Should advertising parental care be honest?

Species with paternal care show less exaggerated sexual ornamentation than those in which males do not care, although direct benefits from paternal care can vastly exceed the indirect benefits of mate choice. Whether condition-dependent handicaps can signal parenting ability is controversial. The good-parent process predicts the evolution of honest signals of parental investment, whereas the differential-allocation model suggests a trade-off between the attractiveness of a mate and his care-provisioning. I show that both alternatives can arise from optimal allocations to advertisement, parental investment and future reproductive value of the male, and that the male''s marginal fitness gain from multiple matings determines which option should apply. The marginal gain is diminishing if opportunities for polygyny or extra-pair copulations are limited. Advertisement is then expected to be modest and honest, indicating genetic quality and condition-dependent parental investment simultaneously. Increasing marginal gains are likely to be related to cases where genetic quality has a significant influence on offspring fitness. This alternative leads to differential allocation with stronger advertisement, more frequent extra-pair copulations, and diminished male care. Reliability is also reduced if allocation benefits have thresholds, e.g. if there is a minimum body condition required for survival, or if females use a polygyny-threshold strategy of mate choice.     

Female sand gobies prefer good fathers over dominant males

Female preference for males successful in male-male competition is generally assumed to result in mating with high quality males. Here I report results from an experiment disentangling the effects of intra- and intersexual selection in the sand goby, Pomatoschistus minutus, a marine fish that exhibits paternal care. I show that large males are successful in male–male competition, but contrary to what one would expect, dominants are not preferred by females and are not better at taking care of the eggs. Female preference, however, correlated with the subsequent hatching success of the eggs. Thus, female choice selects for good parenting. Hence, direct benefits in the form of superior paternal care can explain female choice in this species, supporting a good parent process of sexual selection. However, choosing on the outcome of male–male competition does not enable females to mate with the ''best'' males.     

Baffling: a condition-dependent alternative mate attraction strategy using self-made tools in tree crickets

Intense sexual selection in the form of mate choice can facilitate the evolution of different alternative reproductive strategies, which can be condition-dependent. Tree cricket males produce long-distance acoustic signals which are used by conspecific females for mate localization and mate choice. Our study shows that baffling, an acoustic call amplification strategy employed by male tree crickets using self-made tools, is a classic example of a condition-dependent alternative strategy. We show that though most males can baffle, less preferred males, such as smaller and lower-amplitude callers, predominantly use this alternative strategy. Baffling allows these males to increase their call amplitude and advertisement range, which attracts a higher number of females. Baffling also gives these males a mating benefit because females mate for longer durations with them. Our results suggest that the advantage of baffling in terms of gain in the number of sperm cells transferred while mating is primarily limited to less preferred males, thus maintaining the polymorphism of calling strategies in the population. We summarize that baffling is a condition-dependent strategy used by less preferred tree cricket males to obtain mating benefits.     

Genetic covariance between indices of body condition and immunocompetence in a passerine bird

Background  

Condition-dependence is a ubiquitous feature of animal life histories and has important implications for both natural and sexual selection. Mate choice, for instance, is typically based on condition-dependent signals. Theory predicts that one reason why condition-dependent signals may be special is that they allow females to scan for genes that confer high parasite resistance. Such explanations require a genetic link between immunocompetence and body condition, but existing evidence is limited to phenotypic associations. It remains unknown, therefore, whether females selecting males with good body condition simply obtain a healthy mate, or if they acquire genes for their offspring that confer high immunocompetence.     

Odour cues from suitors’ nests determine mating success in a fish

Animals use a range of sensory cues for finding food, avoiding predators and choosing mates. In this regard, the aquatic environment is particularly suitable for the use of olfactory and other chemical cues. Nevertheless, mate choice research, even on aquatic organisms, has focused on visual signals, while chemical cues relevant in sexual selection have been assumed to be ‘intrinsic’ excretions of mate candidates. Here, using the sand goby Pomatoschistus minutus, a small fish with paternal egg care, we investigated the possibility that ‘extrinsic’ chemical cues in the males’ nests could also have a significant contribution to mating success. We found that females strongly avoided laying eggs into nests subject to the odour of Saprolegnia water moulds (an egg infection) and that this effect was independent of the females’ initial, visually based preference for males. To the best of our knowledge, this is the first study to show that chemical cues related to parental failure can play a large role in sexual selection.     

Relatedness,body size and paternity in the alpine newt,Triturus alpestris

Sexual selection has traditionally been investigated assuming that male quality is as skewed as patterns of male reproductive success can sometimes be. Recently, female choice has been investigated under the model of genetic compatibility, which assumes that each individual female has her own 'best' mate and there is no overall optimal choice for all females. We investigated female mate choice in the newt species Triturus alpestris, a member of a genus where female choice has been investigated only within the context of the optimal male (female choice for condition-dependent traits). We provided females with two males that differed in one condition-dependent trait (body size) and overall genetic composition. Both male body size and female body size did not influence paternity, but the degree of genetic relatedness between females and potential mates did. Two components of fitness (fecundity and hatching success) did not differ between singly and multiply sired clutches, indicating that females do not employ polyandry as a means of increasing offspring fitness through genetic bet-hedging. Instead, we hypothesize that females may mate initially for fertility assurance, but prefer less-related males as the most genetically compatible mates.     

Good vs complementary genes for parasite resistance and the evolution of mate choice

Background  

Female mate choice may be adaptive when males exhibit heritable genetic variation at loci encoding resistance to infectious disease. The Hamilton-Zuk hypothesis predicts that females should assess the genetic quality of males by monitoring traits that indicate health and vigor (condition-dependent choice, or CD). Alternatively, some females may employ a more direct method of screening and select mates based on the dissimilarity of alleles at the major histocompatibility loci (we refer to this as opposites-attract, or OA). Empirical studies suggest that both forms of mate choice exist, but little is known about the potential for natural selection to shape the two strategies in nature.     

MALE MATE CHOICE AND THE EVOLUTION OF FEMALE PLUMAGE COLORATION IN THE HOUSE FINCH

The house finch (Carpodacus mexicanus) is a sexually dichromatic passerine in which males display colorful plumage and females are generally drab brown. Some females, however, have a subdued version of the same pattern of ornamental coloration seen in males. In previous research, I found that female house finches use male coloration as an important criterion when choosing mates and that the plumage brightness of males is a reliable indicator of male nest attentiveness. Male house finches invest substantially in the care of young and, like females, stand to gain by choosing high-quality mates. I therefore hypothesized that a female's plumage brightness might be correlated with her quality and be the basis for male mate choice. In laboratory mate choice experiments, male house finches showed a significant preference for the most brightly plumaged females presented. Observations of a wild population of house finches, however, suggest that female age is the primary criterion in male choice and that female plumage coloration is a secondary criterion. In addition, yearling females tended to have more brightly colored plumage than older females, and there was no relationship between female plumage coloration and overwinter survival, reproductive success, or condition. These observations fail to support the idea that female plumage coloration is an indicator of individual quality. Male mate choice for brightly plumaged females may have evolved as a correlated response to selection on females to choose brightly colored males.     

Pectoral fins and paternal quality in sticklebacks

Sexual selection through female mate choice exerts a strong selection pressure on males' sexual traits, particularly when direct benefits are involved. In species with male parental care, one would expect sexual selection to favour paternal quality, for instance through selection on morphological structures which promote quality. We experimentally studied the influence of pectoral fins on paternal quality in male three-spined sticklebacks (Gasterosteus aculeatus L.). After reductions of fin area to different degrees, similar-sized males had to perform a complete reproductive cycle in enclosures in the field. The collected data on fanning behaviour and egg development showed that a reduction in pectoral fin size affected paternal quality probably through an increased beat frequency of the pectorals. Thus, pectoral fins can potentially signal paternal quality to choosy females.     

An experimental test of mate choice for red carotenoid coloration in the marine copepod Tigriopus californicus

Colorful ornaments are hypothesized to have evolved in response to sexual selection for honest signals of individual quality that provide information about potential mates. Red carotenoid coloration is common in diverse groups, and in some vertebrate taxa, red coloration is a sexually selected trait whereby mates with the reddest ornaments are preferred . Despite being widespread among invertebrates, whether red carotenoid coloration is assessed during mate choice in these taxa is unclear. The marine copepod Tigriopus californicus displays red coloration from the accumulation of the carotenoid astaxanthin. Previous research on copepods has shown that astaxanthin provides protection from UV radiation and xenobiotic exposure and that carotenoid production is sensitive to external stressors. Because of the condition dependency of the red coloration, we hypothesized that Tigriopus would use it as a criterion during mate choice. To test this hypothesis, we conducted trials in which males chose between females that were wild-type red (carotenoid-rich algae diet) or white (carotenoid-deficient yeast diet). To control for dietary differences and to isolate the effect of carotenoid coloration, we also presented males with restored-red females fed a carotenoid-supplemented yeast diet. We found that wild-type red females were weakly preferred over white females. After controlling for diet, however, we found that restored-red females were avoided. Our observations do not support the hypothesis that male copepods prefer the carotenoid coloration of females during mate choice. We hypothesize that algal-derived compounds other than carotenoids play a role in mate choice. Red coloration in copepods appears to be a condition-dependent trait that is not assessed during mating.     

Limited condition dependence of male acoustic signals in the grasshopper Chorthippus biguttulus

In many animal species, male acoustic signals serve to attract a mate and therefore often play a major role for male mating success. Male body condition is likely to be correlated with male acoustic signal traits, which signal male quality and provide choosy females indirect benefits. Environmental factors such as food quantity or quality can influence male body condition and therefore possibly lead to condition-dependent changes in the attractiveness of acoustic signals. Here, we test whether stressing food plants influences acoustic signal traits of males via condition-dependent expression of these traits. We examined four male song characteristics, which are vital for mate choice in females of the grasshopper Chorthippus biguttulus. Only one of the examined acoustic traits, loudness, was significantly altered by changing body condition because of drought- and moisture-related stress of food plants. No condition dependence could be observed for syllable to pause ratio, gap duration within syllables, and onset accentuation. We suggest that food plant stress and therefore food plant quality led to shifts in loudness of male grasshopper songs via body condition changes. The other three examined acoustic traits of males do not reflect male body condition induced by food plant quality.     

Differential Allocation by Female Zebrafish (Danio rerio) to Different-Sized Males – An Example in a Fish Species Lacking Parental Care

Organisms allocate resources to reproduction in response to the costs and benefits of current and future reproductive opportunities. According to the differential allocation hypothesis, females allocate more resources to high-quality males. We tested whether a fish species lacking parental care (zebrafish, Danio rerio) expresses male size-dependent differential allocation in monogamous spawning trials. In addition, we tested whether reproductive allocation by females is affected by previous experience of different-quality males, potentially indicating plasticity in mate choice. To that end, females were conditioned to large, small or random-sized males (controls) for 14 days to manipulate females'' expectations of the future mate quality. Females showed a clear preference for large males in terms of spawning probability and clutch size independent of the conditioning treatment. However, when females experienced variation in male size (random-sized conditioning treatment) they discriminated less against small males compared to females conditioned to large and small males. This might suggest that differential allocation and size-dependent sexual selection is of less relevance in nature than revealed in the present laboratory study.     

Chemoreception, symmetry and mate choice in lizards

Research on fluctuating asymmetry (FA)-mediated sexual selection has focused almost exclusively on visual signals and ignored chemical communication despite the fact that many species rely on chemical signals for attracting mates. Female mate choice based on visual traits appears to be rare in lizards. However, the femoral glands of male lizards produce pheromones which might transmit chemical information about an individual's developmental stability. Therefore, we hypothesized that mate choice may be based on chemical cues. We analysed the effect of the developmental stability levels of males on the attractiveness of males' scents to females in a laboratory experiment with the lizard Lacerta monticola. When we offered two males of similar body size, females preferentially associated with the scents of males with low FA in their femoral pores and also with the scents of males with a higher number of femoral pores. This suggested that the females were able to discriminate the FA of the males by chemical signals alone and that the females preferred to be in areas marked by males of high quality, thus increasing their opportunities of mating with males of high quality. We suggest that the quality and/or amount of male pheromones could communicate the heritable genetic quality of a male to the female and thereby serve as the basis for adaptive female choice in lizards.     

Male mate choice relies on major histocompatibility complex class I in a sex‐role‐reversed pipefish

Mate choice for compatible genes is often based on genes of the major histocompatibility complex (MHC). Although MHC‐based mate choice is commonly observed in female choice, male mate choice remains elusive. In particular, if males have intense paternal care and are thus the choosing sex, male choice for females with dissimilar MHC can be expected. Here, we investigated whether male mate choice relies on MHC class I genes in the sex‐role reversed pipefish Syngnathus typhle. In a mate choice experiment, we determined the relative importance of visual and olfactory cues by manipulating visibility and olfaction. We found that pipefish males chose females that maximize sequence‐based amino acid distance between MHC class I genotypes in the offspring when olfactory cues were present. Under visual cues, large females were chosen, but in the absence of visual cues, the choice pattern was reversed. The use of sex‐role reversed species thus revealed that sexual selection can lead to the evolution of male mate choice for MHC class I genes.     

Direct mate choice for simultaneous acoustic and visual courtship displays in the damselfish, Dascyllus albisella (Pomacentridae)

Acoustic signals are well established as key components of mate selection in terrestrial species, but not in aquatic species. It has long been known that damselfish (Pomacentridae) use a combined visual and acoustic display in their courtship.. This study examined several male qualities including individual size, courtship vigor, territory size and complexity, as well as components of the acoustic call including dominant frequency, pulse characteristics and repetition rate. The objective was to determine which male traits were correlated with mating success. Observations made over ten reproductive cycles revealed that female mate choices were not random and that male mating success was correlated with courtship rate (a simultaneous visual and acoustic cue) and the number of neighboring females, but not with male morphological traits, territory quality, or acoustic call structure. These results suggest that females choose mates based on a condition-dependent trait (courtship rate) that advertises quality of paternal care, which supports good parent models of sexual selection, thereby demonstrating the importance of the combined acoustic/visual display for sexual selection in fishes     

Leptin Manipulation Reduces Appetite and Causes a Switch in Mating Preference in the Plains Spadefoot Toad (Spea bombifrons)

Condition- or context-dependent mate choice occurs when females modify their mate preferences depending on their internal or external environment. While the ecological and evolutionary factors that favor the evolution of such plasticity are emerging, relatively little is known of the mechanisms underlying such choice. Here we evaluated whether leptin, a protein hormone involved in the regulation of appetite, might affect the expression of condition-dependent mate choice decisions. To do so, we administered leptin to spadefoot toads, Spea bombifrons, which exhibit condition-dependent mate choice for males of their own species versus congeneric males of S. multiplicata. In particular, poor-condition S. bombifrons are more likely than are good-condition S. bombifrons to prefer S. multiplicata males, but only in environments where hybridization between the two species is beneficial. We found that our leptin treatment reduced appetite in S. bombifrons adults, as was expected from leptin''s known effects on appetite. However, although we predicted that leptin would reduce female preferences for heterospecific males, we found the opposite. In particular, our leptin treatment generated a consistent, repeatable preference for heterospecifics in an environment where females generally prefer conspecifics regardless of condition. These results indicate that leptin has the potential to affect female mate choice, but that it might do so in non-intuitive ways.     

When are good genes good? Variable outcomes of female choice in wax moths

Female lesser wax moths (Achroia grisella) choose males based on characters of their ultrasonic advertisement signals. Because a female''s opportunity to obtain increased somatic benefits by mating with a particular male is limited, we investigated whether females obtain genetic benefits for their offspring via mate choice. Controlled breeding experiments conducted under favourable food and temperature conditions showed that developmental characters are heritable, that sire attractiveness and offspring survivorship are unrelated, but that females mating with attractive signallers produce offspring who mature faster than the offspring of females mating with non-attractive signallers. However, under some unfavourable food or temperature conditions, it is the offspring of females mating with non-attractive males who mature faster; these offspring are heavier as well. Thus, the relationship between male attractiveness and offspring development is not environmentally robust, and support for a good genes model of mate choice in A. grisella is dependent on conditions. These findings suggest genotype–environment interactions and emphasize the necessity of testing sexual selection models under a range of natural environments.     

Preference for Male Traits Differ in Two Female Morphs of the Tree Lizard,Urosaurus ornatus

Non-random female mating preferences may contribute to the maintenance of phenotypic variation in color polymorphic species. However, the effect of female preference depends on the types of male traits used as signals by receptive females. If preference signals derive from discrete male traits (i.e., morph-specific), female preferences may rapidly fix to a morph. However, female preference signals may also include condition-dependent male traits. In this scenario, female preference may differ depending on the social context (i.e., male morph availability). Male tree lizards (Urosaurus ornatus) exhibit a dewlap color polymorphism that covaries with mating behavior. Blue morph males are aggressive and defend territories, yellow males are less aggressive and defend smaller territories, and orange males are typically nomadic. Female U. ornatus are also polymorphic in dewlap color, but the covariation between dewlap color and female behavior is unknown. We performed an experiment to determine how female mate choice depends on the visual and chemical signals produced by males. We also tested whether female morphs differ in their preferences for these signals. Female preferences involved both male dewlap color and size of the ventral color patch. However, the female morphs responded to these signals differently and depended on the choice between the types of male morphs. Our experiment revealed that females may be capable of distinguishing among the male morphs using chemical signals alone. Yellow females exhibit preferences based on both chemical and visual signals, which may be a strategy to avoid ultra-dominant males. In contrast, orange females may prefer dominant males. We conclude that female U. ornatus morphs differ in mating behavior. Our findings also provide evidence for a chemical polymorphism among male lizards in femoral pore secretions.     

Divergence in Female Duetting Signals in the Enchenopa binotata Species Complex of Treehoppers (Hemiptera: Membracidae)

Sexual communication often involves signal exchanges between the sexes, or duetting, in which mate choice is expressed through response signals. With both sexes acting as signalers and receivers, variation in the signals of males and females may be important for mate choice, reproductive isolation, and divergence. In the Enchenopa binotata species complex – a case study of sympatric speciation in which vibrational duetting may have an important role – male signals are species‐specific, females choose among males on the basis of signal traits that reflect species and individual differences, and female preferences have exerted divergent selection on male signals. Here, we describe variation in female signals in the E. binotata species complex. We report substantial species differences in the spectral and temporal features of female signals, and in their timing relative to male signals. These differences were similar in range to differences in male signals in the E. binotata complex. We consider processes that might contribute to divergence in female signals, and suggest that signal evolution in the E. binotata complex may be influenced by mate choice in both sexes.     

A theoretical study on the evolution of male parental care and female multiple mating: effects of female mate choice and male care bias

Male parental care and female multiple mating are seen in many species in spite of the cost they entail. Moreover, they even coexist in some species though polyandry, by reducing paternity confidence of caregiving males, seems to hinder the evolution of paternal care. Previous studies have investigated the coevolutionary process of paternal care and polyandry under various simplifying assumptions, including random mating and random provision of male care. We extend these models to examine possible effects of female mate choice and male care bias, assuming that (a) monandrous females mate preferentially with caregiving males while polyandrous females compromise their preference in order to mate with multiple males and (b) caregiving males tend to direct their care to offspring of monandrous females. Our models suggest that both the female preference and the male bias always favor caregiving males while they may not always facilitate the evolution of monandry.