Territoriality and male-biased sexual size dimorphism in Argia reclusa (Odonata: Zygoptera)

In Odonata, many species present sexual size dimorphism (SSD), which can be associated with male territoriality in Zygoptera. We hypothesized that in the territorial damselfly Argia reclusa, male–male competition can favor large males, and consequently, drive selection pressures to generate male-biased SSD. The study was performed at a small stream in southeastern Brazil. Males were marked, and we measured body size and assessed the quality of territories. We tested if larger territorial males (a) defended the best territories (those with more male intrusions and visiting females), (b) won more fights, and (c) mated more. Couples were collected and measured to show the occurrence of sexual size dimorphism. Results indicated that males are larger than females, and that territorial males were larger than non-territorial males. Larger territorial males won more fights and defended the best territories. There was no difference between the mating success of large territorial and small non-territorial males. Although our findings suggest that male territoriality may play a significant role on the evolution of sexual size dimorphism in A. reclusa, we suggest that other factors should also be considered to explain the evolution of SSD in damselflies, since non-territorial males are also capable of acquiring mates.     

Pairing success and sperm reserve of male Gammarus pulex infected by Cyathocephalus truncatus (Cestoda: Spathebothriidea)

Manipulative parasites with complex life cycles are known to induce behavioural and physiological changes in their intermediate hosts. Cyathocephalus truncatus is a manipulative parasite which infects Gammarus pulex as intermediate host. G. pulex males display pre-copulatory mate guarding as a response to male-male competition for access to receptive females. In this paper, we tested the influence that C. truncatus-infection might have on male G. pulex sperm number and pairing success. We considered 3 classes of G. pulex males in our experiments: (i) uninfected males found paired in the field, (ii) uninfected males found unpaired in the field, or (iii) infected males found unpaired in the field. Both infected males and uninfected unpaired males paired less with a new female than uninfected paired males did. Furthermore, infected males appear to be at a strong disadvantage when directly competing for females with a healthy rival male, and had fewer sperm in their testes. We discuss the potential effect of male and female mating strategies on such male host mating alteration.     

Timing of maturation and the mating system of the spider,Stegodyphus lineatus (Eresidae): how important is body size?

Selection on life history traits such as the timing of maturation and the size at maturity strongly depends on the mating system. In spiders, the mating system hypothesized to Be determined by spermathecal morphology and the related sperm precedence pattern. In a natural population of the eresid spider Stegodyphus lineatus , predictions concerning the timing of maturation, male mating behaviour and success were tested. Eresid spiders are supposed to show protandry, prematuration mate guarding and strong male-male competition resulting in selection for large body size and early maturation. In contrast to these predictions, male and female maturation overlapped largely. Males did not guard premature females nor was there evidence for male-male competition. Among mating pairs, male did not relate to female size, nor to duration of cohabitation. Evidence for an advantage of first over second or large over small males is weak. In males, body size at maturity and the time of maturation were negatively correlated although a trade off between timing of maturation and the body size reached by then should result in a positive correlation. Possible causes are discussed.     

Territorial male gobies respond aggressively to sneakers but do not adjust their sperm expenditure

We investigated whether mating behavior (sperm expenditure,courtship rate, and nest guarding) varied according to differentlevels of sperm competition in territorial males of two gobyspecies, the grass and the black gobies. We measured sperm expenditure(sperm released after 30 min from the beginning of the spawning),male courtship rate, and nest-guarding behavior in territorialmales of both species during simulated spawnings, in which wevaried the number of attending sneakers. Our results showedthat, in both species, territorial males adjusted their effortin nest guarding to the presence of rival sneakers by increasingthe time spent patrolling the territory and attacking the sneakers.In contrast, sperm expenditure and male courtship rate werenot influenced by the number of attending sneakers. These resultsare in agreement with those reported for other fish with alternativemating tactics and help to interpret previous inconsistenciesbetween theoretical predictions and measured levels of spermreleased at different levels of sperm competition by sneakersof the two gobiids studied here.     

Sexual selection, sexual size dimorphism and Rensch's rule in Odonata

Odonata (dragonflies and damselflies) exhibit a range of sexual size dimorphism (SSD) that includes species with male-biased (males > females) or female-biased SSD (males < females) and species exhibiting nonterritorial or territorial mating strategies. Here, we use phylogenetic comparative analyses to investigate the influence of sexual selection on SSD in both suborders: dragonflies (Anisoptera) and damselflies (Zygoptera). First, we show that damselflies have male-biased SSD, and exhibit an allometric relationship between body size and SSD, that is consistent with Rensch's rule. Second, SSD of dragonflies is not different from unit, and this suborder does not exhibit Rensch's rule. Third, we test the influence of sexual selection on SSD using proxy variables of territorial mating strategy and male agility. Using generalized least squares to account for phylogenetic relationships between species, we show that male-biased SSD increases with territoriality in damselflies, but not in dragonflies. Finally, we show that nonagile territorial odonates exhibit male-biased SSD, whereas male agility is not related to SSD in nonterritorial odonates. These results suggest that sexual selection acting on male sizes influences SSD in Odonata. Taken together, our results, along with avian studies (bustards and shorebirds), suggest that male agility influences SSD, although this influence is modulated by territorial mating strategy and thus the likely advantage of being large. Other evolutionary processes, such as fecundity selection and viability selection, however, need further investigation.     

Sperm production and mating potential of males after a cold shock on pupae of the parasitoid wasp Dinarmus basalis (Hymenoptera: Pteromalidae)

For ectothermic species, temperature is a key environmental factor influencing several aspects of their physiology and ecology, acting particularly on reproduction. To measure the consequences of a severe thermal stress during development on male reproduction, a cold shock (1h at -18 degrees C) was tested on Dinarmus basalis pupae. D. basalis (Hymenoptera: Pteromalidae) is a parasitoid wasp in which sperm management in both male and female is of prime importance. After a cold shock, developmental success was reduced, with a quarter of cold-shocked males not emerging correctly. The stress effects were estimated at the level of sperm stock in seminal vesicles of males at different ages and on the ability of 2-day-old males to access females in single and multiple mating and in male-male competition. Cold-shocked males had a reduced sperm stock compared to control males and this difference persisted with age. The rate of sperm production was similar in both groups. The consequences of a cold shock on male reproductive ability were perceptible in multiple mating and male-male competition but not in single mating. Cold-shocked males were at a disadvantage, inseminating fewer females and copulating less frequently. Finally, male pupae of D. basalis were able to withstand severe temperature stresses and their reproductive functions were partially preserved.     

Competing dwarf males: sexual selection in an orb-weaving spider

Hypotheses for the adaptive significance of extreme female-biased sexual size dimorphism (SSD) generally assume that in dimorphic species males rarely interfere with each other. Here we provide the first multivariate examination of sexual selection because of male-male competition over access to females in a species with 'dwarf' males, the orb-weaving spider Argiope aurantia. Male A. aurantia typically try to mate opportunistically during the female's final moult when she is defenceless. We show that, contrary to previous hypotheses, the local operational sex ratio (males per female on the web) is male-biased most of the season. Both interference and scramble competition occur during opportunistic mating, the former leading to significant selection for large male body size. Male condition and leg length had no effect on mating success independent of size. We discuss these findings in the context of the evolution of extreme female-biased SSD in this clade.     

Seasonal variation in genital and body size, sperm displacement ability, female mating rate, and male harassment in two calopterygid damselflies (Odonata: Calopterygidae)

Sperm competition is a pervasive force. One adaptation is the male ability to displace the rivals' sperm that females have stored from previous copulations. In the damselfly, Calopteryx haemorrhoidalis asturica , males with wider aedeagi displace more spermathecal sperm. The present study documents that the same mechanism operates in another damselfly, Hetaerina americana . However, this genital width in both species decreases along the season, but late-emerging females have more sperm displaced than early-emerging females. Because territorial males mated more and were larger in body and genital size than nonterritorial males, late-season females mated with considerably larger males with respect to female size and this produced higher sperm displacement. Assuming female benefits from storing sperm but that such benefit does not prevail if males displace sperm, it is predicted that, along the season, females will mate less and male harassment (in terms of male mating attempts and oviposition duration) will increase. These predictions were corroborated. In H. americana , it was also tested whether spermathecal sperm became less viable along the season. The results obtained did not corroborate this. This is the first evidence indicating that season affects sperm displacement ability and female mating frequency due to changes in male body and genital size.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 815–829.     

Size-related mating and mate guarding in the orb-web spiderNephila clavata (Araneae,Araneidae)

The orb-web spiderNephila clavata satisfies three conditions for assortative mating proposed by Ridley (The Explanation of Organic Diversity. The Comparative Method and Adaptations for Mating, Clarendon, Oxford, 1983); (1) a large male advantage in male-male competition, (2) a correlation between female size and fecundity, and (3) a long pairing duration. To test Ridley's hypothesis, size assortative mating and guarding were examined in the field. When data were pooled over time, assortative mating was found but this was due to temporal covariation of body sizes of males and receptive females. After controlling for the effect of time, size assortative guarding was not detected, although females guarded by males were larger than those not guarded in the early breeding season. Possible reasons for the absence of size assortative guarding were discussed.     

Size and stability of vertebrate leks

Lek size varies greatly among lekking species. At present there is no explicit theoretical explanation for this diversity. We extend an existing model of optimal lek size that incorporates female mating preferences and male-male contest competition. The model shows that variation in lek size is predicted by the interaction between lek size, overall copulation rate and the proportion of copulations accruing to males of different rank. In species where females prefer to mate on the largest leks and high-ranking males are able to monopolize females irrespective of the size of the lek, the maximum lek size will be large. Conversely, in species where females show weak preference for mating on large leks or increasing lek size quickly results in scramble competition, the maximum lek size will be smaller. Thus, differences between species in lek size may be due largely to differences in the extent to which high-ranking males can monopolize mating opportunities. Leks become unstable and break down when high-ranking males can no longer get their 'expected' copulation success. Therefore, the mechanism that generates male clustering, that is, sexual parasitism of high-ranking males by subordinates, also sets a limit to the largest stable lek size. Copyright 1999 The Association for the Study of Animal Behaviour.     

Male age and sexual experience affect male mating behavior in the hawthorn spider mite,Amphitetranychus viennensis

As is reported, in species with first-male sperm precedence, male age and previous sexual experience play crucial roles in male mating behavior. In the hawthorn spider mite, Amphitetranychus viennensis Zacher, previous studies showed that only females that copulated for the first time could achieve fertilization. Based on this, the effects of male age and mating history on male mate choice and male mate competition were investigated. It was confirmed that males could distinguish virgins from fertilized females but they were unable to discriminate between virgins and unfertilized females. Interestingly, the copulation duration of males mated with fertilized females was much shorter than that of males mated with virgins or unfertilized females. Additionally, for male mating choice, males of all ages and more experienced males preferred 5-day-old virgin females, whereas only less experienced males preferred 1-day-old virgin females. In male-male competition, 3-day-old males were more competitive and obtained more copulations compared with others. Copula duration was closely related to male age. Though no significant differences were observed in mating competition between virgin and mated males, copula duration of males in first copulation was the longest and gradually shortened in subsequent copulations. In all, this investigation demonstrated that male age and sexual experience affected male mate choice and male-male competition, leading to further insight into the influences of male age and sexual experience on the reproductive fitness of both sexes.

     

Male-male competition and mating success in the orb-web spider,Nephila clavata,with reference to temporal factors

Seasonal occurrence patterns of adults of both sexes, intensity of male-male interactions, and mating success in the spider,Nephila clavata, were examined in the field. Adult males began to attend female webs about 2 weeks before female maturation. Large adult males were abundant in the early breeding season, but small males increased later in the season. From the distribution of males among female webs and size relationship of males within a web, male-male interactions seemed to be more intense when most females were still subadult. This was verified by a field experiment in which males were artificially introduced to female webs that were attended by other males. It was found that the probability of introduced males remaining on subadult female webs was lower than that on adult webs. As mating occurred mostly in the period shortly after the female final molt and first male sperm precedence was known in all spiders reported so far, intense male-male competition on subadult female webs seemed to be reasonable. Male longevity had an important influence on the mating success of males with just-molted females. Mating success was also affected by the relative body size of males present in a given period. Since larger males occupied the position closest to females within a web and stayed there longer, relative body size appeared to influence mating success through male-male competition. Female body size at maturation declined with time; hence, males that attained sexual maturity earlier had the advantage of mating with larger and more fecund females. Therefore, early maturation as well as larger size seem to be two important trairs influencing the reproductive success of males.     

The form of sexual selection arising from male-male competition depends on the presence of females in the social environment

Sexual selection arises from social interactions, and if social environments vary so too should sexual selection. For example, male-male competition often occurs either in the presence or in the absence of females, and such changes in the social environment could affect the form and strength of sexual selection. Here we examine how the presence of a female influences selection arising from male-male competition in a leaf-footed cactus bug, Narnia femorata, which has a resource defence mating system. Males compete for territories on cacti because females lay eggs on the cactus plants. Females are not always present when this competition first occurs; however, the presence or absence of the female matters. We found that both the form and strength of selection on male traits, those traits that influenced success in intrasexual competition, depended on the social context. When a female was not present, male size and the area of the sexually dimorphic hind legs was only marginally important to winning a contest. However, males with larger overall size and leg area were more likely to win in the presence of a female. There was also positive quadratic selection on these traits when a female was present with both the largest and the smallest males winning. The implication is unexpected alternative strategies when females are present. Our results support the notion that sexual selection should be studied under all relevant social contexts.     

Parasite-related pairing success in an intermediate host, Caecidotea intermedius (Isopoda): effects of male behavior and reproductive physiology

The acanthocephalan parasite Acanthocephalus dirus develops from the egg to the cystacanth stage inside the freshwater isopod Caecidotea intermedius. We have shown previously that cystacanth-infected male C. intermedius are less likely to initiate mating attempts with females than uninfected males in competitive situations. Here, we used a field-based experiment to examine whether cystacanth-infected males were also less likely to initiate mating attempts with females in noncompetitive situations. We found that infected males were less responsive to females than uninfected males, and we propose that the cystacanth-related change in male mating behavior is mediated by a change in the mating response of males to females rather than male-male competition. We then examined whether cystacanth-related changes in reproductive function, i.e., sperm content and fertilization ability, could explain this variation in male mating behavior. We found that cystacanth-infected males contained both developing and mature sperm and fertilized as many eggs as uninfected males. Thus, we propose that changes in reproductive function are unlikely to explain cystacanth-related variation in male mating behavior in C. intermedius.     

Sexual size dimorphism and assortative mating in Carolina Wrens

ABSTRACT.   Sexual size dimorphism (SSD) may be due to sexual and natural selection, but identifying specific mechanisms that generate such dimorphism in a species is difficult. I examined SSD in Carolina Wrens ( Thryothorus ludovicianus ) by examining (1) the degree of SSD in the population and between pairs using five morphometrics, (2) assortative mating patterns based on size and age, and (3) relationships between size and longevity. Analysis revealed that males were significantly larger than females in all body measurements. For example, mass, bill, and wing measurements yielded a canonical variable that permitted separation of the sexes and linear classification functions correctly determined the sex of 95% (238/250) of all wrens measured. No evidence was found to suggest that SSD was related to resource partitioning. However, assortative mating trends based on morphometrics (e.g., wing length), positive associations between longevity and morphometrics (e.g., wing length in females and body size in males), and intense male-male contests for territorial resources year-round provide evidence that sexual selection may contribute to SSD in Carolina Wrens.     

Operational sex ratio and resource defence as predictors of the mating system in European bitterling

Operational sex ratio (OSR), the ratio of sexually active males to fertilizable females in a population, plays a central role in the theory of mating systems by predicting that the intensity of male–male competition and the degree of sexual selection increases as the OSR becomes increasingly male biased. At high values of OSR, however, resource defence theory predicts the breakdown of territoriality and a shift towards scramble competition with a decrease in sexual selection. The direction that correlations between OSR and resource competition and variance in mating success will take depends on the biology of the species of interest. We investigated the effects of male population density and male‐biased operational sex ratio on male mating tactics shown by a freshwater fish, the European bitterling, Rhodeus sericeus . This species spawns inside living unioneid mussels. Large males defended territories, were aggressive towards conspecifics under equal sex ratios and monopolized pair spawnings with females. The mating tactic, however, changed at high male density where large males ceased to be territorial and instead competed with groups of smaller males to release sperm when females spawned. This change in male behaviour from pair to group spawning has two ramifications for sexual selection. The intensity of sexual selection and variance in male mating success decrease, and the form of sexual competition changes from resource‐ to sperm competition. Thus, the use of alternative mating tactics renders the OSR unable to predict the direction of resource competition and variance in male mating success at high densities.     

Sexual selection in hermit crabs: a review and outlines of future research

The information currently available on sexual selection in hermit crabs is reviewed to identify the role of males and females before, during and after mating. According to this information, possible mechanisms of male–male competition, female choice and/or sexual conflict are suggested. Important male components that may affect mating success include dragging the female shell, rotations of the female's shell and male cheliped palpations, and male size and/or shell characteristics (species and size). Possible female determinants of male mating/fertilization success include size (as an indicator of egg production capacity), signalling of sexual receptivity to males, delay from mate guarding to copulation and mating duration. Avenues for deeper exploration in males include the role of the number and morphometry of male sexual tubes during sperm transfer, and whether ejaculate size and sperm number can be adjusted with variable situations of sperm competition intensity and risk. In females it would be interesting to investigate the chemical and behavioural mechanisms affecting spermatophore breakage for sperm release and the variable duration from sperm transfer to spawning. Given these possibilities, and that sperm is externally deposited on the female's body but inside her shell (except for those species that do not use shells, e.g. Birgus , or species where shells are rather small and do not cover the body totally, e.g. Parapagurus ), we conclude that hermit crabs are unique subjects for separating male and female effects, particularly with respect to the applicability of current ideas in sexual selection such as female choice and sexual conflict. Some practical ideas are provided to disentangle both hypotheses using these animals.     

Male Choice and Male-male Competition in Idotea baitica (Crustacea,Isopoda)

Sexual selection in mate-guarding Crustacea may involve several processes: male choice, male-male competition, and female choice. To evaluate the relative importance of the different processes in mate choice of the aquatic isopod I. baitica we studied 1) the mate-choice criteria of males, 2) effects of sex ratio on the outcome of the mating contest, and 3) the role of size in male-male interactions. When given a choice between a small and a large female, males most often chose the one that matured earlier for parturial ecdysis. Maturity was a more important choice criterion than female size, but these also correlated positively. Large males had a mating advantage in both male- and female-biased sex ratios; pairing was size-assortative only in the male-biased ratio where guarding was also longer. If an extra male was placed with a precopulatory pair, 30 % take-overs occurred, large males surpassing. Present and earlier work suggests that male size is an asset in both intra- and intersexual interactions. There is little or no direct phenotypic sexual selection on female size: sexual selection for large males presumably contributes to the evolution of sexual size dimorphism in I. baitica.     

Body Size Correlates with Fertilization Success but not Gonad Size in Grass Goby Territorial Males

In fish species with alternative male mating tactics, sperm competition typically occurs when small males that are unsuccessful in direct contests steal fertilization opportunities from large dominant males. In the grass goby Zosterisessor ophiocephalus, large territorial males defend and court females from nest sites, while small sneaker males obtain matings by sneaking into nests. Parentage assignment of 688 eggs from 8 different nests sampled in the 2003–2004 breeding season revealed a high level of sperm competition. Fertilization success of territorial males was very high but in all nests sneakers also contributed to the progeny. In territorial males, fertilization success correlated positively with male body size. Gonadal investment was explored in a sample of 126 grass gobies collected during the period 1995–1996 in the same area (61 territorial males and 65 sneakers). Correlation between body weight and testis weight was positive and significant for sneaker males, while correlation was virtually equal to zero in territorial males. That body size in territorial males is correlated with fertilization success but not gonad size suggests that males allocate much more energy into growth and relatively little into sperm production once the needed size to become territorial is attained. The increased paternity of larger territorial males might be due to a more effective defense of the nest in comparison with smaller territorial males.     

The breeding ecology of Cancer gracilis (Crustacea: Decapoda: Cancridae) and the mating systems of cancrid crabs

Brachyuran crabs in the family Cancridae are found in a variety of habitats, ranging from 'open' sandy or muddy grounds to 'structured' rocky bottoms rich in refuge space. Cancer gracilis Dana inhabits shallow open habitats in the north-east Pacific Ocean. Females in all reproductive conditions aggregate in relatively small areas, where mating activity appears most intense. Maximum mating activity was observed towards the end of the spawning/hatching season (August). Females play an active role in mating, often initiating interactions with the prospective male partner. Mating in the field was non-assortative with respect to size. There was no indication of direct mate choice by either males or females; resolution of male-male competitive interactions was size-based. Males below minimum mating size (77 mm carapace width) had spermatophores in the vas deferens but did not react to receptive females.
Sperm competition is likely because sperm can be retained across moults and multiple copulations are possible within a receptive period. Female spermathecae are of the 'ventral' type (contrary to what has been indicated for cancrids), and during intercourse are beyond the reach of male gonopods. Mechanisms to exclude competing sperm include male-produced sperm plugs and post-copulatory mate guarding.
Results are discussed in the context of the 'structured-to-open' habitat gradient, which may underlay much of the diversity in cancrid reproductive ecology. We argue that, by comparison with C. magister (a larger size sympatric species also found in open habitats), (1) reproductive asynchrony and female gregariousness in C. gracilis increase the environmental potential for polygyny, resulting in a female defence polygynous mating system, and (2) reproductive asynchrony inflates the operational sex ratio, leading to stronger sexual selection.