Large Males Research Articles

Endurance rivalry and female choice jointly influence male mating success in the emerald treefrog (Zhangixalus prasinatus), a lek-chorusing anuran

BackgroundEndurance rivalry and female choice are two important mechanisms of sexual selection in lek-breeding species. Endurance rivalry is when males compete for opportunities to mate by spending more time in leks than others (interaction-independent male-male competition). Because high-quality males can afford to have high lek attendance, females have a higher chance of mating with good-quality males even when they mate randomly. The good gene hypothesis proposes that females can pass good genes on to their offspring by choosing males that display elaborate morphological and/or behavioral traits that reflect the males’ genetic quality. The relative importance of lek attendance and female choice to males’ mating success in anurans is rarely evaluated. In this study, we investigated how these two mechanisms might jointly shape males’ morphological traits in the lek-chorusing emerald treefrog Zhangixalus prasinatus.ResultsOur results show that (1) male lek attendance is positively correlated with body size and condition, and males with higher lek attendance have higher mating success, (2) the dominant frequency of males’ advertisement calls are negatively correlated with body size and males producing lower frequency calls have higher mating success, (3) male body size, but not body condition, has a non-significant positive relationship with mating success and (4) females show preference for calls with lower dominant frequencies in two-choice playback.ConclusionsOverall, both endurance rivalry and female choice play an important role in the mating success of male emerald treefrogs in the field and both are influenced by male body size/condition. By mating with males that have higher lek attendance and produce lower frequency calls, selection may indirectly favor larger males.

Multi-modal locomotor costs favor smaller males in a sexually dimorphic leaf-mimicking insect

BackgroundIn most arthropods, adult females are larger than males, and male competition is a race to quickly locate and mate with scattered females (scramble competition polygyny). Variation in body size among males may confer advantages that depend on context. Smaller males may be favored due to more efficient locomotion leading to higher mobility during mate searching. Alternatively, larger males may benefit from increased speed and higher survivorship. While the relationship between male body size and mobility has been investigated in several systems, how different aspects of male body morphology specifically affect their locomotor performance in different contexts is often unclear.ResultsUsing a combination of empirical measures of flight performance and modelling of body aerodynamics, we show that large body size impairs flight performance in male leaf insects (Phyllium philippinicum), a species where relatively small and skinny males fly through the canopy in search of large sedentary females. Smaller males were more agile in the air and ascended more rapidly during flight. Our models further predicted that variation in body shape would affect body lift and drag but suggested that flight costs may not explain the evolution of strong sexual dimorphism in body shape in this species. Finally, empirical measurements of substrate adhesion and subsequent modelling of landing impact forces suggested that smaller males had a lower risk of detaching from the substrates on which they walk and land.ConclusionsBy showing that male body size impairs their flight and substrate adhesion performance, we provide support to the hypothesis that smaller scrambling males benefit from an increased locomotor performance and shed light on the evolution of sexual dimorphism in scramble competition mating systems.

Functional dietary response of Asian black bears to changes in sika deer density

AbstractOmnivores are generally opportunistic foragers and have a flexible dietary response to resource abundance and availability. Their populations may consist of individuals that differ from each other in terms of their trophic positions, which implies that the dietary response to resource fluctuations differs within a population. We investigated how changes in the abundance of sika deer (Cervus nippon) affected dietary variation and body condition in the Asian black bear (Ursus thibetanus). We used fecal analysis, nitrogen stable isotopes (δ15N), and body measurements to determine whether the variation in dietary meat content of Asian black bears is positively related to variations in the density of the sika deer population, whether male bears have a higher trophic position compared to females, and whether dietary meat content is positively related with body mass or body condition of bears. We found a positive correlation between the occurrence of deer remains in bear feces and deer density, suggesting that bears change their diet in response to temporal changes in deer density. Male bears had higher δ15N values than females, and neither values varied when deer density decreased. Males selectively consumed deer after a reduction in deer density, whereas females consistently consumed more plant‐based diet. The δ15N values were positively related with body mass of adult (>4 yr) bears but had no relationship with body condition of bears of either sex or any age class. Deer seem to be an important food source for large adult males, which have an advantage in mating. Thus, increasing herbivore abundance and availability altered the foraging strategy of Asian black bears, but the importance of herbivore on bear diet differs within a population.

Size variation, allometry and mating success in Aotearoa|New Zealand kelp flies (Coelopidae)

The need to respond quickly to the presence of an ephemeral resource required for breeding is often a feature of scramble competition mating systems. Scramble competition mating systems can feature extreme levels of sexual conflict and coercive mating by males. As a result, sexual selection can act on various traits used by males to overcome female resistance behaviours. Selection on these traits may result in significant intra and intersexual size variation and sexual dimorphism. Additionally, traits that influence mating success in males often show positive static allometry. Kelp flies (Coelopidae) are a small family of Diptera which specialise on wrack (beach cast marine macroalgae), a highly ephemeral resource. The mating system of these flies involves high levels of sexual conflict, with females rejecting all male mating attempts. In this study we describe intra and intersexual size variation and static allometry of traits in two of Aotearoa|New Zealand’s species, Coelopella curvipes and Chaetocoelopa littoralis. In addition, we investigate the mating behaviour of C. littoralis under ecologically relevant mating conditions. We found high levels of variation in both species with significant evidence of sexual dimorphism across all traits measured in C. littoralis, and in mid tibia length in C. curvipes. Furthermore, mid tibia length in both species exhibits positive static allometry and is disproportionally larger in larger males, suggesting that this trait in particular may be under strong sexual selection. We found that larger male C. littoralis which attempt to mate are significantly more likely to mate successfully demonstrating a large-size advantage in this species similar to findings across the Coelopidae. However, we only found a non-significant trend towards a mating advantage for males with longer mid-tibiae. We discuss these findings with reference to the population dynamics and ecology of these species.

Biological parameters in a declining population of narwhals (Monodon monoceros) in Scoresby Sound, Southeast Greenland

A decreasing trend in narwhal (Monodon monoceros Linnaeus, 1758) abundance has been identified in a small population in Scoresby Sound, Southeast Greenland. We hypothesize that excessive hunting has affected life history and population dynamics of this population. Biological information and samples collected from the Inuit hunt, from satellite-tagged narwhals and from official hunters’ reports, were used to estimate age, growth, and reproduction. During 2007 through 2019, a decreasing proportion of young and increasing proportion of older whales were harvested. Male and female body length and male tusk length increased significantly, while body mass of both sexes showed a nonsignificant increase. The probability of catching a female decreased significantly, while a nonsignificant decline of catching a pregnant female was observed in both biological samples and hunters’ reports. Narwhal swimming speeds correlated with fluke widths indicated that larger whales attain greater speeds. The decline in juveniles and females is probably due to an opportunistic hunting practice targeting the easiest-to-catch whales, where bigger whales are faster and more difficult to catch. The cumulative effect of overharvest with a declining proportion of females, an overrepresentation of large males, and a lack of calves and juveniles has detrimental implications for this small narwhal population.

Sex-specific influence of communal breeding experience on parenting performance and fitness in a burying beetle.

Communal breeding, wherein multiple conspecifics live and reproduce together, may generate short-term benefits in terms of defence and reproduction. However, its carry-over effects remain unclear. We experimentally tested the effects of communal breeding on parental care and reproduction in burying beetles (Nicrophorus vespilloides), which use carcasses as breeding resources and provide parental care to offspring. We subjected individuals to communal or non-communal breeding (i.e. pair breeding) during their first breeding event and to non-communal breeding during their second breeding event. We measured the parental care of individuals and of groups and the reproductive success of groups during both breeding events. In communal groups, large individuals became dominant and largely monopolized the carcass, whereas small individuals (i.e. subordinates) had restricted access to the carcass. At the first breeding event, large males in communal groups spent more time providing care than large males in non-communal groups, whereas such an effect was not observed for large females and small individuals. Reproductive successes were similar in communal and non-communal groups, indicating no short-term benefits of communal breeding in terms of reproduction. Compared with males from non-communal groups, males originating from communal groups produced a larger size of brood during their second breeding event, whereas such an effect was not observed for females. Our results demonstrate the sex-specific effects of communal breeding experience on parenting performance and fitness.

Male mate choice in mosquitofish: personality outweighs body size

BackgroundDespite its important implications in behavioural and evolutionary ecology, male mate choice has been poorly studied, and the relative contribution of personality and morphological traits remains largely unknown. We used standard two-choice mating trials to explore whether two personality traits (i.e., shyness and activity) and/or body size of both sexes affect mate choice in male mosquitofish Gambusia affinis. In the first set of trials involving 40 males, we tested whether males would prefer larger females and whether the preference would be affected by males’ body length and personality traits, and females’ activity level. In the second set of trials (using another 40 males), we tested whether males would prefer more active females and whether the preference would be affected by males’ body length and personality traits.ResultsBoth shyness and activity in males were significantly repeatable and constituted a behavioural syndrome. No overall directional preference for large (or small) females with the same activity levels was detected because larger males preferred larger females and smaller males chose smaller females. Males’ strength of preference for larger females was also positively correlated with the activity level of larger females but negatively with the activity level of smaller females. Males spent more time associating with active females regardless of their body lengths, indicating males’ selection was more influenced by female activity level than body size. Males’ preference for inactive females was enhanced when females became active. There was no convincing evidence for the effect of males’ personality traits or body length on their preferences for females’ activity level.ConclusionsOur study supports the importance of body size in male mate choice but highlights that personality traits may outweigh body size preferences when males choose mating partners.

Reproductive behaviour of the North African freshwater crab Potamon algeriense (Bott, 1967) (Brachyura: Potamidae)

ABSTRACT Laboratory and field observations of courtship, copulation, spawning, incubation, and maternal care were made on the endemic North African freshwater crab Potamon algeriense. Mating occurred throughout the year but no discernible courtship behaviour was recorded. However, a forced copulation strategy is practiced by the larger males. The occurrence of ovigerous females extended from June to September with a peak in July (38%). Spawning behaviour took place out of the water and the incubation period lasted 45 ± 2 days. Ovigerous females returned to the stream a few hours before hatching of the eggs. The occurrence of brooding females (females carrying juvenile crabs) extended from July to October, with the highest number in August (36%). Posthatching maternal care was recorded, which lasted between 7 to 10 days before juveniles left their mother’s abdominal cavity for an independent life.

Sexual dimorphism and morphometrics in two populations of the Neotropical freshwater turtle Mesoclemmys vanderhaegei (Testudines, Chelidae)

ABSTRACT As commonly observed in turtles, sexual size dimorphism (SSD) is pronounced in the Neotropical freshwater turtle Mesoclemmys vanderhaegei (Bour, 1973), a species in which females are usually larger than males. We studied SSD in two populations of M. vanderhaegei from the Brazilian Cerrado savannah, based on 245 specimens captured between November 2010 and August 2013. The carapace length of the largest male was 201 mm (9.15% shorter than that of the largest female, 220 mm). The mean sizes of males and females did not differ in the two populations. However, a comparison of eight selected morphological variables revealed that the size distribution pattern differed between the populations. Using model selection, seven out of 34 morphometric variables - from the head, plastron, bridge, and tail - were selected as the most suitable ones to distinguish between males and females. The pattern of SSD found in M. vanderhaegei is similar to that found in other chelonian species and may be the result of natural selection rather than ecological factors, since individuals of both sexes use the same habitats.

Does Size Matter? The Case of the Courtship Pyramids in Red Sea Ghost Crabs (Ocypode saratan).

Simple SummaryMale ghost crabs (Ocypode spp.) are known to build sand structures near the entrance to their burrows. It has always been assumed that the structure played a role in sexual selection and mate attraction. We hypothesized that the larger males would make relatively larger pyramids in order to enhance fitness. We studied Red Sea ghost crabs at Eilat, Israel, and found that the larger crabs actually did not put much effort into building pyramids, and it was the medium-sized crabs that invested the most in building the pyramids.Display, wherein males attempt to maximize fitness by attracting sexually mature females to mate, is known to drive speciation by Sexual Selection. We researched the Red Sea Ghost Crab (Ocypode saratan; RSGC), in which males build display pyramids to attract females. The study was conducted at the beach in Eilat, Israel. At each session, we measured the height (in cm) of all pyramids and the dimensions (height, breadth; in cm) of the burrow entrance. We assumed that the size of the entrance represented the relative size of the carapace width of the occupant. The mean (± SE) entrance volume was 230.8 ± 11.7 cm, and the height of the pyramid was 11.8 ± 0.49 cm (n = 54). The results of our study did not support our hypothesis because we had expected to find a linear correlation between body size and pyramid height, i.e., the larger the male, the larger the pyramid. However, our results show that the largest males in the population either built small pyramids or not at all, and the cut-off of the larger crab’s body size appears to be around 350 cm3. We discovered a step-wise function in the data in that crabs with the smallest body size of ca. 250 cm3 constructed the highest pyramids, with a declining tendency between 250–350 cm3 and extremely low pyramids beyond 350 cm3. However, our findings need to be further studied with a stress on the ambiance and elucidate whether the habitats differ in temperature, humidity, prey-base, etc., before concluding as to why the larger males desist from building pyramids. This study underwrites the importance of studying the mating systems of the macro-fauna of the beaches that are fast disappearing owing to anthropogenic development.

Size matters: Antagonistic effects of body size on courtship and digging in a wolf spider with non-traditional sex roles

Body size, nuptial gift characteristics and courtship behaviour, among other traits, can reflect the quality of a potential mate and, thus, might be under sexual selection. To maximize their mating success, individuals can show behavioural plasticity in sexual context. Allocosa senex is a burrow-digging wolf spider that exhibits reversal in courtship roles and in sexual size-dimorphism expected for spiders. Males construct the mating refuge and females prefer males that build longer burrows, which are considered as nuptial gifts because they are delivered to them after mating. This study aims to determine whether male body size and female reproductive status influence burrow dimensions, courtship displays, female preferences and cannibalism rate in A. senex. For that purpose, we allowed males to construct burrows and performed sexual trials under laboratory conditions. Larger males were more courted by females, and in turn, they expressed more vibratory behaviours during courtship. However, and contrary to our expectations, smaller males constructed longer burrows. We suggest that males of A. senex exhibit size-dependent behavioural plasticity, and when they are smaller, they invest more in burrow construction to compensate their lower opportunities of courting intensively as larger males. In addition, females courted differentially according to their reproductive status, overriding male preferences for virgin females. This study opens several doors to future research regarding mutual choice in A. senex and the traits assessed by males and females during courtship, as well as about the influence of intrinsic and extrinsic factors shaping reproductive decision-making in this and other wandering spider species.

Dual function and associated costs of a highly exaggerated trait in a cichlid fish.

Exaggerated secondary sexual characteristics are apparently costly and seem to defy natural selection. This conundrum promoted the theory of sexual selection. Accordingly, exaggerated secondary sexual characteristics might be ornaments on which female choice is based and/or armaments used during male–male competition. Males of many cichlid fish species, including the adaptive radiation of Nicaraguan Midas cichlids, develop a highly exaggerated nuchal hump, which is thought to be a sexually selected trait. To test this hypothesis, we conducted a series of behavioral assays in F2 hybrids obtained from crossing a species with a relatively small hump and one with an exaggerated hump. Mate‐choice experiments showed a clear female preference for males with large humps. In an open‐choice experiment with limited territories, couples including large humped males were more successful in acquiring these territories. Therefore, nuchal humps appear to serve dual functions as an ornament for attracting mates and as an armament for direct contest with rivals. Although being beneficial in terms of sexual selection, this trait also imposes fitness costs on males possessing disproportionally large nuchal humps since they exhibit decreased endurance and increased energetic costs when swimming. We conclude that these costs illustrate trade‐offs associated with large hump size between sexual and natural selection, which causes the latter to limit further exaggeration of this spectacular male trait.

Environmental Drivers of Fine-Scale Predator and Prey Spatial Dynamics in Sydney Harbour, Australia, and Adjacent Coastal Waters

Greater Sydney is the largest coastal city in Australia and is where bull sharks (Carcharhinus leucas) are present every summer and autumn. A decade of acoustic telemetry data was used to identify drivers of space use for bull sharks and their potential prey, according to standardised 6-h intervals using dynamic Brownian bridge movement models. Influences of environmental, physical, and biological variables on the areas of space use, location, and predator–prey co-occurrence were investigated with generalised additive mixed models. Rainfall in the catchment affected space use for all animals (i.e. teleost species and both sexes of sharks), with varying temporal responses. Male sharks responded most promptly to high rainfall moving upstream in < 1 day, followed by teleosts (2 to 7 days), and female bull sharks after 4 days. Environmental luminosity affected male shark dispersal and space use, possibly indicating use of visual cues for foraging. Physical characteristics of habitat were important factors driving spatial overlaps between predator and prey in estuarine areas. In sandy embayments < 10-m deep, males and female bull sharks overlapped with different species, whereas males and silver trevally (Pseudocaranx georgianus) co-occurred in deep holes (> 30 m). Shark size influenced overlap between sexes, with smaller females less likely to co-occur with larger males (~ 50 cm). Variability in space use suggests spatial segregation by sex and size in bull sharks, with individuals targeting similar prey, yet either in different areas or at different times, ultimately enabling them to exploit different resources when in the same habitats.

Bachelor groups form due to individual choices or environmental disrupters in African striped mice

In several mammal species, bachelor groups occur as a regular life history stage between dispersal and becoming the breeding male of a multifemale group. How such groups come into existence and how males that choose this strategy differ in life history traits from other males is rarely investigated. Males of the socially flexible African striped mouse, Rhabdomys pumilio , have been historically reported to adopt one of three alternative tactics: small group-living philopatric males, intermediate solitary roaming males or large group-living territorial males. Here, we describe for the first time, bachelors as a fourth male tactic. A long-term study on 13 generations of striped mice revealed two types of bachelor groups: (1) kin bachelor groups formed when all females disappeared, probably due to predation, and (2) nonkin bachelor groups formed by unrelated males that originated from different groups. Whereas kin bachelor groups occurred more frequently during the nonbreeding season, nonkin bachelor groups were most common in the breeding season and had a shorter tenure, since most of these males soon switched to the solitary roaming tactic or the territorial breeder tactic. Importantly, we found that nonkin bachelors were heavier and older than kin bachelors and that they did not differ from solitary roaming males in body mass and age. We therefore identified two different forms of bachelor groups, small kin bachelors whose female group members disappeared and larger nonkin bachelor groups showing traits of an alternative reproductive tactic. Both types of bachelor groups differed in multiple traits, demonstrating that it is important to follow the life histories of individuals to understand the mechanisms leading to alternative tactics. • We describe bachelor groups as a fourth male tactic in striped mice. • Kin bachelor groups form when all females disappear. • Nonkin bachelor groups are formed by unrelated males from different groups. • Both types of groups differ in body mass, age and seasonality of occurrence.

A theoretical composite model for population sex-specific shell size dynamics in Strombidae (Gastropoda, Neostromboidae)

ABSTRACT Modelling sexual dimorphism has important implications for understanding the evolution of size relationships among and within organisms. We present a composite model for the regulation and evolution of sex-specific inter-population shell size in a family of herbivorous marine molluscs, the Strombidae. In particular, this model postulates that gene flow acts as a size regulator that limits inter-population divergence by not allowing the mean size of each sex to deviate significantly between disparate populations. The mean shell size of individuals within populations is affected by the ecological setting in which the animal is growing. The model considers niche divergence as inconsequential in the regulation of sexual dimorphism, whereas environmental regulation for the mean size of a population is considered a significant driving factor. Sexual competition, where males may dislodge other males during copulation, is also considered a non-significant driver of male body size because of the open mating system. In such a system, smaller males can simply mate before larger males have matured. Selection for earlier mating in males so as to maximise lifetime fecundity favours male maturation at small body sizes. Conversely, greater fecundity at larger body size in females argues for larger females. The postulated composite model presented here fills a knowledge gap by graphically illustrating the factors affecting inter-and intra-specific variation. These differences include the modelling of mean inter-population shell size, as well as the regulation of intra-population sex size ratios. The components of this new model can be applied to a wider range of sexually dimorphic organisms to explain the evolutionary factors involved in regulating the observed sexual dimorphism.

Mate-guarding duration is mainly influenced by the risk of sperm competition and not by female quality in a golden orb-weaver spider.

Males are expected to mate with as many females as possible, but can maximize their reproductive success through strategic mating decisions. For instance, males can increase their own fitness by mating with high quality females that produce more offspring. Additionally, males can adjust mating effort based on the relative distribution of females and male competitors. To test factors that influence male mate choice, we assessed male mating decisions in the golden silk orb-weaver spider, Trichonephila clavipes (Nephilidae), a species in which females are polyandrous, males guard females before and after copulation occurs and large males are the most successful at guarding mates. We tested the hypothesis that males spend more time guarding high quality females that are spatially isolated, and when the risk of sperm competition is higher. We also hypothesized that this effect increases with male body size. We assessed solitary and aggregated female webs in the field and quantified female quality (i.e., female body condition), male size (i.e., male body size), the risk of sperm competition (i.e., number of males in each female web), and mate-guarding duration (i.e., number of days each male spent in each web). We found that mate-guarding behaviour is largely influenced by the presence of male competitors. In addition, male body size seems to moderately influence male guarding decisions, with larger males guarding for a longer time. Finally, female body condition and type of web (i.e., solitary or aggregated) seem to play small roles in mate-guarding behaviour. As mate-guarding duration increased by 0.718 day per each additional male competitor in the web, and guarding behaviour prevents males from seeking additional mates, it seems that guarding females can be considerably costly. We conclude that failing to guard a sexual partner promotes high costs derived from sperm competition, and a male cannot recover his relative loss in fertilization success by seeking and fertilizing more females. In addition, the search for more sexual partners can be constrained by possible high costs imposed by weight loss and fights against other males, which may explain why the type of web only moderately influenced male mate choice. Following the same rationale, if high-quality females are not easy to find and/or mating with a high-quality female demands much effort, males may search females and guard them regardless of female quality. In conclusion, the factor that most influences male mate-guarding behaviour among T. clavipes in the field is the risk of sperm competition.

Catches in abandoned snow crab (Chionoecetes opilio) pots in the Barents Sea

During a 2018 retrieval cruise for abandoned snow crab (Chionoecetes opilio) pots in the Barents Sea, approximately 8600 pots abandoned 1.5 years earlier were recovered. Forty-three percent of a subsample of 1000 pots contained snow crabs, with an average of three crabs per pot. Most of the crabs were alive (~98%) and dominated by large males. Pinch injuries and limb loss were common and tended to decline with increasing crab size. Reflex testing showed that the crabs were vital (i.e. the crabs moved their legs, chelipeds and maxillipeds when stimulated), which was supported by a relatively high meat content. However, energy reserves in the digestive glands (hepatopancreas reserves) were low, indicating overall energy deficiencies. Our results indicate considerable unaccounted mortality due to self-baiting, continued catch and cannibalism. The findings demonstrate that snow crab pots which are lost or abandoned in the Barents Sea fishery maintain huge potential for ghost-fishing impacts.

Breeding behaviour of ectotherms at high latitudes: the case of the natterjack toad Epidalea calamita at its northern range limit

Abstract We explored the breeding behaviour of a threatened amphibian, the natterjack toad, at its northern range limit in Estonia, to determine the extent to which reproduction is affected by harsh and unstable climatic conditions. Using photo identification of specimens, we found that in optimal weather conditions males formed three breeding cohorts, while in adverse conditions only a single cohort occurred and under extreme conditions reproduction was skipped entirely. During the extended breeding season, larger males participated in reproduction throughout the breeding period, while smaller males appeared in later cohorts. Breeding success was related to the calling effort of a male, where larger males had greater mating success than smaller ones. We found that the natterjack toad males exhibit significant plasticity in reproductive behaviour at the northern range limit, which, given the energetic cost of reproduction and the increased risk of predation, allows them to increase their fitness at high latitudes.

Does body size influence mating success? A morphometric study of two Anastrepha (Diptera: Tephritidae) fruit fly species.

Male mating success depends on various traits and factors, and correctly identifying these traits can be key in the context of pest management. For tephritid pests, controlled through the sterile insect technique (SIT) traits, such as male size, can be manipulated through mass-rearing procedures. Thus, it is particularly important to understand whether male size can favor mating success. Here, we evaluated mating success of males of different sizes in Anastrepha ludens and Anastrepha obliqua, two species controlled through SIT. For both species, a morphometric analysis was performed of mated and unmated mass-reared and wild males in field cages. In both A. ludens and A. obliqua, wild females did not discriminate their mates based on male size and mated more frequently with wild males regardless of size. For mass-reared males, we found no evidence of an advantage of the large males compared to small males in mating success. However, we did find differences between the morphometric traits of mass-reared and wild males. In A. ludens, traits associated to mating success were Face Width (FW), Head Width (HW), Thorax Length (TL) and Wing Length (WL), and for A. obliqua were FW, HW, WL and WW (Wing Width). Overall, FW and TL were more consistent predictors of mating success. In conclusion, female choice seems to suggest multivariate selection, confirming that overall body size (expressed as pupal diameter, which is highly correlated with weight) is not a decisive factor in male mating success in these two species. However, morphological traits such as FW, HW, TL, WL and WW may be relevant in mating preference of wild female.

Territoriality is just an option: allocation of a resource fundamental to the resource defense polygyny in the European wool carder bee, Anthidium manicatum (Hymenoptera: Megachilidae)

The wool carder bee Anthidium manicatum is one textbook example of resource defense polygyny among solitary bees, known for intense male–male competition, forced copulations, and the extreme form of interspecific territoriality toward other flower visitors. This mating system depends on the spatial structure of the defended resource and requires several adaptations in males. The allocation of patches with host plants as well as male body size and phenology was analyzed over 3 years in the diverse habitat of a botanical garden. Anthidium manicatum males searched in groups up to 12 individuals a wide diversity of patches with various food plants of foraging females. Territories were established in small high-quality patches only. Males abandoned aggressive and territorial behavior in large patches. Available patches were occupied by males of the various body size fractions independently of each other according to patch profitability. The higher competitive weight of large males in small patches compared to spacious ones was balanced by an opposing correlation of patch profitability. Although the mating system in A. manicatum is clearly a resource defense polygyny, males were found to be plastic in their behavior, and territoriality was not consistently observed. Mate acquiring tactics, be they territory holder (bourgeois), sneaker, floater, or scrambler for mating, can be considered to be different behavioral phenotypes within one environmentally sensitive conditional strategy.Significance statementTerritoriality is a rare and derived pattern in solitary bee mating behavior. In most cases of territoriality, males defend rendezvous places to meet freshly emerged, virgin females. While this type of mating behavior fits still into the framework of ancestral monandry of aculeate Hymenoptera, the continually polyandric resource defense polygyny found in the genus Anthidium is highly derived. Males occupy flower resources exploited for larval provisions and extort copulations from provisioning nesting females. Territoriality in Anthidium does not lead to a monopolization of females, the exclusion of many competitors from reproduction, and a reduction of sperm competition as is typical for resource-based mating systems. Contrary, Anthidium is a highly promiscuous species and both males and females are lifelong engaged in copulations with multiple mates. Also, the allocation of the resource fundamental to the defense polygyny was found to be more fairly balanced than expected. This study diversifies the mating system of anthidiine bees and demonstrates unusually high plasticity in the resource allocation of a territorial species.