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Sex allocation theory in simultaneous hermaphrodites predicts that optimal sex allocation is influenced by local sperm competition, which occurs when related sperm compete to fertilize a given set of eggs. Different factors, including the mating strategy and the ability to self-fertilize, are predicted to affect local sperm competition and hence the optimal SA. Moreover, since the local sperm competition experienced by an individual can vary temporally and spatially, this can favour the evolution of sex allocation plasticity. Here, using seven species of the free-living flatworm genus Macrostomum, we document interspecific variation in sex allocation, but neither their mating strategy nor their ability to self-fertilize significantly predicted sex allocation among these species. Since we also found interspecific variation in sex allocation plasticity, we further estimated standardized effect sizes for plasticity in response to (i) the presence of mating partners (i.e. in isolation vs. with partners) and (ii) the strength of local sperm competition (i.e. in small vs. large groups). We found that self-fertilization predicted sex allocation plasticity with respect to the presence of mating partners, with plasticity being lower for self-fertilizing species. Finally, we showed that interspecific variation in sex allocation is higher than intraspecific variation due to sex allocation plasticity. Our study suggests that both sex allocation and sex allocation plasticity are evolutionarily labile, with self-fertilization predicting the latter in Macrostomum.

Sperm competition drives traits that enhance fertilization success. The amount of sperm transferred relative to competitors is key for attaining paternity. Female reproductive morphology and male mating order may also influence fertilization, however the outcome for sperm precedence under intense sperm competition remains poorly understood. In the polyandrous spider Pisaura mirabilis, males offer nuptial gifts which prolong copulation and increase sperm transfer, factors proposed to alter sperm precedence patterns under strong sperm competition. First, we assessed the degree of female polyandry by genotyping wild broods. A conservative analysis identified up to four sires, with a mean of two sires per brood, consistent with an optimal mating female rate. Then we asked whether intense sperm competition shifts sperm precedence patterns from first male priority, as expected from female morphology, to last male advantage. We varied sexual selection intensity experimentally and determined competitive fertilization outcome by genotyping broods. In double matings, one male monopolised paternity regardless of mating order. A mating order effect with first male priority was revealed when females were mated to four males, however this effect disappeared when females were mated to six males, probably due to increased sperm mixing. The proportion of males that successfully sired offspring drastically decreased with the number of competitors. Longer copulations translated into higher paternity shares independently of mating order, reinforcing the advantage of traits that prolong copulation duration under intense competition, such as the nuptial gift. Sperm competition intensity enhances the impact of competitive sexual traits and imposes multiple effects on paternity.

Whenever resources are limited and indivisible, fighting will evolve as a means to resolve ownership. Among such resources are mates, and individuals (usually males) of many species compete agonistically with rivals in order to gain access to potential mates. However, securing access is not necessarily enough to guarantee a mating or, if a mating is obtained, to guarantee that it is effective for securing reproductive success. Thus, in addition to fighting, individuals participate in a wealth of behaviours to maximize their reproductive success, from courtship to sperm competition to mate guarding. In recent years, the striking parallels between fighting and mating behaviour have become a subject of discussion. In particular, insights have been drawn from the predictions of contest theory to help us understand the use of repetitive signalling in courtship. Here, we take this discussion further, highlighting similarities between fighting and mating in the use of dynamic repeated behaviours, which function to (1) advertise quality and (2) convince or coerce an individual to relinquish the contested resource (gametes in terms of mating). We focus specifically on a performance trait of emerging interest in the field of animal contests, skill. We identify behaviours used throughout the mating process in which skill is likely to be of importance for securing success, and highlight key questions for future study.

When competition between males for mates is intense it is common to find that some males will adopt alternative tactics for acquiring fertilisations, often involving the use of sneak tactics whereby males avoid contests. These alternative tactics are sometimes associated with discrete differences in male morphology, with sneak males investing less in weaponry but more in traits such as testes which will give an advantage in sperm competition. In some cases it appears that males develop into more than two morphs, with a number of examples of tri- and even tetramorphic arthropod species being described. Here we analyse the scaling relations of the dung beetle species Proagoderus watanabei, which expresses two distinct weapon traits: paired head horns and a pronotal horn. We find that males of this species are trimorphic, with alpha males expressing long head horns and a pronotal horn, beta males with long head horns but no pronotal horn, and gamma males with short head horns only. We also find that alpha males invest less in testes than do beta or gamma males, indicating that beta and gamma males in this species probably experience higher risks of sperm competition than do alphas.

Australian lizards are a diverse group distributed across the continent and inhabiting a wide range of environments. Together, they exhibit a remarkable diversity of reproductive morphologies, physiologies, and behaviours that is broadly representative of vertebrates in general. Many reproductive traits exhibited by Australian lizards have evolved independently in multiple lizard lineages, including sociality, complex signalling and mating systems, viviparity, and temperature-dependent sex determination. Australian lizards are thus outstanding model organisms for testing hypotheses about how reproductive traits function and evolve, and they provide an important basis of comparison with other animals that exhibit similar traits. We review how research on Australian lizard reproduction has contributed to answering broader evolutionary and ecological questions that apply to animals in general. We focus on reproductive traits, processes, and strategies that are important areas of current research, including behaviours and signalling involved in courtship; mechanisms involved in mating, egg production, and sperm competition; nesting and gestation; sex determination; and finally, birth in viviparous species. We use our review to identify important questions that emerge from an understanding of this body of research when considered holistically. Finally, we identify additional research questions within each topic that Australian lizards are well suited for reproductive biologists to address.

Female and male reproductive traits co-evolve through pre- and post-copulatory sexual selection and sexual conflict. Although males typically transfer many sperm during copulation, only a small proportion reach the fertilization site because females often actively or passively reduce sperm number in their reproductive tract. Males may transfer accessory substances to protect their ejaculates against female selective processes, which benefits males but can harm females. In turn, females may use accessory gland fluids to control paternity or sperm storage. Female yellow dung flies (Scathophaga stercoraria) have paired accessory glands that produce fluids involved in fertilization and egg laying. One proposed function for these fluids is spermicide. Alternatively, female accessory gland fluid may help keep sperm alive to avoid fertilization failure or encourage sperm competition. Using yellow dung flies, we investigated the interaction of female accessory gland fluid with sperm in vitro. Significantly more sperm remained alive when exposed to accessory gland fluid compared to buffer only (63% vs. 44%). We conclude that female accessory gland fluid in yellow dung flies can help nourish rather than kill male sperm, although selective nourishment of sperm is as consistent with cryptic female choice as is selective spermicide.

Although it is widely stated that both mating behavior and sperm traits are energetically costly for males, we currently lack empirical estimates of the relative costs to males of pre- versus postcopulatory investments. Such estimates require the experimental separation of the act of mating from that of ejaculation, which is a nontrivial logistical challenge. Here, we overcome this challenge using a novel morphological manipulation (gonopodium tip ablation) in the eastern mosquitofish (Gambusia holbrooki) to tease apart investment in mating effort from that in sperm replenishment following ejaculation. We quantified the relative cumulative costs of investing in mating effort and ejaculation by comparing somatic traits and reproductive performance among three types of males: ablated males that could attempt to mate but not ejaculate; unablated males that could both mate and ejaculate; and control males that had no access to females. We show that, after eight weeks, mating investment significantly reduces both body growth and immunocompetence and results in a significant decline in mating effort. In contrast, cumulative investment into sperm replenishment following ejaculation has few detectable effects that are only apparent in smaller males. These minor costs occur despite the fact that G. holbrooki has very high levels of sperm competition and multiple mating by both sexes, which is usually associated with elevated levels of sperm production. Crucially, our study is the first, to our knowledge, to experimentally compare the relative costs of pre- and postcopulatory investment on components of male fitness in a vertebrate.

The level of male-male competition is an important dimension of the male social niche. When males provide parental care, they potentially face a trade-off between obtaining additional fertilizations and providing care to their offspring. In this study, we will investigate how males plastically adjust their reproductive phenotype, specifically their provision of parental care, depending on the opportunity for extra-pair mating and/or level of sperm competition risk. To this aim, we will experimentally manipulate the social environment in which males are breeding. The study complements a previous, preregistered experiment focusing on male phenotypic adjustment of competitive traits in response to different levels of sperm competition risk (see Lilie, N. D., Riyahi, S., Korsten, P., & Schmoll, T., 2019, July 11. 'Male social niche conformance in zebra finches'. Retrieved from osf.io/84z5r). In the current preregistered study, we will expose males of a well-characterized model species, the socially monogamous zebra finch, to two social treatment levels: double-pair housing, inducing an opportunity for extra-pair mating and elevated sperm competition risk, versus single-pair housing with no opportunity for extra-pair mating or sperm competition risk. We will measure the male phenotypic responses to these different social treatments focusing on their contributions to parental care and their plasma levels of the hormones testosterone and corticosterone. In addition, we will quantify the effects of the social treatment and its resulting male responses on offspring growth and survival.

Polyandry, common in mammalian world, results in sperm competition between males. In order to improve chances of siring offspring, males can use different strategies enhancing success in the competition. One of them is adjusting the copulation time to the most favorable conditions relative to ovulation in female. The aim of current study is to examine preferences of male golden hamsters towards females in different stages of oestrous. The predictions are that males would spend more time near the female, which is in more attractive stage of oestrus. Methods, that will be used to test the hypotheses include assessment of vaginal cytology to identify oestrous stage in females, and male preference tests, carried out in Y-maze.

Evolutionary biologists have endeavoured to explain the extraordinary diversity of sperm morphology across animals for more than a century. One hypothesis to explain sperm diversity is that sperm length is shaped by the environment where fertilization takes place (that is, fertilization mode). Evolutionary transitions in fertilization modes may transform how selection acts on sperm length, probably by affecting postcopulatory mechanisms of sperm competition and the scope for cryptic female choice. Here, we address this hypothesis by generating a macro-evolutionary view of how fertilization mode (including external fertilizers, internal fertilizers and spermcasters) influences sperm length diversification among 3,233 species from 21 animal phyla. We show that sperm are shorter in species whose sperm are diluted in aquatic environments (that is, external fertilizers and spermcasters) and longer in species where sperm are directly transferred to females (that is, internal fertilizers). We also show that sperm length evolves faster and with a greater number of adaptive shifts in species where sperm operate within females (for example, spermcasters and internal fertilizers). Our results demonstrate that fertilization mode is a key driver in the evolution of sperm length across animals, and we argue that a complex combination of postcopulatory forces has shaped sperm length diversification throughout animal evolution.

Microsatellite markers have been used extensively in studies of sperm competition and the genetic diversity of organisms. In the present study, we performed transcriptome sequencing for Velarifictorus aspersus and developed simple sequence repeat markers to identify valuable markers for genetic studies. The average total length of the V. aspersus transcriptome sequence was 7.70 Gb and it contained 7878 microsatellite loci. Sixty-four microsatellite loci were randomly selected for primer design and 39 loci were successfully amplified, among which 23 loci were polymorphic. The microsatellites were also tested in six other species from the Gryllidae family. Among the species belonging to the genus Velarifictorus, 22 loci were amplified in Velarifictorus micado, eight loci in Velarifictorus ornatus, and 3–6 loci in the other four cricket species.

Precise mechanisms underlying sperm storage and utilization are largely unknown, and data directly linking stored sperm to paternity remain scarce. We used competitive microsatellite PCR to study the effects of female morphology, copula duration and oviposition on the proportion of stored sperm provided by the second of two copulating males (S2) in Scathophaga stercoraria (Diptera: Scathophagidae), the classic model for sperm competition studies. We genotyped all offspring from potentially mixed-paternity clutches to establish the relationship between a second male’s stored sperm (S2) and paternity success (P2). We found consistent skew in sperm storage across the three female spermathecae, with relatively more second-male sperm stored in the singlet spermatheca than in the doublet spermathecae. S2 generally decreased with increasing spermathecal size, consistent with either heightened first-male storage in larger spermathecae, or less efficient sperm displacement in them. Additionally, copula duration and several two-way interactions influenced S2, highlighting the complexity of postcopulatory processes and sperm storage. Importantly, S2 and P2 were strongly correlated. Manipulation of the timing of oviposition strongly influenced observed sperm-storage patterns, with higher S2 when females laid no eggs before being sacrificed than when they oviposited between copulations, an observation consistent with adaptive plasticity in insemination. Our results identified multiple factors influencing sperm storage, nevertheless suggesting that the proportion of stored sperm is strongly linked to paternity (i.e., a fair raffle). Even more detailed data in this vein are needed to evaluate the general importance of sperm competition relative to cryptic female choice in postcopulatory sexual selection.

Mating systems constitute an important biological characteristic that must be evaluated in commercially important species due to the effects of these systems on population growth rates and genetic ...

The delicate mouse (Pseudomys delicatulus) ranges from Queensland to northern Western Australia. In this study the morphology of the cellular organisation of the testis and cauda epididymal spermatozoa are compared. Individuals from the mainland of the Northern Territory and Western Australia invariably have only 60–70% of the testes composed of seminiferous tubules with the interstitial tissue between the tubules containing abundant lipid rich Leydig cells, whereas the cauda epididymal sperm have highly polymorphic, often pear-shaped, heads and basally attached tails. In individuals from Queensland the seminiferous tubules make up approximately 80% of the testis, whereas in the cauda epididymides the sperm populations are generally less variable and have bilaterally flattened heads with the tail attached to the lower concave surface. These differences in the morphology of the delicate mouse testis and spermatozoa in these two geographic regions suggest differences in intensity of intermale sperm competition, with individuals from northern Western Australia and the mainland of the Northern Territory exhibiting monogamy whereas those from Queensland may exhibit some degree of intermale sperm competition and hence possibly have a polyandrous or promiscuous mating system. These findings support the suggestion that P. delicatulus, as currently recognised, contains at least one cryptic species.

In lizards and snakes, the kidneys produce seminal fluid in addition to having osmoregulatory functions. Therefore, in response to polyandry, kidney mass should be under selection regimes such as those influencing testes. Male red-sided garter snakes deposit a kidney-derived copulatory plug that functions in sperm competition. We first tested for intersexual differences in kidney mass and allometry in one population and found that males had kidneys twice as heavy as those of females, consistent with stronger selection on male kidney mass. Previous studies have shown that male size enhances mating success in one-on-one competition prevalent in small mating aggregations. We then examined the relationship between body size, kidney mass and testes mass in two populations with low (LD) and high (HD) mating aggregation densities. Males from the HD population had heavier testes and heavier kidneys compared with males from the LD population; HD males were also smaller in body size. Our results suggest that the strength of sexual selection, and possibly the balance between pre- and postcopulatory selection, varies in response to population size or density. However, more replication is needed to confirm the generality of these results within red-sided garter snakes and other squamate reptiles.

Coagulum in the semen of some primates plays different roles, depending on the species. In the present study, we examined sperm motility in the coagulum and liquid portions of semen collected from captive individuals from two great ape species: two adult Bornean orangutans (Pongo pygmaeus) (n = 27) and three adult chimpanzees (Pan troglodytes) (n = 14). The results revealed that orangutan sperm remained motile for significantly longer in the coagulum than in the liquid portion (> 18 h). By contrast, chimpanzee sperm motility did not differ significantly over time between the two portions of the semen, although motility was slightly higher in the liquid portion than in the coagulum. The evolution of the seminal coagulum is thought to be related to postcopulatory sperm competition; however, functions of seminal coagulum have not been completely elucidated. Our data from the orangutan semen suggest that in this species, seminal coagulum may strengthen own-sperm survival. This report is the first to provide evidence for this distinctive function of the seminal coagulum. This unique property of orangutan seminal coagulum might be attributable to their reproductive traits, e.g., difficulty in predicting ovulation due to a lack of genital swelling during estrus. The orangutan is a Critically Endangered species, and captive breeding, including artificial insemination (AI), is expected. However, worldwide, only one case of orangutan AI has been successful. Our findings may contribute to an understanding of their basic semen characteristics and help improve the AI method.

When the reproductive interests of males and females conflict, males can evolve traits that are harmful to females, and females can coevolve traits to resist this harm. In the fruit fly, Drosophila melanogaster, there is genetic variation in female resistance traits, which can affect the pre- and post-mating success of males that try to mate with them. However, it is not clear to what extent the expression of these phenotypes can be modified by environmental factors such as sociosexual experience. Here, we tested how the genetic background of a female and her previous mating experience interact to affect the mating success of focal males. In the experience phase, we placed females from 28 distinct genetic backgrounds individually either with a single male (low conflict) or with three males (high conflict) for 48 hr. In the subsequent test phase, we measured the mating and post-mating fertilization success of focal males paired individually with each female. We found that focal males paired with females from the high-conflict treatment were less successful at mating, took longer to mate when they were successful, and had a lower proportion of paternity share. Furthermore, we identified significant female genetic variation associated with male mating success. These results indicate that female experience, along with intrinsic genetic factors, can independently influence different fitness components of her subsequent mates and has implications for our understanding of plastic female mating strategies and the evolution of sexually antagonistic traits in males and females.

Species in the genus Macaca typically live in multimale-multifemale social groups with male macaques exhibiting some of the largest testis: body weight ratios among primates. Males are believed to experience intense levels of sperm competition. Several spermatogenesis genes are located on the Y-chromosome and, interestingly, occasional hybridization between two species has led to the introgression of the rhesus macaque (Macaca mulatta) Y-chromosome deep into the range of the long-tailed macaque (M. fascicularis). These observations have led to the prediction that the successful introgression of the rhesus Y-haplotype is due to functional differences in spermatogenesis genes compared to those of the native long-tailed Y-haplotype. We examine here four Y-chromosomal loci-RBMY, XKRY, and two nearly identical copies of CDY-and their corresponding protein sequences. The genes were surveyed in representative animals from north of, south of, and within the rhesus x long-tailed introgression zone. Our results show a series of non-synonymous amino acid substitutions present between the two Y-haplotypes. Protein structure modeling via I-TASSER revealed different folding patterns between the two species' Y-proteins, and functional predictions via TreeSAAP further reveal physicochemical differences as a result of non-synonymous substitutions. These differences inform our understanding of the evolution of primate Y-proteins involved in spermatogenesis and, in turn, have biomedical implications for human male fertility.

In environments where females mate multiply, males should adjust their behaviour and physiology in response to the perceived level of sperm competition to maximize their fitness. Evidence of such plasticity has been found in several laboratory and field studies, but little is known about the cues stimulating these responses in natural populations. One way in which males appear to assess sperm competition risk is through encounter rates with conspecific males. Such encounter rates may be driven by the spatial distribution of resources required by males (i.e. food patches or potential mates), which in turn affects local density. However, explicit links between resource distribution, male encounter rates and shifts in behaviour related to sperm competition have not been demonstrated. We found that when group size of Drosophila melanogaster males was held constant, a small decrease in the distance between patches of food resources had striking effects on male behaviour. Compared to those from dispersed resources, males on clustered resources had a significantly reduced intermale distance (and hence encounter rate) and subsequently a longer noncompetitive copulation duration, previously shown to be a reliable indicator of male perception of sperm competition risk. The aggregation of resources, operating via increased encounter rate, can stimulate shifts in behaviour affecting male sperm competition performance. Given that the spatial distribution of resources is typically variable in natural populations (and often unpredictable), selection is likely to favour the evolution of plasticity in sexual behaviour where resource aggregation increases the probability of sperm competition.

Sperm velocity is a key trait that predicts the outcome of sperm competition. By promoting or impeding sperm velocity, females can control fertilization via postcopulatory cryptic female choice. In Chinook salmon, ovarian fluid (OF), which surrounds the ova, mediates sperm velocity according to male and female identity, biasing the outcome of sperm competition towards males with faster sperm. Past investigations have revealed proteome variation in OF, but the specific components of OF that differentially mediate sperm velocity have yet to be characterized. Here we use quantitative proteomics to investigate whether OF protein composition explains variation in sperm velocity and fertilization success. We found that OF proteomes from six females robustly clustered into two groups and that these groups are distinguished by the abundance of a restricted set of proteins significantly associated with sperm velocity. Exposure of sperm to OF from females in group I had faster sperm compared to sperm exposed to the OF of group II females. Overall, OF proteins that distinguished between these groups were enriched for vitellogenin and calcium ion interactions. Our findings suggest that these proteins may form the functional basis for cryptic female choice via the biochemical and physiological mediation of sperm velocity.

The expensive tissue hypothesis predicts a trade-off between investments in the brain and other energetically costly organs due to the costs associated with their growth and maintenance within the finite energy resources available. However, few studies address the strength of relationships between brain size and investments in precopulatory (ornaments and armaments) and postcopulatory (testes and ejaculates) sexual traits. Here, in a broad comparative study, we tested the prediction that the relationship between brain size and investment in sexual traits differs among taxa relative to the importance of sperm competition within them. We found that brain size was negatively correlated with sexual size dimorphism (SSD) in anurans and primates, and it tended to decrease with SSD in ungulates and cetaceans. However, brain size did not covary significantly with armaments (e.g., canine length, horn, antler, and muscle mass). Brain size was not correlated with postcopulatory sexual traits (testes and ejaculates). The intensity of covariance between brain size and precopulatory sexual traits decreased with increasing relative testis size.

Conspecific males often disturb copulating mites, thus interrupting the mating and a subsequent mating may occur. However, in natural populations of Tetranychus urticae, virgin or sperm-depleted females exist (3.2% of the females in a population) and a second mating may occur when these females are placed with males of unknown age. How does multiple mating benefit the fitness of T. urticae females? To answer this question and to establish the success of re-mating, we investigated the effects of mating interruption at various time intervals after the start on the validity of fertilization. Female offspring ratios were significantly lower if mating was interrupted at 30, 60 or 90 s, compared with complete (undisturbed) mating. Females accepted the second mating if they could not obtain sufficient sperm from the first mating owing to interruption. The success of the second mating and priority of the second male’s sperm depended on how soon the interruption of the first mating occurred. Proportion of females produced female offspring of the second males when mated at 6 days after the first mating was fewer than those when mated just after the first mating. To know the effect of males of unknown age, of which some were assumed to be sperm-depleted, we used them as the first mating partner and then used virgin males as the second mating partner. Three females (6.4%, n = 47) who mated with males of unknown age at the first mating, produced female offspring of the second males, but no females produced female offspring of the second males when mated with virgin males at the first mating. These findings indicate that sperm depletion, caused by mating interruption or paternal age in the first mating, might indeed affect the fertilizing ability.

Sea slugs are interesting models to study post-copulatory sexual selection in simultaneous hermaphrodites due to the enormous variation of their reproductive systems. However, the knowledge of the functional morphology of their reproductive system is limited to few species, and it is rarely discussed in the context of sexual selection theory. In this study, we investigated the functional morphology of the sperm-containing chambers (i.e., ampulla, seminal receptacle, and bursa copulatrix) of the reproductive system of Okenia polycerelloides (Ortea & Bouchet, 1983), based on light, confocal, and electron microscopy. Although the morphology of the ampulla is similar to other species, indicating that it is a site for autosperm storage, we found some sperm facing the ampullar epithelium, a feature commonly regarded as characteristic of the seminal receptacle of sea slugs. The seminal receptacle of O. polycerelloides showed secretory activity and contained sperm with distribution and orientation suggestive of stratification of allosperm from distinct mating events, a feature that would affect sperm competition. The bursa copulatrix had epithelial cells with secretory and absorptive characteristics, and contained degraded sperm and yolk granules within its lumen. Comparative analyses of the contents of each organ demonstrated that sperm digestion occurs in the bursa copulatrix and affects sperm heads first, changing their morphology from slender and curved to shorter and ellipsoid before complete lysis. Although digestion and absorption of surplus sperm are currently the main hypothesized functions for the bursa copulatrix, its role in cryptic female choice should not be ruled out. The close structural connection between the seminal receptacle and bursa copulatrix, as well as their muscular walls, would enable control over the fate of the sperm received in each mating event, that is, storage or digestion.

Multiple mating by females increases both the risk and intensity of sperm competition for males. The ability to temporarily or permanently inhibit female receptivity to further mating represents a highly adaptive strategy to reduce the risk of sperm competition. Males of the West Indian sweet potato weevil Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae) and the seed bug Togo hemipterus (Scott) (Heteroptera: Lygaeidae) use accessory gland substances to inhibit remating in females. It is known that there is variance in the length of the female refractory period (i.e., the effectiveness of male ejaculate substances that inhibit female remating) within a species. However, it is unclear what types of male traits affect the length of the female refractory period. The mechanism of the female refractory period is not sufficiently revealed. We focused on male body size, which is often a target of sexual selection, and investigated the relationship between male body size and both the refractory period of females and mating duration. Contrary to our thought, male body size did not significantly influence the refractory period of females or mating duration in either species. Smaller male body size was not particularly favored by post-copulatory sexual selection in either species.

Africa hosts a high number of bat species, many of which have been poorly studied. Among African vesper bats (Vespertilionidae), some species are morphologically similar to each other, hampering identification in the field. Consequently, basic information on these species’ population dynamics, distributions, or behaviors is vague and/or incorrectly documented. Among some Vespertilionidae, variation in flaccid penis shape enables species identification. This variation in morphology is thought to have resulted from sexual selection operating on the erect penis. Furthermore, in the context of strong postcopulatory competition, divergence in sperm traits may have evolved among closely related species. These male reproductive characters have hitherto not been investigated in southern African vespertilionid bats. In this study, we present the morphology of flaccid penis, erect penis, and sperm, of six small vesper species, based on the prediction that these characters have evolved sufficiently toward different optima to allow species discrimination. Species identification based solely on the morphological characters of the penis entirely matched species identification based on traditional (craniodental) characters. We illustrate penis morphology of these species to assist with field identifications for future ecological, conservation, or behavioral studies. Sperm morphology was sufficient to discriminate between the different genera, but could not reliably separate Neoromicia nana, Ne. zuluensis, and Ne. capensis. Hence, we demonstrate that these difficult to identify species can be readily distinguished based on traits directly observable in the field and on living animals, which will lead to more focused field studies on these otherwise morphologically cryptic species. Finally, our comparisons of penis morphology support the suggestion that Hypsugo anchietae is better placed within the genus Neoromicia.

Courtship can have multiple, nonexclusive, functions, and it often plays a central role in mate choice. In the polyandrous, cannibalistic spider Argiope keyserlingi, males perform vibratory courtship that suppresses female aggression. Although females appear to have a preference (i.e. they shorten their latency to mate) for males performing courtship displays with specific characteristics (i.e. many shudders with consistent shudder duration), they are not more likely to mate with them. Here, we asked whether in this species courtship quality plays a role in postcopulatory sexual selection, by affecting male fertilization success via cryptic female choice. We quantified male courtship performance in a standardized thread assay with no male–female interactions, which simulates male approach to the female on the web. We allowed each unmated adult female to mate with two males. Prior to mating, one male from each pair was sterilized using an irradiation treatment and offspring paternity was assigned using the sterile male technique. Unexpectedly, we found that irradiated males had a fertilization advantage over nonirradiated males. When we controlled for that bias, males performing courtship with a large number of shudders and long copula durations gained the largest paternity shares. Together with evidence from previous studies, our results suggest that courtship quality mediates cryptic female choice, but further investigations are needed to parse out the influence of female control and sperm competition on paternity.

Phenotypic plasticity can allow animals to adapt their behavior, such as their mating effort, to their social and sexual environment. However, this relies on the individual receiving accurate and reliable cues of the environmental conditions. This can be achieved via the receipt of multimodal cues, which may provide redundancy and robustness. Male Drosophila melanogaster detect presence of rivals via combinations of any two or more redundant cue components (sound, smell, and touch) and respond by extending their subsequent mating duration, which is associated with higher reproductive success. Although alternative combinations of cues of rival presence have previously been found to elicit equivalent increases in mating duration and offspring production, their redundancy in securing success under sperm competition has not previously been tested. Here, we explicitly test this by exposing male D. melanogaster to alternative combinations of rival cues, and examine reproductive success in both the presence and absence of sperm competition. The results supported previous findings of redundancy of cues in terms of behavioral responses. However, there was no evidence of reproductive benefits accrued by extending mating duration in response to rivals. The lack of identifiable fitness benefits of longer mating under these conditions, both in the presence and absence of sperm competition, contrasted with some previous results, but could be explained by (a) damage sustained from aggressive interactions with rivals leading to reduced ability to increase ejaculate investment, (b) presence of features of the social environment, such as male and female mating status, that obscured the fitness benefits of longer mating, and (c) decoupling of behavioral investment with fitness benefits., Phenotypic plasticity can allow animals to adapt their behavior, such as their mating effort, to their social and sexual environment. However, this relies on the individual receiving accurate and reliable cues of the environmental conditions. Here, we explicitly test this by exposing male Drosophila melanogaster to alternative combinations of rival cues and examining reproductive success in both the presence and absence of sperm competition. The results supported previous findings but there was no evidence of reproductive benefits accrued by extending mating duration in response to rivals.

Copying the mate choice of another conspecific by a bystanding observer is a socially mediated alternative mating strategy to independent mate choice. Because increased sperm competition is a potential cost of mate-choice copying, males should be prudent and less likely to copy the observed mate choice of other males when the latter are more sexually attractive than themselves. We tested this hypothesis using the Trinidadian guppy, Poecilia reticulata. Focal observer males were simultaneously presented with a large and a small female as potential mates, either in the presence or absence of a nearby model male. We varied the relative differences in the body length and colour ornamentation, traits that predict male sexual attractiveness and competitiveness in the guppy, of observer and model males in each of three experimental treatments: (1) observer and model males were matched for body length and colour ornamentation; (2) the model male was larger and more colourful than the observer male; (3) the model male was smaller and less colourful than the observer male. We compared the mating preferences of focal observer males in each of the latter treatments with that of focal males in a control treatment, wherein no model male (and thus no copying opportunity) was present. As predicted, focal males copied the staged apparent mate choice of a model male in all three experimental treatments, but not in the control treatment. However, contrary to our a priori hypothesis, there were no significant differences in the likelihood or strength of the copying response in observer males among the experimental treatments. Although male guppies are known to modify their mating tactics in the presence of sexual competitors, we found no evidence for an influence of the relative phenotypes of observer and model males on mate-choice copying behaviour in focal observer males.

Cephalopods are a remarkable group for studies on sexual selection, due to their complex reproductive behavior, pronounced sexual dimorphism, polygamous behavior, as well as cumulative evidence of pre- and postcopulatory male-male competition and female choice in the group. is paper provides a review of sexual selection studies on cephalopods, with an emphasis on the loliginid squid mating system. In this system, males may adopt alternative reproductive tactics (ART) and display male intrasexual dimorphism, with each morph generally exhibiting distinct behaviors and ejaculate traits. Consort males typically transfer ejaculates to the female mantle cavity, while sneaker males typically transfer ejaculates to the female buccal region, where a sperm storage organ is located. Each site provides distinct fertilization environments for the ejaculates of each morph, differing in the availability of eggs, gamete protection, and timing between mating and fertilization. Accordingly, the ejaculates of each morph show putative adaptations for each site, differing in size, morphology, sperm release duration, and sperm behavior. Although solid evidence is lacking for most loliginids, cryptic female choice may occur in this system through, for example, spermatangia removal or egg string manipulation during fertilization. In addition, recent evidence for one loliginid species indicates that adoption of male ART is ontogenetic and may represent two alternative pathways: from immature males (i) directly to consort phenotype or (ii) going through a sneaker phase, then switching to consort morph. The second pathway would, therefore, imply striking change in behavior, physiology, and ejaculates. Moreover, data on the age of this species suggests that the adoption of sneaker tactic early in life may be associated with extending the male mating period by 25%. With such a peculiar system, loliginids should prove useful not only for comprehending the evolution of female promiscuity and ART in cephalopods, but also as an emerging nonvertebrate and nonarthropod model for testing sperm competition models.

Most terrestrial animals live in natural atmospheric conditions, but some are also adapted to low oxygen (hypoxic) or high‐carbon dioxide (hypercapnic) conditions, such as in social insect nests, soil, caves, wood, and decaying material. Although it is possible that individuals adapt their behaviour to the environmental condition of their habitats, mating behaviour under ecologically possible ranges of CO₂ has not been well studied. We compared walking activity, duration of mating behaviour, and sperm transfer ability in the West Indian sweet potato weevil, Euscepes postfasciatus Fairmaire (Coleoptera: Curculionidae), in high CO₂ (10 000 p.p.m.) vs. normal atmospheric conditions (laboratory air, ca. 800 p.p.m.). We found that high CO₂ enhanced walking activity and mounting frequency. Under such circumstances, we predicted that enhanced activities under a high‐CO₂ environment would increase the risk of sperm competition, which induces an extended copulation period and an increase in sperm transfer. However, weevils shortened the mating period and did not alter their sperm transfer ability under high CO₂. These findings are, as far as we are aware, the first report of the effect of ecologically relevant high CO₂ on insect mating behaviour under mass‐rearing conditions. The effect of ambient conditions on mating behaviour and sperm transfer is discussed in relation to the intensity of female refusal behaviour directed against males.

Sexual selection theory provides a framework for investigating the evolution of traits involved in attracting and competing for mates. Given the sexual function of such traits, studies generally focus on individual interactions (i.e., displays and contests) in explaining trait origin and persistence. We show that ecological factors can strongly influence the adaptive value of these traits, and changes to these factors can lead to rapid evolutionary change. We compared sexually selected traits in the small Indian mongoose (Urva auropunctata) between their sparsely populated native range and four tropical islands to which they were introduced within the last 150 years and where, due to a lack of interspecific competition and predation, they have become invasive and densely populated. Because of a likely increase in encounter rate, we predicted that selection on long-distance chemical advertisement by males would relax in the introduced range. Accordingly, male, but not female, anal pads (used in scent marking) decreased in size in relation to both time since introduction and population density, and their relationship to body size and condition weakened. Concurrently, as predicted by intensified sperm competition, testis size increased following introduction. The small Indian mongoose thus experienced an inversion in the relative contributions to fitness of two sexual traits, followed by their rapid evolution in line with ecological changes.

Rats are gregarious and promiscuous and when a female enters the behavioral estrous, two or more males copulate alternately (mount, intromission or ejaculation are performed by any of them). This c...

Male animals often show higher mutation rates than their female conspecifics. A hypothesis for this male bias is that competition over fertilization of female gametes leads to increased male investment into reproduction at the expense of maintenance and repair, resulting in a trade-off between male success in sperm competition and offspring quality. Here, we provide evidence for this hypothesis by harnessing the power of experimental evolution to study effects of sexual selection on the male germline in the seed beetle Callosobruchus maculatus. We first show that 50 generations of evolution under strong sexual selection, coupled with experimental removal of natural selection, resulted in males that are more successful in sperm competition. We then show that these males produce progeny of lower quality if engaging in sociosexual interactions prior to being challenged to surveil and repair experimentally induced damage in their germline and that the presence of male competitors alone can be enough to elicit this response. We identify 18 candidate genes that showed differential expression in response to the induced germline damage, with several of these previously implicated in processes associated with DNA repair and cellular maintenance. These genes also showed significant expression changes across sociosexual treatments of fathers and predicted the reduction in quality of their offspring, with expression of one gene also being strongly correlated to male sperm competition success. Sex differences in expression of the same 18 genes indicate a substantially higher female investment in germline maintenance. While more work is needed to detail the exact molecular underpinnings of our results, our findings provide rare experimental evidence for a trade-off between male success in sperm competition and germline maintenance. This suggests that sex differences in the relative strengths of sexual and natural selection are causally linked to male mutation bias. The tenet advocated here, that the allocation decisions of an individual can affect plasticity of its germline and the resulting genetic quality of subsequent generations, has several interesting implications for mate choice processes. Title in the list of papers of Julian Baur's thesis: Increased male investment in sperm competition results in offspring of lower quality

In the leaf-cutting ant Atta colombica, queens receive ejaculates from multiple males during one single mating event early in their lives. A queen’s fertility and fitness therefore depend on maximizing the number of sperm cells she can store and maintain inside her spermatheca. Previous studies implied significant physiological mating costs, either originating from energetic investments maximizing sperm survival, or from resolving sexual conflicts to terminate male-driven incapacitation of rival sperm via serine proteases found in seminal fluid. Here we conducted an artificial insemination experiment, which allowed us to distinguish between the effects of sperm and seminal fluid within the queen’s sexual tract on her survival and immunocompetence. We found significantly higher mortality in queens that we had inseminated with sperm, independently of whether seminal fluid was present or not. Additionally, after receiving sperm, heavier queens had a higher probability of survival compared to lightweight queens, and immunocompetence decreased disproportionally for queens that had lost weight during the experiment. These findings indicate that queens pay significant physiological costs for maintaining and storing sperm shortly after mating. On the other hand, the presence of seminal fluid within the queens’ sexual tract neither affected their survival nor their immunocompetence. This suggests that the energetic costs that queens incur shortly after mating are primarily due to investments in sperm maintenance and not costs of terminating conflicts between competing ejaculates. This outcome is consistent with the idea that sexually selected traits in social insects with permanent castes can evolve only when they do not affect survival or life-time fitness of queens in any significant way.

Many species of animals engage in sexual reproduction, a mechanism that allows animal species to evolve to keep up/remain in evolutionary arms races with their predators, prey, and parasites. Mate choice is a core Darwinian attribute in sexually reproducing animals attempting to produce optimum genetic combinations among their offspring. Mate choice played a role in the evolution of sex itself in addition to its current role in behavior and evolution. Competition among animals of the same sex for mates (intrasexual selection) can yield traits that conflict with the expectations of natural selection, such as large horns or antlers. Exaggerated traits result from runaway sexual selection and may cause apparent handicaps in nonmating contexts. Mating systems, such as polygyny, polyandry, and monogamy, can influence the outcome of sexual selection, with high variance in mating success resulting in traits that reflect intense levels of sexual selection. The choice of a mate with “good genes” can involve assessment of traits that are associated with developmental rigor, immunocompetence, the ability to resist parasites, and more.

Saimiri are neotropical primates with seasonal reproduction, males develop a seasonal fattening condition that has been suggested as a pre-copulatory sexual selection strategy. Furthermore, females mate with multiple males in the same season. This could also favor the evolution of a postcopulatory sexual strategy by sperm competition. In this study, we tested the hypothesis that there is a relationship between the fatted condition and advantageous seminal characteristics in Saimiri collinsi and evaluated its implications for sperm competition. Adult males (N = 10), aged 5-15 years, housed in mixed or only-male groups, were analyzed from June, 2015 to July, 2016. Measurements of weight, axilla, and arm skinfold, and testicular volume were taken monthly, and semen was collected by electroejaculation. A fattening index was developed to quantify and identify fatted males, biometrics, and seminal parameters were compared between the non-fatted and fatted conditions. The fatted males present a larger testicular volume. This is related to the increase in spermatogenic activity necessary to sustain a high ejaculation frequency during the mating season. An increase in seminal volume and in frequency of semen coagulation were detected in fatted males, advantages related to sperm protection in the female reproductive tract. Age and social context were not significant sources of variation for both morphological and seminal traits. A decrease in response from the fatted males in obtaining semen and an increase in the frequency of azoospermic ejaculates were observed. These unexpected results may be due to intense reproductive activities in a short period. The fattening phenomenon has many implications in the sexual selection of squirrel monkeys, and they are still not entirely unveiled. Our results corroborate the idea that, in S. collinsi, the fatted male condition is related to sexual selection, and we found evidence suggesting it may be also expressed by a post-copulatory component, sperm competition.

Spermatozoal morphology is highly variable both among and within species and in ways that can significantly impact fertilization success. In Drosophila melanogaster, paternity success depends on sperm length of both competing males and length of the female’s primary sperm storage organ. We found that genes upregulated in long sperm testes are enriched for lncRNAs and seminal fluid proteins (Sfps). Transferred in seminal fluid to the female during mating, Sfps are secreted by the male accessory glands (AG) and affect female remating rate, physiology, and behavior with concomitant advantages for male reproductive success. Despite being upregulated in long sperm testes, they have no known function in testis tissue. We found that Sex Peptide and ovulin (Acp26Aa) knockouts resulted in shorter sperm, suggesting that Sfps may regulate sperm length during spermatogenesis. However, knockout of AG function did not affect sperm length, suggesting that AG expression has no influence on spermatogenic processes. We also found that long sperm males are better able to delay female remating, suggesting higher Sfp expression in AG. These results might suggest that long sperm males have a double advantage in sperm competition by both delaying female remating, likely through transfer of more Sfps, and by resisting sperm displacement. However, we also found that this extra advantage does not necessarily translate to more progeny or higher paternity success. Thus, we found that multiple components of the ejaculate coordinate to promote male reproductive success at different stages of reproduction, but the realized fitness advantages in sperm competition are uncertain.Significance StatementThe ejaculate is comprised of sperm produced in the testis and seminal fluid primarily produced in the male accessory glands (AG). These complementary components are both critical for male reproductive success, but they are largely considered to be functionally, genetically, and developmentally independent. In a quest to understand genetic mechanisms of sperm length variation, we found that genes upregulated in long sperm testes are enriched for lncRNAs and seminal fluid proteins (Sfps). Knockout of two Sfps, Sex Peptide and ovulin, results in shorter sperm, though knockout of AG function has no effect. Moreover, long sperm males delay female remating longer. These results suggest sophisticated testis-AG coordination that amplifies male reproductive success, with implications for evolutionary integration of sexually selected traits.

Although equal sex ratio represents an equilibrium in evolutionary theory, cases of sex ratio bias are accumulating, mostly reported in single species. Here, we surveyed progeny sex ratios in 23 species of the nematode genus Caenorhabditis. In experiments where males and females had unlimited access to each other (unlimited mating), we found 15 species out of the 23 species had female(hermaphrodite)-biased sex ratios. Phylogenetic mapping indicates female-bias to represent the ancestral state, with the occurrence of seven independent transitions from female-bias to non-bias along the phylogeny. As sperm competition could underlie the female bias, we also assayed progeny sex ratios in an experiment where mating was limited to a few hours. Of the 15 species that showed female-biased ratios under unlimited mating, six species showed no sex ratio bias when mating was limited. This result is consistent with sperm competition whereby X-bearing sperm take precedence over nullo-X during fertilization, yielding more female progeny when sperm are unlimited but equal sex ratio when sperm are limited. The other nine species showed sex ratio bias in both experiments, but the day-by-day profiles suggest sperm competition may also play a role. Our study shows that sex ratio evolution within Caenorhabditis nematodes is dynamic and that sex ratio bias is common not only in parasites as previously found but also in free-living nematodes. Our results also suggest that sperm competition could be a mechanism underlying sex ratio bias.

Senescence, or the deterioration of functionality with age, varies widely across taxa in pattern and rate. Insights into why and how this variation occurs are hindered by the predominance of lab-focused research on short-lived model species with determinate growth. We synthesize evolutionary theories of senescence, highlight key information gaps, and clarify predictions for species with low mortality and variable degrees of indeterminate growth. Lake trout are an ideal species to evaluate predictions in the wild. We monitored individual males from two populations (1976-2017) longitudinally for changes in adult mortality (actuarial senescence) and body condition (proxy for energy balance). A cross-sectional approach (2017) compared young (ages 4-10 years) and old (18-37 years) adults for (1) phenotypic performance in body condition, and semen quality - which is related to fertility under sperm competition (reproductive senescence), and (2) relative telomere length (potential proxy for cellular senescence). Adult growth in these particular populations is constrained by a simplified food web, and our data support predictions of negligible senescence when maximum size is only slightly larger than maturation size. Negative senescence (aka reverse senescence) may occur in other lake trout populations where diet shifts allow maximum sizes to be much larger than maturation size.

Origin of functional new genes is a basic biological process that has significant contribution to organismal diversity. Previous studies in both Drosophila and mammals showed that new genes tend to be expressed in testis, and avoid the X chromosome presumably because meiotic sex chromosome inactivation (MSCI). Here we analyze the published single-cell transcriptome data of Drosophila adult testis and find an enrichment of male germline mitotic genes, but an underrepresentation of meiotic genes on the X chromosome. This can be attributed to an excess of autosomal meiotic genes that were derived from their X-linked mitotic progenitors, which provides direct cell-level evidence for MSCI in Drosophila. We reveal that new genes, particularly those produced by retrotransposition, tend to exhibit an expression shift toward late spermatogenesis compared to their parental copies, probably due to the more intensive sperm competition or sexual conflict. Our results dissect the complex factors including the age, the origination mechanisms and the chromosomal locations that influence the new gene origination and evolution in testis, and identify new gene cases that show divergent cell-level expression pattern from their progenitors for future functional studies.

In many species, males can rapidly adjust their ejaculate performance in response to changing levels of sperm competition, an ability that is probably mediated by seminal fluid adaptive plasticity. In the black goby, Gobius niger , territorial males attach viscous ejaculate trails to the nest roof, from which sperm are slowly released into the water during the long-lasting spawning events. Sneaker males release their sperm in the vicinity of the nest, and territorial males try to keep them at a distance by patrolling their territory. We show here that territorial males' ejaculate trails released a higher proportion of their sperm in the presence of a single sneaker, but this proportion decreased when there were three sneakers, an effect that is most likely mediated by a change in the seminal fluid composition. Field observations showed that when multiple sneaking attempts occurred, territorial males spent more time outside the nest, suggesting that ejaculation rate and territory defence are traded-off. Altogether, these results suggest that the adjustment of sperm release from the ejaculate may be strategic, guaranteeing a more continuous concentration of the territorial male's sperm in the nest, although at a lower level, when he is engaged in prolonged territory defence outside the nest.

Over the last 50 years, sperm competition has become increasingly recognised as a potent evolutionary force shaping male ejaculate traits. One such trait is sperm swimming speed, with faster sperm associated with increased fertilisation success in some species. Consequently, sperm are often thought to have evolved to be longer in order to facilitate faster movement. However, despite the intrinsic appeal of this argument, sperm operate in a different biophysical environment than we are used to, and instead increasing length may not necessarily be associated with higher velocity. Here, we test four predictive models (ConstantPower Density, Constant Speed, Constant Power Transfer, Constant Force) of the relationship between sperm length and speed. We collated published data on sperm morphology and velocity from 141 animal species, tested for structural clustering of sperm morphology and then compared the model predictions across all morphologically similar sperm clusters. Within four of five morphological clusters of sperm, we did not find a significant positive relationship between total sperm length and velocity. Instead, in four morphological sperm clusters we found evidence for the Constant Speed model, which predicts that power output is determined by the flagellum and so is proportional to flagellum length. Our results show the relationship between sperm morphology (size, width) and swimming speed is complex and that traditional models do not capture the biophysical interactions involved. Future work therefore needs to incorporate not only a better understanding of how sperm operate in the microfluid environment, but also the importance of fertilising environment, i.e., internal and external fertilisers. The microenvironment in which sperm operate is of critical importance in shaping the relationship between sperm length and form and sperm swimming speed.

Mammalian sperm must spend a minimum period of time within a female reproductive tract to achieve the capacity to fertilize oocytes. This phenomenon, termed sperm ‘capacitation’, was discovered nearly seven decades ago and opened a window into the complexities of sperm–female interaction. Capacitation is most commonly used to refer to a specific combination of processes that are believed to be widespread in mammals and includes modifications to the sperm plasma membrane, elevation of intracellular cyclic AMP levels, induction of protein tyrosine phosphorylation, increased intracellular Ca(2+) levels, hyperactivation of motility, and, eventually, the acrosome reaction. Capacitation is only one example of post-ejaculatory modifications to sperm (PEMS) that are widespread throughout the animal kingdom. Although PEMS are less well studied in non-mammalian taxa, they likely represent the rule rather than the exception in species with internal fertilization. These PEMS are diverse in form and collectively represent the outcome of selection fashioning complex maturational trajectories of sperm that include multiple, sequential phenotypes that are specialized for stage-specific functionality within the female. In many cases, PEMS are critical for sperm to migrate successfully through the female reproductive tract, survive a protracted period of storage, reach the site of fertilization and/or achieve the capacity to fertilize eggs. We predict that PEMS will exhibit widespread phenotypic plasticity mediated by sperm–female interactions. The successful execution of PEMS thus has important implications for variation in fitness and the operation of post-copulatory sexual selection. Furthermore, it may provide a widespread mechanism of reproductive isolation and the maintenance of species boundaries. Despite their possible ubiquity and importance, the investigation of PEMS has been largely descriptive, lacking any phylogenetic consideration with regard to divergence, and there have been no theoretical or empirical investigations of their evolutionary significance. Here, we (i) clarify PEMS-related nomenclature; (ii) address the evolutionary origin, maintenance and divergence in PEMS in the context of the protracted life history of sperm and the complex, selective environment of the female reproductive tract; (iii) describe taxonomically widespread types of PEMS: sperm activation, chemotaxis and the dissociation of sperm conjugates; (iv) review the occurence of PEMS throughout the animal kingdom; (v) consider alternative hypotheses for the adaptive value of PEMS; (vi) speculate on the evolutionary implications of PEMS for genomic architecture, sexual selection, and reproductive isolation; and (vii) suggest fruitful directions for future functional and evolutionary analyses of PEMS.

Given that sperm production can be costly, theory predicts that males should optimally adjust the quantity and/or quality of their sperm in response to their social environment to maximize their paternity success. Although experiments demonstrate that males can alter their ejaculates in response to manipulations of the social environment and studies show that ejaculate traits covary with social environment across populations, it is unknown whether individual variation in sperm traits corresponds to natural variation found within wild populations. Using an island population of brown anole lizards (Anolis sagrei), we tested the prediction that sperm traits (sperm count, sperm morphology, sperm velocity) respond to natural variation in the risk of sperm competition, as inferred from the local density and operational sex ratio (OSR) of conspecifics. We found that males living in high-density areas of the island produced relatively larger sperm midpieces, smaller sperm heads, and lower sperm counts. Sperm traits were unrelated to OSR after accounting for the covariance between OSR and density. Our findings broaden the implications of sperm competition theory to intrapopulation social environment variation by showing that sperm count and sperm morphology vary with fine-scale differences in density within a single wild population. Electronic supplementary material The online version of this article (10.1007/s00442-019-04511-z) contains supplementary material, which is available to authorized users.

Mating activity of a wild Mueller’s gibbon group (Hylobates muelleri) was observed in the Danum Valley Conservation Area, Sabah, Malaysia. The purpose of this study was to investigate the function of copulation calls in gibbons. The female emitted copulation calls at the time of intromission and pelvic thrusting. Copulation calls were composed of two notes and one of them was sung only while mating. Approximately half of copulation calls were sung near the range boundary. Mating with copulation calls sometimes occurred while singing. According to the model that female copulation calls have evolved under the selective pressures of risk of infanticide and sperm competition, copulation calls should be rare in species with little female promiscuity. As gibbons usually live in pair-living social organization and have a monogamous mating system, no vocal signals by female gibbons are considered to be needed. However, clear copulation calls were emitted by the female. It suggests that the relationship between paired gibbons is unstable. Copulation calls by the female Mueller’s gibbon may function to increase mate guarding and strengthen the pair bond.