Mechanisms of selective attention in grasshopper choruses: who listens to whom?

In many rhythmically signaling species of acoustic insects and anurans, males form choruses at typical population densities. Recent findings that females may prefer leading calls indicate that the timing of a male's signals relative to those of neighboring choruses is an important component of mate attraction. Within a chorus, however, males cannot time their calls such that they lead all neighbors. Hence, they are expected to show selective attention toward only a subset of the group. We used field playback experiments to determine the incidence of and mechanism(s) responsible for selective attention in choruses of the territorial grasshoppers Ligurotettix coquilletti and L. planum. Our data revealed significant selective attention in both species and are inconsistent with either fixed-threshold or fixed-number mechanisms. Rather, regulation of selective attention by a sliding-threshold mechanism is supported. We discuss these results in the context of the evolution of chorus structure. Received: 24 September 1997 / Accepted after revision: 28 February 1998     

The effect of male coloration on female mate choice in closely related Lake Victoria cichlids (Haplochromis nyererei complex)

We studied the effect of male coloration on interspecific female mate choice in two closely related species of haplochromine cichlids from Lake Victoria. The species differ primarily in male coloration. Males of one species are red, those of the other are blue. We recorded the behavioral responses of females to males of both species in paired male trials under white light and under monochromatic light, under which the interspecific differences in coloration were masked. Females of both species exhibited species-assortative mate choice when colour differences were visible, but chose non-assortatively when colour differences were masked by light conditions. Neither male behaviour nor overall female response frequencies differed between light treatments. That female preferences could be altered by manipulating the perceived colour pattern implies that the colour itself is used in interspecific mate choice, rather than other characters. Hence, male coloration in haplochromine cichlids does underlie sexual selection by direct mate choice, involving the capacity for individual assessment of potential mates by the female. Females of both species responded more frequently to blue males under monochromatic light. Blue males were larger and displayed more than red males. This implies a hierarchy of choice criteria. Females may use male display rates, size, or both when colour is unavailable. Where available, colour has gained dominance over other criteria. This may explain rapid speciation by sexual selection on male coloration, as proposed in a recent mathematical model. Received: 11 April 1997 / Accepted after revision: 27 July 1997     

Sexual selection in the zebra finch Taeniopygia guttata : condition, sex traits and immune capacity

The aim of this study was to test two hypotheses: (1) that condition-dependent secondary sexual traits reflect an individual's immune capacity and (2) that immune capacity and secondary sexual traits covary with primary sex traits, specifically ejaculate quality. We used the zebra finch Taeniopygia guttata as a study species, since the traits that females find attractive in males of this species, song rate and beak colour, are well established. A paired experimental design comprising 31 pairs of brothers was used; for each pair, one male was assigned to a control group provided with ad libitum food and no additional exercise, and the other male was assigned to an experimental group which experienced additional exercise and a reduced rate of food intake. After 11 weeks, the experimental group differed significantly from the control group in a range of variables, including body mass, haematocrit, granulocyte:lymphocyte (G:L) ratio and several primary sex traits, indicating that condition in this group was reduced. Birds in the experimental group showed a differential response to the treatment. We used the rank order in which birds could be captured by an experimenter as an index of condition. Birds easily caught were assumed to be in poorer condition than those which were more difficult to capture. Rank capture order was repeatable and was significantly correlated with the G:L ratio in the experimental group, but not in the control group. In the experimental group, rank capture order was correlated significantly with both secondary sex traits: birds in better condition had redder beaks and a higher song rate. However, beak colour and song rate did not covary significantly, suggesting that these two traits provide different types of information. Secondary sex traits did not covary with primary sex traits or any sperm features. Thus, there was no evidence for Trivers' sexual-competence hypothesis or the phenotype-linked fertility hypothesis. We used four assays of immune capacity, two general (G:L ratio and spleen mass) and two specific [antibody titres to sheep red blood cells (SRBCs) and Brucella abortus (BA)]. The G:L ratio was significantly higher in the experimental group, spleen mass (absolute and relative) did not differ between the groups, anti-SRBC antibody titres were significantly higher in the control group (contrary to expectation), and anti-BA antibody titres were close to being significantly lower in the experimental group. Within the experimental group, there was no evidence that antibody titres covaried with secondary sex traits. Although we demonstrated that beak colour and song rate were condition dependent, our experiment provided no evidence that either of these traits covaried with immune capacity or sperm features. Received: 9 February 1998 / Accepted after revision: 5 September 1998     

Male genital traits and mating interval affect male fertilization success in the water strider Gerris lacustris

Parker's seminal work brought attention to the possibility of postmating sexual selection by non-random fertilization success. Mechanisms for these processes are still only partly understood and there is clearly a need for more studies of intraspecific variation in sperm precedence. Here, we report results from an experimental study of the variation in fertilization success between males of the water strider Gerris lacustris. Genital morphology, male body size, and copulation duration were examined as possible correlates of paternity. The significance of guarding duration was also analysed. Only male genital morphology was correlated to fertilization success. This is one of the first studies showing a relationship between male genital traits and fertilization success, supporting the view that sexual selection may be responsible for the rapid and divergent evolution of genital structures in animals with internal fertilization. The fertilization success of last males varied considerably after double matings with a short mating interval (10 min). Last-male priority ranged from 0 to 100% and usually one of the males involved fertilized almost all the eggs. After double matings with a short mating interval, the proportion of eggs fertilized by the last male averaged 0.68 and was greater than 0.5. In contrast, the average fertilization success was biased towards the first male when the matings were more spread out over time (24 h). These results do not support earlier suggestions of a widespread last-male sperm priority in water striders. Received: 28 July 1998 / Received in revised form: 15 March 1999 / Accepted: 28 March 1999     

Seasonal variation in the relationship between cellular immune response and badge size in male house sparrows (Passer domesticus)

The immunocompetence handicap hypothesis postulates that secondary sexual traits are honest signals of male quality because steroid hormones (such as corticosteroids and sex steroids), which are supposed to favor the development of secondary sexual traits, may also have immunosuppressive effects. Certain secondary sexual traits are not only used as mate choice signals but also play a role as badges of status. In the house sparrow (Passer domesticus), males have a bib of black feathers which is used both as a signal of social status in male-male interactions and by females when choosing a mate. We investigated the relationships between bib size and cellular immune response in male house sparrows during and outside the reproductive season. Males with large badges were found to have lower levels of immunocompetence, as assessed using a T-cell-mediated immunity assay, during the reproductive season, as predicted by the immunocompetence handicap hypothesis. Conversely, in November, the correlation between badge size and cellular immune response was positive, possibly reflecting the better access to trophic resources of large-badged dominant males in winter flocks. Received: 24 September 1998 / Received in revised form: 2 February 1999 / Accepted: 14 February 1999     

Female distribution affects mate searching and sexual selection in male northern water snakes (Nerodia sipedon)

Mating systems and sexual selection are assumed to be affected by the distribution of critical resources. We use observations of 312 mating aggregations to compare mate-searching success of male northern water snakes (Nerodia sipedon) in two marshes in which differences in mating substrate availability resulted in more than fourfold differences in female dispersion. Reproductive males had significantly larger home ranges where females were dispersed than where females were clumped. The number of females encountered by males increased significantly with male home range size where females were dispersed, and decreased significantly where females were clumped. Where females were clumped, males were more likely to encounter other males when they located females. We found no evidence in either population that mate searching was energetically expensive or that males with relatively more energy had larger home ranges. However, males with greater fat reserves at the start of the season participated in more mating aggregations when females were dispersed, suggesting that fat reserves could affect a male’s willingness to attempt mating or to persist in aggregations. When females were dispersed there was weak stabilizing selection acting to maintain male body size (β=–0.14), but strong directional selection favoring larger (β=0.50) and fatter (β=0.37) males. Over 7 years, the intensity of selection favoring larger males varied substantially (β=0.14–1.15), but that variation was not related to variation in the operational sex ratio. We found no evidence of directional selection on either body size (β=0.05) or fat reserves (β=0.10) of males when females were spatially clumped. Overall, the distribution of females had a pronounced effect on male behavior, on the factors that affected male success in locating females, and probably on the extent of sperm competition once females had been located. Received: 23 November 1998 / Received in revised form: 9 August 1999 / Accepted: 18 August 1999     

Is avian humoral immunocompetence suppressed by testosterone?

A key issue in sexual selection theory is how a correlation between male secondary sexual characters and male genetic quality can be maintained. The immunocompetence-handicap hypothesis proposes that testosterone-dependent male characters remain honest signals because of the immunosuppressive effect of elevated steroid hormone levels. The hypothesis requires that physiological levels of testosterone depress immune system function. We quantified testosterone titers and humoral immunocompetence of captive male and female red-winged blackbirds (Agelaius phoeniceus) at three points in the annual cycle (autumn, prebreeding, and breeding). We also conducted an implant experiment on the males to assess the effects of prolonged, above-normal testosterone titers on humoral immune responses. Humoral immunocompetence was measured as secondary antibody production to a non-pathogenic protein antigen, keyhole limpet hemocyanin, using an enzyme-linked immunosorbent assay we developed for A. phoeniceus. Secondary antibody responses of individuals were highly repeatable between sampling periods. Neither physiological nor above-normal levels of plasma testosterone suppressed secondary antibody production. In paired tests of the same individuals between prebreeding and breeding, and between breeding and implant, plasma testosterone increased significantly but secondary antibody responses were unaffected. We are confident in these results because with 80% power, an 11–14% difference in antibody titers would have been detected. There was no relationship between plasma testosterone levels and humoral immunocompetence in free-ranging males tested at the peak of breeding. These results cast doubt on a key assumption of the immunocompetence-handicap hypothesis. Received: 13 April 1998 / Accepted after revision: 7 October 1998     

Exaggerated male eye span influences contest outcome in stalk-eyed flies (Diopsidae)

Evolution of male weapons or status signals has been hypothesized to precede evolution of female mating preferences for those traits. We used staged male fights among three species of Malaysian stalk-eyed flies (Diptera: Diopsidae) to determine if elongated eye span, which is preferred by females in two sexually dimorphic species, influences contest outcome. Extreme sexual dimorphism, with large males possessing longer eye span than females, is shared by Cyrtodiopsis whitei and C. dalmanni. In contrast, C. quinqueguttata exhibits a more ancestral condition – short, sexually monomorphic eye stalks. Videotape analysis of 20-min paired contests revealed that males with larger eye span and body size won more fights in the dimorphic, but not monomorphic, species. To determine if males from the dimorphic species use eye span directly to resolve contests, we competed male C. dalmanni from lines that had undergone artificial selection for 30 generations to increase or decrease eye span. We found that eye span, independently of body size, determines contest outcome in selected-line males. Furthermore, in both dimorphic species, the average encounter duration declined as the eye span difference between contestants increased, as expected if males use eye span to assess opponent size. The number of encounters also increased with age in dimorphic, but not monomorphic, species. Selected-line males did not differ from outbred males in either fight duration or number of encounters. We conclude that exaggerated male eye stalks evolved to influence both competitive interactions and female mating preferences in these spectacular flies. Received: 20 July 1998 / Received in revised form: 2 February 1999 / Accepted: 12 March 1999     

Effects of male dominance and courtship display on female choice in the ring-necked pheasant

Male traits and behaviours acting in mate choice and intrasexual competition are expected to be congruent. When studying their evolution, this often makes it difficult to differentiate between these two components of sexual selection. Studies are therefore needed on mate choice in conjunction with the role of displays and dominance. We present the results from two experiments conducted to investigate the effects of male dominance and courtship displays on female choice in the ring-necked pheasant, Phasianus colchicus, controlling for differences in morphological male traits. We found: (1) different courtship behaviours had different effects on female choice: females were mainly attracted by the feeding courtship behaviour, while another courtship display (the lateral display) was effective in producing the copulation-acceptance response by the females; (2) subordinate males performed the courtship behaviour before females less frequently than dominant males, and females reinforced intrasexual selection by choosing dominant males, and (3) subordinate males in visual contact with a dominant became less attractive to females. The results support the idea (armament-ornament model) that female pheasants may benefit from using traits selected in male-male competition as clues for mate choice. Received: 23 October 1997 / Accepted after revision: 7 October 1998     

Parasite distribution and virulence: implications for parasite-mediated sexual selection

The Hamilton-Zuk hypothesis states that females choosing males with more developed secondary sexual traits, i.e. brighter males, achieve greater fitness if variability in brightness reflects heritable variation in resistance to parasites. However, several factors will affect the likelihood that parasites play a role in sexual selection in given species. Here, using simple models, we show that because of parasite aggregation on a few hosts, only few breeding males would suffer from reductions in brightness due to parasites. Only in cases where parasites are abundant and show low levels of aggregation among their hosts would there be sufficient variability in brightness among breeding males for female choice of bright, resistant males to evolve. In addition, sufficient parasite-induced variability in brightness among breeding males will only occur in host-parasite systems where pathology is linearly related to the number of parasites per host. The presence of males that are uninfected and bright but genetically susceptible to parasites will also influence the fitness advantages obtained by females choosing bright males. If genetic immunity against parasites is rare in the host population, females can probably only benefit from choosing bright males if parasites are common and little aggregated among males. These results greatly limit the generality of the Hamilton-Zuk hypothesis, and suggest that only a small fraction of host-parasite associations could promote the evolution of host mate choice for resistance based on brightness. Correspondence to: R. Poulin