The relationship between multiple mating by queens, within-colony genetic variability and fitness in the ant Lasius niger

Multiple mating has been suggested to benefit social insect queens because high genetic variation within colonies might decrease the load imposed by sterile diploid males, enhance resistance to parasites and pathogens, and lead to a more effective division of labour and/or a wider range of tolerable environmental conditions. We tested these hypotheses in the ant Lasius niger with three population samples from Switzerland and Sweden. We found no diploid males in young or mature colonies suggesting a lack of diploid male load. Colonies with multiply-mated queens were not larger nor did they produce more sexuals than colonies with singly-mated queens. We did find a significantly lower frequency of multiple mating among newly mated queens than among the queens heading mature colonies in one population sample (Switzerland 1997). However, this result was not repeated in the other study population, or in the following year in the Swiss population.     

Population and colony structure and morphometrics in the queen dimorphic harvester ant, <Emphasis Type="Italic">Pogonomyrmex pima</Emphasis>

The North American seed-harvester ant Pogonomyrmex (Ephebomyrmex) pima displays a dimorphism that consists of winged (alate) and wingless (intermorph) queens; both types of queens are fully reproductive. Microsatellite allele frequencies and a mitochondrial phylogeny demonstrate (1) alate and intermorph queens represent an intraspecific wing polymorphism, and (2) an absence of assortative mating and inbreeding by males. Surveys at our field site in southcentral Arizona, USA, demonstrated that only one type of queen (intermorph or dealate) occurred in each colony, including those excavated during the season in which reproductive sexuals were present. Colony structure appeared to vary by queen type as most intermorph colonies contained multiple mated queens. Alternatively, dealate queen colonies rarely contained a mated queen. Our inability to find mated dealate queens in these colonies probably resulted from difficulty in excavating the entire colony and reproductive queen, especially given that these colonies were only excavated over one day. A morphometric analysis demonstrated that intermorph queens are intermediate in size to that of workers and alate queens, but that intermorph queens retain all of the specialized anatomical features of alate queens (except for wings). Some colonies had queens that foraged and performed nest maintenance activities, and these queens sometimes accounted for a significant portion of colony foraging trips. Dissections revealed that these queens were uninseminated; some of these queens produced males in the laboratory. Received 24 October 2006; revised 1 December 2006; accepted 8 December 2006.     

Review. Lifelong commitment to the wrong partner: hybridization in ants

The extraordinary lifelong partner commitment in social insects is expected to increase choosiness in both sexes and therefore to be associated with particularly low hybridization frequencies. Yet, more and more studies reveal that in many ant taxa hybrids are surprisingly common, with up to half of all female sexuals receiving sperm from allospecific males in extreme cases. In a few ant species, hybridization has led to the evolution of reproductively isolated new lineages with a bizarre system of genetic caste differentiation: colonies produce hybrid workers and pure-lineage female sexuals. This requires that colonies either contain multiple queens or that queens mate multiple times. In most other cases, hybridization appears to be an evolutionary dead end and fertile hybrid queens are rarely found. In such cases, haplodiploid sex determination appears to decrease the costs of mating with an allospecific male. As long as hybrid workers are viable, a cross-mated queen can partially rescue its fitness by producing males from unfertilized eggs. Mating with an allospecific partner may thus be an option for queens when conspecific mates are not available. The morphological similarity of most ant males, perhaps resulting from the lack of sexual conflict, may similarly contribute to the commoness of hybridization.     

Queen-queen competition by precocious male production in multiqueen ant colonies

Arriving earlier in the breeding area than his rivals may be beneficial for a male when females mate only once or during a short time span. The timing of a male's entrance is usually determined by the male himself, e.g., through returning early from his winter quarters or through accelerated larval development . Here, we document a surprisingly simple way of "first come, first served" in a species with local mate competition. In multiqueen colonies of a Cardiocondyla ant, mother queens make sure that their own sons are the first to monopolize mating with a large harem of female sexuals by producing extremely long-lived males early in colony life. Whereas queens in newly founded single-queen colonies started to produce male and female sexuals only several weeks after the eclosion of their first worker offspring, queens in multiqueen colonies precociously reared sons long before the first female sexuals and even before the emergence of their first workers. These early males killed all later emerging males in the nest and mated with all female sexuals subsequently produced. Our data document that the patterns of growth and productivity of insect colonies are surprisingly flexible and can be turned upside down under appropriate selection pressures.     

Queen-queen competition and reproductive skew in a <Emphasis Type="Italic">Cardiocondyla</Emphasis> ant

Queens and female sexuals of the Southeast Asian ant Cardiocondyla sp. engage in aggressive interactions. By biting and violently antennating female sexuals, queens appear to prevent them from shedding their wings and presumably also from starting to lay haploid eggs. Aggression among dealate queens apparently results in the establishment of reproductive rank orders with considerable differences in offspring production among individual nestmate queens, as visualized by the pronounced color polymorphism of this taxon. Reproductive skew ranged from complete monopolization of both worker and female sexual production to more or less equal partitioning of reproduction. Division of reproduction was associated with variation in the location of queens close to or away from the center of the brood pile, which again appeared to be caused by queen-queen antagonism. Received 23 February 2007; revised 15 May 2007; accepted 22 May 2007.     

Three queen morphs with alternative nest-founding behaviors in the ant, Temnothorax longispinosus

In many ant species, multiple modes of founding new colonies occur in the same population. These modes include dependent founding, independent founding by haplometrosis (single queen), and independent founding by pleometrosis (multiple queens). In several cases, a dimorphism in queen size has been found, such that each morph specializes in a particular nest-founding behavior. I investigated queen size in the ant Temnothorax longispinosus in several southern Wisconsin populations and found three distinct queen morphs: small queens with very low fat content and short wings, large queens with low fat and long wings, and large queens with high fat and very long wings. Several traits associated with founding behavior correlated with these queen sizes. Small queens were produced in lower numbers, were more common in polygynous nests, and returned to the nest in higher proportions than both large queen morphs. The size ranges and fat levels of each queen morph were similar to those of other species that specialize in either haplometrosis (very large, high fat), pleometrosis (large, low fat), or dependent founding (small and low fat). However, there was extensive overlap in several of the founding behaviors, suggesting that the morphs in these populations have some flexibility in founding behavior. The queen morphs in these populations of T. longispinosus may resemble early stages in the evolution of more specialized dispersal polymorphisms found in other ant species. Received 11 January 2006; revised 15 September 2006; accepted 18 September 2006.     

A newly-discovered mode of colony founding among fire ants

Summary Queen ants start new colonies either unassisted by workers (independent founding), assisted by workers from their natal nest (dependent founding), or assisted by the workers of other species (dependent, socially parasitic). The monogyne form of the fire ant,Solenopsis invicta, founds independently in summer, but in the fall it also produces a few sexuals some of which overwinter, then fly and mate in early spring. These overwintered queens lack the nutritional reserves and behaviors for independent colony founding. Rather, they seek out unrelated, mature, orphaned colonies, enter them and exploit the worker force to found their own colony through intraspecific social parasitism. Success in entering orphaned colonies is higher when these lack overwintered female alates of their own. When such alates are present, orphaning causes some to dealate and become uninseminated replacement queens, usually preventing entry of unrelated, inseminated replacement queens. Such colonies produce large, all-male broods. Successful entry of a parasitic queen robs the host colony of this last chance at reproductive success. Only overwintered sexuals take part in this mode of founding.     

The influence of hybridization on colony structure in the ant species Temnothorax nylanderi and T. crassispinus

The parapatric sibling ant species Temnothorax nylanderi and T. crassispinus hybridize in the contact zone in the Franconian Jura, Southern Germany. Aim of our study was to investigate the impact of hybridization on colony composition and fitness. We classified colonies as either ‘pure’ or containing hybrids by determining their allozyme pattern at GPI, an enzyme that is fixed for different alleles in the two parental species, and quantified their reproductive output. Most colonies with hybrid workers had a T. crassispinus queen. Colonies with heterozygous, hybrid workers produced more young workers than colonies of the parental species but similar numbers of male and female sexuals. Female sexuals from colonies with heterozygous workers had a significantly lower weight than female sexuals from pure colonies. Only a single reproductive queen was found to be heterozygous, suggesting reduced fitness of hybrid queens. As in the parental species, hybrid colonies appear to be frequently taken over by alien queens, which obscures the genetic colony structure. Received 6 April 2006; revised 10 June 2006; accepted 15 June 2006.     

Impact of Thelohania solenopsae (Microsporidia: Thelohaniidae) on polygyne colonies of red imported fire ants (Hymenoptera: Formicidae)

Three studies were conducted to assess the effects of the entomopathogen Thelohania solenopsae on polygynous, red imported fire ant, Solenopsis invicta Buren, colonies. A total of 57 of 122 queens (46.7%) from nine, field-collected, polygyne, S. invicta colonies, was infected with T. solenopsae. Infection rate of queens for each colony ranged from 25 to 75%. Laboratory colonies of polygyne S. invicta, with three to 12 queens, were inoculated and infected with T. solenopsae. Brood levels in all infected colonies declined to 0 after 26-52 wk. Brood did not reappear in any of the colonies after 3-11 wk, even though in two of the eight infected colonies, five fertile queens that were uninfected were recovered. Thus, polygyne, S. invicta colonies infected with T solenopsae, which were confined and isolated under laboratory conditions, did not recover. Field plots that contained polygynous S. invicta colonies, which were infected with T solenopsae, were monitored over a 2-yr period. Infection rates increased during the study and reached a maximum of 9.3%. Fire ant nest density and colony sizes fluctuated over time, with maximum reduction of 63% per plot. In general, fire ant reductions were attributed to smaller colony sizes. T. solenopsae infections in polygynous S. invicta can result in a slow colony decline and death. Under field conditions, the prolonged colony death may mask the impact of T. solenopsae by allowing for concurrent reinfestations.     

Mating frequency and mating system of the polygynous ant, Leptothorax acervorum

Multiple mating by queens (polyandry) and the occurrence of multiple queens in the same colony (polygyny) alter patterns of relatedness within societies of eusocial insects. This is predicted to influence kin-selected conflicts over reproduction. We investigated the mating system of a facultatively polygynous UK population of the ant Leptothorax acervorum using up to six microsatellite loci. We estimated mating frequency by genotyping 79 dealate (colony) queens and the contents of their sperm receptacles and by detailed genetic analysis of 11 monogynous (single-queen) and nine polygynous colonies. Results indicated that 95% of queens were singly mated and 5% of queens were doubly mated. The corrected population mean mating frequency was 1.06. Parentage analysis of adults and brood in 17 colonies (10 monogynous, 7 polygynous) showed that female offspring attributable to each of 31 queens were full sisters, confirming that queens typically mate once. Inbreeding coefficients, queen-mate relatedness of zero and the low incidence of diploid males provided evidence that L. acervorum sexuals mate entirely or almost entirely at random. Males mated to queens in the same polygynous colony were not related to one another. Our data also confirmed that polygynous colonies contain queens that are related on average and that their workers had a mixed maternity. We conclude that the mating system of L. acervorum involves queens that mate near nests with unrelated males and then seek readoption by those nests, and queens that mate in mating aggregations away from nests, also with unrelated males.     

Response of late-instar Blattella germanica (Dictyoptera: Blattellidae) to dietary insect growth regulators

Late-instar German cockroaches, Blattella germanica (L.), were used to evaluate the relative effects of single treatments and combinations of three insect growth regulators (IGRs): pyriproxyfen, fenoxycarb, and diflubenzuron. Groups of 15 males or 15 females were held for 2 wk on food treated with varying amounts of IGRs. After removal, newly mature adults were placed with untreated adults of the opposite sex. Mortality, adult phenotype (normal wings, divergent wings, curly wings, and nymphoids), and reproduction were assessed. Mortality occurred largely in the nymphal stage except when all three IGRs were combined. Neither the average number of nymphs per egg case nor hatch of egg cases from phenotypically normal cockroaches (normal wings, occasional darkening of the body) was affected in single treatments or in combinations of two IGRs, but nymphal numbers were reduced when the three IGRs were combined. Hatch from matings of insects with divergent wings varied. At low concentrations (3-10 ppm), hatch was generally normal; at high concentrations, most mating tests were unproductive. Mating tests of cockroaches with curly wings were almost always unproductive. When males with curly wings were mated, females dropped either unfertilized egg cases (no mating) or partially fertilized egg cases. Curly-wing females either dropped unfertilized egg cases or failed to form egg cases because of deleterious effects on ovarian development. Nymphoids did not mate. Diflubenzuron at 100 ppm had no effect other than causing the appearance of a few insects with divergent wings. Effects on phenotype and reproduction began at 3 ppm of both pyriproxyfen and fenoxycarb and at a comparable concentration in the combination of pyriproxyfen + fenoxycarb (1 ppm each). Female sterility was complete at 100 ppm of pyriproxyfen and fenoxycarb. When pyriproxyfen or fenoxycarb was combined with equal amounts of diflubenzuron, the number of productive matings was not reduced at 6 ppm (3 ppm per each IGR). At 20 ppm (10 ppm per each IGR), a reduction in productive matings coincided with the appearance of curly wings. Complete female sterility occurred only at 600 ppm (300 ppm per IGR). The most severe effects occurred in the experiment with equal amounts of pyriproxyfen, fenoxycarb, and diflubenzuron. In addition to reduced hatch from normal phenotypes, this experiment caused complete male sterility (300 ppm; 100 ppm of each IGR). Female sterility was complete at greater than or equal to 90 ppm (30 ppm of each IGR).     

Production of winged and wingless males in the ant, <Emphasis Type="Italic">Cardiocondyla minutior</Emphasis>

Summary Wingless (ergatoid) males of the tramp ant Cardiocondyla minutior attack and kill their young ergatoid rivals and thus attempt to monopolize mating with female sexuals reared in the colony. Because of the different strength of local mate competition in colonies with one or several reproductive queens, we expected the production of new ergatoid males to vary with queen number. Sex ratios were mostly female-biased, but in contrast to the sympatric species C. obscurior (Cremer and Heinze, 2002) neither the percentage of ergatoid males nor of female sexuals among the first 20 sexuals produced varied considerably with queen number. As in C. obscurior, experimental colony fragmentation led to the production of winged males, whereas in unfragmented control colonies only ergatoid males eclosed.Received 3 December 2003; revised 20 February 2004; accepted 23 February 2004.     

Effect of queen number on the production of sexuals in natural populations of the fire ant, Solenopsis invicta

ABSTRACT. To determine whether the production of sexuals in the fire ant, Solenopsis invicta Buren, is related to queen number in the field, the numbers of sexuals in monogyne (M) and polygyne (P) colonies of this species were compared. Large colonies ( n =25 M and 25 P) were collected at random during spring, summer and fall when sexuals are actively produced, and the numbers of sexuals present were counted. The numbers of alates plus sexual pupae and larvae in M colonies far exceeded those in P colonies in each season. Significant differences between M and P colonies were also found when the numbers of alates and sexual immatures (pupae + larvae) were compared separately in each season. In addition, M colonies contained significantly more female sexuals (alates + pupae) than did P colonies in the summer and fall, and significantly more males in all three seasons. The negative relationship between queen number and number of sexuals provides evidence that queen control over the production of sexuals, previously established in laboratory experiments, also occurs under natural conditions.     

Bait distribution among multiple colonies of Pharaoh ants (hymenoptera: Formicidae)

Pharaoh ant, Monomorium pharaonis (L.), infestations often consist of several colonies located at different nest sites. To achieve control, it is desirable to suppress or eliminate the populations of a majority of these colonies. We compared the trophallactic distribution and efficacy of two ant baits, with different modes of action, among groups of four colonies of Pharaoh ants. Baits contained either the metabolic-inhibiting active ingredient hydramethylnon or the insect growth regulator (IGR) pyriproxyfen. Within 3 wk, the hydramethylnon bait reduced worker and brood populations by at least 80%, and queen reductions ranged between 73 and 100%, when nests were in proximity (within 132 cm) to the bait source. However, these nest sites were reoccupied by ants from other colonies located further from the bait source. The pyriproxyfen bait was distributed more thoroughly to all nest locations with worker populations gradually declining by 73% at all nest sites after 8 wk. Average queen reductions ranged from 31 to 49% for all nest sites throughout the study. Even though some queens survived, brood reductions were rapid in the pyriproxyfen treatment, with reductions of 95% at all locations by week 3. Unlike the metabolic inhibitor, the IGR did not kill adult worker ants quickly, thus, more surviving worker ants were available to distribute the bait to all colonies located at different nest sites. Thus, from a single bait source, the slow-acting bait toxicant provided gradual, but long-term control, whereas the fast-acting bait toxicant provided rapid, localized control for a shorter duration.     

Selection against Aerial Dispersal in Ants: Two Non-Flying Queen Phenotypes in Pogonomyrmex laticeps

The South American seed-harvester ant Pogonomyrmex laticeps has dimorphic queens: ergatoid (permanently wingless) and brachypterous (short, non-functional wings). Surveys in western Argentina indicated that colonies near Chilecito, La Rioja Province, produced only ergatoid queens, while those near Punta Balasto, Catamarca Province (263 km away), produced only brachypterous queens. Brachypterous queens were significantly larger than ergatoid queens for 10 of 11 external characters, but both phenotypes had comparable reproductive potential, i.e., a spermatheca and a similar number of ovarioles. Using normal winged queens of the closely related P. uruguayensis for comparison, we determined that both queen phenotypes in P. laticeps had a full set of dorsal thoracic sclerites, albeit each sclerite was much reduced, whereas workers had a thorax without distinct dorsal sclerites. Sclerites were fused and immobile in ergatoid queens, while they were separable and fully articulated in brachypterous queens. Both phenotypes lacked the big indirect flight muscles, but brachypterous queens retained the tiny direct flight muscles. Overall, this dimorphism across populations indicates that there are alternative solutions to selective pressures against flying queens. We lack field data about colony founding strategy (independent or dependent) for either queen phenotype, but colonies at both sites produced numerous gynes, and we infer that all foundresses initiate colonies independently and are obligate foragers.     

Stenamma debile (Hymenoptera, Formicidae): Facultative polygyny, and an extreme 1997 sex ratio

Summary: Monogynous and a high proportion of truly polygynous colonies of Stenamma debile were found in populations of this small and inconspicuous ant in southern Germany. Moreover, dissections revealed that queens of different age may co-occur in one nest. Thus polygyny is not only an outcome of pleometrosis. Probably young queens sometimes join the mother colonies.¶Practically no female sexuals were reared in the 1997 brood of 29 colonies, thus a population-level sex ratio of &gif1; was recorded. It is likely that such an extreme sex ratio is atypical for this species, suggesting substantial year-to-year variation in sex ratios. Variation of sex ratios over several years has been reported for other ant species, too, but rarely to such an extent. The reason for the failure of queen production in 1997 was perhaps an unusual sequence of warm and cold periods in the spring which may have caused a re-determination of the queen-destined hibernated larvae to become workers, while male larvae were reared in ordinary numbers.     

保幼激素类似物对德国小蠊卵黄发生及繁殖的影响

应用ELISA、单向免疫扩散法,研究了保幼激素类似物（灭幼宝）对德国小蠊Blattella germanica卵黄发生的影响;同时,观察了灭幼宝对德国小蠊繁殖的影响。灭幼宝促进了德国小蠊脂肪体Vg的合成,使血淋巴Vg的浓度明显增高,出现时间早于对照组;且变化规律不同于对照组,呈双峰模式。灭幼宝处理组昆虫卵巢中Vt的含量在第5天、第7天明显高于对照组,而在第9天明显下降,仅为对照组的50%左右,而此时对照组卵巢的Vt含量则上升到处理组第7天水平。灭幼宝处理新羽化的雌虫,导致不育。     

Selective male mortality in the red imported fire ant, Solenopsis invicta

Males in polygyne populations of Solenopsis invicta are primarily sterile diploids and thought to not express the Gp-9 gene coding for a pheromone-binding protein affecting complex social behavior. We examined an aspect of the breeding system hitherto not considered--male Gp-9 genotypes in relation to sperm stored in queens. Four sites with varying frequencies of sympatric monogyne and polygyne colonies were sampled, including sexuals, workers, and broods from four colonies. Most queens were heterozygotes storing B sperm. Although predicted to be common, only 14 of 504 males were B or BB genotypes, suggesting strong selection. Increased frequency of polygyne colonies at each site paralleled increases in queens with b sperm (1.9-32.8%) and of noninseminated queens. The presence of both B and b sperm in 1.9-18.9% of queens, genotype profiles of colonies, and genotypes of offspring from individual queens suggest some frequency of multiple mating. The bb genotype, rather than an obligate, developmental lethal, was present in some queens and common in alates, workers, and brood. Selective mortality of sexuals may affect multiple aspects of the breeding system, including female-mediated dispersal, mating success, and gene flow.     

Identification and action of juvenile hormone III from sexually mature alate females of the red imported fire ant,Solenopsis invicta

Analysis of extracts of hemolymph obtained from sexually mature alate females of Solenopsis invicta from monogyne colonies resulted in identification of juvenile hormone III (JH III). The average amount of JH III was 0.32±0.04 pmol/μmolof hemolymph. Topical application of 0.038 pmol of JH III was sufficient to stimulate alates to shed their wings in the presence of the queen. The time in which alates were induced to dealate decreased linearly with increasing concentrations of JH III from 0.038 to 3.8 pmol. However, higher JH III concentrations deviated from linearity and did not reach dealation times comparable with those that occur after mating flights. Thus, it appears that the mechanism of dealation that occurs when female alates are out of the influence of their queen is different from the one associated with mating flights. Application of 0.42 μmol of precocene II inhibited dealation of alates in queenless colonies. However, this inhibition was reversed after applying 38 pmol JH III to precocene-treated alates. The sizes of corpora allata (CA) from sexuals treated with JH III did not differ from those of controls. However, the sizes of CA were reduced in alates treated with precocene II. The results indicated that JH was important to dealation.     

No inbreeding depression but increased sexual investment in highly inbred ant colonies

Inbreeding can lead to the expression of deleterious recessive alleles and to a subsequent fitness reduction. In Hymenoptera, deleterious alleles are purged in haploid males moderating inbreeding costs. However, in these haplodiploid species, inbreeding can result in the production of sterile diploid males. We investigated the effects of inbreeding on the individual and colony level in field colonies of the highly inbred ant Hypoponera opacior. In this species, outbreeding winged sexuals and nest‐mating wingless sexuals mate during two separate reproductive periods. We show that regular sib‐matings lead to high levels of homozygosity and the occasional production of diploid males, which sporadically sire triploid offspring. On the individual level, inbreeding was associated with an increased body size in workers. On the colony level, we found no evidence for inbreeding depression as productivity was unaffected by the level of homozygosity. Instead, inbred colonies altered their allocation strategies by investing more resources into sexuals than into workers. This shift towards sexual production was due to an increased investment in both males and queens, which was particularly pronounced in the dispersive generation. The absence of inbreeding depression combined with increased reproductive investment, especially in outbreeding sexuals, suggests that these ants have evolved active strategies to regulate the extent and effects of frequent inbreeding.