Facultative paternal investment in the polyphenic beetle Onthophagus taurus: the role of male morphology and social context

Members of a population often differ significantly in theirparental investment. Such variation is generally believed tohave important consequences for mating system evolution andhas been suggested to play an important role in the evolutionof some secondary sexual traits and displays. Recent studiessuggest that individuals are able to adjust the intensity and kindof parental investment they provide according to the breedingconditions they encounter. As a consequence, between-individualvariation in parental investment may depend more on externalconditions than previously thought for these taxa. This mayhave important implications for current perspectives on therole of differential parental investment in the evolution andmaintenance of certain mating systems and sexual selection regimes.Here I quantify patterns of variation in paternal investmentas a function of social conditions in a species of beetle thatis dimorphic for male horn morphology. I demonstrate that undercertain conditions (namely, the absence of other males), paternalassistance covaries with male morphology, with horned males investingsubstantially more time in assisting females than hornless males.I also show that the magnitude of differences in paternal investmentbetween male morphs varies in response to external conditions.In the presence of other males, paternal assistance was negligiblefor both male morphs, who instead invested substantially andequally in mate-securing behaviors. I use my findings to discussthe significance of variation in paternal assistance for onthophaginemating systems and evaluate ideas proposed to explain the evolutionof alternative morphologies in the genus Onthophagus.     

Sexual selection and condition dependence of courtship display in three species of horned dung beetles

Sexual selection has traditionally been divided into competitionover mates and mate choice. Currently, models of sexual selectionpredict that sexual traits are expressed in proportion to thecondition of their bearer. In horned beetles, male contestcompetition is well established, but studies on female preferencesare scarce. Here I present data on male mating success and condition dependence of courtship rate in three species of horn-dimorphicdung beetles, Onthophagus taurus, Onthophagus binodis, andOnthophagus australis. I found that in the absence of malecontest competition, mating success of O. taurus and O. australiswas unrelated to their horn length and body size, whereas inO. binodis horn size had a negative effect but body size hada positive effect on male mating success. Overall, in O. binodismajor morph males had greater mating success than minor morphmales. In all three species male mating success was affectedby courtship rate, and the courtship rate was condition dependent such that when males were manipulated to be in poor conditionthey had lower courtship rates than males that were manipulatedto be in good condition. My findings provide new insight intothe mating systems of horned dung beetles and support an importantassumption in indicator models of sexual selection.     

Proximate determination of male horn dimorphism in the beetle Onthophagus taurus (Coleoptera: Scarabaeidae)

The existence of discrete phenotypic variation within one sex poses interesting questions regarding how such intrasexual polymorphisms are produced and modified during the course of evolution. Approaching these kinds of questions requires insights into the genetic architecture underlying a polymorphism and an understanding of the proximate mechanisms determining phenotype expression. Here we explore the genetic underpinnings and proximate factors influencing the expression of beetle horns – a dramatic sexually selected trait exhibiting intramale dimorphism in many species. Two relatively discrete male morphs are present in natural populations of the dung beetle Onthophagus taurus (Scarabaeidae, Onthophagini). Males exceeding a critical body size develop a pair of long, curved horns on their heads, while those smaller than this critical body size remain essentially hornless. We present results from laboratory breeding experiments designed to assess the relative importance of inherited and environmental factors as determinants of male morphology. Using father–son regressions, our findings demonstrate that horn length and body size of male progeny are not predicted from paternal morphology. Instead, natural variation in an environmental factor, the amount of food available to larvae, determined both the body sizes exhibited by males as adults and the presence or absence of horns. The nonlinear scaling relationship between the body size and horn length of males bred in the laboratory did not differ from the pattern of variation present in natural populations, suggesting that nutritional conditions account for variation in male morphology in natural populations as well. We discuss our results by extending ideas proposed to explain the evolution of conditional expression of alternative phenotypes in physically heterogeneous environments toward incorporating facultative expression of secondary sexual traits. We use this synthesis to begin characterizing the potential origin and subsequent evolution of facultative horn expression in onthophagine beetles.     

The behavioral ecology of threshold evolution in a polyphenic beetle

Facultative expression of alternative male morphologies is thoughtto allow individual males to select the phenotype with the highestfitness gain given their competitive status relative to othermales with which they compete for females. Choice of, or switchingbetween, morphs commonly relies on developmental threshold responses.Evolutionary changes in developmental threshold responses arethought to provide an important avenue for phenotypic diversificationand the evolution of morphological and behavioral novelties.However, the extent to which alternative male phenotypes andtheir underlying threshold responses actually evolve in naturalpopulations is unclear. Likewise, the ecological factors thatshape the evolution of threshold responses in natural populationsare unexplored for most organisms, as are the consequences ofsuch modifications for patterns of morphological diversity.I examined the ecological basis of rapid threshold evolutionin exotic populations of the horn-polyphenic dung beetle Onthophagustaurus. Male O. taurus vary continuously in body size as a functionof larval feeding conditions. Only males that exceed a criticalthreshold body size develop a pair of long horns on their heads,whereas males below this threshold remain hornless. Populationsin two exotic ranges of this species, the eastern United Statesand western Australia, have diverged in the mean threshold bodysize, which has resulted in the evolution of highly divergentand novel horn length–body size allometries in these populations.Populations in a third and previously unstudied exotic rangeof O. taurus in eastern Australia exhibit threshold body sizesroughly intermediate between the eastern U.S. and western Australianpopulations. I tested three hypothesis to explain how differencesin ecological and demographic factors can drive allometric divergencesbetween populations, using data derived from comparative, standardizedsampling of a large number of populations in each exotic range.Results suggest that differences in the intensity of both intra-and interspecific competition have contributed to the evolutionof divergent thresholds in these populations. My results donot support the hypothesis that shifts in threshold body sizesto larger body sizes are a consequence of increases in the meanbody size of competing males. I discuss my results in the contextof Onthophagus mating systems and the evolutionary implicationsof threshold evolution.     

Allometry and proximate mechanisms of sexual selection in photinus fireflies, and some other beetles

Ever since Darwin identified it as the force responsible forthe evolution of exaggerated male characters, sexual selectionhas been the focus of research aimed at understanding the mostbizarre and intriguing morphologies and behaviors in Nature.Two congeneric species in the firefly genus Photinus, P. pyralisand P. macdermotti, afford a unique opportunity to examine theinteraction between sexual and countervailing natural selectionthat act to shape the evolution of mating behavior and bodysize in closely related species with very different courtshipstrategies. Photinus pyralis males emit very bright flashesduring their extended patrolling flights and form ‘loveknots’ of competing suitors, while P. macdermotti males,whose courtships seldom exceed two competing males, produceweaker flashes during a shorter patrolling period. Possibly as a consequence of their scramble competition andlong flights, not only is there an extremely wide range of bodysizes in P. pyralis males, but they also exhibit wing allometricslopes greater than one, and lantern allometries less than one.In contrast, P. macdermotti males do not have allometric slopessignificantly different from one. Small males, when tested inan artificial scramble situation move faster than large males,an advantage in the intense competition that ensues once a femaleis located. Females answer several males in alternation andfail to respond to all male flashes, with the evident consequencethat love knots frequently develop around such fickle females.Allometric relationships in two non-luminescent beetle specieswith non-visual courtship protocols are compared. Visual, sexuallyselected characters showed positive allometric slopes, whilenon-visual characters showed isometry or negative slopes. Datapresented here support the existence of distinct patterns inmodality-specific sexual selection.     

Food, vigilance, and sperm: the role of male direct benefits in the evolution of female preference in a polygamous bird

The adaptive significance of female selection of copulationpartners remains unresolved, particularly in polygamous specieswhere males do not provide paternal care. In these species thepossibility that direct benefits other than paternal care mayplay an important role in the evolution of female choice hasreceived little attention. I tested whether direct benefitsare associated with female choice in the polygamous feral fowl,Gallus g. domesticus, where females prefer socially dominantcopulation partners and males do not care for the young butdo provide females with three commodities: food, vigilance,and sperm. I used a combination of empirical and experimentaldata to show that male propensity to offer food and vigilance,but not sperm, was positively associated with male social status,suggesting that the provision of these resources may be costlyand condition dependent in males. Copulation success was correlatedwith male status but not with the number of feedings a femalereceived from a male, indicating that a female preferred dominantpartners that in general provided any female with more food,rather than partners that provided only her with more food,consistent with the idea that females may use male resourceprovisioning as a proximate mechanism to assess male condition.Together, these results indicate that male resources provisioningis (1) tightly linked to male social status, (2) a potentialindicator of male condition and possibly genetic quality, and(3) a potential criterion for females to select dominant partners,thus playing an important role in the evolution of partner choiceeven in polygamous species lacking paternal care.     

Size, operational sex ratio, and mate-guarding success of the carrion beetle, Necrophila americana

When male insects guard females until oviposition, the benefitsfrom last-male sperm precedence must outweigh the costs of relinquishingadditional fertilizations. The profitability of guarding isincreased when males guard large, fecund females and when femalesare scarce because fewer fertilizations are sacrificed. However,the male reproductive success is not only determined by theprofitability of guarding but also by his ability to maintainguarding. In this study, we used male carrion beetles (Necrophilaamericana) to examine the effects of sex ratio, male relativesize, and female quality on the ability to guard. First, wepresent a model of mate guarding that explores factors, suchas sperm precedence, sex ratio, male size, and female quality,that influence the profitability of postcopulatory riding. Ourmodel predicts that large N. americana males should preferentiallyguard the largest female only when the sex ratio is male biasedand sperm precedence is above 80%. In contrast, small malesgain little from guarding because they are not likely to maintainit and be the last male to mate. Then, we tested these predictionsby manipulating sex ratio, relative male size, and female quality.All males in equal sex ratio and large males in male-biasedsex ratio guarded females significantly longer than did malesin female-biased sex ratio. In male-biased sex ratio, largemales guarded significantly longer and achieved more takeoversthan small males. Large females were guarded longer. The successof guarding males in this beetle depends on their size relativeto other males and the operational sex ratio.     

Endocrine mediation of vertebrate male alternative reproductive tactics: the next generation of studies

In many species of animals, males may achieve reproductive successvia one of several alternative reproductive tactics. Over thepast decade or so, there has been a concerted effort to investigateendocrine mechanisms that underlie such discrete behavioral(and often morphological) variation. In vertebrates, the firstgeneration of studies focused on potential organizational oractivational effects of steroid hormones (Moore, 1991; Mooreet al., 1998). Some of these studies have made it clear that,in addition to circulating hormone levels, one must also considerother aspects of the endocrine system, including hormone receptors,binding globulins and potential interactions among endocrineaxes. In this paper, I review recent work on endocrine mechanismsand suggest possibilities for future investigation. I highlighthow individual variation in sensitivity to environmental conditions,particularly with respect to various stressors, may accountfor the existence of alternative male reproductive phenotypes.Along these lines, I briefly explain the logic behind our workwith male phenotypes of longear sunfish (Lepomis megalotis)that is aimed at determining the tissue-specific distributionand activity of two enzymes that are common to androgen andglucocorticoid metabolism. A major goal of our work is to examinethe potential role of steroidogenic enzymes in the transductionof environmental information to influence the expression ofalternative male reproductive phenotypes.     

Convergent song preferences between female field crickets and acoustically orienting parasitoid flies

Traits that increase the attractiveness of males to femalesoften make them more conspicuous to predators. In the fieldcricket (Gryllus lineaticeps), males are attacked by parasitoidtachinid flies (Ormia ochracea) that locate males through theircalls. Female flies larviposit on crickets and the larvae burrowinto and feed on the cricket, killing the cricket upon emergence.To determine whether traits preferred by females increase amale's risk of attracting a predator, I examined the effectof variation in male singing behavior on mate and predator attraction.Both female crickets and female flies preferred male callingsongs with higher chirp rates, longer chirp durations, and higherchirp amplitudes. In addition, both female crickets and femaleflies preferred male calling songs with higher chirp rates andlonger chirp durations, even when these songs were of loweramplitude. These results suggest that sexual selection by femalechoice will favor the evolution of higher chirp rates and longerchirp durations. However, call types that increase a male'sattractiveness to females also appear to increase a male's riskof attracting parasitoids. Sexual and natural selection appearto have opposing effects on the evolution of male singing behaviorin this species.[Behav Ecol 7: 279-285 (1996)]     

Behavioral plasticity and G × E of reproductive tactics in Nicrophorus vespilloides burying beetles

Phenotypic plasticity is important in the evolution of traits and facilitates adaptation to rapid environmental changes. However, variation in plasticity at the individual level, and the heritable basis underlying this plasticity is rarely quantified for behavioral traits. Alternative behavioral reproductive tactics are key components of mating systems but are not often considered within a phenotypic plasticity framework (i.e., as reaction norms). Here, using lines artificially selected for repeated mating rate, we test for genetic (G × E) sources of variation in reproductive behavior of male Nicrophorus vespilloides burying beetles (including signaling behavior), as well as the role of individual body size, in responsiveness to changes in social environment. The results show that body size influences the response of individuals’ signaling behavior to changes in the social environment. Moreover, there was G × E underlying the responses of males to variation in the quality of social environment experienced (relative size of focal male compared to his rival). This shows that individual variation in plasticity and social sensitivity of signaling behavior can evolve in response to selection on investment in mating behavior, with males selected for high mating investment having greater social sensitivity.     

Experimental Removal of Sexual Selection Reveals Adaptations to Polyandry in Both Sexes

Polyandrous mating is extremely common, yet for many species the evolutionary significance is not fully resolved. In order to understand the evolution of mating systems, it is crucial that we investigate the adaptive consequences across many facets of reproduction. We performed experimental evolution with the naturally polygamous flour beetle Tribolium castaneum subjected to either polyandry or enforced monogamy, creating contrasting selection regimes associated with the presence or absence of sexual selection. After 36 generations, we investigated male and female adaptations by mating beetles with an unselected tester strain to exclude potential effects of male–female coevolution. Reproductive success of focal monogamous and polyandrous beetles from each sex was assessed in separate single male and multiple male experiments emulating the different selection backgrounds. Males and females from the polyandrous regime had more offspring in the experiments with multiple males present than monogamous counterparts. However, in single male experiments, neither females nor males differed between selection regimes. Subsequent mating trials with multiple males suggested that adaptations to polyandry in both sexes provide benefits when choice and competition were allowed to take place. Polyandrous females delayed the first copulation when given a choice of males and polyandrous males were quicker to achieve copulation when facing competition. In conclusion, we show that the expected benefits of evolutionary adaptation to polyandry in T. castaneum depended on the availability of multiple mates. This context-dependent effect, which concerned both sexes, highlights the importance of realistic competition and choice experiments.     

Food availability and parasite infection influence mating tactics in guppies (Poecilia reticulata)

Despite the important effects of diet and parasite infectionon male reproductive behavior, few studies have simultaneouslyaddressed their influence on intrasexual selection (male–malecompetition). We examined the synergistic effects of 2 naturallyvarying environmental factors, lifetime food intake and infection,with the monogenean parasite Gyrodactylus turnbulli on the matingtactics and foraging behavior of male guppies (Poecilia reticulata).We allowed fish to interact directly with each other duringobservations and found that unparasitized males won more intermalecontests, courted females more frequently, and received positiveresponses to courtship displays more frequently than males thathad been infected. Infected males devoted more time to foragingand less time to courtship and competition than uninfected males,suggesting that they were energetically limited and could notincrease reproductive effort despite their reduced expectedlifespan. This interpretation was supported by the observationthat greater food intake ameliorated the negative effects ofparasite infection on courtship effort. Our results have bearingon how natural variation in food availability and parasite prevalenceinfluence geographic variation in reproductive behavior.     

The evolution of mating systems in bark and ambrosia beetles (Coleoptera: Scolytidae and Platypodidae)

The extraordinary array of mating systems in the Scolytidae and Platypodidae has been largely overlooked by researchers interested in the evolution of sexual behaviour. This paper provides the first overview of reproductive behaviour in this important and widespread group, known to most biologists only by the reputations of tree-killing taxa. Referred to generally as ‘bark beetles’, these insects chew egg tunnels inside a variety of (usually dead) plant tissues, though most species are either phloeophagous (breeding in the inner bark of woody plants) or xylomycetophagous (all stages feeding on mutualistic fungi growing on sapwood or heartwood). In most species, permanent records of many aspects of reproductive behaviour are etched in the host; in many, engravings reveal female fecundity, eggs sired per male, hatching success, and offspring survivorship. Each gallery arm represents a good portion of a given female's lifetime reproduction, but in many species females commonly re-emerge to reproduce in one or two additional sites. In most species of bark beetles, each female initiates her own gallery, to be joined later by a male. These monogynous gallery systems are associated with mating systems defined by how long males stay with females: in a few species, males seldom if ever join females under the bark; in the vast majority of species, males stay for part or all of the oviposition period then leave to seek other mates; and a few groups exhibit permanent monogamy, in that both sexes die in their only gallery system. While these patterns emerge from an overview of the world scolytid fauna, the length of male residency has seldom been quantified, and the costs and benefits associated with male mating strategies have not been measured for any bark beetle. Male-initiated monogyny is uncommon in Scolytidae, though the rule in Platypodidae; all instances of which I am aware are summarized from a phylogenetic perspective. Inbreeding polygyny with highly biased sex ratios has arisen at least seven times in Scolytidae. These taxa are usually characterized by males being dwarfed, flightless, and uncommon. Sex determination is known for only a few examples, but both haplodiploidy and diplodiploidy have been reported. Multiple origins of harem polygyny (otherwise rare in invertebrates) add an exciting dimension to the comparative and experimental study of scolytid mating systems. In harem polygynous taxa, males initiate gallery construction. I summarize what little can be learned from the literature about the fine structure of harem polygynous mating systems in bark beetles, and the problem of measuring reproductive success. Data on the nature of harem polygyny in Pityophthorus lautus are presented, illustrating (a) the fluidity of harems; (b) that average eggs laid per gallery arm is relatively unaffected by harem size, but strongly influenced by resource quality; (c) that male egg-gain is strongly correlated with territory quality (a consequence of (b) above); and (d) the temporal patterning of immigration and emigration and its effects on gallery system sex ratios. The second half of this paper is a discussion of the evolution of bark and ambrosia beetle mating Mating systems, emphasizing sexual selection and the role of resources. Male, residence is interpreted as postcopulalory guarding—preventing sexual liaisons with wandering males. Operational sex ratio, encounter rate, synchrony of breeding, ejaculate competition, and spatiotemporal distribution of resources are discussed as evolutionary forces moulding scolytid and platypodid male postmating behaviour. The nature of male male competition is reviewed. The paucity of information on male behaviour in gallery systems is mentioned; whether or not males significantly aid females is not known. Three hypotheses are presented for why females re-emerge, a feature which strongly affects operational sex ratios. Finally, I summarize features of bark beetle existence predisposing them to the evolution of post-inseminative guarding. Male-initiated monogyny presents a puzzle. I propose that most uncontested examples can be explained by monogyny re-evolving from (male-initiated) harem polygyny, and I present an argument for the evolution of harem polygyny leading to the development of male gallery initiation. The evolution of harem polygyny in birds and mammals has attracted considerable attention. The Verner Willson Orians polygyny threshold model is discussed with respect to bark beetles in general and P. lautus in particular. Resource quality is a major factor in P. lautus harem dynamics: the cost to females of joining harems is apparently slight compared to benefits accrued from moving into sites with higher quality inner bark. Female-biased adult sex ratios have been suggested to lead to harem polygyny, and literature and original data pertinent to this hypothesis are examined. The geometric constraints model, based on the polygyny threshold concept but tailored to bark beetles, is proposed to account for the failure of most species to evolve harem polygyny, and testable predictions are derived that interrelate breeding systems, habitat quality, and progeny size. The evolution of Inbreeding is briefly covered, and two routes to inbreeding polygyny are suggested.     

Female-specific color is a signal of quality in the striped plateau lizard (Sceloporus virgatus)

Recent theoretical and empirical studies confirm that male matechoice and/or female–female mate competition can be expressedin the absence of sex-role reversal. Such reproductive patternsmay select for the evolution of female traits that indicatefemale phenotypic or genotypic quality among non–role-reversedspecies. Although attention to the evolution and function offemale ornaments is increasing, additional focus is needed onfemale-specific ornaments (those not expressed in conspecificmales) and on nonavian systems in order to gain a broad understandingof how selection acts directly on ornamentation of female animals.In the striped plateau lizard, Sceloporus virgatus, only femalesdevelop orange throat patches during the reproductive season.The color peaks in expression near the time of ovulation andappears to stimulate male courtship. Here, I examine whetherthis female-specific ornament can be used by males to reliablyassess female phenotypic quality. Using multivariate regressionanalyses, I show that the area of the orange color patch predictsbody condition and mite load, the chroma (i.e., saturation)of the color patch predicts body size, and both patch area andchroma reliably predict average egg mass. Thus, female reproductivecolor may function as a condition-dependent signal, indicatingphenotypic quality to potential mates.     

Geographic variation in male ornamentation and female mate preference in the house finch: a comparative test of models of sexual selection

I compared the mate preferences of female house finches (Carpodacusmexicanus) from populations in which males differ in both plumagecoloration and the extent of ventral pigmentation (patch size).Phylogenetic analysis indicated that the small patch of ventralcoloration displayed by males in some populations is derivedfrom a larger-patched ancestral state. Regardless of the appearanceof males in their own populations, however, females from allpopulations showed a preference for the most brightly coloredmales and males with the largest patches. A reduction in patchsize independent of a change in female mate preference is notconsistent with sensory-bias or reproductive isolation modelsof sexual selection or with general predictions of runaway modelsof sexual selection. In contrast, a lack of congruence betweenfemale mate preference and male trait expression is predictedby the honest advertisement model, with house finches respondingto variation in regional and local access to carotenoid plumagepigments.     

Behavioral correlations across breeding contexts provide a mechanism for a cost of aggression

Identifying correlations among behaviors is important for understandinghow selection shapes the phenotype. Correlated behaviors canindicate constraints on the evolution of behavioral plasticityor may reflect selection for functional integration among behaviors.Obligate cavity-nesting birds provide an opportunity to examinethese correlations because males must defend limited nest cavitieswhile also competing for mating opportunities and providingparental care. Here, I investigated the role of behavioral correlationsin producing a counterintuitive relationship between nest defenseand reproductive success in western bluebirds (Sialia mexicana)such that males that defended their nests most intensely hadthe lowest reproductive success, measured as the number of withinand extrapair offspring that fledged. By experimentally measuringaggression across contexts, I show that this cost of nest defensewas due to the correlated expression of aggression across thecontexts of nest defense and male–male competition coupledwith a trade-off between male–male aggression and parentalcare. In particular, more aggressive males provisioned theirfemales less during incubation and this led to disrupted incubationpatterns and fewer fledged offspring. However, aggressive malesdid not benefit from avoiding parental investment by gainingextrapair fertilizations, and thus, it is unclear how high levelsof aggression are maintained in this population despite apparentcosts. These results suggest that there are constraints to theevolution of plasticity in aggression and emphasize the importanceof considering the integrated behavioral phenotype to understandhow variation in behavior is linked to fitness.     

Multiple signaling in a variable environment: expression of song and color traits as a function of ambient sound and light

Many animals communicate using more than one signal, and several hypotheses exist to explain the evolution of multiple signals. However, these hypotheses typically assume static selection pressures, and previous work has not addressed how spatial and temporal environmental variation can shape variation in signaling systems. In particular, environmental variability, such as ambient lighting or noise, may affect efficacy (e.g., detectability/perception by receivers) of signals. To examine how signal expression varies intraspecifically as a function of habitat characteristics, we evaluated relationships between spatial environmental variation and song and plumage color expression in a tropical songbird, the Red‐throated Ant‐tanager (Habia fuscicauda) in Panama. We recorded male ant‐tanager song, plucked feathers to measure coloration, and quantified the acoustic and light environments from each male's territory. In addition, we took several morphological measurements from each male to assess the potential information content of song and plumage color. We found that males with redder and more saturated crown plumage occurred on darker territories, and males that sang shorter and lower frequency songs occurred on noisier territories. We also found that more colorful males tended to sing longer and lower frequency songs. Finally, we found that song and color correlated similarly with male morphology (e.g., tarsus length, body mass). Altogether, these results indicate that spatial variation in the environment is related to male coloration and song, and that males might be optimizing color and song expression for their particular territorial environment.     

Predation is associated with variation in colour pattern, but not body shape or colour reflectance, in a rainbowfish (Melanotaenia australis)

1. In freshwater fishes, inter-population variation in male phenotype is often associated with differences in predation intensity, but these effects can be difficult to disentangle from environmental influences. 2. The western rainbowfish Melanotaenia australis exhibits marked sexual dimorphism - females are plain with a slender body, while males have striking coloration and are deeper in the body. Male traits differ in expression among populations, but this has not been described or explored in the literature. 3. This paper describes a study designed to test for geographic structuring of male phenotype in M. australis and to determine whether between-population variation in male phenotype is attributable to variation in predation regime, after accounting for environment. 4. We collected data describing habitat, and the size, activity and abundance of predators at sites containing M. australis populations. We then used photography, spectrometry and geometric morphometrics to describe colour pattern, spectral reflectance and body shape in males from these populations. Finally, we used permutation-based multivariate statistics to partition variance in these traits according to environment and predation regime. 5. Downstream environments posed higher predation risk to M. australis. Furthermore, males from these sites consistently exhibited larger cheek spots and fewer coloured lateral stripes than those from upstream sites. Variation in predation regime accounted for a significant proportion of the total variance in these traits (30·9%), after controlling for the effects of environment. 6. Variation in predation regime did not explain variation in reflectance or shape. Environmental variation, however, explained a significant portion of the total variance in reflectance (74·9%), and there was a strong trend towards it explaining a portion of the total variance in body shape (34·9%). 7. We conclude that natural selection by predators may be an important determinant of the evolution of colour pattern variation in M. australis, but not of that of body shape or colour reflectance. 8. Further study of M. australis will complement existing models, which show complex relationships between predation regime, environment and phenotype. Understanding these relationships is prerequisite to predicting the evolution of phenotypic variation in natural systems.     

The Red Queen Visits Sage Grouse Leks

Coevolution between parasites and host is a sufficient althoughnot necessary condition for the evolution of secondary sexualcharacteristics. I review evidence supporting the role of parasitesin the maintenance of lek behavior in Sage Grouse (Centrocercusurophasianus). Males bearing avianmalaria (Plasmodium pediocetii)or lice (Lagopoecus gibsoni or Goniodes centrocerci) have significantlylower reproductive success than noninfected males. Malaria-infectedmales attend leks significantly less frequently and lek attendanceis highly correlated with male reproductive success. Inaddition,males with malaria secured copulations later in the breedingseason with hens that wereyounger, in poorer condition, andless successful than mates of malaria-free males. Lice createhematomas on the air sacs of males which females can detectto avoid lousy males. Results of our field studies are reinforcedby experiments; captive males given antibiotics to reduce parasiteloads are chosen more often by females in arena trials. Ourresults lend empirical support for Hamilton and Zuk's (1982)interpretation of the RedQueen's hypothesis, although many unknownsremain in our understanding of the interaction between parasitesand Sage Grouse.     

Evolutionary reduction in testes size and competitive fertilization success in response to the experimental removal of sexual selection in dung beetles

Sexual selection is thought to favor the evolution of secondary sexual traits in males that contribute to mating success. In species where females mate with more than one male, sexual selection also continues after copulation in the form of sperm competition and cryptic female choice. Theory suggests that sperm competition should favor traits such as testes size and sperm production that increase a male's competitive fertilization success. Studies of experimental evolution offer a powerful approach for assessing evolutionary responses to variation in sexual selection pressures. Here we removed sexual selection by enforcing monogamy on replicate lines of a naturally polygamous horned beetle, Onthophagus taurus, and monitoring male investment in their testes for 21 generations. Testes size decreased in monogamous lines relative to lines in which sexual selection was allowed to continue. Differences in testes size were dependent on selection history and not breeding regime. Males from polygamous lines also had a competitive fertilization advantage when in sperm competition with males from monogamous lines. Females from polygamous lines produced sons in better condition, and those from monogamous lines increased their sons condition by mating polygamously. Rather than being costly for females, multiple mating appears to provide females with direct and/or indirect benefits. Neither body size nor horn size diverged between our monogamous and polygamous lines. Our data show that sperm competition does drive the evolution of testes size in onthophagine beetles, and provide general support for sperm competition theory.