Decision-making conditions for intra- or inter-nest mating of winged males in the male-dimorphic ant <Emphasis Type="Italic">Cardiocondyla minutior</Emphasis>

Only winged male and female ants generally mate through nuptial flight during the reproductive season. In the ants of Cardiocondyla, the males show wing dimorphism and their reproductive strategies differ depending on the differences in wing morphology. It has been suggested that wingless “ergatoid” males bearing very similar external morphologies to workers mate within natal nests, whereas winged males bearing typical ant male morphology disperse from their nests to mate. However, some behavioral observations suggest that the winged males of some Cardiocondyla ants such as C. obscurior and C. minutior may mate within natal nests before dispersion. We evaluated the factors affecting the mating behaviors of the winged males of C. minutior under laboratory conditions. We found that (1) the winged males remained and mated with virgin females in natal nests when either virgin winged females or the relatively mature pupae of winged females (i.e., at least 10 days) were present in the nest, (2) the winged males dispersed to adjacent nests with virgin winged females when only mated queens and the relatively young pupae of winged females (i.e., <9 days) were present in the nest, and (3) all winged males were accepted by the workers of non-natal nests irrespective of the distance from the natal nests in the field. Although most ergatoid males were accepted by the workers of close non-natal nests, they were all attacked and killed by the workers of distant non-natal nests. These results suggest that intra-nest mating and the dispersion of the winged males of C. minutior are facultatively determined by the condition of winged females (virginity and relative pupal age) in natal nests. Furthermore, our results suggest that winged males are likely to seek mating partners chemically and to mate with virgin winged females.     

Genetic structure and reproductive strategy of the ant Cardiocondyla elegans: strictly monogynous nests invaded by unrelated sexuals

Cardiocondyla elegans is a Mediterranean ant that nests on river banks. It rears only wingless (ergatoid) males that live peacefully in the same nest as opposed to other species of the same genus, which have both peaceful, winged and mutually aggressive 'ergatoid' males. Using microsatellite analysis, we investigated the genetic structure of 21 colonies from three different locations as well as the parentage of sexuals of two colonies of C. elegans. We show that C. elegans is strictly monogynous, and that its nests can contain foreign sexuals. The presence of alien sexuals inside ant nests is described for the first time and probably counteracts inbreeding resulting from matings between siblings. In the laboratory, aggression tests showed that workers only allow alien males to enter their nests, while all winged female sexuals attempting to enter were attacked. Nevertheless, the presence of alien female sexuals in nests in the field seems to result from active carrying behaviour by workers during the reproductive period.     

深圳市红火蚁婚飞规律研究

本研究利用自制红火蚁Solenopsis invicta Buren收集装置对红火蚁婚飞规律进行研究,结果表明:本试验过程中共观察到婚飞273次,无论大、中、小蚁巢均有有翅蚁婚飞现象;共收集到有翅蚁10 618头,雌雄比为2.1:1;单次婚飞有翅蚁均为雌虫的有167次,均为雄虫的有25次,单次婚飞有翅蚊既有雌虫又有雄虫(雌虫+雄虫)的共81次;同一蚁巢可以发生多次婚飞,同一蚁巢不同次婚飞的有翅蚁性别比也会发生变化;除2009年11月和2010年2月未收集到有翅蚁,其余各月均可观察到有翅蚁婚飞,婚飞最高峰期为2010年4月和5月,其次是2009年6-9月,婚飞次数和收集到的有翅蚁数量成明显的正向相关关系;8:30-15:00均有有翅蚁婚飞,而婚飞高峰期为11:30-13:30,婚飞一般持续30～150 min.婚飞一般发生在雨后晴天或相对低温后气温快速回升以后.     

Observations on the foraging activity of Hospitalitermes umbrinus (Haviland), (Isoptera: Termitidae) in the Gunong Mulu National Park, Sarawak

Abstract. 1. A single nest of the black surface-foraging nasute termite Hospitalitermes umbrinus was studied for 5 days in July 1978. Foraging parties of workers and soldiers tend to leave the nest in a continuous stream in the evening and return carrying food balls during the following morning.
2. An estimated maximum of 500000 individuals were involved in the foraging excursions. At a density of 3 nests ha^-1, the total foraging population is estimated as being at least 180 m^-2.
3. The colony brought back an estimated 46.4 g (d.w.) of food in 4 days. The biomass of the colony was estimated at 349 g (d.w.) ( c. 1396 g w.w.) excluding alates and the consumption at 8.3 mg (d.w.) g^-1 (w.w.) d^-1.
4. The food balls were found to be of two types, light and dark coloured. The light balls contained wood constituents, bryophytes, blue-green algae and fungal hyphae while the dark balls consisted mainly of lichens and their spores.
5. Swarming began at 14.30 hours on 9 July and ended with the nuptial flight at 17.32 hours. About 3000 alates were released. They climbed to 9 m on a tree before taking off.     

Multiple functions of fire ant Solenopsis invicta mandibular gland products

Alarm pheromones of social insects are best known for their role in the defence and maintenance of colony integrity. Previous studies with the fire ant Solenopsis invicta Buren (Hymenoptera: Formicidae) demonstrate that the mandibular glands of workers (sterile females) and male and female sexuals produce an alarm pheromone, 2‐ethyl‐3,6‐dimethylpyrazine. The function of alarm pheromones in worker ants is well understood and divergent from the function of these compounds in the winged sexual forms. The present study quantifies the amount of pyrazine in the mandibular glands from male and female alate sexuals, as well as queens. Pyrazine production in female alates starts in the late pupal stage and increases until they reach mating flight‐ready maturity; however, after mating flight participation, the pyrazine level declines by >50%. Interestingly, mature male alates lose >85% of their mandibular gland pyrazine during mating flight activity. The results of the present study indicate that male and female sexuals use mandibular gland secretions for mating flight initiation and during mating flights. Furthermore, the ontogeny of mandibular gland products (pyrazine as the marker) from newly‐mated queens to mature colony queens shows a more than two‐fold increase in the amount of pyrazine by 6 months after mating. However, this is followed by a decline to trace amounts in mature colony queens (>2 years old), suggesting a function for mandibular gland products during colony development. Multifunctional use of social insect pheromones is well documented and data are reported in the present study suggesting new roles for mandibular gland products in fire ants.     

Competition and opportunity shape the reproductive tactics of males in the ant Cardiocondyla obscurior

Context-dependent adjustment of mating tactics can drastically increase the mating success of behaviourally flexible animals. We used the ant Cardiocondyla obscurior as a model system to study adaptive adjustment of male mating tactics. This species shows a male diphenism of wingless fighter males and peaceful winged males. Whereas the wingless males stay and exclusively mate in the maternal colony, the mating behaviour of winged males is plastic. They copulate with female sexuals in their natal nests early in life but later disperse in search for sexuals outside. In this study, we observed the nest-leaving behaviour of winged males under different conditions and found that they adaptively adjust the timing of their dispersal to the availability of mating partners, as well as the presence, and even the type of competitors in their natal nests. In colonies with virgin female queens winged males stayed longest when they were the only male in the nest. They left earlier when mating partners were not available or when other males were present. In the presence of wingless, locally mating fighter males, winged males dispersed earlier than in the presence of docile, winged competitors. This suggests that C. obscurior males are capable of estimating their local breeding chances and adaptively adjust their dispersal behaviour in both an opportunistic and a risk-sensitive way, thus showing hitherto unknown behavioural plasticity in social insect males.     

Natal Dispersal Patterns of a Subsocial Spider Anelosimus cf. jucundus (Theridiidae)

Species that alternate periods of solitary and social living may provide clues to the conditions that favor sociality. Social spiders probably originated from subsocial‐like ancestors, species in which siblings remain together for part of their life cycle but disperse prior to mating. Exploring the factors that lead to dispersal in subsocial species, but allow the development of large multigenerational colonies in social species, may provide insight into this transition. We studied the natal dispersal patterns of a subsocial spider, Anelosimus cf. jucundus, in Southeastern Arizona. In this population, spiders disperse from their natal nests in their penultimate and antepenultimate instars over a 3‐mo period. We tracked the natal dispersal of marked spiders at sites with clustered vs. isolated nests. We found that most spiders initially dispersed less than 5 m from their natal nests. Males and females, and spiders in patches with different densities of nests, dispersed similar distances. The fact that both sexes in a group dispersed, the lack of a sex difference in dispersal distance, and the relatively short distances dispersed are consistent with the hypothesis that natal dispersal results from resource competition within the natal nest, rather than inbreeding avoidance in competition for mates. Additionally, an increase in the average distance dispersed with time and with the number of spiders leaving a nest suggests that competition for nest sites in the vicinity of the natal nest may affect dispersal distances. The similar distances dispersed in patches with isolated vs. clustered nests, in contrast, suggest that competition among dispersers from different nests may not affect dispersal distances.     

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.     

Social Control over the Survival and Selection of Winged Virgin Queens in an Ant without Nuptial Flight: Iridomyrmex humilis

Laboratory and field experiments show that the number of winged virgin queens of the Argentine ant Iridomyrmex humilis allowed to remain alive in the nest is under social control. When reared in their natal nest, survival of virgin queens depends on the presence of males. In the presence of mature alate males or even male pupae only 20% of the gynes are executed before they are 4 days old. By contrast, if pupal and adult males are absent, about 55% of the gynes are executed. The workers tend to kill the lightest ones. Although the Argentine ant is a unicolonial species exhibiting complete acceptance of foreign conspecific workers and mated queens, regardless of whether the recipient nest is queenright or queenless, the acceptance of gynes is variable: they are executed by foreign queenless workers but widely accepted in queenright nests. These results are discussed with regard to possible existence of queen and gyne pheromones allowing recognition and to the lack of nuptial flight in this species.     

Fighting and mating behaviors of dimorphic males in the ant

Colony composition inCardiocondyla wroughtoni and the fighting and mating behaviors of 2 types of males, alates and ergatoids, are described. This species is polygynous, with a mean of 7.0 queens per nest, and forms polycalic colonies. Within nests, ergatoid males fight with each other, leading to the death of all but one in single nests. On the other hand, alate males exhibit no aggressive behavior towards any of their colony members. Both types of males conduct intranidal mating with their sisters, though the alate males also conduct nuptial flights. Many alate females leave their maternal nest even if they have already been inseminated by intranidal mating.     

红火蚁的巢间相容性

在室内条件下通过将红火蚁Solenopsis invicta Buren异巢不同品级的个体置于同一容器中,对红火蚁的巢间相容性进行研究。结果表明,蚁后与异巢幼蚁共处时,异巢幼蚁的羽化率为33.3%,与对照处理的38.3%(蚁后与同巢幼蚁共处)无显著差异。工蚁与异巢幼蚁共处时,异巢幼蚁的羽化率为53.7%,与对照处理的72.2%(工蚁与同巢幼蚁共处)无显著差异。蚁后与异巢工蚁共处时,异巢蚁后的存活率为45%,与对照处理的61.7%(工蚁与同巢蚁后共处)无显著差异。有翅繁殖雌蚁与异巢工蚁共处时,异巢工蚁的存活率为86.7%与对照处理的99.4%(工蚁与同巢有翅繁殖雌蚁共处)存在显著差异;异巢有翅雌蚁的存活率为22.2%,与对照处理的88.9%(工蚁与同巢有翅繁殖雌蚁共处)存在显著差异。由此可见,同区域内红火蚁异巢各品级间存在一定程度的相容性,并巢行为有可能发生。     

Ants under the conditions of an extremely hot summer

Specific features of ant behavior during the extremely hot summer of 2010 were studied, as well as the aftereffects of this season on simple and complex family units of ants in 2011–2012. Simultaneous studies were carried out in southern taiga (Moscow Province, Verkhnaya Klyazma myrmecological protected area) and northern taiga (Arkhangelsk Province, Pinezhskii Nature Reserve). Ants of the genus Formica responded to the extreme heat by (1) changing their foraging patterns; (2) redesigning their nests; (3) rearranging the spatial and functional structure of the colonies. They switched to a bimodal activity pattern with maxima in the morning and in the evening and a prolonged daytime intermission. Along ant roads, there were underground pavilions with dense roofs built of conifer needles and soil. Covers of the same kind appeared over root aphid colonies. The most radical improvement was the construction of a battery of brood chambers underneath the mound. The soil excavated was used for strengthening the mound and restricting the convective heat exchange between the brood chambers and the external environment. The anthill surface was covered with a smooth crusted layer of soil and fine plant debris particles, which protected the nest from the inflow of hot air from without. A portion of the inhabitants of large nests moved to newly built extensions and auxiliary nests. It was only active, non-damaged colonies that could afford these measures and thus survive the heat with minimal loss. Depressed colonies lost the major part of their brood during this time. Furthermore, small secondary colonies that emerged as the result of destructive activity of animals also failed to reassemble due to the summer heat of 2010. The months of heat were followed by a long rainy and cool period, and the colonies that had already been depressed faced critical conditions for preparing for winter. The ants were unable to accumulate lipid reserves sufficient for spring nest heating and rearing of sexuals and workers, which turned out to be an important aftereffect of the 2010 season the following year. Two first generations of workers were absent in almost all the nests. Mass oviposition in F. aquilonia commenced only late in May 2011, whereas the flight of alates did not occur at all because alates of this species are only reared in spring. Consequently, the F. aquilonia colonies had not recovered even by the end of 2012. In F. lugubris and F. polyctena, species that rear sexuals twice a year, there was only late-season alate flight, and yet it allowed their colonies to replenish the pool of ovipositing females and restore the initial colony size by the end of 2012. Other ants, represented by Lasius niger and Myrmica rubra, also modified their nests and foraging patterns during the period of heat. M. rubra suffered the most, so that its abundance and activity remained very low in 2011 as well. Only L. niger, which remained practically unscathed by the heat, successfully completed its annual cycle in 2010. Therefore, one of the 2010 year’s results was a drastic shift in the ratio of nests built by the abovementioned species in favor of L. niger.     

Reproductive allocation in multinest colonies of the ponerine ant Pachycondyla goeldii

Abstract.  1.  Pachycondyla goeldii constitutes the only recorded case of a monogynous (i.e. one queen per colony) polydomous (i.e. several nests per colony) species in the Ponerinae subfamily. This study examines the impact of polydomy on reproductive allocation between nests (also called 'calies' in polydomous society) in Pachycondyla goeldii Forel, by: (i) recording the number of workers and sexuals in 67 nests belonging to 21 colonies; (ii) dissection of workers in nine nests containing a queen (QR nests), nine nests without a queen but associated to a QR nest (QL nests) and five nests belonging to colonies that permanently lost the queen (OR nests); and (iii) measuring the length of all eggs present in the nests (our laboratory study shows that queen-laid eggs were significantly longer than worker-laid eggs).
2. The number of workers was significantly higher in QR nests than in QL nests, while the number of virgin queens was significantly higher in QL nests compared with QR nests.
3. Worker ovarian activity is inversely related to queen proximity: highest in OR nests, intermediate in QL nests, and lowest in QR nests.
4. Egg length was highest in QR nests, where the queen was most likely the primary egg-layer, intermediate in QL nests, where eggs could have originated from both the queen and workers, and lowest in OR nests, where workers were the sole egg-layers.
5. We postulate that the proximal mechanism explaining differences between QR and QL nests is the pheromonal absence of the queen from QL nests and that the evolutionary reasons of these divergences between nest types are likely to originate from the different conflicts occurring in ant colonies.     

Observation of plugging behaviour reveals entrance‐guarding schedule of morphologically specialized caste in Colobopsis nipponicus

The success of social insects is often attributed to the specialized morphologies and behaviours of workers. One of the most elaborate specializations in ant species is the workers’ plugging behaviour, in which their uniquely shaped heads are used for nest defence. These species generally nest in tree cavities. Without morphologically specialized workers (major workers) plugging the nest entrance, nests are easily attacked and replaced by conspecific and heterospecific cavity‐dwelling ants and other predators. Therefore, major workers are expected to defend the nest at all times. However, empirical evidence on the social time investment for nest defence is still lacking. Here, we investigated how major workers of Colobopsis nipponicus achieve nest defence (plugging behaviour) in terms of their work schedule. Our results showed that the nests had more major workers than entrances. The observation of artificial plastic nests over 6 days in the laboratory revealed that the nest entrance was guarded almost continuously by multiple major workers. The entrance‐guarding schedule was not equally divided among major workers, and there were no clear shift patterns in the timing of changeovers. We highlight the importance of investigating the time investment of workers to fully understand the defence strategies of this morphologically specialized caste.     

The biology in Jamaica of the adults of the sphecid wasp Sceliphron assimile Dahlbom

Abstract. 1. The adult biology of the solitary mud-wasp Sceliphron assimile Dahlbom was studied in Jamaica.
2. Adults were active from sunrise until they finally settled in roosting groups a little before sunset. Nesting took place during an 8.5 h period commencing about 3 h after sunrise but individuals rarely spent > 4 h nesting per day, the remainder being spent in resting and feeding.
3. Males sought females in all places in the habitat.
4. The number of cells per nest was positively correlated to high cell density per unit area but negatively correlated to illuminance.
5. The number of pellets used to build a cell, but not cell length, was negatively correlated to the body length of the builder. There is evidence that females that build multiple nests locate them within a few metres of each other.
6. Incompletely stored cells were closed with an externally concave lamella of mud at the onset of rain or at 16.00—17.00 hours E.S.T. and never reopened the same day.
7. Like other species of Sceliphron studied in this respect S. assimile collected spiders belonging to the Argiopidae (particularly), and also mainly to the Thomisidae, Salticidae and Oxyopidae.
8. The consequences of the nest architecture on mortality and the method of building on distribution are discussed.
9. Protarrhenotoky and proterandry occur and probably influence the sex ratio through differential mortality and fecundity. A suggested interaction between the flight motor and the ovaries could regulate egg production.     

Courtship with two spoons—Anatomy and presumed function of the bizarre antennae of Cardiocondyla zoserka ant males

Mating in ants often occurs on the wing during nuptial flights or on the ground when scattered female sexuals attract males by pheromones. In both scenarios, there is little opportunity for males to engage in prolonged aggressive competition or elaborate courtship displays. Male morphology is therefore adapted to locating female sexuals and mating, and it lacks specific weapons or other traits associated with courtship. In contrast, sexuals of the ant genus Cardiocondyla typically mate in their natal nests. As a consequence, in many species winged males have been replaced by wingless fighter or territorial males, which kill or expel rival males with their strong mandibles and show complex mating behavior. However, no wingless males are known from Cardiocondyla zoserka from West Africa, and instead, winged males have evolved a bizarre secondary sexual trait: uniquely shaped antennae with spoon‐like tips that show heavily sculptured ventral surfaces with numerous invaginations. We here report on the courtship behavior of C. zoserka males and describe antennal glands with class 3 gland cells, which presumably secrete a close range sex pheromone. Antennal glands have not yet been found in males of other ant species, including a close relative of C. zoserka, suggesting that in ants with intranidal mating sexual selection can rapidly lead to highly divergent adaptations and the evolution of novel structures.     

THE EVOLUTION OF ALTERNATIVE REPRODUCTIVE TACTICS IN MALE CARDIOCONDYLA ANTS

Alternative reproductive tactics are often associated with discontinuous variation in morphology but may evolve independent from each other. Based on life‐history data and a phylogeny we examine how male morphology and reproductive behavior are linked in the evolution of the ant genus Cardiocondyla. Wingless Cardiocondyla males engage in lethal fighting for access to female sexuals, whereas winged males disperse and mate away from the nest. This basic pattern shows considerable variation across species. A phylogeny based on ～3 kbp sequence data shows that male diphenism and lethal fighting are ancestral traits tightly linked in evolution. Winged males were lost convergently in several species groups, apparently in response to the low probability of encountering female sexuals in nests without a resident fighter male. An early dichotomy separates two clades with alternative male morphologies and fighting behavior, but phenotype and fighting strategy are not correlated with the presence of winged males.     

Colony Populations and Biomass in Nests of the Amazonian Forest Termite Anoplotermes banksi Emerson (Isoptera: Termitidae)

The arboreal nests of the termite Anoplotermes banksi are abundant in Central Amazonian primary rain forests. Colony size of 7 nests (weight 92–6891 g) varied between 2,593 and 39,256 individuals/nest (1.5 – 22.1 g termites/nest). Average body fresh weight was 0.9 mg for workers and 2.1 mg for alates. Queens weighed 10–30 mg. No relationship between nest weight and maturity was detected, as the ratio of workers to larvae was 1:1, independent of nest size, and alates were found in nests weighing less than 200 g. Nests of A. banksi (12–18 ha -1) accounted for 12–15% of the nest density of all Isopteran species, but the calculated fresh weight of the termites of this species (15–23 mg/m 2) represented only 0.2–0.4% of the total termite biomass in the study area.     

Nest architecture and colony size of the fungus-growing ant <Emphasis Type="Italic">Mycetophylax simplex</Emphasis> Emery, 1888 (Formicidae,Attini)

Detailed measurements on nest architecture and colony size of the fungus-growing ant Mycetophylax simplex Emery, 1888 (Formicidae, Attini) are reported for the first time, based on excavations of 55 nests from two sites in southern Brazil. All nests were subterranean, with a single entrance hole. Most nests consisted of two chambers, an upper and a lower chamber, but one and three-chamber nests were also found. The chambers were more cone-shaped than rounded, and located at a depth ranging from 4.0 cm to 32.5 cm below the nest entrance. The chamber dimensions generally increased as the depth of the chambers increased, and the lower chamber was mostly wider than the upper one. The fungus garden was always found resting on the chamber floor. The average colony size was 264.1 workers, ranging from 67 to 610 workers. Colonies produced most sexuals during the summer (from December to March) and a few during the winter (July). Direct observations showed that colonies were mostly monogynous, but more than one queen was recorded in two nests, suggesting that polygyny may also occur in this species. Received 30 November 2006; revised 20 April 2007; accepted 23 April 2007.     

Mate guarding and alternative reproductive tactics in the ant Hypoponera opacior

In many animals, males have evolved weapons, elaborate courtship displays, or costly ornaments to increase their reproductive success. Ants, in contrast, commonly mate during nuptial flights, in which males do not profit from fighting or attempting to monopolize females. However, where mating occurs in the nest, males can use other reproductive tactics. We found that wingless (apterous) males of Hypoponera opacior sat on top of queen cocoons, inserted their genitalia into the cocoons and remained in copula with cocooned queens for up to 40 h. These males were tolerant of each other; fighting was never recorded. Our observations therefore suggest that wingless males of H. opacior ensure reproduction by copulatory mate guarding. This strategy, although time consuming, presumably reduces the likelihood of subsequent inseminations by other males. Apterous H. opacior males have only a limited amount of sperm available: histological preparations showed that, in contrast to Cardiocondyla fighter males, the testes degenerate in early adult life. Males ofH. opacior have relatively few mating opportunities. Although some wingless males were reproductively active for more than 3 weeks, we observed a maximum of only six matings per male, with a mean slightly above one. SomeH. opacior males used an alternative reproductive tactic of dispersal and outbreeding. We found colonies headed by single, dealate queens, which did not rear wingless sexuals but presumably reproduced through winged reproductives that mate in nuptial flights. The social structure of those colonies contrasted with nests containing wingless reproductives, which were highly polygynous and polydomous.