Sex roles and sex ratios in animals

In species with separate sexes, females and males often differ in their morphology, physiology and behaviour. Such sex-specific traits are functionally linked to variation in reproductive competition, mate choice and parental care, which have all been linked to sex roles. At the 150th anniversary of Darwin's theory on sexual selection, the question of why patterns of sex roles vary within and across species remains a key topic in behavioural and evolutionary ecology. New theoretical, experimental and comparative evidence suggests that variation in the adult sex ratio (ASR) is a key driver of variation in sex roles. Here, we first define and discuss the historical emergence of the sex role concept, including recent criticisms and rebuttals. Second, we review the various sex ratios with a focus on ASR, and explore its theoretical links to sex roles. Third, we explore the causes, and especially the consequences, of biased ASRs, focusing on the results of correlational and experimental studies of the effect of ASR variation on mate choice, sexual conflict, parental care and mating systems, social behaviour, hormone physiology and fitness. We present evidence that animals in diverse societies are sensitive to variation in local ASR, even on short timescales, and propose explanations for conflicting results. We conclude with an overview of open questions in this field integrating demography, life history and behaviour.     

SPECIES RECOGNITION AND SEXUAL SELECTION AS A UNITARY PROBLEM IN ANIMAL COMMUNICATION

We investigated patterns of mating call preference and mating call recognition by examining phonotaxis of female túngara frogs, Physalaemus pustulosus, in response to conspecific and heterospecific calls. There are four results: females always prefer conspecific calls; most heterospecific calls do not elicit phonotaxis; some heterospecific calls do elicit phonotaxis and thus are effective mate recognition signals; and females prefer conspecific calls to which a component of a heterospecific call has been added to a normal conspecific call. We use these data to illustrate how concepts of species recognition and sexual selection can be understood in a unitary framework by comparing the distribution of signal traits to female preference functions.     

Evolution of female mating preferences in stalk-eyed flies

Sensory exploitation predicts that female mate preferences existbefore the evolution of exaggerated male ornaments. We testedthis prediction by estimating female preference functions, rematingintervals, and copulation durations for three species of stalk-eyedflies. Two species, Cyrtodiopsis whitei and C dalmanni, exhibitextreme sexual dimorphism in eye span, with eye stalks exceedingbody length in large males. In contrast, C quinqucguttata ofboth sexes possess short eye stalks. Maximum parsimony analysisof 437 basepairs of the 16S mitochondrial ribosomal RNA genefrom 6 Malaysian diopsids reveals that short, sexually monomorphiceye stalks are plesiomorphic in Cyrtodiopsis. Observations ofmultiple copulations by females in paired-choice mating chambersindicated that female C whitei and C. dalmanni exhibit relativepreferences for longer eye stalks such that preference intensityincreases linearly with the difference in eye stalk length betweenmales. Females from the sexually monomorphic species showedno detectable preference for male eye stalk length. Female matingpreferences of bodi sexually dimorphic species exhibited significantrepeatability, as expected if genetic variation underlies thepreference. In addition, female C whitei and C. dalmanni exhibitedshorter copulations, mated more frequently, and rejected fewermating attempts than female C quinqueguttata. Thus, opportunitiesfor sperm competition have increased with acquisition of femalepreferences. We conclude that female sensory bias for maleswith long eye span did not exist in a common ancestor to thesespecies. Instead, female preference and remating propensityeither coevolved with eye span dimorphism or evolved after maleeye stalks elongated.     

INTRASEXUAL COMPETITION ALONE FAVORS A SEXUALLY DIMORPHIC ORNAMENT IN THE RUBYSPOT DAMSELFLY HETAERINA AMERICANA

I studied the sex-limited red spots on the wings of male rubyspot damselflies (Hetaerina americana) in relation to territoriality and fitness in the wild. Both observational and experimental (wing spot manipulation) studies indicated that wing spots were selected through competition among males for mating territories, not through female choice or direct competition for females. Males with naturally or artificially large wing spots were more successful at holding territories and consequently mated at higher rates than males with relatively small wing spots. In contrast, sexual selection on male body size appeared to operate among nonterritorial males at the clasping stage of the mating sequence, perhaps because larger males were better at clasping females forcibly. Of four models proposed to explain the evolution of ornaments through territory competition, only the agonistic handicap model makes predictions consistent with the results of this study.     

Role of male-male competition and female choice in the development of breeding coloration in pupfish (Cyprinodon pecosensis)

The function of male breeding coloration and the conditionsunder which it is expressed were investigated using two experimentalprotocols that differed in the degree of control of social factors.In tank experiments, focal males were separated by opaque orclear partitions or allowed to interact freely without partitions.In lake experiments, males were introduced into opaque or clearenclosures in the lake from which they were collected. Thisdesign tested the relative importance of breeding substrate,social interactions with other males, and presence of femalesand juveniles on the expression of breeding coloration and agonisticand mating behavior in pupfish. Breeding coloration was notexpressed in the absence of conspecifics, even in the presenceof suitable breeding substrate, or in the presence of juveniles.Breeding coloration was poorly developed when only males werepresent, was more intense in the presence of females, and darkenedsignificantly after a spawning bout Males in clear enclosuresin their natural lake habitat were in visual contact with conspecificsof both sexes and developed more intense blue breeding colorationthan males in the tank experiments that were in visual contactonly with other males. Uncon-fined males with territories inthe lake developed the most intense breeding coloration. Breedingcoloration of male pupfish is a facultative trait that is welldeveloped when breeding females are present and plays a majorrole in the attraction and courtship of mates. However, maximumdevelopment of breeding coloration by territorial males suggeststhat intrasexual and intersexual selection act concordantlyon this trait.[Behav Ecol 7: 431-437 (1996)]     

A Comparative Bayes tactic for mate assessment and choice

Models of mate choice tactics have assumed that females randomlyencounter males when collecting information and the informationis perfect. Empirical observations of four bird species showthat females selectively visit males and repeat visits to malesbefore mating. This suggests that the assumptions of previousmodels have been too restrictive. An alternative model of informationgathering and mate choice, which relaxes the assumptions ofrandom encounters and perfect information, is presented. Inthis Comparative Bayes model, the decision of when and fromwhom to collect information is made using Bayesian estimatesof each male's quality. Predictions from the model are that:(1) the occurrence of mate assessment will increase as initialuncertainty about the quality of males increases, as the costof gathering information decreases, and as the signal perceivedby the female becomes a better representation of males' actualqualities; (2) the occurrence of repeat visits to males willbe highest when signals from males are of medium reliability;and (3) the decision of which male to assess will depend onthe estimated qualities of males, prior certainty about eachmale's quality, the reliability of each male's signal, and thecosts of assessment. Simulations compare the fitness outcomesof the Comparative Bayes tactic to other mate choice tactics.The fitness from the Comparative Bayes tactic is significantlyhigher than from the fixed threshold tactic and than from thebest-of-n tactic when the cost of assessment is low. [BehavEcol 7: 451–460 (1996)]     

Host stress influences oviposition preference and performance of a pine sawfly

Abstract.

• 1 The effect of ponderosa pine (Pinus ponderosa Doug]. ex Laws) water stress, measured by xylem water potential, on oviposition preference and egg development of Neodiprion fulviceps was determined by two laboratory experiments in 1987 and 1988.
• 2 In the first experiment, adult females were allowed to choose between foliage from water-stressed, watered, and untreated control trees.
• 3 Significant differences were found between treatments in xylem water potential in 1987 but not in 1988.
• 4 Females chose water-stressed trees and a positive correlation was found between female preference and xylem water potential in 1987 but not in 1988.
• 5 In the second experiment, females were placed in globe cages with a single branch from each treatment and allowed to oviposit.
• 6 Significant differences were found among treatments in egg survival in 1987 but not in 1988.
• 7 These results indicate that sawflies can distinguish between stressed and non-stressed trees and that this behaviour is adaptive in that egg survival was greatest on the preferred treatment.

     

Differential male reproductive success in Douglas fir

Summary Differential male reproductive success was studied in clones at two seed orchards of Douglas fir, Pseudotsuga menziesii (Mirb.) Franco. The performance of tester pollen parents was compared in controlled pollinations with two-parent pollen mixes. Marker pollen homozygous for a rare IDH allele was the genetic marker in each pollen mix. The resulting seeds were analyzed electrophoretically. At both seed orchards, the proportion of seeds sired by tester pollen significantly varied among the tester pollen parents. Tester pollen parents did not perform the same across all seed parents. The significant interaction effect was evidence of male-female complementarity. These results suggest a genetic basis to differential male reproductive success in Douglas fir.