Stronger compensatory growth in a permanent-pond Lestes damselfly relative to temporary-pond Lestes

Compensatory growth where animals compensate for time stress or transient nutritional or thermal stress by accelerating their growth rate is widespread. We know, however, relatively little about the evolution and ecological correlates of compensatory growth. For this we need studies on congeneric species with known phylogenetic relationships that also focus on the associated largely understudied costs. Here we tested for compensatory growth and associated costs in response to time stress (manipulated by photoperiod) and a transient period of starvation or cooling in larvae of the permanent-pond damselfly Lestes eurinus , and compare the results with former studies on temporary-pond Lestes . Larvae showed full compensation in body mass at emergence for all combinations of time stress and starvation or cooling. Unexpectedly, compensatory growth to starvation or cooling was not stronger under time stress. Instead, males under time stress delayed emergence after these transient stressors. In line with a stronger compensatory growth response to time stress than to the other stressors, physiological costs in terms of a reduced investment in immune response (measured as phenoloxidase activity) and energy storage (measured as fat content) were detected only under time stress. Compared to temporary-pond Lestes , L. eurinus showed stronger compensatory growth to time stress. We hypothesize that the stronger compensatory (growth) response in permanent-pond Lestes co-evolved with their derived slower lifestyle when they invaded permanent ponds.     

Sexual size dimorphism in anurans

Several hypotheses have been proposed to explain the direction and extent of sexual size dimorphism in anurans (in which males are usually smaller than females) as a result of sexual selection. Here, we present an analysis to test the hypothesis that sexual dimorphism in anurans is largely a function of differences between the sexes in life-history strategies. Morphological and demographic data for anurans were collected from the literature, and the mean size and age in each sex were calculated for 51 populations, across 30 species and eight genera. Comparisons across 14 Rana species, eight Bufo species and across the genera showed a highly significant relationship between size dimorphism, measured using the female-male size ratio, and mean female-male age difference. A comparison of a subset of 17 of these species for which phylogenetic information was available, using the method of independent contrasts, yielded a similar result. These results indicate that most of the variation in size dimorphism in the anura can be explained in terms of differences in the age structure between the sexes in breeding populations. If sexual selection has an effect on size dimorphism in anurans, it is likely to be only a secondary one.     

Sexual selection explains sex-specific growth plasticity and positive allometry for sexual size dimorphism in a reef fish

In 1950, Rensch noted that in clades where males are the larger sex, sexual size dimorphism (SSD) tends to be more pronounced in larger species. This fundamental allometric relationship is now known as ‘Rensch''s rule’. While most researchers attribute Rensch''s rule to sexual selection for male size, experimental evidence is lacking. Here, we suggest that ultimate hypotheses for Rensch''s rule should also apply to groups of individuals and that individual trait plasticity can be used to test those hypotheses experimentally. Specifically, we show that in the sex-changing fish Parapercis cylindrica, larger males have larger harems with larger females, and that SSD increases with harem size. Thus, sexual selection for male body size is the ultimate cause of sexual size allometry. In addition, we experimentally illustrate a positive relationship between polygyny potential and individual growth rate during sex change from female to male. Thus, sexual selection is the ultimate cause of variation in growth rate, and variation in growth rate is the proximate cause of sexual size allometry. Taken together, our results provide compelling evidence in support of the sexual selection hypothesis for Rensch''s rule and highlight the potential importance of individual growth modification in the shaping of morphological patterns in Nature.     

Sexual size dimorphism in parasitoid wasps

Sexual dimorphism in body length and proportion of overlap between the ranges of body length for males and females were estimated for 361 species of parasitoid wasps from 21 families. In most species, females are generally larger than males, though the range of male and female sizes overlap. Species in the family Ichneumonidae differ significantly from species in other families in three ways: (1) ichneumonids on average are larger, (2) in most species, females are generally smaller than males, and (3) on average, proportion overlap between the ranges of body length for males and females is greater. At present, there is a paucity of life history data on parasitoid wasp species for which size dimorphism is known. Thus it is not clear why ichneumonids differ from species in other families. Possible evolutionary explanations for variation in dimorphism among parasitoid wasp species are discussed.     

Sexual dimorphism of tooth size in anthropoids

We have examined the size of the canine and postcanine teeth of cebid and catarrhine primates in relation to each other, to jaw size and to body weight. We have found that the canine size of males is large enough to be limited by jaw shape and size. A large contribution of P4 to the postcanine row is associated with smaller canines in males. Neither factor seems to limit canine size in females. The females of a small number of species possess enlarged canines. Much of the variation of the postcanine row can be described by the ratio of the (nominal) crown areas of M1 to M3. This ratio is monomorphic which conforms with the general lack of dietary dimorphism in primates. A brief discussion of the evolution of canine size is offered with a new suggestion to account for canine reduction in male hominids.     

Sexual size dimorphism in Asian colobines revisited

Asian colobines exhibit a wide range of sexual dimorphism in body mass. Some species are monomorphic, whereas others are strongly dimorphic. Strong sexual dimorphism is generally viewed as the consequence of intense male contest competition over access to mates, but this idea appears not to explain variation in sexual dimorphism in Asian colobines. Our results show that modular colobines, i.e. species in which social units aggregate into higher‐level bands or often associate, have significantly higher levels of sexual dimorphism in body mass than the nonmodular ones. This finding was corroborated by means of phylogenetically controlled methods and multiple regression analyses. The results suggest that living in a modular society intensifies the contest competition among males, which is further exacerbated by the continuous presence of all‐male units. Am. J. Primatol. 71:609–616, 2009. © 2009 Wiley‐Liss, Inc.     

Sexual size dimorphism and sexual selection in primates

1. In most animals, females are larger than males. Paradoxically, sexual size dimorphism is biased towards males in most mammalian species. An accepted explanation is that sexual dimorphism in mammals evolved by intramale sexual selection. I tested this hypothesis in primates, by relating sexual size dimorphism to seven proxies of sexual selection intensity: operational sex ratio, mating system, intermale competition, group sex ratio, group size, maximum mating percentage (percentage of observed copulations involving the most successful male), and total paternity (a genetic estimate of the percentage of young sired by the most successful male).
2. I fitted phylogenetic generalised least squares models using sexual size dimorphism as the dependent variable and each of the seven measures of intensity of sexual selection as independent variables. I conducted this comparative analysis with data from 50 extant species of primates, including Homo sapiens, Pan troglodytes, and Gorilla spp.
3. Sexual dimorphism was positively related to the four measures of female monopolisation (operational sex ratio, mating system, intermale competition, and group sex ratio) and in some cases to group size, but was not associated with maximum mating percentage or total paternity. Additional regression analyses indicated that maximum mating percentage and total paternity were negatively associated with group size.
4. These results are predicted by reproductive skew theory: in large groups, males can lose control of the sexual behaviour of the other members of the group or can concede reproductive opportunities to others. The results are also consistent with the evolution of sexual size dimorphism before polygyny, due to the effects of natural, rather than sexual, selection. In birds, the study of molecular paternity showed that variance in male reproductive success is much higher than expected by behaviour. In mammals, recent studies have begun to show the opposite trend, i.e. that intensity of sexual selection is lower than expected by polygyny.
5. Results of this comparative analysis of sexual size dimorphism and sexual selection intensity in primates suggest that the use of intramale sexual selection theory to explain the evolution of polygyny and sexual dimorphism in mammals should be reviewed, and that natural selection should be considered alongside sexual selection as an evolutionary driver of sexual size dimorphism and polygyny in mammals.

     

Sexual size dimorphism and Rensch's rule in Canidae

The size variation between males and females of a species is a phenomenon known as sexual size dimorphism (SSD). The observed patterns of variation in SSD among species has led to the formulation of Rensch's rule, which establishes that, in species showing a male size bias, SSD increases with an increase in the body size of the species. However, for species in which there is a female size bias, the SSD would decrease when the body size of the species increases. In the present study, we examined the variation in body size and SSD of 33 species of canids from estimates of body mass and body length. We studied its relationship with life‐history characteristics and tested Rensch's rule using phylogenetic generalized least squares and phylogenetic reduced major axis regressions, respectively. We observed the existence of correlation between body mass and body length, although the SSDs from these estimators are uncorrelated. SSD did not show the pattern predicted by Rensch's rule. SSD also did not show any correlation with life‐history traits. It is likely that the low SSD observed in canids is related to the monogamy observed in the family, which is a rare situation in mammals.     

Sexual size dimorphism in parasitic oxyurid nematodes

As in many invertebrates, female oxyurids are larger than male. Sexual size dimorphism (SSD) of oxyurid nematodes (the hosts of which are both invertebrate and vertebrate), is investigated regarding body size of both host and parasite. SSD of parasites appeared to be weakly, but not significandy, correlated with invertebrate and vertebrate host body size. However, this study reveals a different pattern for SSD with respect to either type of host. SSD does not increase in tandem with body size in vertebrate parasites either at the level of species or genus. SSD is much more pronounced in Syphaciidae than in Heteroxynematidae, two families of vertebrate parasites exhibiting different modes of transmission (members of the Syphaciidae are transmitted through perianal contamination). SSD is investigated in one monophyletic group of parasites of primates, for which a phylogeny is known. Independent comparisons method is used and we find that the body size of female parasite is strongly correlated with that of the male. The hypoallometry (slope<1) of the relationship suggests that the SSD is not linked to an increase of parasite body size. Moreover, there is no influence of host body size on parasite SSD. The pattern in parasites of invertebrates is different. First, SSD has been found to increase with parasite body size in two groups of invertebrate parasites: the oxyurids of Dictyoptera and Coleoptera. Second, female body size of invertebrate parasites is not correlated with male body size either at genus or species level. Finally, the evolution of SSD is discussed in relation to the demographic patterns of invertebrate parasites and the haplodiploid mode of reproduction of these parasitic nematodes.     

Sexual size dimorphism and male combat in snakes

Summary This paper reviews published literature on snakes to test the hypothesis that large male size, relative to female size, evolves because of the advantage it confers in male combat. Analysis of the data reveals a high correlation between the occurrence of male combat, and sexual dimorphism in which the male is the larger sex. This correlation holds (i) within the total sample of snake species (n=224), (ii) within the family Colubridae (n=134), and (iii) in a comparison between the eight families of snakes for which data are available. These results strongly support the hypothesis that large male size is an adaptation to intrasexual competition. The analysis also shows that females are larger than males in about 66% of snake species, that male combat is known in only about 15% of species, and that both sexual size dimorphism and the incidence of male combat tend to be distributed along taxonomic lines.     

Sexual selection and sexual size dimorphism in animals

Sexual selection is often considered as a critical evolutionary force promoting sexual size dimorphism (SSD) in animals. However, empirical evidence for a positive relationship between sexual selection on males and male-biased SSD received mixed support depending on the studied taxonomic group and on the method used to quantify sexual selection. Here, we present a meta-analytic approach accounting for phylogenetic non-independence to test how standardized metrics of the opportunity and strength of pre-copulatory sexual selection relate to SSD across a broad range of animal taxa comprising up to 95 effect sizes from 59 species. We found that SSD based on length measurements was correlated with the sex difference in the opportunity for sexual selection but showed a weak and statistically non-significant relationship with the sex difference in the Bateman gradient. These findings suggest that pre-copulatory sexual selection plays a limited role for the evolution of SSD in a broad phylogenetic context.     

Sexual dimorphism and human tooth size differences

Current male/female differences in tooth size are due to the male/female differences in body bulk that exist in any given human population. These differences are residues of the sexual dimorphism that was maintained for adaptive reasons during the Middle Pleistocene. Late in the Pleistocene the development of food processing techniques led to the reduction of both male and female dental dimensions. Dental sexual dimorphism, however, was maintained until the very end of the Pleistocene when the hunting of large game animals by crude techniques was replaced by a focus on great numbers of small game caught by more sophisticated means and by an increasing utilization of plant foods. The subsequent reduction in dimorphism represents the actions of the Probable Mutation Effect operating under conditions of relaxed selection. The conclusion offered is that the smallest degree of sexual dimorphism visible in the modern world is to be found among those populations that are separated by the greatest interval of time from precursors who depended for their survival on a Pleistocene big game hunting mode of subsistence.     

Effect of lamellae autotomy on survival and foraging success of the damselfly Lestes sponsa (Odonata: Lestidae)

Damselfly larvae can autotomize their caudal lamellae to escape predation. Costs of caudal lamellae autotomy were investigated by directly manipulating lamellae condition of Lestes sponsa in laboratory experiments. Larvae without lamellae had higher predation mortality in the presence of Notonecta. Both lamellae loss and larval density increased the probability of being cannibalized. The results suggest that the increased vulnerability after lamellae loss resulted from a reduced escape performance. Larvae were less mobile after lamellae loss or in the presence of a predator, but the decrease was no longer significant when both factors were combined. This indicates that larvae compensate for the increased predation risk with a fixed response. Both lamellae loss and predator presence reduced hunting success, but the decrease after lamellae loss was only significant in the absence of a predator. The fitness consequences of these effects for both the larval and adult stages are discussed. In general, the data strongly suggest that lamellae autotomy plays a role in population regulation of damselflies. Received: 1 April 1998 / Accepted: 28 August 1998     

Sex ratio, sex-specific chick mortality and sexual size dimorphism in birds

It has been suggested that sexual size dimorphism (SSD) may influence sex ratios at different life stages. Higher energy requirements during growth associated with larger body size could lead to a greater mortality of the larger sex and ultimately to an overproduction of the smaller sex. To explore the associations between SSD and hatching and fledging sex ratio we performed a species-level analysis and a phylogenetically controlled analysis, based on 83 bird species. Overall, there was a significant inverse relationship between the degree of SSD and the proportion of males at hatching and fledging. Sex-specific mortality related to SSD showed a weak but persistent negative tendency, suggesting a mortality bias towards the larger sex. These results suggest that changes in relation to SSD may take place mainly at the conception stage, but could be adjusted during growth. However, conclusions should be treated cautiously as these relationships weaken when additional variables are considered.     

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.     

Female-biased sexual size dimorphism in the yellow-pine chipmunk (Tamias amoenus): sex-specific patterns of annual reproductive success and survival

Sexual size dimorphism is ultimately the result of independent, sex-specific selection on body size. In mammals, male-biased sexual size dimorphism is the predominant pattern, and it is usually attributed to the polygynous mating system prevalent in most mammals. This sole explanation is unsatisfying because selection acts on both sexes simultaneously, therefore any explanation of sexual size dimorphism should explain why one sex is relatively large and the other is small. Using mark-recapture techniques and DNA microsatellite loci to assign parentage, we examined sex-specific patterns of annual reproductive success and survival in the yellow-pine chipmunk (Tamias amoenus), a small mammal with female-biased sexual size dimorphism, to test the hypothesis that the dimorphism was related to sex differences in the relationship between body size and fitness. Chipmunks were monitored and body size components measured over three years in the Kananaskis Valley, Alberta, Canada. Male reproductive success was independent of body size perhaps due to trade-offs in body size associated with behavioral components of male mating success: dominance and running speed. Male survival was consistent with stabilizing selection for overall body size and body size components. The relationship between reproductive success and female body size fluctuated. In two of three years the relationship was positive, whereas in one year the relationship was negative. This may have been the result of differences in environmental conditions among years. Large females require more energy to maintain their soma than small females and may be unable to maintain lactation in the face of challenging environmental conditions. Female survival was positively related to body size, with little evidence for stabilizing selection. Sex differences in the relationship between body size and fitness (reproductive success and survival) were the result of different processes, but were ultimately consistent with female-biased sexual size dimorphism evident in this species.     

Sexual size dimorphism and phylogeny in North American minnows

Sexual size dimorphism (SSD) is predicted to vary across mating systems. A previous study examined a model of SSD in fishes as it relates to three mating system variables: probability of sperm competition, male territorial guarding, and male-male contest. I tested the ability of these variables to predict SSD in North American freshwater minnows, after controlling for phylogenetic effects by an independent contrasts method. Across 58 species only male territorial guarding was significandy related to SSD in a stepwise multiple regression. When tested for 26 genera and subgenera, both male territorial guarding and male-male contest were significant in the model. The concentrated-changes test revealed that character changes in SSD (from males the same size or smaller than females, to males larger than females) were more concentrated on branches with presence of male guarding (similar results were found for changes in SSD and presence of sperm competition), at the species and genus levels. Both comparative approaches demonstrated that male guarding and male-male contest variables are linked to SSD in minnows.     

Sexual size dimorphism in three North Sea gadoids

Existing biological data on whiting Merlangius merlangus, cod Gadus morhua and haddock Melanogrammus aeglefinus from a long‐term international survey were analysed to address sexual size dimorphism (SSD) and its effect on their ecology and management. Results show that SSD, with larger females of the same age as males, is a result of higher growth rates in females. A direct consequence of SSD is the pronounced length‐dependent female ratio that was found in all three gadoids in the North Sea. Female ratios of the three species changed from equality to female dominance at specific dominance transition lengths of c. 30, 35 and 60 cm for M. merlangus, G. morhua and M. aeglefinus, respectively. An analysis by area for M. merlangus also revealed length dependence of female ratios. SSD and length‐dependent female ratios under most circumstances are inseparable. Higher overall energy demand as well as a higher energy uptake rate must result from the observed SSD and dimorphism in growth rates. Potential processes related to feeding, locomotion and physiology are proposed that could balance the increased energy investment of females. Potential consequences of SSD and length dependency of female ratios are the reduction of the reproductive potential of a stock due to size‐selective fishing and biased assessment of the true size of the female spawning stock that could distort decisions in fisheries management.     

Sexual dimorphism in the size of nuclei with dioecious plants

Measurements of the size of the nuclei of dioecious plants showed that the nuclei of male and female plants differ in agreement with the larger quantity of chromatin. The male. plants ofRumex acetosella andMelandrium album had larger nuclei, their Y chromosome being larger than the X chromosome, the same is true forRumex acetosa where the Y chromosome is smaller but there are two in the set.Ginkgo biloba had larger female nuclei because the Y chromosome was smaller than the X. The curves obtained by grouping all the nuclei of both sexes never had two peaks with regard to the small differences between the classes of maximum frequency.