Male Reproductive Behavior of the Social Wasp Polistes fascatus (Hymenoptera: Vespidae)

Populations of male Polistes fuscatus simultaneously exhibit two different mate-locating tactics. Some males defend territories in female nesting and hibernation sites. These males frequently do not occupy the same territory each day, and they drag their gasters over perches, which may function to apply a secretion to the perch. Another segment of the population patrols large overlapping areas in female foraging sites. In contrast to territorial males, patrolling males do not rub their gasters on perches, and males seen on more than one day are tenacious to one area. Males in both sites are aggressive to other males and attempt to copulate with females. A laboratory study indicates that large males have an advantage in male-male competition. The mean size of patrollers is smaller than that of territorial males, indicating that patrollers are competitively inferior males. Yet there is considerable size overlap of males between the two sites, suggesting that there is also overlap in the range of probability of mating success between the two sites.     

Variability of GnRH secretion in two goby species with socially controlled alternative male mating tactics

Male reproductive phenotypic plasticity related to environmental-social conditions is common among teleost fish. In several species, males adopt different mating tactics depending on their size, monopolizing mates when larger, while parasitizing dominant male spawns when smaller. Males performing alternative mating tactics are often characterized by a strong dimorphism in both primary and secondary reproductive traits. According to studies on sex-changing species and on species where only one male morph is reproductively active, male alternative phenotypes are expected to vary also in gonadotropin-releasing hormone (GnRH) neurons in forebrain preoptic area (POA). Here, we compared the intra- and inter-sexual variations in number and size of GnRH neurons, along with gonads and male accessory structure investment, in two goby species, the grass goby, Zosterisessor ophiocephalus, and the black goby, Gobius niger, characterized by male alternative mating phenotypes. In both species, older and larger males defend nests, court and perform parental care, while younger and smaller ones try to sneak territorial male spawning. We found that grass goby and black goby have different patterns of GnRH expression. Grass goby presents a clear intra-sexual dimorphism in GnRH expression, related to the occurrence of alternative mating tactics, while in the black goby, only inter-sexual differences are observed. The inter- and intra-specific variability in the GnRH neurons in these two goby species is discussed in light of the differences in migratory behavior, nest type, and mating system.     

SEXUAL SELECTION AND FITNESS VARIATION IN A POPULATION OF SMALLMOUTH BASS,MICROPTERUS DOLOMIEUI (PISCES: CENTRARCHIDAE)

Monogamy is often presumed to constrain mating variance and restrict the action of sexual selection. We examined the reproductive patterns of a monogamous population of smallmouth bass (Micropterus dolomieui), and attempted to identify sources of within-season fitness variation among females and known-age males. Many males did not acquire a nest site, and many territorial males were unsuccessful in acquiring a mate. The likelihood that territorial males mated depended on several aspects of nest sites. Mated males of age three were larger than the average size of age-three males in the population. The mean sizes of age-four and age-five mated males were not different from the average of same-age males in the population. Thus, selection resulting from the acquisition of a mate favored large size among only age-three males. Timing of nest construction and breeding among territorial males was negatively related to male size and did not depend on male age after taking male size into account. Indirect evidence (numbers of eggs deposited in nests) suggests that the timing of spawning among females was also negatively related to female size. Fertility selection favored early reproduction within the season by males of all ages, but large male size was favored among only age-four males. The combined early breeding of fecund females and female mate choice of large males may explain the positive correlation between the size of age-four males and the number of eggs acquired. Despite large differences of female fecundity, however, the variance of relative mate number contributed about two times more than the variance of relative fertility among females to the total variance of relative fitness within each sex.     

Body Size Correlates with Fertilization Success but not Gonad Size in Grass Goby Territorial Males

In fish species with alternative male mating tactics, sperm competition typically occurs when small males that are unsuccessful in direct contests steal fertilization opportunities from large dominant males. In the grass goby Zosterisessor ophiocephalus, large territorial males defend and court females from nest sites, while small sneaker males obtain matings by sneaking into nests. Parentage assignment of 688 eggs from 8 different nests sampled in the 2003–2004 breeding season revealed a high level of sperm competition. Fertilization success of territorial males was very high but in all nests sneakers also contributed to the progeny. In territorial males, fertilization success correlated positively with male body size. Gonadal investment was explored in a sample of 126 grass gobies collected during the period 1995–1996 in the same area (61 territorial males and 65 sneakers). Correlation between body weight and testis weight was positive and significant for sneaker males, while correlation was virtually equal to zero in territorial males. That body size in territorial males is correlated with fertilization success but not gonad size suggests that males allocate much more energy into growth and relatively little into sperm production once the needed size to become territorial is attained. The increased paternity of larger territorial males might be due to a more effective defense of the nest in comparison with smaller territorial males.     

The relative costs and benefits of territorial defense and the two conditional male mating tactics in the Comanche Springs pupfish (Cyprinodon elegans)

We examined the aggressive costs and reproductive benefits of territorial defense and its alternatives in a population of the Comanche Springs pupfish (Cyprinodon elegans). The breeding system was characterized by three different male mating tactics: territorial defense, satellite positioning, and sneak spawning. The mating tactic adopted by males reflected the males' sizes. Territorial residents were the largest, satellites were medium-sized, and sneakers were the smallest adult males observed. Consistent with the hypothesis that primary mating tactics are relatively high-cost, high-benefit strategies, we found that territorial males engaged in a number of aggressive encounters but had higher reproductive successes than any other males. However, our observations did not support the premise that conditionally breeding males engage in relatively low-benefit, low-cost tactics. Medium satellites and small sneakers acquired fewer spawns than did territorial males, but both satellites and sneakers were involved in as much aggression as territorials. That is, the data supported the prediction that satellite or sneaker males with the inability to compete for territories would attempt to accrue some reproductive opportunities in the presence of territorial males despite the high costs of spending time on the breeding grounds. Adopting conditional tactics appeared to allow satellites and sneakers to make the best of a bad situation. During a subsequent breeding season, large males were absent from the population, and medium-sized males established territories. The aggressive behavior exhibited by medium residents was similar to the previous year, but these males did not acquire higher reproductive successes than medium satellites had. The implications of switching tactics and the influence of operational sex ratios on the costs and benefits of the male tactics are briefly addressed. Electronic Publication     

Determinants of male mating success in a natural population of a stream goby of the genus Rhinogobius

The mating success of individually identified males of a stream goby, Rhinogobius sp. CB (cross‐band type), was recorded over one breeding season. One to 3 year‐old males were active in mating, but 3 year‐old males, which accounted for 36% of the population, amounted to c . 70% of the total males guarding eggs in the nest. Three quarters of the breeding males had only one brooding cycle, but the others had two or three. All the latter males changed their nest sites between cycles within a riffle. In 1 and 2 year‐old males, the number of brooding cycles contributed more to the mating success than egg mass size in one brooding cycle. For mating success of 3 year‐old males, the egg mass size in one brooding cycle, which can be enhanced by spawning with a large female or multiple females, was as important as the number of brooding cycles. These male reproductive tactics could be attributed to the age‐related ability of nest construction and mate acquisition.     

Mating Systems and Male Size in Australian Hylaeine Bees (Hymenoptera: Colletidae)

The mating systems of seven previously unstudied members of the colletid bee genus Hylaeus Fabricius and one of Hyleoides Smith are described. Male mating tactics can be categorized as territorial (perched males defend flowers or other sites that attract receptive females) or non-territorial (patrolling males search for receptive females at flowering plants). The four species in which some territorial males occur are characterized by: 1. grappling fights among males for preferred perches; 2. territorial control by larger males; 3. the possession of prominent spines or other projections on the venter of the abdomen in larger males; and 4. the occurrence of some males that are as large as, or larger than, the largest females of their species (the ‘large-male phenomenon’). In contrast, the four species that lack territorial males are distinctive in that males: 1. do not engage in grappling contests; 2. lack abdominal weaponry; and 3. are smaller than the largest females of their species. In addition, we searched for the large-male phenomenon in museum collections of four species of Hylaeus that exhibit male abdominal spines (presumed to be the weapons used by territorial individuals) and two other species that lack these attributes (presumed non-territorial patrolling species). The results tend to support the sexual-selection-for-fighting-ability hypothesis, which argues that the evolution of unusually large males is a selective consequence of aggressive male—male competition for access to mates. The limitations of the present data set as a comparative test of this hypothesis are discussed.     

The mating system of a bee fly (Diptera: Bombyliidae). II. Factors affecting male territorial and mating success

Males of an undescribed bombyliidfly (Comptosia sp.)occupy traditional territories on a Southeast Queensland hilltop, to which females come solely for the purpose of mating. Territorial fights between males involve aerial collisions during which modified spines on the wing margins produce scars on the bodies of opponents. Territory owners and mating males are not different in size or age from the remainder of the male population. Although residency is related to fighting success, the strength of the effect is ambiguous. Consequently, our data do not appear to fit predictions from game theoretical models for fighting protocol. Hilltop males lacked the extensive population variation typically found in territorial species, and thus, the presumed advantages of traits such as large size may be suppressed. Hilltop males were larger than males at a nonhilltop, resource-based mating site and the possibility of alternative mating tactics is discussed.     

Sex, drugs and mating role: testosterone-induced phenotype-switching in Galapagos marine iguanas

Males of many vertebrate species have flexible reproductivephenotypes and must decide before each mating season whetherto adopt sneaker, satellite, or territorial mating tactics.How do males gauge their abilities against others in the population?We tested experimentally whether hormone–behavior feedbackloops allow Galapagos marine iguana males to activate theirthree behavioral phenotypes as predicted by the relative plasticityhypothesis. Territorial males defended small mating areas andhad significantly higher plasma testosterone (T) levels (75± 11 ng/ml) than did satellite males that roamed aroundterritories (64 ± 8 ng/ml) or sneaker males that behavedlike females within territories (43 ± 11ng/ml). In territorialmales, temporary pharmacological blockade of T slowed head-bobpatrolling, decreased territory size threefold, and reducedthe number of females on territories 20-fold. This supportsprevious data that females may gauge male attractiveness byusing head-bob patrolling, here shown to be a T-dependent trait.Control-treated neighbors reacted to the weakening of T-blockedmales by increasing head-bob rate fivefold and territory size1.6-fold, and female numbers increased 2.5-fold. Unmanipulatedor control-injected males remained unchanged. Behavioral effectswere partly reversed after 7 days. T injections induced satellitemales to establish temporary territories, even at unconventionallocations. Some T-boosted satellite males suffered serious fightinginjuries. T-injected sneakers left female clusters and behavedlike larger satellite males that roam around territories. Thus,territorial and mating tactics are activated by T, but experimental(de-) activation at the wrong ontogenetic stage is costly: manipulatedmales switched phenotype but thereby lowered their access tofemales. We hypothesize that T levels of males that are basedon early-season behavioral interactions influence a males' subsequentphenotypic role.     

Territorial male gobies respond aggressively to sneakers but do not adjust their sperm expenditure

We investigated whether mating behavior (sperm expenditure,courtship rate, and nest guarding) varied according to differentlevels of sperm competition in territorial males of two gobyspecies, the grass and the black gobies. We measured sperm expenditure(sperm released after 30 min from the beginning of the spawning),male courtship rate, and nest-guarding behavior in territorialmales of both species during simulated spawnings, in which wevaried the number of attending sneakers. Our results showedthat, in both species, territorial males adjusted their effortin nest guarding to the presence of rival sneakers by increasingthe time spent patrolling the territory and attacking the sneakers.In contrast, sperm expenditure and male courtship rate werenot influenced by the number of attending sneakers. These resultsare in agreement with those reported for other fish with alternativemating tactics and help to interpret previous inconsistenciesbetween theoretical predictions and measured levels of spermreleased at different levels of sperm competition by sneakersof the two gobiids studied here.     

Competition for landmark territories among male Polistes canadensis (L.) (Hymenoptera: Vespidae): large-size advantage and alternative mate-acquisition tactics

Alternative male mating tactics of insects at landmarks (leks)have only rarely been investigated. Some males of the paperwasp, Polistes canadensis (L.), were territorial at small treesalong the crests of dry ridges in Santa Rosa National Park,Costa Rica. Territories did not contain nests or resources forwhich females foraged. Contrary to other "hilltopping" species,male P. canadensis competed most intensely for territories insaddles along these ridges rather than at the highest points.Nonterritorial males patrolled small areas of the ridge line,following a path that took them to a number of territories.Many males switched between territoriality and patrolling, suggestingthat both size-related tactics belong to one conditional strategy.Males that were territorial on 2 or more days were larger thanthose that were territorial on only 1 day, and these in turnwere larger than permanent patrollers. Moreover, the mean sizeof territorial males was positively correlated with two measuresof territory attractiveness, suggesting that larger males monopolizepreferred sites. Mean age of territorial males was also relatedto territory attractiveness, but males of intermediate age claimedthe most attractive territories.     

The Resource-Based Mating System of the Mediterranean Pollen Wasp Ceramius fonscolombei Latreille 1810 (Hymenoptera, Vespidae, Masarinae)

Males of C. fonscolombei patrol and perch at water collection sites or at plants of Reseda, both of which are important resources for female brood care. The mating system can be classified as resource defense polygyny modified by the existence of alternative male mating tactics. Occupying temporary territories at watercollection sites constitutes the primary tactic which is more profitable for larger males. The secondary tactic of patrolling at flowers provides a nonaggressive alternative through which smaller males gain at least some mating success. Males at water collection sites occupy considerably smaller ranges but spend a higher proportion of time patrolling than males at flowers. They frequently grapple with other males, an activity that is absent at flowers. Males at water collection sites copulate about 2.5 times more frequently than males at flowers. The copulation frequency of the males at water collection sites is positively correlated with their body size, while copulation frequency is negatively correlated with body size at flowers. Males patrolling at water collection sites and males patrolling at flowers do not differ in body size, indicating that the decision between alternative mating tactics is not made relative to body size but is influenced by other factors.     

Mate-locating behavior of the butterfly Lethe diana (Lepidoptera: Satyridae): do males diurnally or seasonally change their mating strategy?

The mate-locating behavior of male butterflies has been classified into two major types, territorial and patrolling. Territorial males defend a specific site, whereas patrolling males fly around a wider area without having to defend a site. In this study, I investigated the use of these tactics by males of the satyrine butterfly, Lethe diana. A previous study suggested that the males of L. diana change their mate-locating behavior during the day (they patrol in the morning and defend territories in the afternoon) and that patrolling is the primary mating strategy, whereas defending territories is a supplementary one. In the present study, I found that the daily activity pattern of the males of L. diana was similar to that described in the previous study: males often flew around in the morning and competed for territories in the afternoon. However, contrary to the previous study, all courtships and copulations were performed within male territories during their territorial activity. Closer observations revealed that copulations found in male territories were achieved by the owner of the territory. Males tended to feed in the morning, suggesting that the males flying in the morning searched for food rather than females. I conclude that territory holding is the primary male matelocating tactic in L. diana. I further found that, in summer, males exhibited territorial behavior later than in spring or autumn, which may be a strategy for preventing heat stress.     

On the song resumption, polyterritorial behaviour and their population context in the Sedge Warbler Acrocephalus schoenobaenus

Mating tactics in species with facultative polygyny seem to be very flexible and to depend on local environmental conditions. We analysed the habitat and population contexts of territorial behaviour, associated with polygyny, in a population of Sedge Warblers Acrocephalus schoenobaenus inhabiting natural wetlands. Nearly one-third of all breeding males (46 individuals) resumed song after completing their mating with the first female, in order to attract another one. Resuming males were usually the earliest arrivals. There was a continuity in the territorial behaviour between resuming song on first territory and polyterritorial behaviour. Fifty-nine percent of resuming males set up second territories, clearly separated from the first. The second territory was usually located close to the primary female activity area. The quality of the first and second territories were correlated, with second territories being significantly inferior. However, their quality was not significantly different from the territories of non-breeding males. The quality of the second territories was also negatively related to the distance from the first territories. Polyterritorialism was influenced by population numbers: the higher the number of territorial males, the lower the number of second territories settled and the lower their average quality. Although the polygyny frequency in the studied population was very low, polyterritorial males were significantly more likely to be polygynous. We concluded that polygyny frequency can be significantly influenced by population numbers, which might be one of the main factors responsible for the variability in the mating system in this species.     

Operational sex ratio and resource defence as predictors of the mating system in European bitterling

Operational sex ratio (OSR), the ratio of sexually active males to fertilizable females in a population, plays a central role in the theory of mating systems by predicting that the intensity of male–male competition and the degree of sexual selection increases as the OSR becomes increasingly male biased. At high values of OSR, however, resource defence theory predicts the breakdown of territoriality and a shift towards scramble competition with a decrease in sexual selection. The direction that correlations between OSR and resource competition and variance in mating success will take depends on the biology of the species of interest. We investigated the effects of male population density and male‐biased operational sex ratio on male mating tactics shown by a freshwater fish, the European bitterling, Rhodeus sericeus . This species spawns inside living unioneid mussels. Large males defended territories, were aggressive towards conspecifics under equal sex ratios and monopolized pair spawnings with females. The mating tactic, however, changed at high male density where large males ceased to be territorial and instead competed with groups of smaller males to release sperm when females spawned. This change in male behaviour from pair to group spawning has two ramifications for sexual selection. The intensity of sexual selection and variance in male mating success decrease, and the form of sexual competition changes from resource‐ to sperm competition. Thus, the use of alternative mating tactics renders the OSR unable to predict the direction of resource competition and variance in male mating success at high densities.     

Social organization in the pheasant (Phasianus colchicus): harem formation, mate selection and the role of mate guarding

Harem formation and mate selection were studied in the pheasant in order to determine the advantages of territorial harem defence polygyny to the two sexes. We investigated the factors affecting harem size and the advantage to a female in remaining with one territorial male during breeding.
Female group size declined during late March and early April as females moved from large overlapping ranges into smaller, more widely dispersed breeding ranges. The proportion of female groups accompanied by males increased during this period.
Some males had a disproportionate share of females. Settled females were monogamous but, because a female's nest was generally outside the male's territory, her home range was larger than his territory.
Harem members were usually from the same winter group. Harem size was not related to territory quality in terms of food supply or nesting cover. Females were loyal to one male in more than one year even if his territory position changed. Older, territory-owning males had more females, both adult and immature, than males with newly-established territories. Harem size was not correlated with territory size.
We conclude that the mating system of the pheasant is based on mate guarding which protects females not only from the risk of predation or injury, but also from excessive energy expenditure incurred through being chased by other males. When escorted by a territorial male, females spent three times as much time feeding, one-fifth as much time running, and one-tenth as much time alert, as they did when not guarded.     

A note on group sizes of oribi (Ourebia ourebi, Zimmermann, 1783) from two contrasting sites in Zambia, with and without predation

The oribi (Ourebia ourebi, Zimmermann, 1783) is a small species of antelope widely represented across open grasslands of sub‐Saharan Africa. Although largely territorial, differences in group size and mating systems (monogamy/polygamy), have been linked to habitat conditions and population density. At high population densities, additional males are recruited to assist in territorial defence. Other factors which might impact on group size, include predation threat, especially at low population densities. In this study (1995–98), we recorded group sizes of oribi across two contrasting areas (c. 300 km² each) of Zambia with (Kafue), and without (Bangweulu), large vertebrate predators. A total of 412 groups was recorded across the two sites, with mean sizes of 2.31 (n = 217) at Bangweulu and 2.33 (n = 195) at Kafue. The modal group size was two throughout (range 1–6). Groups were more variable in size and sex composition at Kafue than at Bangweulu, but there was no significant difference in estimates of population density between sites. Mean estimates were 2.02 and 1.90 km^?2 for Bangweulu and Kafue, respectively. Sex ratios (males : females) were biased towards females (1 : 1.72 and 1 : 3.55, respectively) at both sites. Adult males were particularly scarce at Kafue. However, values of density and group size, were both unexceptional for the species. Whilst a modal group size of two is consistent with a monogamous mating system, variations in group composition and size were more difficult to explain. Adult females appeared to retain or recruit additional animals to groups, irrespective of the needs of territorial defence. This suggests that vigilance is an important function of groups on large open plains, especially in the presence of predators. By extension, reversed sexual dimorphism in oribi could reflect increased selection for vigilance duties in females. Predators had no apparent effect on population density, but oribi groups were more variable in size and sex composition in their presence. Adult males may be particularly vulnerable to predation when defending territorial borders at low population densities. However, further work is needed to define group dynamics in this species and to identify causes of mortality amongst adult males.     

Ecological and phylogenetic perspectives on wing stiffness in nine Theclini species (Lepidoptera: Lycaenidae)

We studied the relationship between wing stiffness and butterfly ecology and phylogeny. Nine species belonging to the tribe Theclini of the family Lycaenidae were selected and examined for the wing stiffness of dried specimens by a three‐point bending test. It was found in Japonica lutea that the wing stiffness was not affected by the humidity to which it had been exposed, but was strongly affected by wing size and sex. Comparisons of sexual differences in four species indicated that females of patrolling species had stiffer wings than conspecific males, but that males of territorial species had stiffer wings than conspecific females. Finally, the wing stiffness was compared among males of nine species that use different mate‐locating tactics, and the results revealed a tendency that males of territorial species have stiffer wings than males of patrolling species. These results, though including a few exceptional cases, are discussed from the perspective of ecological requirements and phylogenetic constraints on the species.     

Male–male competition for large nests as a determinant of male mating success in a Japanese stream goby, Rhinogobius sp. DA

Males of the stream goby Rhinogobius sp. DA (dark type) court females in deep pools and care for the eggs under stones in shallow riffles. We studied male–male competition for access to females and nest sites to understand how male size influences the mating success of this species. In field observations, larger males won in fighting with other males. However, large males did not tend to monopolize courtship opportunities, and the frequency of successful courtships, after which males led the females to the nests, was not related to male body size. The fact that courted females always escaped from the fighting sites once males began fighting likely explains why male size was not positively related to courtship success. Large males occupied large nest stones, and the number of eggs received in the nest was correlated positively with nest size. In aquarium experiments with two tiles of different sizes provided as nesting materials, males always chose the larger nest and, when two males were introduced simultaneously, the larger one occupied the larger nest. These results suggested that male mating success of this goby is determined by male–male competition for large nests rather than for access to females. Received: June 9, 2000 / Revised: September 2, 2000 / Accepted: October 4, 2000     

Reproductive success in male sunbleak, a recent invasive fish species in the U.K.

The spawning behaviour of male nest guarding sunbleak Leucaspius delineatus , a recent invasive species in southern England, was studied and quantified for the first time. In the absence of physical differences between territorial and non‐territorial males ( i.e . colour, size, etc .), the reproductive behaviour of territorial males was analysed and related to reproductive success. The results showed that females preferred high‐courting and highly aggressive males. The initial cue in female mate choice, however, was based on courtship, while aggression was the decisive behavioural trait in influencing mate choice, providing a direct signal of physical condition and 'paternal competence'. Some males picked nest sites which were subsequently preferred by other males taking over the nest of a previous male ('communal nest'), with the new territorial male adopting the eggs already present at the nest. It appears that either due to female preference for nests already containing eggs or lower rates of sired egg predation by dilution among unrelated eggs, sunbleak males have adopted the mating strategy of allopaternal care.