Competition For Mates Research Articles

The Role of Bioacoustic Signals in Koala Sexual Selection: Insights from Seasonal Patterns of Associations Revealed with GPS-Proximity Units.

Despite being a charismatic and well-known species, the social system of the koala (Phascolarctos cinereus—the only extant member of the family Phascolarctidae) is poorly known and much of the koala’s sociality and mating behaviors remain un-quantified. We evaluated these using proximity logging-GPS enabled tracking collars on wild koalas and discuss their implications for the mating system of this species. The frequency and duration of male-female encounters increased during the breeding season, with male-male encounters quite uncommon, suggesting little direct mating competition. By comparison, female-female interactions were very common across both seasons. Body mass of males was not correlated with their interactions with females during the breeding season, although male size is associated with a variety of acoustic parameters indicating individuality. We hypothesise that vocal advertising reduces the likelihood of male-male encounters in the breeding season while increasing the rate of male-female encounters. We suggest that male mating-season bellows function to reduce physical confrontations with other males allowing them to space themselves apart, while, at the same time, attracting females. We conclude that indirect male-male competition, female mate choice, and possibly female competition, mediate sexual selection in koalas.

Social structure and space use of Amur tigers (Panthera tigris altaica) in Southern Russian Far East based on GPS telemetry data.

To better understand the spatial structure of Amur tigers (Panthera tigris altaica) at the southern edge of their range we fitted 14 tigers (6♀♀ and 8♂♂) with 15 GPS-Argos collars between 2008 and 2011 in 2 study sites: the Ussuriskii Reserve of southern Sikhote-Alin and the Land of the Leopard National Park in southwest Primorye, Russian Far East. Fixed kernel estimates of male home ranges were larger than those of female home ranges (P < 0.05 [mean 95% fixed kernel(♀) = 401 ± 205 km(2) ; mean 95% fixed kernel(♂) = 778 ± 267 km(2)]). The home range size of females varied greatly, but on average was similar to estimates derived from earlier work further north. Low overlap of adjacent home ranges suggested that females retained exclusive territories. Real core areas of females overlapped only slightly, and remained stable over multiple years. The home ranges of adult males were smaller than those of males to the north, and in contrast to previous studies, high overlap among males indicated the absence of territoriality. Nonetheless, real core areas of males did not overlap, suggesting some spatial separation. In comparison to other tiger populations and other areas of the Russian Far East, the sex ratio in our 2 study areas was highly skewed towards males. We believe this skewed sex ratio resulted in the dissolution of territoriality of males due to an inability to defend individual females, with males resorting to scramble competition for mates. Continued monitoring of these sites to determine whether shifts in the sex ratio might result in a return to male territoriality would provide confirmation of our tentative hypothesis.

The largest relative testis size among primates and aseasonal reproduction in a nocturnal lemur, Mirza zaza.

Testis size is an indirect indicator of a species' mating system, along with sexual size and canine dimorphism, existence and usage of mating and advertisement calls as well as the spatial distribution of males and females ready to mate in solitary species. Upon its recent discovery, the northern giant mouse lemur Mirza zaza was suggested to have a polygynandrous mating system and to exhibit seasonal breeding. We tested these predictions in a field study in Sahamalaza National Park, NW Madagascar. We caught 12 M. zaza, before and during the suggested mating period and obtained standard field morphometric measurements, including testes size. We show that M. zaza has the highest relative testis volume among primates, indicating strong sperm competition and polygynandrous mating. In addition, based on inferred age of captured animals, observed mating behavior, a female caught in met-estrus and data from captivity, we suggest M. zaza to be one of the few lemurs that breed aseasonally. Future field work on this endangered species is required to illuminate the causes and consequences of intense promiscuity and aseasonal breeding despite strong habitat seasonality, which distinguish M. zaza from most other nocturnal lemurs.

Cognitive ability is heritable and predicts the success of an alternative mating tactic.

The ability to attract mates, acquire resources for reproduction, and successfully outcompete rivals for fertilizations may make demands on cognitive traits--the mechanisms by which an animal acquires, processes, stores and acts upon information from its environment. Consequently, cognitive traits potentially undergo sexual selection in some mating systems. We investigated the role of cognitive traits on the reproductive performance of male rose bitterling (Rhodeus ocellatus), a freshwater fish with a complex mating system and alternative mating tactics. We quantified the learning accuracy of males and females in a spatial learning task and scored them for learning accuracy. Males were subsequently allowed to play the roles of a guarder and a sneaker in competitive mating trials, with reproductive success measured using paternity analysis. We detected a significant interaction between male mating role and learning accuracy on reproductive success, with the best-performing males in maze trials showing greater reproductive success in a sneaker role than as a guarder. Using a cross-classified breeding design, learning accuracy was demonstrated to be heritable, with significant additive maternal and paternal effects. Our results imply that male cognitive traits may undergo intra-sexual selection.

Copulation rate declines with mating group size in dusky dolphins (Lagenorhynchus obscurus)

Males in polygamous species often engage in intrasexual competition for mates. If females actively evade mating attempts, it may benefit males to cooperate to restrict female movement, as has been found in some mammals. We tested if male dusky dolphins (Lagenorhynchus obscurus (Gray, 1828)) cooperate or compete during group mating chases. If they cooperate, then the per-male probability of copulating should increase with group size; if they compete, then the probability should decrease. We followed mating groups by boat during the breeding season (October 2013 – January 2014) off Kaikoura, New Zealand. The copulation rate per male decreased with increasing group size and with the number of noncopulating males in proximity to a copulating female. Male dusky dolphins have multiple mates and appear to use sperm and exploitative scramble competition. Males may remain in mating groups despite competition because there are alliances within the groups, they are unable to exclude rivals from joining a group, the time and energy costs of searching for unescorted females exceed the costs of reduced mating opportunities in a group, they receive other direct or indirect benefits that offset the costs of reduced mating opportunities, or they are in the group largely for social learning rather than procreation.

To be or not to be horned—Consequences in cattle

In cattle, both males and females of horned breeds have permanent horns which are directly connected with the frontal sinuses of the skull and are growing during their entire life. To date a large proportion of cattle is disbudded or dehorned, or increasingly bred for polledness. This review explores the possible consequences of cattle having horns or not, both for the cattle themselves and for the farmers. During natural selection horns may have provided advantages concerning predator defence or in competition for mates and resources. Additionally, there are indications that horns serve as honest signals in mate selection, as they reflect individual health. Presence or absence of horns will likely affect quality and quantity of social interactions as well as social relationships in a herd. It clearly affects the way of cattle fighting. Only few studies deal with social behaviour in horned herds in comparison to hornless herds, and they rely on single herd observations or need to be interpreted with caution because of confounding factors. However, it appears that horned cattle attempt to keep greater inter-individual distances and resort less to physical interactions than hornless cattle, leading to more stable social relationships under suitable environmental conditions and management. Cattle may, furthermore, use horns for self-grooming of body regions which are otherwise out of reach. Even less is known on possible physiological functions of horns. While there is good evidence regarding a thermoregulatory function, other claimed functions, e.g. concerning digestion, have not been scientifically investigated yet. Although farmers in favour of keeping horned cattle, for instance for ethical reasons, are able to reduce the increased risk of animal injuries and human accidents by providing improved housing and management conditions and maintaining a good human–animal relationship, it may be economically advantageous to keep hornless cattle. If disbudding/dehorning is carried out, the consequences in terms of stress and pain can be alleviated by a combination of local anaesthesia and anti-inflammatory treatment, as well as sedation of animals insufficiently used to handling. However, possible negative side effects of sedation should be addressed in future studies. Furthermore, the possibility of longer lasting pain after disbudding/dehorning is not sufficiently investigated yet. Finally, it is an open question whether the removal of horns alters the social behaviour of the animals, or is a mere symptomatic measure to adjust them to husbandry conditions that are insufficiently adapted to the species-specific needs of cattle.

Sex allocation theory reveals a hidden cost of neonicotinoid exposure in a parasitoid wasp.

Sex allocation theory has proved to be one the most successful theories in evolutionary ecology. However, its role in more applied aspects of ecology has been limited. Here we show how sex allocation theory helps uncover an otherwise hidden cost of neonicotinoid exposure in the parasitoid wasp Nasonia vitripennis. Female N. vitripennis allocate the sex of their offspring in line with Local Mate Competition (LMC) theory. Neonicotinoids are an economically important class of insecticides, but their deployment remains controversial, with evidence linking them to the decline of beneficial species. We demonstrate for the first time to our knowledge, that neonicotinoids disrupt the crucial reproductive behaviour of facultative sex allocation at sub-lethal, field-relevant doses in N. vitripennis. The quantitative predictions we can make from LMC theory show that females exposed to neonicotinoids are less able to allocate sex optimally and that this failure imposes a significant fitness cost. Our work highlights that understanding the ecological consequences of neonicotinoid deployment requires not just measures of mortality or even fecundity reduction among non-target species, but also measures that capture broader fitness costs, in this case offspring sex allocation. Our work also highlights new avenues for exploring how females obtain information when allocating sex under LMC.

Mating competition and entrepreneurship

We used a natural experiment in Taiwan to test whether mating competition is a major motive for entrepreneurship. After being defeated in Mainland China, more than half million solders retreated to Taiwan in 1949. The soldiers were not allowed to get married until the military marriage ban was lifted in 1959. The policy change created a severe marriage squeeze for local men, triggering their animal spirits to engage in entrepreneurial activities. Empirical results show that local men exposed to high sex ratios as a result of the removal of the marriage ban were more likely to become entrepreneurs later on.

Population density influences dispersal in female white-tailed deer

Dispersal behavior in white-tailed deer (Odocoileus virginianus) predominantly occurs in 1-year-old males; however, females of the same age also disperse. The timing of female dispersal during fawning season and low dispersal rates suggest that competition for mates and reduced inbreeding are not ultimate causes of female dispersal, as suggested for males. We proposed that female dispersal is the result of competition for space when pregnant females seek to isolate themselves before and after parturition. To test this hypothesis, we conducted a meta-analysis of female dispersal rates from 12 populations of white-tailed deer and predicted dispersal rate and distance were positively related to deer density. We found a positive relationship between dispersal rate and deer per forested km 2 and between dispersal distance and deer per forested km 2 . These results are consistent with the hypothesis that female dispersal is density-dependent and caused by the exclusion of subordinate 1-year-olds as adult females seek isolation before and after parturition.

Pheromones mediating copulation and attraction in Drosophila

Intraspecific olfactory signals known as pheromones play important roles in insect mating systems. In the model Drosophila melanogaster, a key part of the pheromone-detecting system has remained enigmatic through many years of research in terms of both its behavioral significance and its activating ligands. Here we show that Or47b-and Or88a-expressing olfactory sensory neurons (OSNs) detect the fly-produced odorants methyl laurate (ML), methyl myristate, and methyl palmitate. Fruitless (fru(M))-positive Or47b-expressing OSNs detect ML exclusively, and Or47b- and Or47b-expressing OSNs are required for optimal male copulation behavior. In addition, activation of Or47b-expressing OSNs in the male is sufficient to provide a competitive mating advantage. We further find that the vigorous male courtship displayed toward oenocyte-less flies is attributed to an oenocyte-independent sustained production of the Or47b ligand, ML. In addition, we reveal that Or88a-expressing OSNs respond to all three compounds, and that these neurons are necessary and sufficient for attraction behavior in both males and females. Beyond the OSN level, information regarding the three fly odorants is transferred from the antennal lobe to higher brain centers in two dedicated neural lines. Finally, we find that both Or47b- and Or88a-based systems and their ligands are remarkably conserved over a number of drosophilid species. Taken together, our results close a significant gap in the understanding of the olfactory background to Drosophila mating and attraction behavior; while reproductive isolation barriers between species are created mainly by species-specific signals, the mating enhancing signal in several Drosophila species is conserved.

Mate competition and evolutionary outcomes in genetically modified zebrafish (Danio rerio).

Demonstrating relationships between sexual selection mechanisms and trait evolution is central to testing evolutionary theory. Using zebrafish, we found that wild-type males possessed a significant advantage in mate competition over transgenic RFP Glofish® males. In mating trials, wild-type males were aggressively superior to transgenic males in male-male chases and male-female chases; as a result, wild-type males sired 2.5× as many young as did transgenic males. In contrast, an earlier study demonstrated that female zebrafish preferred transgenic males as mates when mate competition was excluded experimentally. We tested the evolutionary consequence of this conflict between sexual selection mechanisms in a long-term study. The predicted loss of the transgenic phenotype was confirmed. More than 18,500 adults collected from 18 populations across 15 generations revealed that the frequency of the transgenic phenotype declined rapidly and was eliminated entirely in all but one population. Fitness component data for both sexes indicated that only male mating success differed between wild-type and transgenic individuals. Our predictive demographic model based on fitness components closely matched the rate of transgenic phenotype loss observed in the long-term study, thereby supporting its utility for studies assessing evolutionary outcomes of escaped or released genetically modified animals.

Target-site resistance mutations (kdr and RDL), but not metabolic resistance, negatively impact male mating competiveness in the malaria vector Anopheles gambiae.

The implementation of successful insecticide resistance management strategies for malaria control is currently hampered by poor understanding of the fitness cost of resistance on mosquito populations, including their mating competiveness. To fill this knowledge gap, coupled and uncoupled Anopheles gambiae s.l. males (all M form (Anopheles coluzzii)) were collected from mating swarms in Burkina Faso. This multiple insecticide resistant population exhibited high 1014F kdrR allele frequencies (>60%) and RDLR (>80%) in contrast to the Ace-1R allele (<6%). Kdr heterozygote males were more likely to mate than homozygote resistant (OR=2.36; P<0.001), suggesting a negative impact of kdr on An. coluzzii mating ability. Interestingly, heterozygote males were also more competitive than homozygote susceptible (OR=3.26; P=0.006), suggesting a heterozygote advantage effect. Similarly, heterozygote RDLR/RDLS were also more likely to mate than homozygote-resistant males (OR=2.58; P=0.007). Furthermore, an additive mating disadvantage was detected in male homozygotes for both kdr/RDL-resistant alleles. In contrast, no fitness difference was observed for the Ace-1 mutation. Comparative microarray-based genome-wide transcription analysis revealed that metabolic resistance did not significantly alter the mating competitiveness of male An. coluzzii mosquitoes. Indeed, no significant difference of expression levels was observed for the main metabolic resistance genes, suggesting that metabolic resistance has a limited impact on male mating competiveness. In addition, specific gene classes/GO terms associated with mating process were detected including sensory perception and peroxidase activity. The detrimental impact of insecticide resistance on mating competiveness observed here suggests that resistance management strategies such as insecticide rotation could help reverse the resistance, if implemented early.

Wild chacma baboons (Papio ursinus) remember single foraging episodes.

Understanding animal episodic-like memory is important for tracing the evolution of the human mind. However, our knowledge about the existence and nature of episodic-like memory in non-human primates is minimal. We observed the behaviour of a wild male chacma baboon faced with a trade-off between protecting his stationary group from aggressive extra-group males and foraging among five out-of-sight platforms. These contained high-priority food at a time of natural food shortage. In 10 morning and eight evening trials, the male spontaneously visited the platforms in five and four different sequences, respectively. In addition, he interrupted foraging sequences at virtually any point on eight occasions, returning to the group for up to 2h. He then visited some or all of the remaining platforms and prevented revisits to already depleted ones, apparently based on his memory for the previous foraging episode about food value, location, and time. Efficient use of memory allowed him to keep minimal time absent from his group while keeping food intake high. These findings support the idea that episodic-like memory offers an all-purpose solution to a wide variety of problems that require flexible, quick, yet precise decisions in situations arising from competition for food and mates in wild primates.

Sexual Stimulation and Sexual Selection.

Sexual Stimulation and Sexual Selection.

Toward a Biology of Collectivism

The signs of mating competition are written into the physiology of the human male, but they are not written equally into the physiology of all racial groupings of human males. It seems that Asian males are different, different in that they are more fully dissimilar from the gorilla than are other races, showing less sexual dimorphism, muscularity, and less marked secondary sexual characteristics, and different in that they are more fully dissimilar from the chimpanzee than are other races, showing less sexual drive and activity as well as smaller testicles and lower sperm counts. It is presently argued that such anatomical differences are a testament to a more peaceably monogamous mating history. In turn, it is then argued that such physiological markers are directly associated with the collectivist ethos that has been historically, anthropologically, and sociologically observed among the Asian people.

Women’s Competition for Mates: Experimental Findings Leading to Ethological Studies

There has been an explosion of survey-based and experimental work pertaining to women’s intrasexual competition for mates. Rather than extensively review this growing and vast body of literature, the goal of this paper is instead to call for ethological studies on this topic. I propose that, in general, non-ethological studies should cause us to question the reliability of findings, how frequently, and in what contexts competitive strategies are used. After a condensed overview of the evolutionary theory of female intrasexual competition, the paper is organized around three central problems that are faced by researchers who want to use an ethological approach. First, I will briefly review how female intrasexual competition involves multiple strategies that are often indirect or covert. Second, I will discuss how female intrasexual competition is dynamic, and changes depending on particular variables, such as hormonal status and audience. Third, I will argue that the context for examining competition matters, such that the reach of competitive views and attitudes is far wider than previously considered. I support this third point by presenting the results of a preliminary study where women appeared to engage in competition after merely being primed to think about potential threats to their romantic relationships.

Wolbachia-based population control strategy targeting Culex quinquefasciatus mosquitoes proves efficient under semi-field conditions.

In mosquitoes, the maternally inherited bacterial Wolbachia induce a form of embryonic lethality called cytoplasmic incompatibility (CI). This property can be used to reduce the density of mosquito field populations through inundative releases of incompatible males in order to sterilize females (Incompatible Insect Technique, or IIT, strategy). We have previously constructed the LR[wPip(Is)] line representing a good candidate for controlling field populations of the Culex quinquefasciatus mosquito in the islands of the south-western Indian Ocean. The main purpose of the present study was to fill the gap between laboratory experiments and field implementation, i.e. assessing mating competitiveness of these incompatible males under semi-field conditions. In a first set of experiments, we analyzed crossing relationships between LR[wPip(Is)] males and La Réunion field females collected as larvae in 19 distinct localities throughout the island. This investigation revealed total embryonic mortality, confirming the strong sterilizing capacity of LR[wPip(Is)] males. Subsequently, mating competitiveness of LR[wPip(Is)] males was assessed under semi-field conditions in the presence of field males and females from La Réunion. Confrontations were carried out in April and December using different ratios of LR[wPip(Is)] to field males. The results indicated that the LR[wPip(Is)] males successfully compete with field males in mating with field females, displaying even higher competitiveness than field males in April. Our results support the implementation of small-scale field tests in order to assess the feasibility of IIT against Cx. quinquefasciatus in the islands of southwestern Indian Ocean where this mosquito species is a proven competent vector for human pathogens.

Effects of autodisseminated Metarhizium guizhouense PSUM02 on mating propensity and mating competitiveness of Bactrocera cucurbitae (Diptera: Tephritidae)

We investigated the effects of inoculation by Metarhizium guizhouense PSUM02 on mating propensity and mating competitiveness of Bactrocera cucurbitae, with a view on pest management. On day 4 postinoculation, the M. guizhouense-treated male flies had significantly lowered mating propensity and mating competitiveness, while the treated female flies had reduced mating propensity on day 4 and reduced mating competitiveness on day 5. The mating propensity and competitiveness of treated male and female flies then further declined until death. Kaplan–Meier survival analysis of treated male and female flies gave average survival times (AST) of 6.2 ± 0.2 and 5.4 ± 0.3 days in the mating propensity assay, and about 5.0 ± 0.1 and 4.4 ± 0.2 days in the mating competitiveness assay. The AST of untreated flies ranged from 12.8 ± 0.1 to 14.7 ± 0.2 days for comparison (observation up to 15 days). Untreated flies had decreased AST and mating characteristics when exposed to contact with treated male flies, indicating transmission of the fungal infection by such contact also to untreated male flies. Surprisingly, contact with treated female flies did not affect the AST of untreated males or females in the same cage. These results corroborate the potential for pest control by autodissemination with treated male flies, which transmit the fungus to a healthy population better than the treated female flies.

Sexual behavior and male volatile compounds in wild and mass‐reared strains of the Mexican fruit fly Anastrepha ludens (Diptera: Tephritidae) held under different colony management regimes

We compared the calling and mating behavior and volatile release of wild males Anastrepha ludens (Loew) with males from 4 mass-reared strains: (i) a standard mass-reared colony (control), (ii) a genetic sexing strain (Tap-7), (iii) a colony started from males selected on their survival and mating competitiveness abilities (selected), and (iv) a hybrid colony started by crossing wild males with control females. Selected and wild males were more competitive, achieving more matings under field cage conditions. Mass-reared strains showed higher percentages of pheromone calling males under field conditions except for Tap-7 males, which showed the highest percentages of pheromone calling males under laboratory cage conditions. For mature males of all strains, field-cage calling behavior increased during the last hour before sunset, with almost a 2 fold increase exhibited by wild males during the last half hour. The highest peak mating activity of the 4 mass-reared strains occurred 30 min earlier than for wild males. By means of solid phase microextraction (SPME) plus gas chromatography-mass spectrometry (GC-MS), the composition of volatiles released by males was analyzed and quantified. Wild males emitted significantly less amounts of (E,E)-α-farnesene but emitted significantly more amounts of (E,E)-suspensolide as they aged than mass-reared males. Within the 4 mass-reared strains, Tap-7 released significantly more amounts of (E,E)-α-farnesene and hybrid more of (E,E)-suspensolide. Differences in chemical composition could be explained by the intrinsic characteristics of the strains and the colony management regimes. Characterization of calling behavior and age changes of volatile composition between wild and mass-reared strains could explain the differences in mating competitiveness and may be useful for optimizing the sterile insect technique in A. ludens.

Female song and vocal interactions with males in a neotropical wren

Bird song is thought to function primarily in same-sex competition, mate attraction, and reproductive stimulation of a partner. However, these conclusions are based largely on studies of the song of male birds in north-temperate species. We investigate female song in a Neotropical wren, Thryophilus pleurostictus, using observations and experiments to test the function of female song. Female banded wrens sang much less often than males, their songs were shorter, and their repertoire of song types was smaller. Females did not seem to sing for same-sex competition for resources or mates: female song rate did not increase in response to simulated intrusion, and females sang in response to less than one-third of playbacks simulating territorial intrusion by either unpaired or paired females. Territorial defense is important for both survival and reproduction in species that occupy all-purpose territories year-round, but female involvement in territorial defense was limited. Females were more likely to approach simulated intruders when their partner approached more closely, and were closer to their partner during playback simulating a pair of intruders, perhaps contributing to defense jointly with their partner. Females did not appear to use song to attract males for mating: only 25% of females sang in response to playback simulating an unpaired male during the nest-building period, and they were less likely to sing shortly before laying when they were more likely to be fertile. Female song in banded wrens seemed to be used primarily for communicating with their breeding partner: female songs overlapped or began within one second of a song by their partner more often than expected by chance, and male vocal behavior changed in response to song by their partner. However, the low rate of female song in banded wrens suggests this function does not select for song elaboration, consistent with the view that same-sex competition is the main driver of female song elaboration.