Recruitment to food by the German yellowjacket, Vespula germanica

The hypothesis that Vespula germanica foragers can recruit nestmates to food resources was tested using a protocol that controlled for the biasing effects of social factors at the resource, including local enhancement and food-site marking substances. Foragers from an observation colony in the field were trained to visit a dish of scented corn syrup solution 15?m east of the nest. A second feeding station, 22?m northeast of the nest, offered incoming foragers a choice between food with the training scent and food with a control scent. Significantly more naive foragers arriving at that station chose the food with the training scent. We conclude that the German yellowjacket is able to recruit nestmates to carbohydrate food sources, and that recruits use food odor to locate the source of food being brought into the nest.     

Effects of urban greenspace configuration and native vegetation on bee and wasp reproduction

Pollinator welfare is a recognized research and policy target, and urban greenspaces have been identified as important habitats. Yet, landscape-scale habitat fragmentation and greenspace management practices may limit a city's conservation potential. We examined how landscape configuration, composition, and local patch quality influenced insect nesting success across inner-city Cleveland, Ohio (U.S.A.), a postindustrial legacy city containing a high abundance of vacant land (over 1600 ha). Here, 40 vacant lots were assigned 1 of 5 habitat treatments (T1, vacant lot; T2, grass lawn; T3, flowering lawn; T4, grass prairie; and T5, flowering prairie), and we evaluated how seeded vegetation, greenspace size, and landscape connectivity influenced cavity-nesting bee and wasp reproduction. Native bee and wasp larvae were more abundant in landscapes that contained a large patch (i.e., >6 ha) of contiguous greenspace, in habitats with low plant biomass, and in vacant lots seeded with a native wildflower seed mix or with fine-fescue grass, suggesting that fitness was influenced by urban landscape features and habitat management. Our results can guide urban planning by demonstrating that actions that maintain large contiguous greenspace in the landscape and establish native plants would support the conservation of bees and wasps. Moreover, our study highlights that the world's estimated 350 legacy cities are promising urban conservation targets due to their high abundance of vacant greenspace that could accommodate taxa's habitat needs in urban areas.     

Control of reproduction in social insect colonies: individual and collective relatedness preferences in the paper wasp, Polistes annularis

Social insect colonies often have one or a few queens. How these queens maintain their reproductive monopoly, when other colony members could gain by sharing in the reproduction, is not generally known. DNA microsatellite genotyping is used to determine reproductive interests of various classes of colony members in the paper wasp, Polistes annularis. The relatedness estimates show that the best outcome for most individuals is to be the reproductive egg-layer. For workers, this depends on the sex of offspring: they should prefer to lay their own male eggs, but are indifferent if the queen lays the female eggs. The next-best choice is usually to support the current queen. As a rule, subordinates and workers should prefer the current queen to reproduce over other candidates (though subordinates have no strong preference for the queen over other subordinates, and workers may prefer other workers as a source of male eggs). This result supports the theory that reproductive monopoly stems from the collective preferences of non-reproductives, who suppress each other in favor of the queen. However, we reject the general hypothesis of collective worker control in this species because its predictions about who should succeed after the death of the present queen are not upheld. The first successor is a subordinate foundress even though workers should generally prefer a worker successor. If all foundresses have died, an older worker succeeds as queen, in spite of a collective worker preference for a young worker. The results support the previous suggestion that age serves as a conventional cue serving to reduce conflict over queen succession. Received: 3 May 1996 / Accepted after revision: 22 September 1996     

Mating in bighorn sheep: frequent male reproduction via a high-risk “unconventional” tactic

Rocky Mountain bighorn rams use three distinct tactics in competition for mates. Two tactics (tending and blocking) feature defense and cooperative mating over a relatively prolonged consort period (up to 3 days). In the coursing tactic, subordinate rams fight dominants for temporary copulatory access (lasting seconds) to defended ewes. By combining population-wide genetic (microsatellite) exclusion of paternity, behavioral data and a model of bighorn reproductive competition, we estimated that coursing rams fathered 44% of 142 lambs assigned paternity in two natural populations. In one population, the probability of successful defense against coursing was lowest among rams that had many female consorts and held highest dominance rank. Even so, per capita annual male reproductive success was positively associated with social rank in both herds when measured in terms of fall conceptions. The proportion of coursing versus defending ram matings in each population (0.36 and 0.39) was similar to the corresponding fraction of lambs (0.43 and 0.47) fathered by coursing rams, suggesting that sperm competition approximated a fair lottery. Male traits important in gaining social status and obtaining cooperative consorts with ewes were different and potentially in conflict with those needed to defend against (and practise) coursing. Although the concussive weapons (horns) of rams are less dangerous than, for example, the piercing weapons of other bovids, injury from falls and horn blows during coursing brawls may cause death, handicap future mating competition or increase the risk of predation. Coursing is a rare example of an unconventional alternative mating tactic that is high-gain and high-risk. Received: 17 April 1996 / Accepted after revision: 15 March 1997     

Queen and worker policing in the tree wasp Dolichovespula sylvestris

Insect societies are sometimes exploited by workers who reproduce selfishly rather than help to rear the queens offspring. This causes a conflict-of-interest with the mother queen and, frequently, with the non-reproductive workers as well. One mechanism that can reduce conflict is policing, whereby either the queen or other workers aggress egg-laying workers or destroy worker-laid eggs. Here we present the first direct observations of queen and worker policing in natural, unmanipulated colonies of a social insect, the tree wasp Dolichovespula sylvestris. Worker reproduction was common, with workers producing 50% of all male eggs. However, most worker-laid eggs, 91%, were policed within 1 day, whereas most queen-laid eggs, 96%, remained unharmed. The workers were responsible for 51% of all policing events and the queen for 49%. The workers and mother queen also commonly aggressed ovipositing workers, and successfully prevented them from depositing eggs in 14% and 6% of all attempted ovipositions. Hence, both queen policing and worker policing occur and policing acts via two distinct mechanisms: selective destruction of worker-laid eggs and aggression of ovipositing workers. At a general level, our study shows that both centralized and decentralized control can act together to suppress conflict within social groups.Communicated by R. Page     

Mating systems in the sea anemone genus Epiactis

Four morphologically similar species in the sea anemone genus Epiactis exhibit overlapping distributions on the Pacific coast of North America; E. prolifera, E. lisbethae, E. ritteri and E. fernaldi. All brood their offspring up to the juvenile stage, but each has a different combination of internal versus external brooding and hermaphroditism versus gonochory (separate sexes). Specimens were collected from sites ranging from British Columbia to southern California between December 1988 and July 1992. Mating systems were inferred from genetic comparisons of mothers and offspring histological analyses of sex expression and observations on brooding and spawning behavior. Allozyme and multilocus DNA fingerprint analyses of the gynodioecious hermaphrodite E. prolifera showed that offspring were all identical to their mothers, a result consistent with either asexual reproduction, self-fertilization or extreme biparental inbreeding. In the gonochore E. lisbethae, mothers and offspring were also electrophoretically identical, but variation in DNA fingerprints indicated cross-fertilization. Similar DNA fingerprint differences between mother and offspring in the gonochore E. ritteri implied that cross-fertilization also occurs in this species. No mother-offspring comparisons were performed on E. fernaldi, as this species was not observed brooding offspring during this study. Although incomplete, the results of this study increase our knowledge of the very unusual combination of reproductive modes in the genus Epiactis, and argue for further investigations of the evolution and genetic consequences of mating systems in these species.     

Mating behaviour in the penaeid shrimp Penaeus vannamei

Mating behaviour of Penaeus vannamei was observed during January, 1986. Mating behaviour was divided into four phases: (1) approach, (2) crawling, (3) chasing, and (4) mating. Male mating and spermatophore transfer to the mature female take place at intermolt stage C₄. The glutinous spermatophore emitted from the male can be transferred onto the female open thelycum during the ventralto-ventral position in mating in P. vannamei. Though male P. vannamei often chase males or immature females with undeveloped ovaries, no males were ever mated and mating only occurred in females with ripe ovaries.     

Mating in the reef white-tip shark Triaenodon obesus

Mating reef white-tip sharks, Triaenodon obesus, were observed and photographed in shallow waters of the Hawaiian Islands. The sharks copulated in a stationary parallel orientation with their heads on the bottom and bodies angled upwards approximately 45° into the water column. Pair contact during copulation was maintained by the male's left clasper anchored in the female's vent and by his tight oral grasp of the female's left pectoral fin. Results show that reposed sedentary modes of copulation are not limited to small species. In a reproductive context, biting behavior by male sharks functions as a premating releaser for female cooperation and for pair-contact maintenance during copulation.     

Cuticular hydrocarbons and reproductive status in the social wasp Polistes dominulus

Most species of social insect are characterized by a reproductive division of labor among morphologically specialized individuals. In contrast, there exist many species where all individuals are morphologically identical and dominance relationships determine which individuals mate and/or reproduce. In newly founded multiple-foundress associations of the social wasp Polistes dominulus, foundresses establish dominance hierarchies where the top-ranked (alpha) female monopolizes egg laying. The possibility that chemical cues are used for recognition of egg-laying individuals has not been explored in this wasp. Using non-destructive techniques, we examined the relationship between ovarian activity and the proportions of cuticular hydrocarbons of three female types (dominant and subordinate foundresses and workers) in 11 colonies. Immediately after nest foundation, no differences were found between alpha and subordinate females. In contrast, at worker emergence, alpha females were statistically distinguishable from both subordinates and workers. We experimentally removed the alpha female in 5 of the original nests and reanalyzed hydrocarbon proportions of the new dominant individual. Replacement individuals were all found to acquire a cuticular signature characteristic of the alpha female. This suggests that cuticular hydrocarbons are used as cues of ovarian activity in P. dominulus, and we discuss our results in terms of a switch from behavioral dominance to chemical signaling in this wasp.     

Solitary nesting and reproductive success in the paper wasp Polistes aurifer

Female Polistes paper wasps are capable of independent nesting, yet many populations demonstrate a mixture of solitary and cooperative nest foundation. Previous studies of Polistes have found survival and/or productivity advantages of cooperative nest foundation compared to solitary nesting, and reproductive skew models have been designed to predict the dynamics of such flexible cooperation. In this paper, we examine the success of different nesting strategies in a previously unstudied population of Polistes aurifer in southern California. The colony cycle of this population is less synchronous than that of other temperate species, and the frequency of solitary nesting averages 86.2%. Our data suggest that this low rate of cooperative nest founding is adaptive, as demonstrated by the lack of survival or productivity advantages for cooperative foundress associations. Due to foundress turnover and nest foundation later in the season, many nests produce only one set of offspring. This results in a loss of the eusocial nature of some nests in the population. Data from a small sample of multifoundress nests show significant positive reproductive skew, despite concession model predictions that skew should be low in populations with low ecological constraints on independent nesting. This lack of support for the concessions skew model reflects a diminished incentive for cooperation.Communicated by L. Keller     

Intraspecific comb usurpation in the social wasp Polistes fuscatus

Summary Incidents of usurpation were observed in colonies of Polistes fuscatus nesting on farm buildings (1977–79) and in nestboxes (1980–84) in Johnson County, Iowa, USA. Most usurpations (84.8%) occurred in the latter half of the preworker phase of the colony cycle, which coincided with periods of high predation of combs by vertebrates. Usurpers were probably displaced single foundresses which did not join neighbors or refound colonies after comb loss. Most (89–100%) usurpers of known relatedness to the foundresses they replaced were cousins or less related to them. Usurpation was a significant source of nest loss (19.6%) among single foundresses, but was rare (2.2%) in multiple-foundress colonies and colonies with workers (3.5%). Usurpers often destroyed younger brood (eggs and larvae in instars 1–3) in host colonies, while older larvae and pupae were usually spared. Brood destruction was more pronounced in more advanced host combs. Usurper survivorship after workers eclosed was lower than than of queenright single foundresses (61.5% vs 87.0%). Reproductive success by usurpers was less than that of queenright single foundresses, but greater than that of foundresses which initiated colonies late in the preworker colony cycle.     

Mating behavior and sperm competition in the fly,Dryomyza anilis

Summary Mating behavior in D. andis was studied both in nature and in the laboratory. Each mating in nature consisted of several repeated copulation and oviposition bouts during which the male stayed mounted. One copulation bout included an intromission in the beginning followed by several tapping sequences. During one tapping sequence the male touched the external genitalia of the female with his claspers several times. The experimental matings with irradiated sterile males contained one copulation bout after which the female laid her eggs. In these matings the percentage fertilization for the last male to mate with the female increased with increasing number of tapping sequences. The percentage fertilization was affected positively also by the time between the end of the copulation bout and the start of oviposition, but the time spent in the copulation bout was not important. Furthermore, there was a negative correlation between the length of intromission and the percentage fertilization. After the copulation bout, the percentage fertilization was higher among the first eggs laid. This difference between the first and later eggs laid increased with the increasing number of tapping sequences, suggesting that copulating and ovipositing in several bouts further increase the male percentage fertilization. In matings recorded in nature, both the number of tapping sequences and the number of copulation and oviposition bouts varied greatly. Of the variables recorded during these matings, individual size and the number of eggs the female contained in the beginning of a mating affected the number of copulation bouts and the number of tapping sequences. Males also mated differently with females containing immature or mature eggs. The male behavior resulting in sperm precedence and the variability of matings in nature in relation to male sperm competition is discussed.     

Sex ratio and host size in a parasitoid wasp

Summary Parasitoid wasps often lay male eggs in small hosts and female eggs in larger hosts. The selective advantage of this strategy can be explained by assuming wasp fitness increases with host size and that this fitness increase is greater in females than in males. I conducted experiments to test a model based on this explanation and found the results generally supported the model with one exception; unlike what the model assumed, these wasps were unable to adjust their offspring sex ratios in each generation to different host size distributions. This finding suggests an alternate view as to how selection might operate in the evolution of parasitoid sex ratios.     

Mating behavior of the marine copepodOithona davisae

Oithona davisae Ferrari and Orsi were collected from the innermost region of Tokyo Bay, Japan, in 1980 and 1981. The mating behavior of this marine cyclopoid copepod consists of several steps, starting with the paddling of the male in random search of a mate. This behavior is followed by a spiraling movement in pursuance of a mate. Next is the copulatory grasp during which the male grasps the fourth swimming legs of the female partner by means of his first antennae. Whilst in this position, the male's urosome vibrates to allow the spermatophores to extrude from his genital openings. The mating behavior then culminates in the spermatophore transfer. Males do not grasp the uro-some or caudal setae of the mating partner before proper copulatory grasp. The specialized setae of the female's fourth legs may help the male to grasp her legs firmly. Spiraling occurs when the male approaches or traverses the trail of a female that is ready to copulate and that presumably emits a sex-attractant pheromone. The turning radius reduces gradually from more than 1 mm to ca. 0.25 mm as the male approaches the mate female. Females may register spiraling as a mate (male)-approaching signal. Spiraling may lead the male to locate a pheromone source more accurately, and to promote diffusion of the pheromone to prevent other males from pursuing the source. This swimming strategy can increase the copulatory chance of mature virgin females.     

Mating systems,parasites and plumage dimorphism in waterfowl

Summary Despite a tendency for males of polygynous bird species to show bright or elaborate plumage, comparative analyses have failed to show any consistent relationship between male brightness or plumage dimorphism and the form of breeding systems. Here we argue that this may be partly because the opportunity for sexual selection varies between species showing serial or seasonal monogamy and life-long monogamy. In waterfowl, both the brightness of male plumage and sexual dimorphism in colouration vary between these categories of monogamy. Other ecological factors related to male brightness or plumage dimorphism include male assistance in protecting young, latitude and an index of parasite sharing. The adaptive significance of these trends is discussed. Offprint requests to: D.K. Scott     

Mating system shift in a pronghorn population

Summary At the National Bison Range (western Montana, USA), pronghorn (Antilocapra american) maintained a territorial mating system from as early as 1965 through 1978. Preliminary observations in 1981 suggested that the mating system had changed. Data from the ruts, 1982–1984 revealed a progressive decay in territoriality. In this paper, data from the territorial years 1969–1978 are contrasted with data from the decay years 1982–1984, with results as follow: 1. In 1982–84, fewer territories were defended than in 1969–78. This was attributable to a smaller proportion of males defending territories in 1982–84, not to a smaller number of males. 2. In 1982–84, most territory owners either abandoned their territories early in ruts, or lost control of females on them following frequent, persistent intrusions by non-territorial males. Abandonment and loss of control did not occur in 1969–78. 3. In 1982–84, territorial males that maintained control of females on their territories did so by shrinking their zones of defense to small areas around female groups. 4. In 1982–84, following the disruption or severe disturbance of all territories, many females left territories, and mated elsewhere, with non-territorial males. In 1969–78, most females remained on territories throughout rut, and mated with territory owners. —The mating system change followed catastrophic winter mortality, 1978–79, that removed 75% of the males, including all males older than 5 years, and all male fawns, from the population. In 1982–84, the number of males present was not different from the number of males in 1969–1978, but the frequency distribution of male ages was strongly shifted toward younger ages. The small number of older males, 1982–84, likely resulted in smaller proportions of males initially defending territories, and in less effective territory defense. When females then clustered on the few territories where defense was at first successful, they attracted large numbers of non-territorial males. The resulting high rates of raids on these territories, coupled with reduced defense radii by territorial males, allowed females only 12% reclining time (summer percentage was 39%). This increased energy cost, plus an apparently greater risk of injury on weakly defended territories, appeared to prompt many females to seek calmer matings elsewhere. Also, if female pronghorn practiced mate selection based upon horn or body size, they may have reduced their efforts to remain on territories in 1982–84. Males from 1969–78 were larger than males from 1982–84, and showed greater variance in horn size. At least two conditions appear to influence the tendency of males to be territorial. First, males must be at least three years old before they attempt to defend a territory. Second, the declining proportion of males defending territories, 1982–84, that coincided with an increasing number of males three years and older, suggests that males also decide whether or not to attempt territory defense based upon the frequency of territorial defense in the population.     

Male influence on sex allocation in the parasitoid wasp Nasonia vitripennis

Sex allocation is an important reproductive decision for parents. However, it is often assumed that females have substantial control over sex allocation decisions, and this is particularly true in haplodiploid insects, in which females apparently determine sex by deciding whether to fertilise an egg (and produce a diploid daughter) or not (and produce a haploid son). Mechanisms by which males may influence sex allocation are not so straightforward, and their potential influence on sex ratios has been somewhat neglected. Here, we test whether males influence offspring sex ratios in the parasitoid wasp Nasonia vitripennis. We show that some of the variation in observed sex ratios can be attributed to males when comparing the affect of male strain on sex ratio. We did not find among-male variation in sex ratio with a less powerful experiment using males from only one strain or an effect of male mating environment. Our data suggest that males can influence female sex ratios and contribute to the variation around the sex ratios optimal for females. However, the influence is not large, suggesting that females have more influence on sex allocation than do males. We conclude by considering whether male influences on sex ratio represent differences in male reproductive competence or deliberate attempts by males to increase their fitness by influencing daughter production.