Sex‐specific selection on plant architecture through “budget” and “direct” effects in experimental populations of the wind‐pollinated herb,Mercurialis annua

Sexual selection may contribute to the evolution of plant sexual dimorphism by favoring architectural traits in males that improve pollen dispersal to mates. In both sexes, larger individuals may be favored by allowing the allocation of more resources to gamete production (a “budget” effect of size). In wind‐pollinated plants, large size may also benefit males by allowing the liberation of pollen from a greater height, fostering its dispersal (a “direct” effect of size). To assess these effects and their implications for trait selection, we measured selection on plant morphology in both males and females of the wind‐pollinated dioecious herb Mercurialis annua in two separate experimental common gardens at contrasting density. In both gardens, selection strongly favored males that disperse their pollen further. Selection for pollen production was observed in the high‐density garden only, and was weak. In addition, male morphologies associated with increased mean pollen dispersal differed between the two gardens, as elongated branches were favored in the high‐density garden, whereas shorter plants with longer inflorescence stalks were favored in the low‐density garden. Larger females were selected in both gardens. Our results point to the importance of both a direct effect of selection on male traits that affect pollen dispersal, and, to a lesser extent, a budget effect of selection on pollen production.     

Intrasexual competition underlies sexual selection on male breeding coloration in the orangethroat darter,Etheostoma spectabile

Elaborate, sexually dimorphic traits are widely thought to evolve under sexual selection through female preference, male–male competition, or both. The orangethroat darter (Etheostoma spectabile) is a sexually dichromatic fish in which females exhibit no preferences for male size or coloration. We tested whether these traits affect individual reproductive success in E. spectabile when multiple males are allowed to freely compete for a female. The quality and quantity of male coloration were associated with greater success in maintaining access to the female and in spawning as the primary male (first male to participate). On the other hand, sneaking behavior showed little correlation with coloration. Male breeding coloration in E. spectabile may therefore demonstrate how intrasexual competition can be a predominant factor underlying the evolution of male ornaments.     

No evidence for MHC‐based mate choice in wild giant pandas

Major histocompatibility complex genes (MHC), a gene cluster that controls the immune response to parasites, are regarded as an important determinant of mate choice. However, MHC‐based mate choice studies are especially rare for endangered animals. The giant panda (Ailuropoda melanoleuca), a flagship species, has suffered habitat loss and fragmentation. We investigated the genetic variation of three MHC class II loci, including DRB1, DQA1, and DQA2, for 19 mating‐pairs and 11 parent‐pairs of wild giant pandas based on long‐term field behavior observations and genetic samples. We tested four hypotheses of mate choice based on this MHC variation. We found no supporting evidence for the MHC‐based heterosis, genetic diversity, genetic compatibility and “good gene” hypotheses. These results suggest that giant pandas may not use MHC‐based signals to select mating partners, probably because limited mating opportunities or female‐biased natal dispersal restricts selection for MHC‐based mate choice, acknowledging the caveat of the small sample size often encountered in endangered animal studies. Our study provides insight into the mate choice mechanisms of wild giant pandas and highlights the need to increase the connectivity and facilitate dispersal among fragmented populations and habitats.     

FREQUENCY‐DEPENDENT SOCIAL DOMINANCE IN A COLOR POLYMORPHIC CICHLID FISH

A mechanism commonly suggested to explain the persistence of color polymorphisms in animals is negative frequency‐dependent selection. It could result from a social dominance advantage to rare morphs. We tested for this in males of red and blue color morphs of the Lake Victoria cichlid, Pundamilia. Earlier work has shown that males preferentially attack the males of their own morph, while red males are more likely to win dyadic contests with blue males. In order to study the potential contribution of both factors to the morph co‐existence, we manipulated the proportion of red and blue males in experimental assemblages and studied its effect on social dominance. We then tried to disentangle the effects of the own‐morph attack bias and social dominance of red using simulations. In the experiment, we found that red males were indeed socially dominant to the blue ones, but only when rare. However, blue males were not socially dominant when rare. The simulation results suggest that an own‐morph attack bias reduces the social dominance of red males when they are more abundant. Thus, there is no evidence of symmetric negative frequency‐dependent selection acting on social dominance, suggesting that additional fitness costs to the red morph must explain their co‐existence.     

Sexual selection in mushroom-forming basidiomycetes

We expect that sexual selection may play an important role in the evolution of mushroom-forming basidiomycete fungi. Although these fungi do not have separate sexes, they do play female and male roles: the acceptance and the donation of a nucleus, respectively. The primary mycelium (monokaryon) of basidiomycete fungi, growing from a germinating sexual spore, is hermaphroditic, but it loses female function upon the acceptance of a second nucleus. The resulting dikaryon with two different nuclei in each cell retains a male potential as both nuclei can fertilize receptive mycelia. We tested the occurrence of sexual selection in the model species of mushroom-forming basidiomycetes, Schizophyllum commune, by pairing monokaryons with fully compatible dikaryons. In most pairings, we found a strong bias for one of the two nuclei although both were compatible with the monokaryon when paired alone. This shows that sexual selection can occur in mushroom-forming basidiomycetes. Since the winning nucleus of a dikaryon occasionally varied depending on the receiving monokaryon, we infer that sexual selection can operate through choosiness of the receiving individual (analogous to female choice). However, in other cases the same nucleus won, irrespective of the receiving monokaryon, suggesting that competition between the two nuclei of the donating mycelium (analogous to male–male competition) might also play a role.     

Males that drop a sexually selected weapon grow larger testes

Costly sexually selected weapons are predicted to trade off with postcopulatory traits, such as testes. Although weapons can be important for achieving access to females, individuals of some species can permanently drop (i.e. autotomize) their weapons, without regeneration, to escape danger. We capitalized on this natural behavior to experimentally address whether the loss of a sexually selected weapon leads to increased testes investment in the leaf‐footed cactus bug, Narnia femorata Stål (Hemiptera: Coreidae). In a second experiment, we measured offspring production for males that lost a weapon during development. As predicted, males that dropped a hind limb during development grew significantly larger testes than the control treatments. Hind‐limb autotomy did not result in the enlargement of other nearby traits. Our results are the first to experimentally demonstrate that males compensate for natural weapon loss by investing more in testes. In a second experiment we found that females paired with males that lost a hind limb had 40% lower egg hatching success than females paired with intact males, perhaps because of lower mating receptivity to males with a lost limb. Importantly, in those cases where viable offspring were produced, males missing a hind limb produced 42% more offspring than males with intact limbs. These results suggest that the loss of a hind‐limb weapon can, in some cases, lead to greater fertilization success.     

Protein pheromone expression levels predict and respond to the formation of social dominance networks

Communication signals are key regulators of social networks and are thought to be under selective pressure to honestly reflect social status, including dominance status. The odours of dominants and nondominants differentially influence behaviour, and identification of the specific pheromones associated with, and predictive of, dominance status is essential for understanding the mechanisms of network formation and maintenance. In mice, major urinary proteins (MUPs) are excreted in extraordinary large quantities and expression level has been hypothesized to provide an honest signal of dominance status. Here, we evaluate whether MUPs are associated with dominance in wild‐derived mice by analysing expression levels before, during and after competition for reproductive resources over 3 days. During competition, dominant males have 24% greater urinary MUP expression than nondominants. The MUP darcin, a pheromone that stimulates female attraction, is predictive of dominance status: dominant males have higher darcin expression before competition. Dominants also have a higher ratio of darcin to other MUPs before and during competition. These differences appear transient, because there are no differences in MUPs or darcin after competition. We also find MUP expression is affected by sire dominance status: socially naive sons of dominant males have lower MUP expression, but this apparent repression is released during competition. A requisite condition for the evolution of communication signals is honesty, and we provide novel insight into pheromones and social networks by showing that MUP and darcin expression is a reliable signal of dominance status, a primary determinant of male fitness in many species.     

Pre‐ and postcopulatory sexual selection act in concert to determine male reproductive success in Tribolium castaneum

To capture how sexual selection shapes male reproductive success across different stages of reproduction in Tribolium castaneum (Coleoptera, Tenebrionidae), we combined sequential sperm defence (P1) and sperm offence (P2) trials with additional trials where both males were added simultaneously to the female. We found a positive correlation between the relative paternity share in simultaneous male–male competition trials and the P2 trial. This suggests that males preferred by females as sires achieve superior fertilization success during sperm competition in the second male position. In simultaneous male–male competition trials, where pre‐, peri‐ and postcopulatory sexual selection were all allowed to act, the relative paternity share of preferred males was more than 20% higher than in P2 sperm competition trials where precopulatory female choice was disabled. Additional behavioural observations revealed that mating with more attractive males resulted significantly more frequently in offspring production than mating with less attractive males. Thus, by comparing male fertilization success in trials where precopulatory choice was turned off with more inclusive estimates of fertilization success where pre‐ and pericopulatory choice could occur, we show that female mate choice may effectively inhibit sperm competition. Female mate choice and sperm competition (P2) are positively correlated, which is consistent with directional sexual selection in this species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 67–75.     

No evidence for a trade‐off between sperm length and male premating weaponry

Male ornaments and armaments that mediate success in mate acquisition and ejaculate traits influencing competitive fertilization success are under intense sexual selection. However, relative investment in these pre‐ and post‐copulatory traits depends on the relative importance of either selection episode and on the energetic costs and fitness gains of investing in these traits. Theoretical and empirical work has improved our understanding of how precopulatory sexual traits and investments in sperm production covary in this context. It has recently also been suggested that male weapon size may trade off with sperm length as another post‐copulatory sexual trait, but the theoretical framework for this suggestion remains unclear. We evaluated the relationship between precopulatory armaments and sperm length, previously reported in ungulates, in five taxa as well as meta‐analytically. Within and between taxa, we found no evidence for a negative or positive relationship between sperm length and male traits that are important in male–male contest competition. It is important to consider pre‐ and post‐copulatory sexual selection together to understand fitness, and to study investments in different reproductive traits jointly rather than separately. A trade‐off between pre‐ and post‐copulatory sexual traits may not manifest itself in sperm length but rather in sperm number or function. Particularly in large‐bodied taxa such as ungulates, sperm number is more variable interspecifically and likely to be under more intense selection than sperm length. We discuss our and the previous results in this context.     

Multiple Plumage Ornaments as Signals of Intrasexual Communication in Golden‐Winged Warblers

Avian plumage represents some of the greatest diversity in integument coloration of all animals. Plumage signals are diverse in function, including those that allow for assessing potential mates or the mitigation of agonistic interactions between rivals. Many bird species possess multiple ornamental traits that have the potential to serve as multiple or redundant signals. For example, male golden‐winged warblers (Vermivora chrysoptera) have brilliant carotenoid‐based yellow crowns, melanin‐based black throats, and structurally based white patches on their outer tail feathers. Using a correlative approach, we investigated whether plumage ornaments have the potential to reliably signal ability to acquire higher quality territory, aggressive response to simulated territorial intrusions, and reproductive success. We found that both crown chroma and tail brightness were significantly related to habitat quality and aggression; more ornamented birds held territories with higher quality habitat and were less aggressive toward simulated conspecific stimuli. Older birds sang less threatening songs than younger birds and were more likely to sing their mate attraction song type (type 1) rather than songs typically reserved for agonistic interactions (type 2). Finally, despite our previous research demonstrating that habitat strongly predicts reproductive success in this warbler population, we found no evidence of a direct link between ornamentation and reproductive success. Overall, these data suggest that younger males, and those with lower quality ornaments, compensate with more aggressive behaviors. Additional research is needed to investigate the dynamics between behavioral traits and ornaments to better understand complex signaling and how golden wing signals function in conspecific interactions (male–male interactions and mate‐choice).     

Intrasexual selection and testis size in strepsirhine primates

The main objective of this study was to investigate two predictionsof sexual selection theory concerning interspecific variationin testis size among strepsirhine primates (Lemuriformes andLorisiformes). First, the unique evolutionary history of lemursprovides an opportunity for an independent test of the predictionsof sperm competition theory regarding the relationship betweenmating system and relative testis size. Second, I examined theevolutionary relationship between the morphological correlatesof pre- and postcopulatory competition (i.e., between sexualdimorphism and testis size) because polygamous lemurs, in contrastto other polygamous primates, lack sexual dimorphism. Basedon measurements from 174 captive strepsirhines from 24 species,I found that multi-male species had significantly larger testesthan pair-living ones, but that they did not differ significantlyfrom solitary species. This result deviates from theoreticalexpectations, but may be the result of yet-unknown heterogeneityin mechanisms of male-male competition in both multi-male andsolitary species. There was no difference in relative testissize between nonmonogamous lemurs and lorises, indicating thatpresumably lower levels of precopulatory competition are notnecessarily compensated by more intense sperm competition. Bodysize and phylogenetic effects were also found to considerablyaffect interspecific variability in testis size. Analyses ofindependent contrasts revealed that evolutionary changes inmating system, testis size, sexual size, and canine dimorphismwere not, or only weakly, associated in this monophyletic groupof primates. Additional comprehensive comparative studies ofsexual dimorphism, testis size, mating system, and copulatorybehavior in these and other taxa are indicated to illuminategeneral patterns and causes of covariation among these traits     

Intra‐community coalitionary lethal attack of an adult male southern muriqui (Brachyteles arachnoides)

We report on the first evidence of intra‐community coalitionary lethal aggression in muriquis (Brachyteles). The event occurred in southern muriquis (Brachyteles arachnoides) during a long‐term study (>15 years) of two social groups inhabiting mostly pristine Atlantic forest habitat in the Parque Estadual Carlos Botelho, southern São Paulo State, Brazil. The attack took place deep in the core area of the Group Caetê home range. Tense agonistic behaviors and vocalizations preceded the lethal coalitionary attack, and the tension increased over a 36–48 hr period. One adult female and two unidentified individuals also took part in a coalition led by six adult males. The members of the coalition collectively approached, embraced, immobilized and repeatedly bit the entire body of an adult male, resulting in severe bleeding injuries and the victim's death in less than 1 hr after the attack commenced. Combined ecological, behavioral and spatial data related to the event indicate that this was an intra‐community attack and suggest social tensions related to mating competition as the proximate trigger of the coalitionary killing. The attack resembled those reported for chimpanzees, with clear numeric superiority and a low risk of injury to aggressors, resulting in the death of a lone conspecific victim. This observation (n=1) is suggestive of a capacity for escalated aggression in muriquis and reinforces arguments for the potential adaptive significance of intra‐community aggression in male philopatric societies, as reported for spider monkeys and chimpanzees. These characteristics challenge the view of the muriquis as a peaceful primate and support the general hypothesis that imbalances of power contribute to intra‐specific killing in primates, such as chimpanzees and humans. Am. J. Primatol. 71:860–867, 2009. © 2009 Wiley‐Liss, Inc.     

Sexual selection drives asymmetric introgression in wall lizards

Hybridisation is increasingly recognised as an important cause of diversification and adaptation. Here, we show how divergence in male secondary sexual characters between two lineages of the common wall lizard (Podarcis muralis) gives rise to strong asymmetries in male competitive ability and mating success, resulting in asymmetric hybridisation upon secondary contact. Combined with no negative effects of hybridisation on survival or reproductive characters in F1‐hybrids, these results suggest that introgression should be asymmetric, resulting in the displacement of sexual characters of the sub‐dominant lineage. This prediction was confirmed in two types of secondary contact, across a natural contact zone and in two introduced populations. Our study illustrates how divergence in sexually selected traits via male competition can determine the direction and extent of introgression, contributing to geographic patterns of genetic and phenotypic diversity.     

Condition,not eyespan,predicts contest outcome in female stalk‐eyed flies,Teleopsis dalmanni

In contests among males, body condition is often the key determinant of a successful outcome, with fighting ability signaled by so‐called armaments, that is, exaggerated, condition‐dependent traits. However, it is not known whether condition and exaggerated traits function in the same way in females. Here, we manipulated adult condition by varying larval nutrition in the stalk‐eyed fly, Teleopsis dalmanni, a species in which eyespan is exaggerated in both sexes, and we measured the outcome of contests between females of similar or different body condition and relative eyespan. We found that females in higher condition, with both larger bodies and eyespan, won a higher proportion of encounters when competing against rivals of lower condition. However, when females were of equal condition, neither eyespan nor body length had an effect on the outcome of a contest. An analysis of previously published data revealed a similar pattern in males: individuals with large relative eyespan did not win significantly more encounters when competing with individuals of a similar body size. Contrary to expectations, and to previous findings in males, there was no clear effect of differences in body size or eyespan affecting contest duration in females. Taken together, our findings suggest that although eyespan can provide an honest indicator of condition, large eyespans provide no additional benefit to either sex in intrasexual aggressive encounters; body size is instead the most important factor.     

Contrasting selection pressure on body and weapon size in a polygynous megaherbivore

Sexual size dimorphism (SSD) is a common morphological trait in ungulates, with polygyny considered the leading driver of larger male body mass and weapon size. However, not all polygynous species exhibit SSD, while molecular evidence has revealed a more complex relationship between paternity and mating system than originally predicted. SSD is, therefore, likely to be shaped by a range of social, ecological and physiological factors. We present the first definitive analysis of SSD in the common hippopotamus (Hippopotamus amphibius) using a unique morphological dataset collected from 2994 aged individuals. The results confirm that hippos exhibit SSD, but the mean body mass differed by only 5% between the sexes, which is rather limited compared with many other polygynous ungulates. However, jaw and canine mass are significantly greater in males than females (44% and 81% heavier, respectively), highlighting the considerable selection pressure for acquiring larger weapons. A predominantly aquatic lifestyle coupled with the physiological limitations of their foregut fermenting morphology likely restricts body size differences between the sexes. Indeed, hippos appear to be a rare example among ungulates whereby sexual selection favours increased weapon size over body mass, underlining the important role that species-specific ecology and physiology have in shaping SSD.     

Mate choice in a changing world

Human activities by altering environmental conditions are influencing the mate choice of animals. This is by impacts on: (i) the production and expression of traits evaluated by mate choosers; (ii) the transmission of information about potential mates to choosers; (iii) the reception and processing of the information by choosers; and (iv) the final mate choice. Here, I first discuss how these four stages of the mate‐choice process can be altered by environmental change, and how these alterations, in turn, can influence individuals, populations, and communities. Much evidence exists for human‐induced environmental changes influencing mate choice, but the consequences for the fitness of courters and choosers are less well known, and even less is known about the impact on population dynamics, species interactions and community composition. More evidence exists for altered mate‐choice systems influencing interspecific matings and thereby community composition and biodiversity. I then consider whether plastic adjustments and evolutionary changes can rescue adaptive mate‐choice systems, and reflect on the possibility of non‐adaptive mate‐choice systems becoming less maladaptive under environmental change. Much evidence exists for plastic adjustments of mate‐choice systems, but whether these are adaptive is seldom known, as is the contribution of genetic changes. Finally, I contemplate the possibility of mate‐choice systems rescuing populations from decline in changing environments. I explain how this is context dependent with both positive and negative outcomes possible. In summary, while much evidence exists for human‐induced environmental changes influencing mate‐choice systems, less is known about the consequences for ecological and evolutionary processes. Considering the importance that mate choice plays in determining individual fitness and population viability, the effects of environmental change on mate‐choice systems should be considered in studies on the ecological and evolutionary consequences of human disturbances to habitats.