient addition on several measures of fitness and abundance over 6 years. Despite being a very effective inductive cue, when repeated frequently over 6 years, clipping plus surface litter removal significantly reduced flowering in both species. This negative effect was reduced to some extent by nutrient addition in Muhlenbergia expansa (which exhibited higher reproductive investment following clipping and fire than did Ctenium aromaticum). Frequent clipping resulted in an increasing numerical advantage of C. aromaticum over M. expansa with time. There was evidence of a modest release of C. aromaticum from competition with M. expansa in response to annual clipping. Responses suggest that suppressing flowering until after fire reduces the cost of flowering and maintains shoot densities, at least in M. expansa. Differences in the responses of these two species to repeated clipping and nutrient addition suggest that, despite their both exhibiting fire-stimulated flowering, each species is favored by slightly different fire frequencies. Moderate variation in fire frequency could maintain their coexistence in the long term.     

Patterns of Sperm Transfer in the Golden Orb‐Weaver Nephila edulis

Sperm competition studies typically identify copulation duration as an important predictor of paternity as it may determine the quantity of sperm transferred and thus paternity success. This study explores the relationship between copulation duration, male body size, male age and sperm transfer in the golden orb‐weaving spider, Nephila edulis. Paternity in this species is strongly associated with the relative frequency and duration of copulation, which is also influenced by male size. We determined the number of sperm transferred during copulation, by performing sperm counts in both the male copulatory organs (palps) and female sperm receptacle (spermatheca) of recently mated pairs. The total number of sperm recorded (the sum in the male palps and female spermathecae) was greater for younger males than older males, but did not vary with male body size. In general, younger males transferred more sperm and a greater proportion of their sperm supplies than older males and, among these younger males, larger individuals transferred more sperm. However, there were no significant size effects for older males. More sperm was transferred with longer copulations, but in contrast with previous studies, we found that larger males copulated for longer. The rate of sperm transferred was negatively correlated with the duration of copulation, suggesting that the variation in copulation duration in N. edulis may represent strategic investment by males to alter patterns of paternity, in addition to transferring additional sperm.     

Unique mating behavior,and reproductive biology of a simultaneous hermaphroditic marine flatworm (Platyhelminthes,Tricladida, Maricola)

Flatworms generally are simultaneous hermaphrodites that exhibit various kinds of mating behavior. Here we report on the mating behavior and reproductive biology of the planarian Paucumara falcata. We recognized three phases in its mating behavior: a courtship, copulation, and postcopulatory phase. During the last‐mentioned phase, the partners showed a unique and very characteristic behavior in which their bodies intertwined, forming a spiral. Histological study of partners in copula revealed that the sclerotic tip of the musculo‐parenchymatic organ pierces the body wall of the partner and then becomes lodged in its parenchyma, suggesting that this organ may act as an anchor, thus stabilizing the worms during copulation. Similar organs in other species of marine triclad may also perform a stabilizing role during copulation. During copulation in individuals of P. falcata, sperm transfer was reciprocal or only unilateral. Copulation duration ranged 13–35 min (average 20 ± 5 min), irrespective of whether the mating was successful (i.e., resulted in the production of fertile cocoons). The spiraling phase lasted on average 10 min; some worms did not show the postcopulatory spiraling phase during their mating behavior. After successful copulation, an individual worm produced 1–12 fertile cocoons over a period of 1–17 days; from a cocoon hatched either one young (in 70% of the cases), or two young worms.     

Molecular and morphological evidences reveal a cryptic species in the Vinaceous Rosefinch Carpodacus vinaceus (Fringillidae; Aves)

Wu, H.‐C., Lin, R.‐C., Hung, H.‐Y., Yeh, C.‐F., Chu, J.‐H., Yang, X.‐J., Yao, C.‐J., Zou, F.‐S., Yao, C.‐T., Li, S.‐H. & Lei, F.‐M. (2011). Molecular and morphological evidences reveal a cryptic species in the Vinaceous Rosefinch Carpodacus vinaceus (Fringillidae; Aves). —Zoologica Scripta, 40, 468–478. The Vinaceous Rosefinch (Carpodacus vinaceus) is endemic in East Asia with two recognized subspecies –C. v. vinaceus, distributed along the eastern edge of the Tibetan Plateau and the Himalayas, and C. v. formosanus, restricted to Taiwan’s Central Mountain Range. As reflected in a controversial taxonomic history, this vastly disjunctive distribution pattern suggests that the subspecies, having been isolated from each other for a long time, might have diverged, challenging the current taxonomic treatment and calling for possible species delimitation. Sequences of two mitochondrial fragments (mtDNA) and two Z‐linked nuclear loci (zDNA) were used to reconstruct the intraspecific phylogeny of C. vinaceous. The mtDNA tree shows that the two subspecies of the vinaceous rosefinch form two exclusively monophyletic clades. All but one zDNA sequences from the nominate subspecies and C. v. formosanus also formed exclusively monophyletic clades (the exceptional zDNA sequence from C. v. vinaceous formed a weakly supported clade with two outgroup species). Moreover, by conducting quantitative comparisons of morphometric traits and male plumage coloration, we found that the two subspecies exhibit distinguishable morphological differences. All the evidence therefore suggests that C. v. formosanus is a cryptic species and that its taxonomic status should be restored to full species. Molecular dating suggests that the two sibling rosefinches split 1.7 ± 0.2 million years ago, providing a point estimate for the historical connectivity of biota between eastern Tibet‐Himalayas and montane Taiwan.     

Body Size and Morph as Drivers of Copulation Duration in a Male Dimorphic Damselfly

Copulation duration is often highly variable within and among species. Here, we explore the roles of body size, male morph, morph frequency, and alternative reproductive tactics to explain copulation duration in the damselfly Paraphlebia zoe. P. zoe has two male morphs (pigmented or hyaline wings) which differ in reproductive tactics (territorial or non‐territorial behaviors). We also analyze the effects of season as the frequencies of both morphs tend to vary along the reproductive season. In the first non‐experimental year, we found that the relationship between body size and copulation duration depended on the time of year. Early in the season, body size positively correlated with copulation duration, while late in the year, body size negatively correlated with copulation duration. In the second experimental year (when we reversed the frequency of male morphs in the middle of the season: making pigmented males less frequent than hyaline males), size influenced copulation duration as well as morph – body size positively correlated with copulation duration, and hyaline males mated for longer than pigmented males. Contrary to our prediction, changes to the relative abundances of morphs did not influence copulation duration. Hyaline males may be under selection for longer copulation durations to compensate for their reduced access to females, as long copulations potentially lead to more rival sperm to be removed from the female sperm storage organs and/or increased mate guarding. We do not discard, however, other explanations that drive variation in copulation duration such as cryptic female choice and/or predation.