Changes in sperm stores, ejaculate size, fertilization success, and sexual motivation over repeated matings in the common toad, Bufo bufo (Anura: Bufonidae)

Fertilization is of central importance in the determination of reproductive success for both males and females. In species where males have the chance to mate repeatedly within a short period of time, sperm stocks may become depleted and males may have to carefully economize on their sperm reserves. Also, intensive intrasexual competition for females and repeated matings may lead to exhaustion on the behavioural level. To determine whether the reproductive potential of males is limited and if such a limitation is due to behavioural exhaustion or sperm depletion, we experimentally investigated changes in sperm stores, sperm expenditure, fertilization success, and sexual motivation over three repeated matings in the common toad, Bufo bufo , where the breeding season is short and sequential polygyny occurs. At the end of the breeding season, the number of sperm stored in the testes of males mated repeatedly was close to 50% lower than in testes of unmated males. Ejaculate size, which was estimated by applying a novel method allowing direct quantification, decreased by 88% from first to third matings. We also observed a drop in fertilization success from the first two to third matings by 65%, which was largest in males that had started the reproductive season in bad body condition. Some of these males also showed a decreased interest in females in the third mating round. Our results suggest that sperm depletion and loss of sexual motivation may together set a limit to the reproductive potential of common toad males. The present study draws attention to a limitation in reproductive potential, which may occur more often than currently anticipated and has the potential to strongly influence several aspects of reproductive behaviour.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 361–371.     

Effects of mating order and male size on embryo survival in a pipefish

In species that provide parental care, individuals should invest adaptively in their offspring in relation to the pre‐ and post‐zygotic care provided by their partners. In the broad‐nosed pipefish, Syngnathus typhle L., females transfer large, nutrient‐rich eggs into the male brood pouch during mating. The male broods and nourishes the embryos for several weeks before independent juveniles emerge at parturition. Given a choice, females clearly prefer large partners. Yet, females provide protein‐richer eggs when the same individual mates with a smaller than a larger male. In the present study, we allowed each female to mate with one small and one large male, in alternated order. We found a strong effect of female mating order, with larger clutches and higher embryo mortality in first‐ than second‐laid broods, which may suggest that eggs over‐ripen in the ovaries or reflect the negative effects of high embryo density in the brood pouch. In either case, this effect should put constraints on the possibility of a female being selective in mate choice. We also found that small and large males produced embryos of similar size and survival, consistent with the reproductive compensation hypothesis, suggesting that, in this species, larger males provide better nourishment to the embryos than smaller males. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 639–645.     

Does male secondary sexual trait size reveal fertilization efficiency in Australian Drosophila bipectinata Duda (Diptera: Drosophilidae)?

Males developing relatively large, costly sexually selected traits may be of superior body condition compared to small-ornamented males. Thus, males developing the largest secondary sexual trait in a given environment may also be able to augment their investment into ejaculate quality, and fertilize a larger proportion of a female's eggs. We tested the prediction that the degree of expression of a condition-dependent secondary sexual trait, the male sex comb, in a Cape Tribulation (northeastern Australia) population of Drosophila bipectinata Duda, reveals male ability to fertilize eggs in the absence of sperm competition. This test permitted us also to evaluate whether pre-copulatory sexual selection and fertilization efficiency might act additively to influence male reproductive success because a previous study of the same population demonstrated a positive association between comb size and copulation probability. The results obtained indicate that, although genotypes developing smaller sex combs collectively had a significantly higher rate of insemination failure compared to larger comb genotypes, the hatch rate and the number of eggs laid by females inseminated by the two genotypic categories were not statistically different. The results fail to support the prediction that comb size reveals noncompetitive fertilization efficiency of males in this Australian population.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 406–413.     

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.     

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

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

Increased sperm allocation delays body growth in a protandrous simultaneous hermaphrodite

Sex allocation theory for simultaneous hermaphrodites predicts that hermaphrodites have a fixed amount of reproductive resources to allocate to both sex functions and that they trade off their allocation to both sexes depending on environmental conditions. Ophryotrocha diadema is a simultaneous hermaphroditic polychaete worm which has a protandrous phase prior to the hermaphroditic phase. The ability of adolescent males to compete with mature hermaphrodites for egg fertilization and the costs of an increase in male expenditure during the protandrous phase were tested in experiments where adolescent males had the possibility to fertilize hermaphrodites' eggs. Results document that adolescent males were able to compete with hermaphrodites for egg fertilization and caused the hermaphrodites fitness losses of 31.4%. Adolescent males which fertilized hermaphrodites' eggs had a slower growth rate than males which had no possibility to fertilize eggs. This result indicates that an increased male expenditure is a cost for adolescent males and that, during the protandrous phase, resources are allocated to reproduction at the expense of somatic development.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 149–154.     

Sperm competition in frogs: testis size and a 'sterile male' experiment on Chiromantis xerampelina (Rhacophoridae)

The allometric relationship between body mass and testis mass was calculated using data from 16 genera (37 species) of African and Japanese frogs. Having controlled for body mass, the relative testis mass of Chiromantis xerampelina, Rhacophorus arboreus and R. schlegelli was considerably heavier than predicted (3.8–14.6 times more). All three species have multi-male breeding. Although the result is consistent with sperm competition having selected for increased sperm production in anurans, the phylogenetic distribution of well documented multi-male spawning is confined to the Rhacophoridae. Thus, multi-male mating may have arisen only once effectively reducing the analysis to two data points. However, in the four foam-nesting Rhacophorids whose breeding behaviour has been studied there is also a correlation between relative testis mass and the intensity of sperm competition. This suggests that even within the Rhacophoridae, sperm competition leads to larger testes. Direct evidence for sperm competition in C. xerampelina is provided by a 'sterile male' experiment, which shows that peripheral males are capable of fertilizing eggs.     

Size matters: the (negative) allometry of copulatory duration in mammals

The amount of time taken to copulate varies enormously among mammals. Because copulation likely exposes animals to an increased risk of predation, and uses time and energy that could be spent on foraging, smaller mammals (which are vulnerable to more predators and have a shorter time-to-starvation than larger mammals) should spend less time copulating than do larger mammals. Furthermore, if extended copulation reflects competition among males, then the duration of mating (after correction for body size) should be greater in mammals in which females mate with more than one male. We tested these predictions using comparative data from 113 mammalian species in 85 genera, 40 families, and 14 orders, while controlling for the effect of phylogeny. We found: (1) the relationship between duration of copulation and body size to be negative, not positive; (2) no relationship with inferred multiple mating by females (based upon relative testes mass). We suggest that small mammals may find the sustained maneuvering and body positioning of copulation easier than do large mammals. This hypothesis is supported by an apparently isometric relationship between duration of copulation and ratio of power to mass.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 185–193.     

Convergence and parallelism among cicadas of Argentina and the southwestern United States (Hemiptera: Cicadoidea)

Cicadas of the genera Derotettix in Argentina and Okanagodes in the south-western United States resemble each other in colour, live on salt-tolerant plants ( Atriplex spp. etc.), nearly match the colour of their respective host plants and produce songs above the range of avian hearing. The Argentine cicadas are smaller, but have nearly identical thermal limits for activity measured by the minimum temperature for flight (20–24 °C) and a body temperature at heat torpor (48–49 °C). The species shift activity from basking sites to shade at temperatures above 37 °C, although O .  gracilis rises to a significantly higher temperature (40.7 °C) than its congener (38.2 °C) or Derotettix (37.2 °C). The thermal tolerances are the highest reported for cicada species. A third group using halophytes in Argentina ( Babras sonorivox ) has similar temperature tolerances and is cryptically coloured. The genera are convergent with respect to morphology, coloration, body size, behaviour, habitat choice and host plant selection. The similarities of thermal tolerances and their influence on behaviour can be viewed as parallelism because the underlying mechanisms are the same in all species studied.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 281–288.     

Sperm release and use at fertilization by yellow dung fly females (Scathophaga stercoraria)

Females of many species mate multiple times and store transferred sperm in storage organs. The mechanisms underlying sperm release from the stores at fertilization remain poorly understood, although they are central to an understanding of the female influence on post-copulatory male competition. Using double-mated females of the yellow dung fly, we counted the sperm sticking to the surface of deposited eggs of two successive clutches to obtain insight into the physiological processes associated with fertilization. The number of sperm released to fertilize an egg decreased between the first and second clutches, as well as within clutches from early to late eggs. These results indicate that: (1) sperm are lost from the stores over time independent of egg laying and (2) the number of sperm released depends on the amount of sperm stored. The lower number of sperm on eggs of the second clutches was accompanied by a strong increase of the proportion of sperm adhering to the micropyle region, suggesting that sperm use is more efficient and sperm release better controlled when sperm supply is substantially reduced. Finally, our approach indicates that sperm storage capacity of the female is higher than assumed from counts of spermathecal sperm.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 511–518.     

Covariation between predation risk, body size and fin elaboration in the green swordtail, Xiphophorus helleri

Natural and sexual selection can have either opposing or synergistic effects on the evolution of traits. In the green swordtail Xiphophorus helleri , sexual selection arising from female choice is known to favour larger males and males with longer swords. We examined variation in male and female size and fin morphology among 15 populations that varied in their predation environments. Males and females from populations in which piscivorous fishes were present had longer and deeper bodies than did males and females from populations in which piscivorous fishes were absent. Controlling for a positive effect of body size on sword length, males from populations in which piscivores were present had relatively shorter swords than did males from populations in which piscivores were absent. The associations between morphology and predation environment may be due to direct effects of predation, indirect effects of predation, other sources of selection that covary with predator presence, or other environmental effects on trait expression. These results suggest that while sexual selection favours longer swords, natural selection may have an opposing effect on sword length in populations with predators. Natural selection on body size, however, may act synergistically with sexual selection in populations with predators; both may favour the evolution of larger body size. The body size results for X. helleri contrast with related taxa that have become model systems for the study of life history evolution.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 87–100.     

Inbreeding in three‐spined sticklebacks (Gasterosteus aculeatus L.): effects on testis and sperm traits

Mating between relatives often results in inbreeding depression, and is assumed to have a strong effect on fitness traits such as fertility and gonad/gamete quality. However, data concerning this topic are contradictory and particularly scarce in fishes. Three‐spined sticklebacks (Gasterosteus aculeatus L.) show inbreeding depression in fertilization and hatching success, survival rates, body symmetry and behavioural traits. To date, any knowledge of the impact of inbreeding on males' gonads and gametes is lacking in this species. In the present study, testis and sperm traits were quantified in outbred and inbred males. Overall, these traits were not generally impaired by inbreeding, and this result was not changed by a second/third generation of brother–sister matings. However, testes brightness, a potential measure of oxidative stress, was negatively correlated with sperm number. Additionally, inbred males with higher body condition had significantly brighter testes, whereas their sperm number was significantly negatively correlated with sperm quality (as estimated by head volume). Such a trade‐off did not appear in outbred males. The comparatively small impact of inbreeding on testis and sperm traits might be explained by the low number of inbred individuals that reached the reproductive phase. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 510–520.     

Size-dependent mating strategies and the risk of cannibalism

The role of sexual selection in determining the nature and direction of sexual size dimorphism may depend upon the timing of sexual selection, and this may also influence the variation in male size. For example, selection through sperm competition favours smaller males in the highly sexually size dimorphic orb-weaving spider Nephila edulis , whereas larger males are better able to exclude their smaller rivals from the central hub of the web where mating takes place. We investigate experimentally the role of body size and hub tenure in determining male fertilization success when males of different sizes compete for a single female over a 24-h period that includes a period of darkness. Our results confirm that small and large males obtain similar paternity share but that, in contrast with previous studies, hub tenure does not translate into greater paternity share. Unexpectedly, smaller males are at greater risk of postmating sexual cannibalism than larger males, suggesting that natural selection through sexual cannibalism may place a lower limit on male size.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 355–363.     

Body size in proboscideans, with notes on elephant metabolism

Mass estimates for a number of fossil proboscideans were computed using regression analyses on appendicular bones to body mass, for seven specimens of modern elephants, for which body masses had been recorded prior to death. The marked differences in physical proportions between extant Loxodonta and Elephas , implying substantial differences in body mass at any given shoulder height, were not present in their long bone parameters. Length and least circumferences proved to be the best parameters for prediction of body mass. Some extinct proboscideans, notably certain Mammuthus and Deinotherium , were much larger than extant elephants. Both the basal and the field metabolic rates of extant elephants are lower than predicted for a hypothetical mammal, in accordance with their body size and subsistence on low-quality foods. The feeding quantities often ascribed to extant wild elephants are exaggerated, and would in fact have sufficed to nourish much larger species.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 523–549.     

Sperm quality, secondary sexual characters and parasitism in roach (Rutilus rutilus L.)

According to sperm competition models, a male spawning in a disfavoured role should have spermatozoa with higher velocity but shorter longevity compared with a male spawning in a favoured role. Moreover, immunosuppressive androgens are needed to produce both secondary sexual characters and sperm cells. The 'sperm protection' hypothesis suggests that the immunosuppressive action of androgens has evolved to protect haploid spermatozoa, which are antigenic, from autoimmune attacks. Therefore, a male with high sexual ornamentation may be more susceptible to diseases but may possess better quality ejaculate than his less ornamented rival. We studied sexual ornamentation (breeding tubercles), ejaculation quality (sperm concentration, longevity and spermatozoal velocity) and intensity of parasitism in the cyprinid, Rutilus rutilus . Sperm longevity and spermatozoal velocity were positively correlated. Males having elaborated sexual ornamentation had longer-lived sperm and more Myxobolus mülleri parasites in the liver compared with males with low ornamentation. However, no difference was found between males with different degrees of ornamentation with respect to sperm concentration, spermatozoal velocity or other parasites. Since the highly ornamented males had higher sperm longevity, the present results are partly consistent with the predictions of the sperm competition models and the 'sperm protection' hypothesis.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 111–117.     

Condition-dependent spermatophore size is correlated with male's age in a bushcricket (Orthoptera: Phaneropteridae)

During mating, male bushcrickets transfer a spermatophore that consists of a sperm-containing ampulla and a product of the accessory glands, the spermatophylax, which females directly ingest. In the present study, we demonstrate male spermatophore allocation in the bushcricket Poecilimon zimmeri . Males of this species show condition-dependent spermatophore investment. This investment depended upon the age at first mating of males, with older individuals transferring larger spermatophores than younger ones of the same body mass. Independently of age, heavier males transfer larger spermatophores, but the size of males (as measured by femur length) was not a good predictor. Heavier males allocate a lower proportion of their mass to spermatophores and reach their maximal investment point earlier than less heavy males. Spermatophylax production levelled off to a species specific maximum earlier than that of sperm investment (measured as ampulla mass), suggesting that males face high levels of sperm competition.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 354–360.     

Life history reaction norms to time constraints in a damselfly: differential effects on size and mass

Optimality models predict that, under a time constraint, organisms should accelerate development, and preferably so by increasing growth rate, to keep size at emergence constant. Unfortunately, most tests did neglect genetic constraints and interchanged mass with body size which may explain mixed support for some of the models' predictions. We imposed time constraints on full sibling larvae of the damselfly Enallagma cyathigerum by manipulating day length regimes. Under a time constraint, larval development and growth rate based on size indeed were faster. This made it possible to keep size at emergence constant, despite the shorter development time. Interestingly, under a time constraint, growth rate based on mass was not increased and larvae had a lower mass at emergence. We see two reasons for this difference between body mass and size. First, size is fixed at emergence, while mass can still increase in many insects. Secondly, genetic constraints may have contributed to different responses for size and mass. In general, our results strongly suggest discriminating between size and mass when testing life history responses.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 187–196.     

Who's your daddy? Paternity testing reveals promiscuity and multiple paternity in the carnivorous marsupial Dasyurus maculatus (Marsupialia: Dasyuridae)

Female promiscuity is common among mammals but its advantages, particularly for marsupials, remain unclear. Using microsatellite DNA from pouch young of known mothers, we identified the most likely fathers of 25 wild spotted-tailed quolls ( Dasyurus maculatus ) from six litters. We aimed to determine whether young within the same litter had different fathers, and whether breeding success of males was associated with large body mass (consistent with inter-male competition) or scrotal width (consistent with sperm competition). We also explored the possible influence of promiscuity on relatedness within litters. Finally, we used data on paternity and relatedness to make inferences regarding movement and dispersal.
Four litters were sired by more than one male, and three males sired offspring in more than one litter. Known fathers had higher body mass, but not scrotal width, than males of unknown paternity status, suggesting that males may compete for access to females. Sires were less related to dams than expected by chance, and litters with multiple paternity had lower relatedness than litters sired by a single male.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 1–7.