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Landmark Territoriality in the Neotropical Paper Wasps Polistes canadensis (L.) and P. carnifex (F.) (Hymenoptera: Vespidae)
Abstract
Male Polistes canadensis and P. carnifex aggregate along crests of prominent ridges in Santa Rosa National Park, Costa Rica. At these sites males of both species defend territories (trees and shrubs) by chasing conspecific rivals. Territories do not contain nests or resources that are collected by females. Chasing by territorial males reduces the amount of time spent by intruders in a territory. I describe and contrast male territorial behavior of both species. Some male P. canadensis are territorial while others in the same area exhibit patrolling behavior, flying from one occupied territory to another. Males of P. carnifex exhibit territoriality only. Patrolling in P. canadensis is an outcome of relatively high male density along the ridge, rendering territories in short supply, as shown by the observation that experimentally vacated territories are seized rapidly by formerly patrolling males. Due to a high intraspecific intrusion rate, territorial male P. canadensis spend less time perching and more time flying and chasing intruders from their territories than do male P. carnifex. Males of these two species also differ in the placement of their territories along the ridgeline; P. canadensis occupy territories in saddles while P. carnifex occupy those at peaktops. I show that this divergent spatial pattern is not maintained by competitive exclusion of either species by the other, and I discuss alternative explanations for their separate spatial distributions. Comparative data suggest that males are territorial because females restrict matings to within territories, and I discuss alternative hypotheses to explain this bias in female behavior.
... All sugar and food was obtained in batches and haphazardly split between colonies to ensure equal food quality provided to the adults to prevent any longevity variability as a result of adult nutrition (Johanowicz & Mitchell, 2000;Harvey et al., 2012). In addition to food, in each nest-box we provided plastic artificial planting in the form of a strip of 7 cm × 7 cm × 2.5 cm grass sp. and a 5 cm × 5cm × 5 cm plastic Hedera sp. for environmental enhancement to provide shelter from female aggression for males (Polak, 2010). The nest boxes were cleaned regularly with distilled water without disturbing wasps or nest. ...
Insects have been used as an exemplary model in studying longevity, from extrinsic mortality pressures to intrinsic senescence. In the highly eusocial insects, great degrees of variation in lifespan exist between morphological castes in relation to extreme divisions of labour, but of particular interest are the primitively eusocial insects. These species represent the ancestral beginnings of eusociality, in which castes are flexible and based on behaviour rather than morphology. Here we present data on the longevity of the primitively eusocial Neotropical paper wasp P. canadensis, in a captive setting removed of environmental hazards. Captive Polistes canadensis had an average lifespan of 193 ± 10.5 days; although this average is shorter than most bee and ant queens, one individual lived for 506 days in the lab-longer than most recorded wasps and bees. Natal colony variation in longevity does exist between P. canadensis colonies, possibly due to nutritional and genetic factors. This study provides a foundation for future investigations on the effects of intrinsic and extrinsic factors on longevity in primitively eusocial insects, as well as the relationship with natal group and cohort size.
... Male mason bees, Hoplitis anthocopoides Schenck, compete for and defend territories for mate encounter, learn their location, and return to (Eickwort, 1977). Similarly, males of paper wasp, Polistes spp., maintain mate-encounter territories, depart at night and return each morning (Polak, 1993), obviously having learned their location. ...
The ability of insects to learn locations of future resources has rarely been studied. Here, we show that males of the solitary parasitoid wasp Pimpla disparis Viereck (Hymenoptera: Ichneumonidae) learn locations of future mates. Male P. disparis reportedly arrest on parasitized pupae of wax moth, Galleria mellonella L. (Lepidoptera: Pyralidae), and gypsy moth, Lymantria dispar L. (Lepidoptera: Erebidae), when mate emergence is imminent. We tested the hypothesis that male P. disparis identify, memorize, and revisit the location(s) of parasitized host pupae as a strategy to attain mates. We colour‐coded P. disparis males in the field and noticed that they revisit parasitized moth pupae on consecutive days, and arrest on those pupae with a near‐emergence P. disparis parasitoid. In a laboratory experiment with two large corrugated cardboard cylinders (CCCs) as surrogate trees, each CCC bearing two parasitized moth pupae with a near‐emergence P. disparis parasitoid or two pupae not parasitized, males on day 1 of the experiment visited parasitized pupae more often than pupae not parasitized. On day 2, when each CCC had been replaced and now carried pupae that were not parasitized, males returned to the same CCC, or the same micro‐location on that CCC, which on day 1 had carried parasitized pupae. Field and laboratory data combined indicate that male P. disparis learn the location of future mates. With female P. disparis being haplodiploid and capable of reproducing without mating experience, the onus to find a mate is on males. They accomplish this by detecting parasitized pupae, learning their location, revisiting them frequently, and then arresting on them when the prospective mate nears emergence, taking a 50% chance that it is indeed a female.
... The mating systems of social wasps reveal geographic variability. For example, the males of Polistes canadensis (L.) demonstrate territorial behavior on the nests in eastern Colombia (West-Eberhard, 1983), whereas in Costa Rica they demonstrate territoriality outside the nest, patrolling prominent elements of the landscape (Polak, 1993). At the same time, the males of P. dominulus (Christ) have similar reproductive strategies both in southern Italy and in southern Ukraine. ...
The reproductive strategies of Polistes nimphus (Chris) and P. gallicus (L.) males in the Low Dnieper area are described. In the P. nimphus population, territorial and migrating males were observed. The territorial males of P. gallicus formed swarms or patrolled their areas singly. The male color variants and size were shown to correlate with their reproductive
strategies. The territorial males of P. nimphus had a paler coloration and smaller head and wings than migrating males. The swarming territorial males of P. gallicus were paler and had a larger head and wings than single territorial males. The geographic variability of precopulatory behavior
of males of the studied species is discussed.
... Although body size plays some role in mating behaviour of some paper wasps (Polak, 1993), it is not surprising that our second hypothesis was also supported and that neither males nor females chose larger or smaller partners as body size does not seem to play a significant role in any part of social life of R. marginata (Gadagkar, 2001). ...
We investigated the effect of nestmateship and body size on mate selection through a choice based assay in the primitively eusocial wasp Ropalidia marginata. A recent study has shown that male and female R. marginata mate with their nestmates and non-nestmates with equal probability if no choice is available. That study could also not detect any influence of body size on mating probability in the absence of choice. To confirm that the same results can be obtained even when the wasps have a choice, we offered a choice of two virgin partners either to a virgin test male or to a virgin test female and measured the probability that the test individual would mate with any particular partner based on nestmateship or body size. We show here that even when a choice is available, neither male nor female test wasps base their mate choice on the nestmateship or body size of the partner. We therefore suggest that the natural mating habit of these wasps is sufficiently promiscuous and not constrained by such factors as nestmateship and body size.
Little is known about the roles of sex pheromones in mate-finding behavior of social wasps (Vespidae). Working with the aerial yellowjacket, Dolichovespula arenaria (Fabricius), baldfaced hornet, Dolichovespula maculata (L.), western yellowjacket, Vespula pensylvanica (Saussure), southern yellowjacket, Vespula squamosa (Drury), and Vespula alascensisPackard, we tested the hypotheses (1) that gynes produce an airborne sex pheromone attractive to males, and (2) that males are more strongly attracted to non-sibling gynes based on olfactory cues. A field experiment provided the first definitive evidence that D. arenaria gynes attract males. Surprisingly, we did not find such evidence in similar field experiments for sexual attractiveness of gynes of V. squamosa, V. pensylvanica, V. alascensis, or D. maculata. In Y-tube olfactometer experiments with three of these species (D. arenaria, D. maculata, V. pensylvanica), only D. maculata gynes attracted males, provided they were non-siblings, implying an olfactory-based mechanism of nestmate recognition and inbreeding avoidance. Lack of sex attraction responses for V. pensylvanica, V. alascensis, and V. squamosa in this study does not rule out pheromone-mediated sexual communication. Instead, it highlights the possibility that pheromonal signaling may be dependent on the presence of appropriate contextual cues.
The macroparasite Xenos vesparum affects both the behaviour and the physical traits of its host, the social wasp Polistes dominulus. Female wasps, if parasitized, do not perform any social tasks and desert the colony to gather at specific sites, where the parasite mates; at the end of summer they form prehibernating clusters joined by healthy future queens to overwinter. Parasitized wasps become highly gregarious. In April, healthy wasps leave the aggregations to found new colonies, while parasitized wasps remain in overwintering groups and release parasites to infect wasp larvae only later in the season. We studied the prolonged gregarious behaviour of parasitized wasps and analysed the morphology of parasitized and healthy wasps in aggregations collected over a 7-year period to determine whether the parasite affects host size, wing symmetry, ovarian development and lipid stores. All parasitized wasps were smaller and had undeveloped ovaries and more wing fluctuating asymmetry than unparasitized wasps, irrespective of time of year, parasite load and parasite sex. If infected only by one or two X. vesparum females, the wasps had large fat bodies, which could facilitate their overwintering. In contrast, wasps infected by at least one male parasite had little lipid and died at the end of the summer. Thus, X. vesparum, may play a role in the fate of its host, by exploiting wasps' tendency to form aggregations outside the colony and by altering its caste system, nutrient allocation, diapause timing and life span to achieve its own reproduction and dispersal.
Social wasps need an efficient communication system to coordinate their members in the numerous activities of the colony. In this regard, the chemical channel is the most utilized by social wasps to transfer information in intraspecific (pheromones) and interspecific (allomones) communication. In this chapter, we reviewed the main chemical substances which mediate recognition between colony members and coordinate nest defense, alarm and recruitment. Due to their central role in the colonial life, the majority of pheromones have been identified and their functions have been deeply investigated in many species. On the contrary, sex pheromones which are the most studied in insects, have been quite neglected in social wasps.
Insects of the order Hymenoptera are biologically and economically important members of natural and agro ecosystems and exhibit diverse biologies, mating systems, and sex pheromones. We review what is known of their sex pheromone chemistry and function, paying particular emphasis to the Hymenoptera Aculeata (primarily ants, bees, and sphecid and vespid wasps), and provide a framework for the functional classification of their sex pheromones. Sex pheromones often comprise multicomponent blends derived from numerous exocrine tissues, including the cuticle. However, very few sex pheromones have been definitively characterized using bioassays, in part because of the behavioral sophistication of many Aculeata. The relative importance of species isolation versus sexual selection in shaping sex pheromone evolution is still unclear. Many species appear to discriminate among mates at the level of individual or kin/colony, and they use antiaphrodisiacs. Some orchids use hymenopteran sex pheromones to dupe males into performing pseudocopulation, with extreme species specificity.
Males of the red admiral butterfly Vanessa atalanta (Nymphalidae) establish and defend territories in the late afternoon on a central Arizona hilltop. Resident males engage conspecific rivals in lengthy chases but respond less aggressively toward males of 3 congeneric species. A comparison between V. atalanta and 2 conge-ners, V. annabella and V. cardui, shows that the degree of site tenacity is correlated with the density of rivals. Males of V. annabella, the least abundant species, are most site tenacious, as measured by mean duration of residency and frequency of return to the peak. In contrast, males of V. cardui experience the highest hilltop densities and are the most ephemeral. Males of V. atalanta occur in intermediate density and exhibit intermediate site tenacity. Although male V. atalanta readily respond to any flying insect, the duration and complexity of these interactions is reduced. The various species also express different perch site preferences, which may aid in reducing the frequency of non-productive congeneric encounters.
Mate choice in noneconomic mating systems has been considered paradoxical because, relative to economic systems, females were thought to have "highly developed' preferences, despite males' having little to offer. Efforts to resolve this paradox have generally searched for genetic benefits of choice through either "good genes' or "runaway' coevolution. This paper emphasizes natural selection acting directly on females and their offspring. Although females are expected to pay lower costs in noneconomic mating systems, this need not translate into examining fewer males or spending less time in this activity. Furthermore, various direct (nongenetic) benefits may accrue. In species in which males offer benefits that are more variable, such as territories or parental care, females should evolve toward greater investment in mate choice, especially when these resources cannot be shared among females. Any tendency for females to be more selective in noneconomic mating systems, despite lower benefits, can probably be explained if the much lower costs of search, and thus net benefits, are considered. Therefore, there may be no lek paradox. -Authors
A study of hilltopping behavior on a forested hilltop in Tennessee has revealed a vertical zone for Papilio glaucus separate from other Papilioninae. Separate hilltop vertical zones for species occupying the same horizontal habitat increase the species-packing capacity of the habitat and may increase the likelihood of successful mate location by reducing the interspecific encounter rate. -from Author
Territories of birds, usually defended against conspecific individuals, are sometimes defended against individuals of other species. Since such behavior is demanding both of time and energy, natural selection should favor ecological should favor ecological divergence, the establishment of overlapping territories, and the reduction of aggression. Lack of divergence in modes of exploitation could mean that insufficient time has elapsed for the changes to be completed or that the environment imposes some limitation preventing the evolution of the required degree of divergence. Such environmental limitation can be predicted in (a) structurally simple environments, (b) when feeding sites are strongly stratified in structurally complex vegetation, or (c) when the presence of other species in the environment prevents divergence in certain directions. The known cases of interspecific territoriality in birds are analyzed and shown to be largely in accordance with these predictions, although several cases of overlapping territories in situations where interspecific territoriality has been predicted provide relationships worthy of further study. We suggest that Darwinian selection at the level of the individual permits an understanding of the known structure of avian communities and that there is no need at present to invoke new selective mechanisms at the level of the community or ecosystem.
Three species of Leucorrhinia dragonflies show extensive temporal and spatial overlap in their use of mating grounds. This sets the stage for potentially costly interspecific interactions during mating and subsequent oviposition. Males frequently attempt to mate with heterospecific females, but prolonged interspecific matings are uncommon. Males guard ovipositing mates against takeovers by intruding males. Guarding males were as likely to chase heterospecific intruders as conspecific intruders during the oviposition period. Males incurred considerable reproductive cost from this lack of species discrimination; 29% of the successful takeovers by conspecifics occurred while males were chasing heterospecific intruders. One hypothesis for lack of species discrimination in this group is that effective discrimination would require a time investment by the guarding male while he assessed the species identity of the intruder. During this assessment period, the guarding male would risk losing his female to the intruder.
Alternative male mating tactics of insects at landmarks (leks) have only rarely been investigated. Some males of the paper wasp, Polistes canadensis (L.), were territorial at small trees along the crests of dry ridges in Santa Rosa National Park, Costa Rica. Territories did not contain nests or resources for which females foraged. Contrary to other ''hilltopping'' species, male P. canadensis competed most intensely for territories in saddles 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, suggesting that both size-related tactics belong to one conditional strategy. Males that were territorial on 2 or more days were larger than those that were territorial on only 1 day, and these in turn were larger than permanent patrollers. Moreover, the mean size of territorial males was positively correlated with two measures of territory attractiveness, suggesting that larger males monopolize preferred sites. Mean age of territorial males was also related to territory attractiveness, but males of intermediate age claimed the most attractive territories.
Males of the tarantula hawk wasp Hemipepsis ustulata defend conspicuous palo verde trees on mountain ridges. A comparison of data from two flight seasons (1981 and 1980) reveals that the preference rankings of males for a set of perennial territories on one ridge remained highly consistent from generation to generation. This is the first thoroughly documented demonstration for insects that landmarks used as mating encounter sites retain the same relative attractiveness from year to year. The stability of male preferences is indirect evidence that access to females is related to the ability to acquire high-ranking territories. Moreover, in both years, (1) the flight season was 2.5 to 3 months long, (2) the range of body sizes in this highly variable species remained the same, and (3) the ratio of territorial males to non-territorial patrollers was constant. A key difference was a strong reduction in the total number and density of males found in the study site in 1981. The wasps responded to the decrease in competition for perch territories by failing to occupy some low-ranking sites that were taken in 1980.
Insects of many groups form mating aggregations on hilltops. Males of some hilltopping species behave like certain classic lekking vertebrates, notably in their defense of territories that are devoid of resources useful to females but which are visited by females solely for mating. But even within a sample of 11 hilltopping insects found in the same habitat in central Arizona, there is considerable diversity with respect to resources transferred to mates by males, the degree to which males are aggregated and territorial, and the extent to which females can choose freely among hilltopping males. Thus ‘hilltopping’ is not a single mating system type; although some species conform closely to the traditional definition of a lek, others are very different. The ecological basis for the diversity in the behaviour of male hilltopping insects appears linked to differences in population density, which affect the costs of territoriality, and differences in the nature of female choice, which are little understood as yet.
SUMMARY 1. Flies of the genus Syrphus aggregated at specific sites in the field ('leeks'). Flies at leeks were always capable of 'instant' take-of, even at ambient temperatures of 10 °C or less. 2. The flies regulated their thoracic temperature by a combination of basking and shivering. During hovering flight in sunshine thoracic tempera- ture rose 12-14 °C above the ambient temperature. 3. The flies engaged in frequent brief chases while at the leeks. 4. At an air temperature > 18 °C the flies at the leek remained in hover- ing flight most of the time. 5. The vibration frequencies of the thorax during shivering and flight ranged from about iooto 200 Hz at 10-27 °C, though at a given temperature and spike frequency the vibration rate during warm-up was higher than the wing-beat frequency (assumed to be the same as thoracic vibration frequency) during flight. 6. During shivering, but not in flight, there is a tendency for the indirect flight muscles to be activated in synchrony.
1. The biology of the eumenid mud-wasp Pachodynems nasidens (Latr.) was studied in trapnests.
2. Females nested in traps placed in the shade and some nested in the same traps from which they had emerged.
3. The nesting behaviour and nest structure were generally similar to those of other eumenids that nest in borings, but a temporary crescent-shaped plug at the nest entrance and the use of both dry soil and mud for nest-building seems unique to P.nasidens.
4. Like most other eumenids, P.nasidens collected caterpillars, mainly Pyral-idae, Olethreutidae, Alucitidae and Thyrididae.
5. In linear nests, the innermost cells contained females and the outermost cells males, resulting in proterandry. One female can make several nests, each showing proterandry.
6. Female immatures were larger and took more time to develop than males. Also, their cells were larger and stocked with more food than male cells, hence needing more time to be provisioned.
7. The greater ‘cost’ to produce a female and a sex ratio biased toward females in short traps and about unity in longer traps, leads to a comment on parental investment.
Male Grey hairstreak butterflies employed two different tactics, centred on palo verde trees growing on hilltops in central Arizona. During much of the spring flight season of about four months single males repelled all others from a perch tree. Interactions between the territorial resident in a tree and visiting intruder males were frequent but usually brief. One male retained possession of his perch for two weeks, although changes in ownership were generally much more frequent. But territorial behaviour was abandoned during periods when the number of intruders increased sharply, at which time perched males switched to non-aggressive patrolling about the preferred palo verdes. Females occasionally visited the palo verdes and interacted with males but no matings were observed. The mating system of the Grey hairstreak, like that of other similar landmark-defending insects, appears to have evolved in response to a diffusely distributed population of receptive females.
Males of the paper wasp Polistes commanchus aggregate in large numbers on hilltops in central and southeastern Arizona where individual males defend perch sites on shrubs, trees or rocks. Residents chase intruders away quickly and exhibit strong site fidelity at their perches. Takeovers are rare as are prolonged battles for possession of a site. Males have only weak preferences for peaktop perch sites as opposed to those lower on hillsides, and a similarly weak preference for perches in shrubs as opposed to those on rocks. Receptive females appear occasionally at the male aggregation sites to mate with perch holders. The relatively weak territoriality exhibited by P. commanchus may stem in part from the high density of males at peaktops and the correspondingly high rate of interactions. The evolution of hilltopping in a species whose reproductive females are clumped spatially at nest sites poses a puzzle for the ecological analysis of mating systems.
and Summary
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.
1. In order to understand fully the evolution of a behavioural trait one must not only consider whether it is adaptive in its present environment but also whether it originated as an adaptation to existing selective forces or as a fortuitous consequence of selection for a different role in other environments (i.e., as a pre‐adaptation) or of selection for different traits (e.g., as a pleiotropic effect). In this paper interspecific territorialism is examined in species of humming‐birds, sun‐birds, tropical reef fishes, stingless bees, stomatopods, crayfish, and limpets as a means of determining its adaptiveness and its origins.
2. Humming‐birds form complex assemblages with species sorted out among the available resources. Dominant species establish feeding territories where flowers provide sufficient nectar. A few large, dominant species, usually uncommon, are marauders on others' territories. Subordinate species establish territories where flowers are more dispersed or produce less nectar, or they fly a circuit from nectar source to nectar source when flowers are even more dispersed, a foraging pattern called ‘traplining’, or they steal nectar from the territorial species by being inconspicuous while foraging. Two species, Amazilia saucerottei and Selasphorus sasin , subordinate in one‐to‐one encounters, are able to take over rich resources by establishing several small territories within a territory of a dominant and forcing it to forage elsewhere.
3. Among humming‐birds, territorial individuals attacked not only subordinate competitors but marauding humming‐birds and some insects, which stayed in the territory and foraged at will, and seemingly inappropriate targets, such as non‐competitors. This suggests that the stimulus for aggression is ‘any flying organism near the food resources’, regardless of its appearance. The behaviour rather than the identity of the intruder is the stimulus.
4. Sun‐birds resemble humming‐birds to the extent that dominants establish territories on rich nectar sources and subordinates establish territories on less rich nectar sources or steal from the territories of dominants. The diversity of foraging patterns is not so great as in humming‐birds, perhaps because so few species of sun‐birds have been studied. However, the advantage of territorialism has been measured in the sun‐bird Nectarinia reichenowi. Individuals with territories lose much less nectar to competitors than do those without territories.
5. Field work on three species of tropical reef fishes involved a single aggressive species whose individuals attacked a wide range of species intruding on their territories. The stimulus for aggression in Pomacentrus jenkinsi seemed to be an “object moving through [its] territory”. As suggested for humming‐birds, the stimulus is the behaviour rather than the identity of the intruder.
6. The relationships found in stingless bees, stomatopods, crayfish, and limpets are simpler. The dominant and subordinate species divide the resources in their habitat, the dominants' aggression preventing the subordinates from using resources that were otherwise available to them.
7. A general pattern emerges. Mutual interspecific territorialism occurs between species that (i) have different geographic ranges, (ii) occupy different habitats, or (iii) use different resources within the same habitat. Examples of two species holding separate territories on the same resources within the same habitat are rare and occur when the dominant species is rare relative to the available resources. These observations are contrary to the usual view that interspecific territorialism is an adaptation that permits co‐existence of potential competitors within the same habitat.
8. Interspecific territorialism is sometimes adaptive and sometimes maladaptive, depending upon the species and the situation.
9. The general pattern of occurrence of the behaviour and the general nature of the stimulus for aggression, i.e., the behaviour rather than the identity of the intruder, suggest that interspecific territoriality is a fortuitous consequence of selection for intraspecific territorialism, the latter being not only an adaptation to the presence of conspecific competitors but a pre‐adaptation to the presence of competitors of other species, should they occur.
In summer, males of Polistes dominulus form large aggregations at sunny landmarks. We identified two size-correlated behavioural categories: residents (R) and transient (T). R males, which constitute 20%–25% of the total population, are larger than T males, territorial, aggressive, and more site-faithful, while T males range more widely, are non-aggressive, and show little site tenacity. Field and laboratory data suggest that R males have an advantage in mating, particularly if they engage in frequent flights while on their territories. These alternative mating tactics within the same population are combined with behavioural flexibility in some individuals, which switch from one option to the other.
In lek-mating species, males defend territories lacking in resources attractive to females and females visit these sites only to mate (Bradbury, 1977). Emlen and Oring (1977) predicted that species are likely to possess a lek-mating system when their breeding season is long and when females or resources critical to female reproduction are widely dispersed in space and time, that is, when costs to males of searching for females are prohibitively high. In contrast, resourceand female-defense polygyny are more likely to evolve when resources or females are concentrated so that the energetic expenditure associated with defense falls within the capabilities of individual males. In Polistes paper wasps, males of some species mate-search at sites where females are concentrated, consistent with the predictions of Emlen and Oring (1977). For example, male P. fuscatus defend territories at female hibernation sites (Post and Jeanne, 1983a), and male P. erythrocephalus intercept females tending newly initiated nests (West Eberhard, 1969). In West Eberhard's (1969) study, 25% of females at new nests were virgin: thus there appears to be a strong incentive for males to mate-search at these sites. P. canadensis males defend territories along the summit of dry ridges in northwest Costa Rica from approximately May to August (Polak, 1992). These hilltop territories contain no resources or nests. Because females concentrate at nests away from the hilltop, the lek behavior of male P. canadensis seems to pose an exception to the predictions of Emlen and Oring (1977). Matthes-Sears
1.
Some males of the megachilid bee Anthidium maculosum establish territories at patches of flowering Monarda and may remain there for up to at least 21 days. The frequency of male-male interactions and territorial takeovers appears partly dependent upon population density.
2.
Males that do not hold territories tend to be smaller than territorial individuals. They either visit many territories repeatedly as wandering intruders or occupy a corner of one territory as satellite males. They always flee when challenged by territory owners but often return sooner or later. Although most matings are performed by territory owners, non-aggressive subordinates do occasionally copulate.
3.
Females may mate dozens of times in the span of a week. Multiple mating may be an adaptation designed to cope with male harassment while they exploit the food resources in territories. It may save time and energy and reduce foraging interruptions to copulate passively for 30 s rather than to try to evade or repel the large, aggressive males.
4.
A male may mate with the same female at intervals of about 6 min and may guard her against intruders during her visit to his territory. These traits may help a territorial male to be the last male to copulate with a female prior to oviposition. If sperm precedence occurs in A. maculosum only this last male is likely to fertilize her egg.
Male Leucorrhinia dragonflies defend territories from conspecific and heterospecific intruders. Defense against heterospecifics is surprising, as mating and oviposition are the only activities that occur on the territories, and heterospecific males are not expected to pose a reproductive threat. L. frigida and L. intacta males respond aggressively with equal frequency and intensity against intrusions by conspecifics and heterospecifics. In contrast, L. proxima males respond more aggressively against conspecifics. The apparent lack of species discrimination shown by L. frigida and L. intacta males may result because territorial males that assess intruders (as do L. proxima) suffer a tactical disadvantage from hesitating when an intruder flies in. This assessment process may lead to reduced fighting success by the territorial male. Thus there is a tradeoff between assessment cost incurred when the intruder is a conspecific and benefit gained from avoiding conflict when the intruder is a heterospecific. Given this assessment cost, males of species that only rarely encounter conspecific intruders are more likely to evolve species discrimination than males of species that commonly encounter conspecific intruders.
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.
Females of the solitary eumenid wasps Ancistrocerus adiabatus and Euodynerus foraminatus control the adult size of their offspring by the amount of food provded to the larvae. For both species, larger females provision more offspring and collect more food than smaller females. Males of E. foraminatus, upon emergence as adults, fight for control of the nest area. The winning brother remains at the nest and mates with his sisters as they emerge several days later. Males of A. adiabatus are not as agonistic as males of E. foraminatus so that a number of males may remain near the natal nest. When females nest near each other, male-male interactions increase, and the likelihood of sibling mating decreases. Under conditions of clumped nesting, females make larger offspring and invest more in males than when nesting in isolation.
Polistes major males mark perches and defend patrol routes much like other male vespids, but are unusual in the enlarged ectal mandibular glands and mandibular applicator brushes. Facial rubbing is a prominent behavior in addition to the more wide spread sternal rubbing behavior. Glandular size and the volatile character of the product suggest adaptation for long distance communication. The pheromone of the gland is apparently without aphrodisiac properties and appears to be the same compound in two subspecies collected from distant localities. Laboratory reared wasps do not discriminate against kin in mating.Les mles dePolistes major marquent leurs perchoirs et dfendent leurs parcours de patrouille comme les autres mles de la familleVespidae, mais leurs glandes mandibulaires latrales sont agrandies d'une faon peu commune ainsi que leurs brosses mandibulaires. Ils frottent frquemment leurs perchoirs avec leur face de mme qu'avec le sternum, ce dernier comportement tant plus largement rpandu dans la famille. La taille de la glande et les caractres des produits volatils suggrent une adaptation la communication grande distance. Il semble que la phromone de la glande n'ait pas de proprits aphrodisiaques et qu'elle possde la mme composition chez deux sousespces rcoltes dans deux localits loignes. Au laboratoire, les gupes s'accouplent sans distinction de partenaires, incluant donc l'inceste.
1.
The breeding season of H. ustulata extends over a 2 1/2 month period in the Sonoran desert near Phoenix, AZ. Individual males are active on mountain ridges for periods averaging about 3 weeks for those wasps captured in the first half of the flight season. Many males live more than 1 month.
2.
Some palo verde trees chosen as landmark territories by male wasps are occupied earlier in the flight season and more consistently thereafter than others. Preferred territories tend to be located higher on ascending ridges.
3.
There is considerable turnover in territory ownership at some trees during the course of the flight season. The average duration of residence by a male wasp was 8 days for all occupants of 18 trees followed over the breeding period. Some changes in ownership occur because an intruder defeats the resident in an aerial contest. Small males are replaced by large ones at preferred territories over the course of the flight season.
4.
Smaller males can adjust their behavior to their social evironment, claiming low-ranking territories if excluded from favored trees or holding prime territories during days when large males are absent. Alternatively, they may abandon territoriality altogether to patrol a series of trees along a ridge.
5.
Despite an abundance of males (>350 marked), females were rarely seen and only two copulations were observed. Male territoriality was not focused directly on emerging or nesting females nor was it focused on food resources attractive to females. The breeding season was prolonged (2 1/2 months) and the operational sex ratio heavily skewed toward males. The wasp's mating system resembles lek polygyny in the sense of Emlen and Oring (1977).
At the end of summer, males of Polistes gallicusfly in swarms around vertical landmarks and land in clusters on their favorite perches, where they drag their legs and abdomen. Here males occasionally crowd around a perched female; they make no effort to defend an exclusive mating territory but instead attempt to copulate by displacing rivals from the female. In this work we describe this spatial-nuptial system, which entails site fidelity without territoriality, unisexual swarms, common patrol routes, collective sexual approaches, and scramble competition polygyny. Mating success is evaluated in relation to the familiarity with flight paths (routine patrollers versus newcomers), to the type of sexual approach (single males versus in- group males), and, in the laboratory, to the individual activity level.
Hymenop-tera: Vespidae): large size advantage and alternative mate-acquisition tactics Male reproductive behavior of the social wasp Polistes fuscatus (Hymenoptera: Vespidae) Costs and benefits of female mate choice: Is there a lek paradox?
- Post
- D C Jeanne
- R L
- J D Reynolds
- M R Gross
_ _ 1993: Competition for landmark territories among male Polistes canadensis (L.) (Hymenop-tera: Vespidae): large size advantage and alternative mate-acquisition tactics. Behav. Ecol., in press. POST, D. C. & JEANNE, R. L. 1983: Male reproductive behavior of the social wasp Polistes fuscatus (Hymenoptera: Vespidae). Z. Tierpsychol. 62, 157-1 71. REYNOLDS, J. D. & GROSS, M. R. 1990: Costs and benefits of female mate choice: Is there a lek paradox? Am. Nat. 136, 23C-243.
Hilltopping behavior of Polistes commancbus navajoe MURRAY, B. G. 1981 : The origins of adaptive interspecific territorialism
- Michal Polak
- P Maithes-Sears
- W Alcock
MICHAL POLAK, Landmark Territoriality in Polistes canadensis and P. carnifex MAITHES-SEARS, W. & ALCOCK, J. 1986: Hilltopping behavior of Polistes commancbus navajoe MURRAY, B. G. 1981 : The origins of adaptive interspecific territorialism. Biol. Rev. 56, 1-22.
Parasitology: The Biology of Animal Parasites Interspecific territories in birds
- E R Noble
- G A Noble
- Lea
- Febiger
- G H Orians
- M F Willson
NOBLE, E. R. & NOBLE, G. A. 1982: Parasitology: The Biology of Animal Parasites. Lea & Febiger, ORIANS, G. H. & WILLSON, M. F. 1964: Interspecific territories in birds. Ecology 45, 7 3 6 7 4 5. PAJUNEN, V. I. 1964: Aggressive behaviour in Leucorrbiniu caudalis Charp. (Odon., Libellulidae).
The Evolution of Insect Mating Systems Vertical stratification of hilltopping behavior in swallowtail butterflies (Papilionidae)
- R Thornhill
- J Alcock
- Cambridge
- J Turner
THORNHILL, R. & ALCOCK, J. 1983: The Evolution of Insect Mating Systems. Harvard Univ. Press, Cambridge. TURNER, J. 1990: Vertical stratification of hilltopping behavior in swallowtail butterflies (Papilionidae). J. Lep. SOC. 44, 174-179.