Sperm competition in tropical versus temperate zone birds

Sperm competition represents an important component of post-copulatory sexual selection. It has been argued that the level of sperm competition declines in birds towards the equator. However, to date, sperm competition estimates have been available mainly for avian species inhabiting the northern temperate zone. Here we apply a novel approach, using the coefficient of between-male variation (CV_bm) in sperm size as an index for sperm competition risk, in a comparative analysis of 31 Afrotropical and 99 northern temperate zone passerine species. We found no difference in sperm competition risk between the two groups, nor any relationship with migration distance. However, a multivariate model indicated that sperm competition risk was highest in species with a combination of low body mass and few eggs per clutch. The effect of clutch size was most pronounced in tropical species, which indicates that sperm competition risk in tropical and temperate species is differently associated with particular life-history traits. Although tropical species had lower sperm competition risk than temperate zone species for overlapping clutch sizes, the idea of a generally reduced risk of sperm competition in tropical birds was not supported by our analysis.     

Autosomal variation for male body size and sperm competition phenotypes is uncorrelated in Drosophila melanogaster

Correlations between male body size and phenotypes impacting post-copulatory sexual selection are commonly observed during the manipulation of male body size by environmental rearing conditions. Here, we control for environmental influences and test for genetic correlations between natural variation in male body size and phenotypes affecting post-copulatory sexual selection in Drosophila melanogaster. Dry weights of virgin males from 90 second-chromosome and 88 third-chromosome substitution lines were measured. Highly significant line effects (p<0.001) documented a genetic basis to variation in male body size. No significant correlations were identified between male body size and the components of sperm competitive ability. These results suggest that natural autosomal variation for male body size has little impact on post-copulatory sexual selection. If genetic correlations exist between male body size and post-copulatory sexual selection then variation in the sex chromosomes are likely candidates, as might be expected if sexually antagonistic coevolution was responsible.     

Pitfalls in experiments testing predictions from sperm competition theory

As females of many species mate with more than one male, ejaculates often face competition from the sperm of other males. In recent years, numerous papers have been published on theoretical predictions of evolutionary, behavioural and physiological responses to variation in the strength of sperm competition (SC). These theoretical predictions have also been extensively tested. However, although predictions from SC theory are relatively straightforward, extra caution has to be paid in the design of experiments testing them. One difficulty is for example to disentangle immediate and mean SC risk and intensity. Without carefully designed experiments, it is also very easy to simultaneously increase SC risk and the probability of intense SC--a situation for which we currently have no clear predictions, as the theoretical models to date only assume variation in either SC risk or intensity. In this paper, we discuss these and some other pitfalls related to manipulations of SC risk and intensity and suggest how to avoid them.     

High multiple paternity and low last-male sperm precedence in a hermaphroditic planarian flatworm: consequences for reciprocity patterns

It is difficult to predict a priori how mating success translates into fertilization success in simultaneous hermaphrodites with internal fertilization. Whereas insemination decisions will be determined by male interests, fertilization will depend on female interests, possibly leading to discrepancies between insemination and fertilization patterns. The planarian flatworm Schmidtea polychroa, a simultaneous hermaphrodite in which mating partners trade sperm was studied. Sperm can be stored for months yet individuals mate frequently. Using microsatellites, maternity and paternity data were obtained from 748 offspring produced in six groups of 10 individuals during four weeks. Adults produced young from four mates on average. Reciprocal fertilization between two mates was found in only 41 out of 110 registered mate combinations, which is clearly less than what is predicted from insemination patterns. Multiple paternity was high: > 80% of all cocoons had two to five fathers for only three to five offspring per cocoon. Because animals were collected from a natural population, 28% of all hatchlings were sired by unknown sperm donors in the field, despite a 10-day period of acclimatization and within-group mating. This percentage decreased only moderately throughout the experiment, showing that sperm can be stored and used for at least a month, despite frequent mating and sperm digestion. The immediate paternity a sperm donor could expect to obtain was only about 25%. Male reproductive success increased linearly with the number of female partners, providing support for Bateman's principle in hermaphrodites. Our results suggest that hermaphrodites do not trade fertilizations when trading sperm during insemination, lending support to the view that such conditional sperm exchange is driven by exchange of resources.     

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.     

Male age, germline mutations and the benefits of polyandry

The number of de novo mutations in the germline can be expected to increase with age in males, therefore females might decrease mutation load in their progeny by avoiding mating with older males. Here, I propose that female polyandry can be more effective in decreasing the risk of genetic disorders in progeny than pre‐copulatory mate choice, particularly if sperm competitiveness declines more steeply with age than other traits affecting chances of males to mate. If faster ageing of spermatogenic tissue causes older males to transfer inadequate numbers of functional sperm, polyandry would also benefit females directly.     

Larger testes are associated with a higher level of polyandry, but a smaller ejaculate volume, across bushcricket species (Tettigoniidae)

While early models of ejaculate allocation predicted that both relative testes and ejaculate size should increase with sperm competition intensity across species, recent models predict that ejaculate size may actually decrease as testes size and sperm competition intensity increase, owing to the confounding effect of potential male mating rate. A recent study demonstrated that ejaculate volume decreased in relation to increased polyandry across bushcricket species, but testes mass was not measured. Here, we recorded testis mass for 21 bushcricket species, while ejaculate (ampulla) mass, nuptial gift mass, sperm number and polyandry data were largely obtained from the literature. Using phylogenetic-comparative analyses, we found that testis mass increased with the degree of polyandry, but decreased with increasing ejaculate mass. We found no significant relationship between testis mass and either sperm number or nuptial gift mass. While these results are consistent with recent models of ejaculate allocation, they could alternatively be driven by substances in the ejaculate that affect the degree of polyandry and/or by a trade-off between resources spent on testes mass versus non-sperm components of the ejaculate.     

Biochemical and egg size evolution of freshwater fishes in the Rhinogobius brunneus complex (Pisces, Gobiidae) in Okinawa, Japan

Many colour morphs have been recognized in the Rhinogobius brunneus complex. A recent electrophoretic study has revealed that some of the colour morphs are well differentiated from each other genetically. In Okinawa, egg-size and life-history variation has been found in addition to the colour variation in this species complex. To clarify the nature of this life-history variation, gobies of a fluviatile morph with large eggs as well as three other amphidromous morphs with small eggs, inhabiting a single stream sympatrically in Okinawa, were analysed by electrophoresis and the females were also used for comparison of egg and clutch sizes. Electrophoretic data for 34 loci showed that each morph was separated by a fixed-allele difference for at least one locus, indicating that the four morphs which occurred sympatrically are reproductively isolated from each other. However, Nei's genetic distance between the fluviatile morph and one of the amphidromous morphs was much smaller (0.026) than distances among amphidromous gobies (0.323-0.480). Egg size of the former was by far the largest among the four. These results imply that speciation of the fluviatile morph accompanied by the egg-size increase has been completed rapidly without considerable genetic differentiation.     

A panel of microsatellite markers to study sperm precedence patterns in the emperor dragonfly Anax imperator (Odonata: Anisoptera)

Odonates were the first group of organisms where sperm competition and last male sperm precedence have been identified. With the development of 10 microsatellites for the emperor dragonfly Anax imperator, the function and priority patterns of the multiple sperm storage organs of females can be studied and compared between species in natural populations. In addition, two microsatellite loci developed for the sister species Anax parthenope, are also highly polymorphic in A. imperator. For the presented 12 microsatellite loci, the number of alleles per locus ranged from two to 24. Observed heterozygosity ranged from 0.07 to 0.88.     

Female presence influences sperm velocity in the guppy

As sperm production is costly, males are expected to strategically allocate resources to sperm production according to mating opportunities. While sperm number adjustments have been reported in several taxa, only a few studies investigated whether sperm quality shows adaptive plasticity as well. We tested this prediction in the guppy, Poecilia reticulata. A total of 46 males were initially stripped of all retrievable sperm before being randomly allocated to one of two treatments simulating different levels of mating opportunities (visual contact with females or female deprived). After 3 days, males were stripped and sperm velocity was assayed using Computer Assisted Sperm Analysis. Males in the presence of females produced significantly faster sperm than their counterparts. Implications for the evolution of this ejaculate plasticity in the light of results of sperm competition studies are discussed.     

Ejaculate production schedule and the degree of protandry in fishflies (Megaloptera: Corydalidae)

1. To examine the relationship between male–female emergence patterns and ejaculate dynamics, patterns producing sperm and seminal fluids in male internal reproductive organs, the size of a spermatophore transferred at mating and the fate of spermatophore contents moved into female storage organs were compared among the fishflies (Megaloptera: Corydalidae: Chauliodinae), Parachauliodes continentalis, P. japonicus and Neochauliodes sinensis .
2. Spermatophore contents moved into female storage organs decreased rapidly in P. japonicus and N. sinensis , but hardly at all in P. continentalis . This suggests that the females of the former two species may remate sooner than the latter species as it is known in insects that material remaining in the storage organs mechanically inhibits receptivity to mating.
3. Male P. japonicus and N. sinensis increased in internal reproductive organ mass continuously after adult eclosion, and the spermatophore size produced at the first mating increased with male age. In contrast, the internal reproductive organs of P. continentalis were relatively small and did not increase in mass after emergence. P. continentalis transferred a constantly smaller spermatophore at any copulation than the former two species.
4. Males of P. japonicus and N. sinensis emerged earlier than females, while P. continentalis showed a nearly simultaneous emergence pattern between the sexes. It seems that males of P. japonicus and N. sinensis (more polyandrous than P. continentalis ) inhibit female receptivity for a longer time period by emerging earlier and transferring larger ejaculates. In these two species, the number of sperm ejaculated at the first mating also increased with male age. The increase in sperm number by emerging earlier may be adaptive for the males owing to numerical sperm competition when the female remates.