Mating Flight Research Articles

First Autonomous Rendezvous Using Relative GPS Navigation by ETS-VII

ABSTRACT: Engineering Test Satellite-VII (ETS-VII) successfully performed the world's first autonomous rendezvous docking (RVD) flight using relative GPS (RGPS) navigation. ETS-VII is a test satellite designed to perform in-orbit demonstration of autonomous rendezvous docking technology. RGPS navigation is a form of differential GPS (DGPS), and is used as a primary means in the relative approach phase of flight (farther than 500 m from the target satellite). Three methods were used to evaluate the in-orbit performance of the ETS-VII RGPS system during mating flights and rendezvous flights. The accuracy of RGPS navigation was found to be about 10 m in position and 3 cm/s in velocity. This paper describes the evaluation of the in-orbit performance of RGPS during the first autonomous RVD experiment.

Burrowing mayflies (Hexagenia) as indicators of ecosystem health

Abstract Three State of the Lakes Ecosystem Conferences have been held since 1996 to encourage the development of Great Lakes indicators of ecosystem health for use in reporting on progress in restoring and maintaining the chemical, physical and biological integrity of the Great Lakes ecosystem. Here we report on the development of an indicator based on burrowing mayflies, Hexagenia (Ephemeroptera: Ephemeridae), using production and biomass as the indicator metrics. Burrowing mayflies were selected because they (1) were historically abundant in unpolluted, soft-bottomed mesotrophic habitats throughout the Great Lakes, (2) are intolerant of and were extirpated by pollution in most of those habitats during the 1940s to1950s, (3) have shown the ability to recover in one of those habitats following pollution abatement, (4) are ecologically important as bioturbators of lakebed sediments and as trophic integrators that link detrital energy resources directly to fishes that feed preferentially on them, and (5) have highly visible mating flights, which carry the message directly to an informed public that the source water body is healthy. In addition, their annual production can be estimated from their mean annual biomass by the sizefrequency method. Productivity and biomass can also could be estimated with a ‘cohort-direct’ method, using the biomass of mature nymphs collected in May or early June from the cohort that is about to emerge as subimagos in late June or early July. Although both the size-frequency and cohort-direct methods provide reliable estimates of productivity and biomass, the latter method greatly reduces sample collection and processing effort and thus makes it feasible to use Hexagenia as an indicator of ecosystem health in surveys requiring the collection of large numbers of samples.

Colony size and reproduction in the western harvester ant, Pogonomyrmex occidentalis

We report data from a four-year field study on the relationship between colony size and reproduction in the western harvester ant, Pogonomyrmex occidentalis. In all years, the likelihood of reproduction significantly increased with increasing size in both field censuses during naturally-occurring mating flights and experimentally-watered colonies whose entire reproductive output was collected. However, the total amount of reproductive biomass was unrelated to colony size. We describe the size threshold for reproduction in P. occidentalis and show that it varies across years. Once colonies become reproductively mature, they reproduce consistently although not in every year. We describe a method for collecting the entire reproductive output for desert ants whose reproductive flights are cued by rainfall.

No Behavioral Control over Mating Frequency in Queen Honey Bees (Apis melliferaL.): Implications for the Evolution of Extreme Polyandry

Ever since the seminal papers of Hamilton (1964), a common focus of sociobiology has been the genetic relationships among individuals of insect societies. At issue are agents that reduce the average relatedness among colony members since they are difficult to interpret in light of the evolution of sociality by kin selection. One such factor is polyandry or female multiple mating. Low levels of polyandry are common among the social Hymenoptera, occurring across a broad range of taxa and social systems (Page and Metcalf 1982; Page 1986; Crozier and Pamilo 1996). In spite of that, high mating frequencies (‚2 mates/ female) are relatively rare among the highly eusocial Hymenoptera, limited to only one or a few genera in each of the ants, bees, and wasps (Boomsma and Ratnieks 1996). These extraordinary levels of polyandry have been of particular interest among students of social insect research since their adaptive significance is still in question and highly debated (Kraus and Page 1998; Sherman et al. 1998). The genus Apis has been studied in depth for both queen reproductive behavior and mating frequency, facts that allow insight into the evolution of extreme polyandry in these species. The reproductive behavior of honey bee queens (Apis mellifera) occurs early in their lifetimes. Virgin queens initiate mating behavior when they are about 1 wk old by briefly exiting the colony and orienting to local landmarks (Ruttner 1956). After such orientation * To whom correspondence should be addressed; e-mail: drtarpy@

Genetic structure of colonies and a male aggregation in the stingless bee Scaptotrigona postica , as revealed by microsatellite analysis

Stingless bee queens have for long been assumed to mate once on a nuptial flight, early in life. To evaluate critically monandry in one stingless bee, Scaptotrigona postica, worker offspring (adults or brood) were genetically analysed with microsatellite loci, five of which were developed specifically for the species. Marker loci were highly variable; unbiased estimates of heterozygosity were > 0.5. "Foreign" workers, either those having drifted from other colonies (circa 2%) or those of a replacement queen, were identified with the genetic markers and removed from further analysis. Worker genotypes were consistent with some queens having mated once and others having mated with up to six different males. Scaptotrigona postica queens are therefore facultatively polyandrous. Effective mating frequencies, m e , were generally lower than the number of patrilines observed. Relatedness estimates of nestmates from individual colonies concurred with those derived from direct counts of the number of patrilines and their proportional representation. Putative genotypes of a colony's queen and her mates were deduced from those of her workers. Queens were generally not related to their mates. For one polyandrous queen, her six mates were related to each other, possibly because of numerically biased representation of males from different colonies at mating sites. However, males at an aggregation outside a colony came from numerous colonies.

Colony reproduction in Tetragonisca angustula (Apidae, Meliponini)

Colony multiplication of the neotropical stingless bee Tetragonisca angustula started with the finding of a new nesting site by scout bees, followed by the cleaning of the cavity. Cerumen was flown in from the mother colony for the sealing of crevices and the construction of an entrance tube. Major investment of the mother colony in the daughter nest was in workers and cerumen. Almost negligible amounts of pollen were transported from the mother colony after arrival of the gyne in the founding colony. Small quantities of honey were transferred as well. Workers from the daughter nest foraged actively for pollen. Some 500 to 1000 workers entered in the daughter nest, following the virgin queen. This is only a relatively small proportion of the population of the mother colony. Two to four days after the nuptial flight the young queen started egg-laying, and storage pots, mainly filled with honey and some pollen were encountered. In the mother nest, ample pollen and honey stores were present during the months preceding the swarming process. Shortly before swarming took place brood (and adult) populations were 20 to 50% higher than in the other non-swarming colonies, whereas the production of royal cells and males was not different. Data were compared with those got in literature.

Mating flights and sperm transfer in the dwarf honeybee Apis andreniformis (Smith, 1858)

Mating flights of 3 virgin queens of Apis andreniformiswere observed at their natural nest- ing site. They initiated mating flights between 12.33 and 12.50 h. The flight duration was between 19 and 23 minutes. The sting chamber of the returning queens contained the orange-colored secretion from the cornual gland of the drone's endophallus. Immediately after the mating flights, the queens were dissected. No sperm was detected in the oviducts, but spermatozoa were found in the sper- mathecae. In 2 queens, the spermathecae contained 0.09 million spermatozoa, which corresponds to about 75% of the sperm of 1 drone. The third queen had 0.31 million spermatozoa. The spermatozoa in the spermatheca were observed to be moving, and formed an undulating thread. These results suggest that sperm is transferred not into the oviducts but directly into the spermatheca (via the sper- maduct). Seven egg-laying queens of unknown age had between 0.33 and 1.26 million spermatozoa in their spermathecae. The mode of sperm transfer is discussed in relation to the number and sequence of the spermatozoa received from each drone in the spermatheca. Apis andreniformis / queen / mating flight / sperm transfer/ spermatozoa count

Observations on gynes and drones around nuptial flights in the stingless bees Tetragonisca angustula and Melipona beecheii (Hymenoptera, Apidae, Meliponinae)

The nuptial flight of gynes of Tetragonisca angustulaand Melipona beecheiiwas stud- ied. The moment of nuptial flight was found to be related to the ambient temperature, and the dura- tion of the nuptial flight for M. beecheii was longer in November (rainy season) than in March (dry season). A repeated mating flight was recorded for two gynes of T. angustula. Three of five T. angus- tula queens and all six M. beecheii queens were mated successfully. Behavioural data of drones and gynes shortly before and after the nuptial flight are presented. Drones of T. angustula participated in a congregation for up to three days. The importance of pheromones for the attraction of drones and gynes is discussed. An hypothesis explaining the observed seasonal occurrence of male congregations near nests of T. angustula is presented.

Unusual Behavior of Polygyne Fire Ant Queens on Nuptial Flights

This study reports previously undescribed behavior of fire ant queens (Solenopsis invicta) on their nuptial flights. We captured large numbers of alate (winged) queens flying at low altitudes in dense swarms that were virtually devoid of males. We assayed the genotypes of these alate queens at the locus Gp-9, which exhibits strong genotype frequency differences between monogyne (single-queen) and polygyne (multiple-queen) populations, and found that almost all of these low-flying queens originated from polygyne colonies. Comparisons of mtDNA haplotype distributions of these queens to those of alates leaving polygyne nests suggest that the flying queens had not dispersed more than a few hundred meters. Moreover, the proportion of flying queens that were mated did not differ significantly from the proportion of reproductive queens that were mated within the same sites. Thus the flight behavior appears to occur subsequent to mating. We suggest that the flying queens are sampling the local environment in order to select a suitable landing site. Such a site would contain established polygyne nests into which the queens may be adopted as new reproductives.

Behavioural development and abdomen inflation of gynes and newly mated queens of Melipona beecheii (Apidae, Meliponinae)

The behaviour of gynes of Melipona beecheii in queen-deprived colonies was studied. The period after emergence until acceptance is characterized by agonistic behaviour of workers towards the gynes. The gynes escaped from this worker aggression by hiding in the periphery of the nest, by performing rapid turn-arounds once grabbed by a worker, and "feigning death". Between acceptance and nuptial flight, gynes spent most of their time in pushing, hiding, and antennal contact with workers, and self-grooming or food solicitation. After the nuptial flight the queen's behavioural repertoire shifted to less pushing and food solicitation, to an increase in standing, tapping and antennal contact.¶Accepted gynes had a significantly more prolonged abdomen inflation than gynes that were eliminated.¶An hypothesis is presented to explain how abdomen enlargement and behavioural development influences the acceptance of gynes and the establishment of a dominance relation with workers under queenless conditions.

Queen Dispersal Strategies in the Multiple‐Queen Form of the Fire AntSolenopsis invicta

abstract: Newly produced queens in the multiple‐queen (polygyne) form of the fire ant Solenopsis invicta show dramatic variation in dispersal patterns, and this variation is influenced by genotypic variation at a single locus associated with the genetic marker Gp‐9. Heavy, homozygous Gp‐9BB queens exhibit the highest vagility among polygyne queens and are strongly attracted to the open, disturbed‐habitat patches that characteristically attract queens of the single‐queen (monogyne) form (all of which possess genotype Gp‐9BB). Intermediate weight, heterozygous Gp‐9Bb queens exhibit a mixed dispersal strategy: some remain in the area near their natal nest, while others disperse to land in the same disturbed‐habitat patches as Gp‐9BB queens. Light, homozygous Gp‐9bb queens appear to lack the energy reserves needed to take part in mating flights in substantial numbers. Most queens that disperse from their natal nest site apparently fail to infiltrate mature nests to reproduce. However, consistent with the expe...

Queen Dispersal Strategies in the Multiple-Queen Form of the Fire Ant Solenopsis invicta

Newly produced queens in the multiple‐queen (polygyne) form of the fire ant Solenopsis invicta show dramatic variation in dispersal patterns, and this variation is influenced by genotypic variation at a single locus associated with the genetic marker Gp‐9. Heavy, homozygous Gp‐9BB queens exhibit the highest vagility among polygyne queens and are strongly attracted to the open, disturbed‐habitat patches that characteristically attract queens of the single‐queen (monogyne) form (all of which possess genotype Gp‐9BB). Intermediate weight, heterozygous Gp‐9Bb queens exhibit a mixed dispersal strategy: some remain in the area near their natal nest, while others disperse to land in the same disturbed‐habitat patches as Gp‐9BB queens. Light, homozygous Gp‐9bb queens appear to lack the energy reserves needed to take part in mating flights in substantial numbers. Most queens that disperse from their natal nest site apparently fail to infiltrate mature nests to reproduce. However, consistent with the expectations of game‐theoretical models for the evolution of dispersal, the low realized success of dispersing queens does not prevent relatively large numbers of them from dispersing. Furthermore, the results presented here are consistent with the hypothesis that the reproductive syndrome that characterizes polygyny in S. invicta is largely controlled by a single locus.

Ovipositional Phenology and Behavior of the Spruce Cone Fly, Strobilomyia neanthracina (Diptera: Anthomyiidae)

The behavior of Strobilomyia neanthracina Michelsen, a phytophage infesting spruce (Picea spp.) seed cones, was observed at a field site in northern Ontario and in cages in a greenhouse to investigate spatiotemporal aspects of mating, host location, and oviposition. In the field, adults emerged from 21 to 24 May 1996, which coincided with bud burst of Picea glauca (Moench) Voss seed cones. For 4 days following emergence, Strobilomyia flies could no longer be seen at a monitored P. glauca tree and may have been on a mating or dispersal flight. Subsequently, females but not males were seen again and the oviposition period of ca. 3 weeks began. Mating was observed only in the greenhouse, mostly (i.e., 65%) at age 5–9 days. Although copulations lasted 11–45 min, these females laid infertile eggs only, beginning at age 4 days. No sperm was found in the spermathecal capsules of females, suggesting that no sperm had been transferred during these copulations. In both the field and the greenhouse, ovipositional sequences that resulted in egg deposition occurred throughout the day but few sequences were observed before 1000, probably because flies were not very active at air temperatures below 14°C (most sequences occurred at 25–27°C). In the greenhouse, the typical ovipositional sequence lasted an average of 7 min and consisted of landing on the cone and examining it with the proboscis and sometimes the ovipositor, egg deposition, and postovipositional behaviors such as tapping (touching the cone surface with the flabellum ca. 5 times s−1), which possibly represents a host marking behavior. In the field, tapping was seen less frequently than in the greenhouse but occurred significantly (P = 0.014) more often after sequences that resulted in egg deposition than after sequences that did not. Eggs hatched after 4–5 days at 20°C. In the greenhouse, the median longevity of females and males was 24 and 17 days, respectively.

Observations on multiple mating flights of Apis dorsata queens

Les vols de fecondation ont ete etudies chez les reines d'Apis mellifera, A. cerana, A. florea, A. koschevnikovi et A. andreniformis, mais on sait peu de choses en ce qui concerne les reines d'A. dorsata. Cette etude visait a observer le comportement d'accouplement des reines d'A. dorsata pour obtenir des donnees permettant une etude comparative de la biologie de la reproduction chez les differentes especes d'Apis. Le recherche a ete faite en 1996 dans la foret submergee de Malaleuca du sud du Vietnam. La, de nombreuses colonies nidifient a faible hauteur sur des supports en bois suspendus par les apiculteurs pour attirer les colonies, ce qui rend possible des observations detaillees des vols de fecondation des reines. Habituellement, de telles observations sont impossibles car cette espece a tendance a nidifier dans des endroits eleves. Six reines recemment ecloses ont ete marquees et introduites dans des nuclei. Nous avons observe les reines de chaque colonie de 18 a 19 h et note leur vols. Les reines ont ete dissequees apres leur dernier vol nuptial et les spermatozoides presents dans la spermatheque ont ete denombres. Les cinq reines observees ont accompli leur premier vol 6 ± 1 j (moyenne ± ecart type) apres leur emergence. Quatre reines ont fait des vols d'orientation inferieurs a 3 min et une reine s'est envolee pour s'accoupler sans avoir fait de vol d'orientation (tableau I), Les vols de fecondation ont eu lieu au moment du coucher du soleil (tableau II) et ont dure 15,4 ± 4,3 min (n = 14). Les reines ont fait entre deux et quatre vols de fecondation et possedaient en moyenne 5,5 ± 0,9 (n = 5) millions de spermatozoides dans leur spermatheque apres le dernier vol de fecondation (tableau I). Cette etude confirme, d'une part, la forte polyandrie deja mentionnee chez A. dorsata dans de precedents rapports et, d'autre part, les hypotheses portant sur l'evolution de la forte polyandrie chez le genre Apis.

Queen mating and paternity variation in the antLasius niger

We have electrophoretically analysed the variation in queen mating and worker paternity across and within seven populations of the antLasius nigerin northwestern Europe. Populations were panmictic and not genetically differentiated (FST= 0.003 ± 0.004; rangec.1000 km). Queens (n= 535) were shown to mostly mate with a single male, but double mating occurred in all populations and triple mating was found in one case (the total number of worker offspring analysed for paternity was 4825). The genetically effective queen mating frequency was 1.16 on average across populations (range 1.04–1.42). Double sampling of six out of the seven populations showed that most of the variation in queen mating occurred among populations and not within populations among years. Also, paternity skew in colonies with double‐mated queens was relatively constant per site but varied across populations. Paternity skew was high in populations with low frequencies of double queen‐mating, low in populations with intermediate frequencies of double queen mating and ambiguous in a single population where more than half of the queens mated multiply. Double‐mated queens were only collected halfway through a nuptial flight, suggesting that double mating is time consuming and that the mating swarm sex ratio may affect the likelihood of multiple mating towards the end of a flight. No difference in fresh weight between single‐ and double‐mated queens from the same population was found.

Relatedness among honeybees (Apis mellifera) of a drone congregation

The honeybee (Apis mellifera) queen mates during nuptial flights, in the so-called drone congregation area where many males from surrounding colonies gather. Using 20 highly polymorphic microsatellite loci, we studied a sample of 142 drones captured in a congregation close to Oberursel (Germany). A parentage test based on lod score showed that this sample contained one group of four brothers, six groups of three brothers, 20 groups of two brothers and 80 singletons. These values are very close to a Poisson distribution. Therefore, colonies were apparently equally represented in the drone congregation, and calculations showed that the congregation comprised males that originated from about 240 different colonies. This figure is surprisingly high. Considering the density of colonies around the congregation area and the average flight range of males, it suggests that most colonies within the recruitment perimeter delegated drones to the congregation with an equal probability, resulting in an almost perfect panmixis. Consequently, the relatedness between a queen and her mates, and hence the inbreeding coefficient of the progeny, should be minimized. The relatedness among the drones mated to the same queen is also very low, maximizing the genetic diversity among the different patrilines of a colony.

Polygyny and Monoandry in the Ant Formica japonica (Hymenoptera: Formicidae)

Queen number, mating frequency and nest kin-structure of the ant Formica japonica were studied in the field and the laboratory. Nest excavation in the study site, the east slope of Mt. Fuji, Gotenba, Japan, revealed that F. japonica is weakly polygynous all year round and the queen number increases after the nuptial flight season, suggesting the adoption of newly mated queens by established nests. Dissection and laboratory rearing demonstrated that nearly all queens in polygynous nests had mated and were fertile with mature oocytes in their ovaries. Multilocus DNA fingerprinting was used to examine kin relationships among ants found in the same nests. The fingerprint band patterns were apparently governed by a simple genetic rule and suggested monoandry (single mating per queen). The mean band sharing score of DNA fingerprints among full sisters was 0.90, and the mean value between queens and their daughters was 0.75. Comparison of DNA fingerprints of adult and pupal workers with pupal gynes suggested that multiple queens in a nest may contribute unequally to gyne (new queen) production.

A case of multiple mating in stingless bees (Meliponinae)

In several stingless bee species many males aggregate in the vicinity of a nest when a virgin queen is present in the colony and is preparing for the nuptial flight. We report such male assemblage for Tetragonisca angustula. The departure of a virgin queen from the colony and the subsequent mating could be video-recorded, because the queen and the males that had mounted her fell to the ground. Since at least two males had lost their genitalia, multiple mating seems to have occurred. This is in contrast with the prevailing view found in literature concerning the mating biology of stingless bees.

Sexual Calling by Workers Using the Metatibial Glands in the Ant, Diacamma sp., from Japan (Hymenoptera: Formicidae)

Several species of the ant genus Diacamma reproduce through mated workers (gamergates). Such gamergates have no wings and therefore are unable to conduct nuptial flight. Instead they perform a sexual calling behavior by standing outside the nest and rubbing the tibiae of their hindleg over the surface of the arched gaster. In a series of exclusion experiments we demonstrate that secretions from the metatibial gland are the most important component in making the virgin female attractive to the males.

Nuptial Flight in two Ponerine Ants, Pachycondyla Impressa and P. Fauveli, Overlooking Machu Picchu, Peru

We report on a simultaneous nuptial flight of ants of two sympatric ponerine species, Pachycondyla impressa and P. fauveli, on the top of Huayna Picchu (2700 m), Peru. After flying in and landing on the flat, hare rocks of the summit, the conspecific sexuals approached, antennated and mated without any previous female calling behaviour. During one hour of observation, the number of approximately 30 ♂ ♂ and ♀ ♀ Pachycondyla impressa and 70–86 of P. fauveli on the site remained constant because of individuals continuously arriving and leaving. Although nuptial flight seems to be the most common type of mating system in ants, this is one of the very few reports on the occurrence of this behaviour in ponerine species.