THE EVOLUTION OF ALTERNATIVE REPRODUCTIVE TACTICS IN MALE CARDIOCONDYLA ANTS

Alternative reproductive tactics are often associated with discontinuous variation in morphology but may evolve independent from each other. Based on life‐history data and a phylogeny we examine how male morphology and reproductive behavior are linked in the evolution of the ant genus Cardiocondyla. Wingless Cardiocondyla males engage in lethal fighting for access to female sexuals, whereas winged males disperse and mate away from the nest. This basic pattern shows considerable variation across species. A phylogeny based on ～3 kbp sequence data shows that male diphenism and lethal fighting are ancestral traits tightly linked in evolution. Winged males were lost convergently in several species groups, apparently in response to the low probability of encountering female sexuals in nests without a resident fighter male. An early dichotomy separates two clades with alternative male morphologies and fighting behavior, but phenotype and fighting strategy are not correlated with the presence of winged males.     

Between fighting and tolerance: reproductive biology of wingless males in the ant Cardiocondyla venustula

Male reproductive tactics vary widely across the species of the ant genus Cardiocondyla, from obligatory lethal combat among co‐occurring males to complete mutual tolerance. The African species C. venustula Wheeler, 1908 has an intermediate phylogenetic position between taxa with fighting males and taxa with tolerant males and also shows an intermediate male behavior. Males from 2 native populations in South Africa and a population introduced to Puerto Rico attacked and killed freshly eclosing rivals but rarely engaged in deadly fights with adult competitors. Instead, several males per colony established small “territories” in their natal nests and defended them against other males. Males with a stable territory had more contact with female sexuals than nonterritorial males and more frequently engaged in mating attempts. In controlled choice experiments, female sexuals did not show any preference for particular males. We suggest that male territoriality in C. venustula is an adaptation to the seasonal production of large numbers of female sexuals by multiple mothers.     

First Evidence of Inclusive Sexual Selection in the Ant Cataglyphis cursor: Worker Aggressions Differentially Affect Male Access to Virgin Queens

Sexual selection typically shapes the traits of individuals capable of reproduction. However, in social species, it may also affect the behaviour of non‐reproductive individuals if this increases their inclusive fitness. We examined the potential role of workers during mating in the polyandrous ant Cataglyphis cursor. In this species, mating takes place on the ground near the nest entrance, and workers chase away foreign males trying to mate with young queens (gynes). We investigated the effects of worker aggression on male mating attempts by placing three or four males in the presence of one foreign gyne and 20 of her nestmate workers and contrasted this with a second group of 3–4 males in the presence of another gyne without nestmate workers (n = 8 paired replicates). Workers strongly attacked males (more than 20 aggressions per minute), although this did not affect the total number of mating attempts by the 3–4 males or the skew in the number of mating attempts among these males. However, there was an interaction between worker aggression and relative male weight, as relatively heavy males attempted to mate more often than lighter males in the absence of worker whereas the reverse occurred in the presence of workers. Our study is the first to experimentally test the potential role of non‐reproductive individuals in sexual selection. It shows that workers could potentially change male mating success by disfavouring heavier males.     

Male Social Behavior and Dominance Hierarchy in the Sulawesi Crested Black Macaque (Macaca nigra)

In a 6-week study of the social behavior of wild Sulawesi crested black macaques (Macaca nigra), we found a linear and transitive dominance hierarchy among the six adult males in one social group. Dominance rank, as determined by the direction of supplantations, correlated strongly with percentage of time near more than four neighbors, frequency of grooming received from adult females, and percentage of time with an adult female as nearest neighbor. These results suggest that high-ranking males are socially attractive. Adult females sexually solicited high-ranking males more often than low-ranking males, but frequency of copulation was not correlated with dominance rank. Frequency and intensity of aggression between males are strongly correlated with rank distance, but aggression toward females was greatest for mid-ranking males. Males of all rank displayed significantly more aggression toward sexually receptive females than toward females in other estrous states. These data indicate that male Sulawesi crested black macaques display a social organization similar to that reported for multimale groups in other macaque species rather than the egalitarian social organization described for female Sulawesi macaques.     

Stealthy invaders: the biology of Cardiocondyla tramp ants

Many invasive ant species, such as the Argentine ant or the red imported fire ant, have huge colonies with thousands of mass-foraging workers, which quickly monopolise resources and therefore represent a considerable threat to the native ant fauna. Cardiocondyla obscurior and several other species of this myrmicine genus have similarly been transferred throughout the tropics by human activities. However, because their colonies are tiny and workers forage solitarily, Cardiocondyla are often not recognized as successful invaders. Here, we document that the life history of Cardiocondyla closely resembles that of the more conspicuous tramp species, with polygyny, intranidal mating, budding, worker sterility, low genetic variability, and possibly also unicoloniality. Given that introduced Cardiocondyla may locally reach a very high population density, the effects of these stealthy invaders on the native arthropod fauna should receive more attention. Received 18 May 2005; revised 23 September 2005; accepted 29 September 2005.     

Sex-specific aggression and sex ratio in wintering finch flocks: serins and siskins differ

Males are dominant over females in many bird species. This may lead to male monopolisation of resources whenever food is scarce or clumped and secondarily to lower female survival rates. As a result of the consequent male-biased sex ratio in the adult population, competition may arise either (1) between males and females, as males attempt to exclude females from feeding patches, or (2) between males because females do not pose a competitive threat. We recorded agonistic interactions between males and females in wintering foraging flocks of serins (Serinus serinus) and siskins (Carduelis spinus) to test for inter-specific differences. Most of the aggressive interactions in serins were between males and females, whereas in siskins they were between males. We also compared sex ratios for each species during the winter, determined from separate trapping efforts over an 11-year period, to test whether the direction of aggression by males (i.e. male/male; male/female) relates to variations in female survival rates. The proportion of females was smaller in winter than in autumn for serins, but differences in siskins were negligible. Results are interpreted in relation to the social organization displayed by both species studied.     

The Influence of Pedipalp Autotomy on the Courtship and Mating Behavior of Pardosa milvina (Araneae: Lycosidae)

Male Pardosa milvina wolf spiders use their pedipalps both for copulation and courtship. Pedipalp loss is significantly more common among adult males compared to females. We measured the courtship and mating effects associated with the loss of one or both pedipalps among adult male P. milvina. Pedipalp loss significantly reduced courtship intensity, but had no influence on mounting success. Intact males were less likely to be cannibalized and suffered fewer predatory attacks by females than autotomized males. Loss of the left pedipalp resulted in significantly less intense courtship, higher female aggression levels, and delayed onset of courtship whereas loss of the right pedipalp minimally affected male and female behavior relative to intact males. Pedipalp displays may function in reducing female aggression rather than increasing female receptivity.     

Live and let die: why fighter males of the ant Cardiocondyla kill each other but tolerate their winged rivals

Unlike most social insects, many Cardiocondyla ant species havetwo male morphs: wingless (ergatoid) males, who remain in thenatal nest, and winged males who disperse but, strangely, beforeleaving may also mate within the nest. Whereas ergatoid malesare highly intolerant of each other and fight among themselves,they tend to tolerate their winged counterparts. This is despitethe fact that these winged males, like ergatoid males, representmating competition. Why should ergatoid males tolerate theirwinged rivals? We developed a mathematical model to addressthis question. Our model focuses on a number of factors likelytoinfluence whether ergatoid males are tolerant of winged males:ergatoid male–winged male relatedness, number of virginqueens, number of winged males, and the number of ejaculatesa winged male has (winged males are sperm limited, whereas ergatoidmales have lifelong spermatogenesis). Surprisingly, we foundthat increasing the number of virgin queens favors a kill strategy,whereas an increase in the other factors favors a let-live strategy;these predictions appear true for C. obscurior and for a numberof other Cardiocondyla species. Two further aspects, unequalinsemination success and multiple mating in queens, were alsoincorporated into the model and predictions made about theireffects on toleration of winged males. The model is applicablemore generally in species that have dimorphic males, such assome other ants, bees, and fig wasps.     

A mathematical model for wing dimorphism in maleCardiocondyla ants

Several species ofCardiocondyla ants have dimorphic males: wingless (ergatoid) and winged (alate) males, while otherCardiocondyla species includingC. nuda have only wingless males. We made an evolutionarily stable strategy model for explaning the male polymorphism and the ratios of wingless males in the genusCardiocondyla. Wingless males emerge earlier than winged males in each reproductive season. Females (F ₁) which have emerged before winged males copulate only with wingless males, and females (F ₂) which emerge after winged males copulate with both wingless and winged males. Wingless males have a lower copulation ability (b _n) than winged males (b _w). The reproductive success of females which copulate at the early stage (v ₁) is assumed to be larger than that of females which copulate at the late stage (v ₂). The model predicts that there are 3 different evolutionarily stable states: 2 monomorphic states of wingless and winged males, and a dimorphism of the 2 types of males. In the dimorphic state, the rate of wingless males increases as the survival rate of wingless males (s) increases,v ₁/v ₂ increases,F ₁/F ₂ increases andb _n/b _w increases. For dimorphism to exist,s b _n/b _w<1 must be satisfied, and this condition corresponds to the value of observed data. The value ofv ₁/v ₂ would be difficult to be obtained by actual data, but we can estimate this value with the model.     

Fine‐ and Broad‐Scale Approaches to Understanding the Evolution of Aggression in Crickets

Male field crickets frequently engage in agonistic contests to establish dominance in social interactions and gain access to mate attraction territories. Crickets (Orthoptera: Gryllidae) are often used as a model taxon to study aggression, but limited documentation of aggression in some cricket species hinders our understanding of its evolutionary costs and benefits. Our study investigated cricket aggression at two scales: the within‐species scale for two cricket species, Gryllus assimilis and G. veletis, whose aggression had not been adequately documented and the among‐species scale to detect evolutionary patterns in species’ levels of aggression. In both G. veletis and G. assimilis, winners spent more time being aggressive than losers, but they were not larger or heavier. Collectively, our results reveal that G. veletis males are more aggressive than G. assimilis. Male G. veletis had higher aggression scores that male G. assimilis. The majority of G. veletis contests escalated to grappling (a highly aggressive behavior), while less than one quarter of G. assimilis contests escalated to grappling. Further, G. veletis males transitioned between two of the most aggressive behaviors most often while G. assimilis transitioned between two of the least aggressive behaviors most often. We integrate this new information on aggression for G. assimilis and G. veletis with previously documented aggression data for many cricket species to investigate aggression in a broader evolutionary context than previously possible. Within a phylogenetic context, we test the hypothesis that species whose males use burrows from which to call and attract females are more aggressive than species with non‐burrowing males. We found evidence consistent with this hypothesis; species with burrowing males tended to be more aggressive than species with non‐burrowing males. Together, our study provides fine‐scale understanding of aggression in two cricket species and broad‐scale evolutionary context for aggression across cricket species.     

Assortative aggression among males in a sympatric pair of Labeotropheus from Lake Malaŵi,Africa

Female mate choice has strong experimental support as a diversifying force in the speciation of the haplochromine cichlid fishes of Lake Malaŵi, Africa. Somewhat less understood is the role that male–male aggression might have played in the evolution of new species of these fishes. In the rock-dwelling haplochromines of Lake Malaŵi, primarily territory-holding males successfully court females; by determining which males gain territories, male–male aggression could support speciation by excluding less-fit males from the breeding population. To test the hypothesis that males should direct more aggression towards conspecific rivals, the aggression directed towards conspecific and heterospecific opponents was compared in a sympatric pair of cichlids of the genus Labeotropheus Ahl 1927 (Labeotropheus fuelleborni Ahl 1927 and Labeotropheus trewavasae Fryer 1956). It was found that when presented with a pair of rivals, males of both species did direct more aggression towards the conspecific opponent, and the amount of aggression increased when the conspecific opponent was larger than the heterospecific opponent. In addition, this study found a difference in the behavioural repertoire of the species: L. fuelleborni tends to rely on displays to intimidate opponents, whereas L. trewavasae employs more physical attacks to drive away opponents. Males of both species can thus recognize conspecifics and assess an opponent's relative threat to their ability to successfully reproduce, and use species-specific strategies to intimidate opponents.     

Pillar Function in the Fiddler Crab Uca beebei (I): Effects on Male Spacing and Aggression

Males of 14 species of fiddler crabs (genus Uca) are known to build structures out of mud or sand at the entrances to burrows they court from and defend. A study of spacing, space use and aggression between courting male Uca musica (Zucker 1974, 1981) suggested that the hoods males build reduce territorial overlap and rates of aggression between neighboring males. Thus, each male may have more time to court females during limited lunar, diurnal and tidal mating periods. I studied the courtship and aggressive behavior of male Uca beebei in the field to determine if the pillars males of this species build affect male behavior as do the hoods of U. musica. U. beebei occurs sympatrically with U. musica on the Pacific coast of Panama and is broadly similar in its ecology and mating behavior. Unlike the hoods of U. musica, pillars did not focus a male's activity space away from its closest neighbor nor did they reduce either overlap with neighbors' activity spaces or rates of aggressive interaction among neighbors. Pillar builders courted more but also fought their neighbors more than did males that did not build pillars. The pillars of U. beebei and the hoods of U. musica affect male behavior differently and probably have different functions.     

Abortion following the immigration of an adult male baboon (Papio cynocephalus)

Three abortions and the death of a vigorous infant occurred in a baboon (Papio cynocephalus) group in Amboseli National Park, Kenya, within the two weeks following the immigration of an aggressive adult male. The immigrant male attained top-ranking dominance status in the group prior to these events. Circumstantial observations suggest that the reproductive losses were related to persistent intense aggression from the immigrant adult male. In the past ten years, only three other miscarriages have occurred in Amboseli. Inducement of abortion is discussed as a reproductive tactic that may be available to males of mammalian species whose life histories favor the evolution of infant-killing by males.     

Aggression and Mating Success in Three Species of Drosophila1

Three closely related species of Drosophila: D. virilis, D. americana, and D. novamexicana, are known to differ in levels of male-male aggression. Through direct observation in the laboratory, we attempted to determine and characterize the relationships between intrasexual aggression and mating success in males of each of these species. Our results indicated that the most important determinant of male mating success was not the amount of aggression performed by a male, but rather the amount of aggression directed towards him.     

Territory Defence in Black Redstarts,Phoenicurus ochruros: Effects of Intruder and Owner Age?

European black redstarts, Phoenicurus ochruros, vigorously defend all-purpose territories and exhibit delayed plumage maturation, most subadult males looking exactly female-like in their first breeding season. We tested the hypotheses that such dull subadult male plumages are beneficial in order to reduce aggression of adult males either by deception or by honest signalling of subordinance status, and that, in turn, conspicuous (adult) plumage colorations are able to intimidate contenders because they act as a signal of fighting ability and aggressive motivation. Adult and dull yearling black redstart territory owners were confronted with intruders mimicked by stuffed mounts of either a conspicuous adult or a dull subadult male. Our results do not support the hypotheses tested: dull plumages of young intruders did not reduce aggression from adult territory owners and aggressiveness towards intruders was significantly higher in yearling territory owners as compared with adult owners. Conspicuous intruders did not deter dull territory owners and we found no indications that conspicuous male coloration is a signal of fighting ability or aggressive motivation in this species. Instead, the amount of aggressive response to intruders showed considerable individual variance within age classes regardless of the plumage of the intruder and the status of the owner.     

Social Relationships in Free-Ranging Male Macaca arctoides

Macaque social relationships differ greatly between species. Based on captive studies that focus mainly on females, researchers have classified stumptail macaque (Macaca arctoides) social relationships as tolerant, as indicated by a high rate of affiliation, frequent aggression, and symmetrical conflicts. To accumulate more data on male social relationships, which are relatively understudied, and to generate comparative data, we investigated male social relationships in a provisioned group of 68 free-ranging, naturally dispersing stumptail macaques in southern Thailand. We collected continuous focal animal and ad libitum data on 7 adult and 2 subadult males, recording social behavior during 283 contact hours between December 2006 and March 2007. Stumptail macaques of this population were less tolerant than predicted based on previous studies on captive groups: Rates of spatial proximity, affiliation, and aggression were low, most males directed affiliative behavior toward higher-ranking males, and conflicts were generally of low intensity and relatively asymmetrical. Thus, male stumptail macaques of the focal group appear to differ in their social style from a previous study of a captive group that mainly comprised of females. In some traits, they are even more intolerant than rhesus macaques, an intensively studied intolerant macaque species. We also compare our data on stumptail macaque males to those on other male macaques, but available data are too sparse to draw final conclusions.     

Do Kin Selection and Intra-Sexual Selection Operate in Spider Mites?

Between the two subsocial spider mites, Schizotetranychus longus Saito and S. miscanthi Saito, a big difference exists in male reproductive behavior. The males of the former species have an extraordinarily mild relationship with conspecific males, whereas those of the latter species show mortal aggression against conspecific males. Field and experimental observations on the structure of mating populations showed that S. longus is under inbreeding conditions because of a lack of males in early spring, whereas S. miscanthi is under outbreeding conditions. Therefore, I hypothesized that the difference in male antagonism between the two species may reflect the difference in relatedness of males, that is, it has evolved by kin selection. The recent discovery of two clinal trends in male–male aggression in S. miscanthi provided evidence in favour of this hypothesis. Furthermore, a correlation analyses between experimentally evaluated male aggression and distribution patterns of males in the field indicated that the variation in male antagonism is actually reflected in field populations. Based on these studies, I discuss the solidity of the kin-selection hypothesis. Lastly I address how spider mites are fruitful model animals for conducting behavioral, ecological and genetic studies to understand the evolution of haplo-diploidy.     

Social correlates of fecal testosterone in male ursine colobus monkeys (Colobus vellerosus): the effect of male reproductive competition in aseasonal breeders

Male testosterone (T) levels are thought to be linked with the mating system, degree of parental care, and male–male aggression in reproductive contexts (The ‘challenge hypothesis’; Wingfield et al., 1990). In many species though, T increases associated with mating behavior cannot be separated from those associated with male–male aggression. We tested the challenge hypothesis on aseasonally breeding ursine colobus (Colobus vellerosus), where male–male competition is intense outside of mating contexts. Fecal samples (N = 109) were collected from > 27 subadult and adult males in seven groups during 13-months of research in Ghana in 2004–2005. Fecal T (fT) levels were determined by enzyme immunosorbant assays. Behavioral data was collected using focal-animal and ad libitum sampling. The number of receptive females in each group did not positively correlate with male fT. There was a trend for adult males to have higher fT than subadult males; however there was no effect of rank on fT. The level of male–male aggression experienced was positively correlated with fT and individual males showed higher mean fT during ‘challenge’ than during ‘non-challenge’ periods. The number of male incursions experienced positively correlated with fT whereas the number of between-group encounters did not. Males attempt to gain reproductive opportunities during incursions, thus these results support the ‘challenge hypothesis’ in C. vellerosus. Outside of mating contexts, higher male fT levels are associated with increased aggression. Male parental investment in the form of infant defense was associated with increased fT, rather than the decline expected from other forms of paternal care.     

REPRODUCTIVE CONFLICT IN BUMBLEBEES AND THE EVOLUTION OF WORKER POLICING

Worker policing (mutual repression of reproduction) in the eusocial Hymenoptera represents a leading example of how coercion can facilitate cooperation. The occurrence of worker policing in “primitively” eusocial species with low mating frequencies, which lack relatedness differences conducive to policing, suggests that separate factors may underlie the origin and maintenance of worker policing. We tested this hypothesis by investigating conflict over male parentage in the primitively eusocial, monandrous bumblebee, Bombus terrestris. Using observations, experiments, and microsatellite genotyping, we found that: (a) worker‐ but not queen‐laid male eggs are nearly all eaten (by queens, reproductive, and nonreproductive workers) soon after being laid, so accounting for low observed frequencies of larval and adult worker‐produced males; (b) queen‐ and worker‐laid male eggs have equal viabilities; (c) workers discriminate between queen‐ and worker‐laid eggs using cues on eggs and egg cells that almost certainly originate from queens. The cooccurrence in B. terrestris of these three key elements of “classical” worker policing as found in the highly eusocial, polyandrous honeybees provides novel support for the hypothesis that worker policing can originate in the absence of relatedness differences maintaining it. Worker policing in B. terrestris almost certainly arose via reproductive competition among workers, that is, as “selfish” policing.     

Male tree weta are attracted to cuticular scent cues but do not discriminate according to sex or among two closely related species

Recognition of conspecifics is an essential precursor of successful mating. Where related species coexist, species discrimination might be important, but because related species are similar, species signal recognition may actually be low. Chemical cues such as cuticular hydrocarbons (CHCs) are frequently used by insects to identify suitable sexual partners. We predicted that New Zealand tree weta (Hemideina spp.), a genus of nocturnal ensiferan Orthoptera that live both allopatrically and sympatrically, use chemical signals from either frass or CHCs to find mates. In a series of six laboratory trials using both H. thoracica and H. crassidens, we found that male tree weta, but not female tree weta, occupied cavities primed with female cuticular cues more often than cavities without. However, males did not discriminate between chemical cues of male and female conspecifics, or between conspecifics and heterospecifics. In field trials, tree weta did not occupy artificial cavities primed with either female frass or female cuticular cues more often than unscented cavities. However, in both trials weta preferentially returned to cavities that had already been occupied earlier in the trials. A final field trial confirmed the presence of mixed species harems during the mating season in one region of sympatry. Our results suggest that selection on sex and species specific chemical cues that could be used to find conspecific mates is weak. Mixed species aggregations suggest that identification of conspecific mating cues has not evolved to be species specific. We infer that for male tree weta, the cost of mating with heterospecifics is likely less than not mating at all.