Testing alternative vicariance scenarios in Western Mediterranean discoglossid frogs

Dated molecular phylogenies are often used to interpret evolutionary history with respect to paleogeographic events. Where more than one interpretation is possible, it is desirable but difficult to assess the alternatives in an objective manner. The present work demonstrates a formalized method for testing molecular clock calibrations and biogeographic scenarios based on them. We assessed the plausibility of several previously published biogeographic hypotheses, using the frog genera Alytes, Discoglossus, and Bombina as model groups. Our data set comprised ca. 900bp of partial mitochondrial 16S and 12S rRNA gene sequences (both genes evolved in a clock-like manner across genera) from nearly all the species and subspecies in the three genera. We tested several calibrations of a molecular clock, which resulted in competing temporal settings for the evolution of taxa. Although only one scenario was in complete accordance with paleogeographic data, statistical testing did not reject the alternatives. Limitations encountered with the present approach may be overcome by more comprehensive analyses in future.     

SEXUALLY SELECTED TRAITS EVOLVE POSITIVE ALLOMETRY WHEN SOME MATINGS OCCUR IRRESPECTIVE OF THE TRAIT

Positive allometry of secondary sexual traits (whereby larger individuals have disproportionally larger traits than smaller individuals) has been called one of the most pervasive and poorly understood regularities in the study of animal form and function. Its widespread occurrence is in contrast with theoretical predictions that it should evolve only under rather special circumstances. Using a combination of mathematical modeling and simulations, here we show that positive allometry is predicted to evolve under much broader conditions than previously recognized. This result hinges on the assumption that mating success is not necessarily zero for males with the lowest trait values: for example, a male who lacks horns or antlers might still be able to copulate if encountering an unguarded female. We predict the strongest positive allometry when males typically (but not always) compete in large groups, and when trait differences decisively determine the outcome of competitive interactions.     

Safer sex with feeding females: sexual conflict in a cannibalistic spider

Mating strategies are to a large degree shaped by conflictsbetween the sexes, causing a rapid antagonistic coevolutionof traits involved in reproduction. The view that sexual cannibalismrepresents a form of sexual conflict leads to the predictionof male traits that facilitate escape from cannibalistic females.A variety of traits have been suggested to serve this functionin spiders, where sexual cannibalism is comparatively common.Empirical evidence, however, is virtually absent. Here we showexperimentally that opportunistic mating with feeding females,which has been reported from several species of orb-weavingspiders, greatly reduces the risk of cannibalism and injuryfor males in the spider Nephila fenestrata. This has directconsequences for a male's fertilization success because survivingmales can reduce the female's remating probability by guardingher against rivals. Although copulation with previously matedfemales sometimes appears to be mechanically impossible, secondmales that do copulate can expect to fertilize on average 64%of a female's eggs. Our results support the view that opportunisticmating may have evolved as a male tactic in a context of sexualconflict over sexual cannibalism.     

Sperm allocation strategies and female resistance: a unifying perspective

The classical viewpoint in sperm competition theory, which holds that males evolve sperm allocation strategies in response to a given degree of sperm competition, has recently been challenged by an alternative viewpoint, which holds that the degree of sperm competition is itself a consequence of these same strategies. Here, we present a game theory model that unites these alternative views as the endpoints of a continuum. Based on the recognition that female control over mating may limit the extent to which male strategies affect the degree of sperm competition, we investigate sperm allocation strategies in a setting where females can resist excessive matings more or less successfully. We discuss how conflicting predictions made by previous theory relate to implicit assumptions about female resistance behavior. Moreover, we show that female resistance, while being highly relevant to the predicted relationship between ejaculate size and the degree of sperm competition, has little effect on the predicted positive correlation between relative testis size and the degree of sperm competition. This result strengthens one of the central predictions of sperm competition theory and is in accordance with empirical findings from a wide range of taxa.     

The evolution of sex roles in mate searching

Searching for mates is a critical stage in the life cycle of most internally, and many externally, fertilizing species. Males usually invest more in this costly activity than females, but the reasons for this are poorly understood. Previous models have shown that female‐biased parental investment, including anisogamy, does not by itself select for male‐biased mate searching, so it requires additional explanations. Here, we correct and expand upon earlier models, and present two novel hypotheses that might explain the evolution of male‐biased mate searching. The “carry‐over hypothesis” states that females benefit less from searching if the associated costs affect other stages of the life cycle, rather than arising only while searching. It is relevant to the evolution of morphological traits that improve searching efficiency but are also expressed in other contexts. The “mating window hypothesis” states that females benefit less from searching if their life cycle includes intervals during which the exact timing of mating does not matter for the appropriate timing of reproduction (e.g., due to sperm storage or delayed embryo implantation). Such intervals are more likely to exist for females given the general pattern of greater female parental investment. Our models shed new light on classic arguments about sex role evolution.     

EVOLUTION OF MATE CHOICE FOR GENOME-WIDE HETEROZYGOSITY

The extent to which indirect genetic benefits can drive the evolution of directional mating preferences for more ornamented mates, and the mechanisms that maintain such preferences without depleting genetic variance, remain key questions in evolutionary ecology. We used an individual-based genetic model to examine whether a directional preference for mates with higher genome-wide heterozygosity ( H ), and consequently greater ornamentation, could evolve and be maintained in the absence of direct fitness benefits of mate choice. We specifically considered finite populations of varying size and spatial genetic structure, in which parent–offspring resemblance in heterozygosity could provide an indirect benefit of mate choice. A directional preference for heterozygous mates evolved under broad conditions, even given a substantial direct cost of mate choice, low mutation rate, and stochastic variation in the link between individual heterozygosity and ornamentation. Furthermore, genetic variance was retained under directional sexual selection. Preference evolution was strongest in smaller populations, but weaker in populations with greater internal genetic structure in which restricted dispersal increased local inbreeding among offspring of neighboring females that all preferentially mated with the same male. These results suggest that directional preferences for heterozygous or outbred mates could evolve and be maintained in finite populations in the absence of direct fitness benefits, suggesting a novel resolution to the lek paradox.     

Emasculation to plug up females: the significance of pedipalp damage in Nephila fenestrata

Female multiple mating selects for male adaptations that maximizefertilization success in a context of sperm competition. Whilemale mating strategies usually reflect a trade-off between presentand future reproduction, this trade-off is largely removed insystems where the maximum number of matings for males is verysmall. Selection may then favor extreme mechanisms of paternityprotection that amount to a maximal investment in a single mating.Males in several arthropod taxa break off parts of their copulatoryorgans during mating, and it has frequently been suggested thatmutilated males can thus secure their paternity. Nevertheless,such a mechanism has rarely been confirmed directly. Here westudy the golden orb spider Nephila fenestrata, which has amating system with potentially cannibalistic, polyandrous females,and males that are often functionally sterile after mating withone female only. We show that males in this species can indeedprotect their paternity by obstructing the female's genitalopenings with fragments of their copulatory organs.     

Monogynous Mating Behaviour and its Ecological Basis in the Golden Orb Spider Nephila fenestrata

Males, especially in species where they provide little or no parental investment, usually have high potential reproductive rates and are expected to maximize their fitness by mating with several females. This view is challenged, however, by species in which males provide no parental investment, but nevertheless mate with one female only. Male monogamy (monogyny), associated with an extreme investment in paternity protection, appears to be comparatively common in web‐building spiders, and has recently been subject to experimental and theoretical studies. To date, however, studies approaching this issue from an ecological perspective are rare. Theory predicts that the evolution of a monogynous mating strategy is favoured by a male‐biased sex ratio, but not necessarily by a high mortality risk for mate‐searching males. To test these predictions, we conducted a field study on the golden orb spider Nephila fenestrata, which has a mating system with potentially cannibalistic, polyandrous females, and males that are often functionally sterile after mating with one female only. Based on daily observations of marked individuals, we confirm that, consistent with laboratory findings, monogyny is common in N. fenestrata. Nevertheless, observations of male movements between females raise the possibility that a proportion of males may mate with two females. We show that the sex ratio in our study population is male‐biased, and that males incur only a relatively moderate mortality risk during mate‐search. These findings provide insights into the ecological basis for the evolutionary maintenance of monogyny.     

Is the evolution of inaccurate mimicry a result of selection by a suite of predators? A case study using myrmecomorphic spiders

Several hypotheses have been put forward to explain the evolution of inaccurate mimicry. Here we investigated the novel hypothesis that inaccurate mimicry (in color and shape) is maintained by opposing selective pressures from a suite of different predators: model-aversive visually oriented predators and model- and mimic-specialized predators indifferent to mimetic cues. We hypothesize that spiders resembling ants in color and shape escape predators that typically avoid ants but fall prey to ant-eating predators. We tested whether inaccurate myrmecomorphic spiders are perceived as their models by two types of predators and whether they can escape from these predators. We found that model-specialized (ant-eating) predators captured mimics significantly less frequently than their ant models, because mimics changed their behavior by fleeing predatory attacks. The fastest escape was found in less accurate mimics, indicating a negative association between visual resemblance and effectiveness of defenses. In trials with spider-eating predators, mimics were not captured more frequently than their models. The quality of defensive mechanisms appears to result from opposing selection forces exerted by the predator complex: mimics are more accurate (in color and shape) in microhabitats dominated by model-aversive predators and less accurate in microhabitats with model- and mimic-specialized predators.     

A mate to die for? A model of conditional monogyny in cannibalistic spiders

Monogynous males in various species actively limit themselves to mating with a single female in their lifetime. Whereas previous models have considered monogyny as an obligate mating strategy, here we explore the potential of monogyny to evolve as a context‐specific (conditional) behavior. Using a state‐dependent dynamic game model based on the biology of the cannibalistic spider Argiope bruennichi, we confirm that conditional monogyny can evolve under broad conditions, including an even sex ratio. We predict that males should make a terminal investment when mating with large, virgin females, especially if population density is low and the encounter occurs late in the season. We encourage empirical tests for the existence of conditional monogyny in all species where monogyny occurs in the absence of strict morphological constraints that would make it obligatory.