Colony social organization of <Emphasis Type="Italic">Lasioglossum malachurum</Emphasis> Kirby (Hymenoptera,Halictidae) in southern Greece

Summary We studied the nesting and social biology of two aggregations of the obligately eusocial halictine bee Lasioglossum malachurum at Agios Nikolaos Monemvasias (ANM) in southern Greece. Observations and nest excavations carried out from May to June 2000, revealed social and demographic variation between aggregations and years at ANM, as well as notable differences between these and other European populations. In southern Greece, the colony cycle includes multiple broods: the first two broods comprise only workers, whereas the third brood comprises workers, males, and gynes. Although pleometrosis is unknown in other populations, in the ANM region, as many as 10% of nests have more than one foundress. Newly emerged workers and gynes exhibit non-overlapping size distributions, but a few queens are worker-sized, indicating that workers occasionally overwinter and become foundresses. Although the vast majority of workers are unmated and most exhibit no ovarian development, an increase in worker ovarian development at the time of male production suggests that many males may develop from worker-laid eggs. Worker reproduction seems to be inhibited by the presence of queens, and annual variation in queen mortality may underlie annual variation in worker ovarian development. Across Europe, the major demographic and social differences among L. malachurum populations are in the number of worker broods and the extent of worker ovarian development. This contrasts with the results of a principal components analysis of social traits among 15 social L. (Evylaeus) populations, which shows that interspecific social variation is defined by the proportion of males in the early brood, the proportion of workers mated, queen-worker size dimorphism, gyne overwintering locale, and the proportion of workers with developed ovaries.Received 17 June 2002; revised 16 January 2002; accepted 27 January 2003.     

Levels of polyandry in <Emphasis Type="Italic">Apis laboriosa</Emphasis> Smith from Nepal

Summary Actual and effective number of matings of Apis laboriosa Smith queens were estimated by analysing 40–192 workers or worker pupae of five colonies using microsatellites. Effective paternity frequencies varied from 13.63 to 31.57, with a mean of 20.19 (SD 6.07). Our results demonstrate that the Himalayan Giant honeybee A. laboriosa shows high levels of polyandry, typical of the genus.Received 30 January 2002; revised 24 October 2003; accepted 3 December 2003.     

Queen number in a supercolony of the invasive garden ant, <Emphasis Type="Italic">Lasius neglectus</Emphasis>

Summary We have analysed the distribution of queens under stones at the core and at the periphery of a supercolony of Lasius neglectus that occupies 14 ha at Seva (NE Spain). Queens were not found alone, but rather within worker groups. Density at the center (mean ± s.d.: 1.38 ± 2.87 queens/stone; n = 100 stones; range 0–14) was not different from density at the periphery (1.18 ± 2.38; range 0–12). The estimate of the number of queens found under stones for the whole colony is about 35500. Egg-laying rates for queens from these two zones were obtained in the laboratory, at three different temperatures, and there were no differences detected. The presence of brood stages, from eggs to cocoons, was also similar in both zones. The homogeneous distribution of colony components may indicate that the area occupied by L. neglectus has already reached saturation. With a different technique – soil core extraction – we could estimate the density of workers in the soil: 800 workers per m². Soil cores had 6.28 ± 20.0 workers/core (range: 0–173), giving a rough estimate of 1.12 × 10⁸ workers in the soil, for the entire colony. Though few, some queens were also recovered from soil cores. Queen numbers for the supercolony, based on queens found in the soil, reaches the astounding level of 360000. Numbers are consistent with previous predictions.Received 10 September 2003; revised 1 December 2003; accepted 9 December 2003.     

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.     

Effects of reproductive resource supplementation on space-use patterns in Dendrobates pumilio

Summary I increased the availability of tadpole-rearing sites (bromeliads) and oviposition sites (leaf litter) and examined the effects on space-use patterns in adult Dendrobates pumilio at the La Selva Biological Reserve in northeastern Costa Rica. I made 4158 observations of 986 marked adults during the seven-month study period. The addition of leaf litter did not result in increased density, but the addition of bromeliads did. The addition of leaf litter resulted in increased home-range size, but the effect depended on bromelial addition and sex. Home-range size was greater for females than males, and may be related to parental care and/or mate selection. Home-range size was not correlated with density. Spatio-temporal overlap was not affected by resource supplementation but the proportion of females captured in sites shared with males varied temporally. Dispersion was no affected by bromeliad supplementation but males on litter-addition plots were less clumped than males on other plots. These results indicate that the density increase observed on bromeliad-addition plots was accommodated by frogs utilizing previously unoccupied space. Frogs were captured more frequently at bromeliads on bromeliad-supplemented plots than on litter-addition and control plots. The size of the area defended by territorial males is smaller than home-range size indicating the absence of home-range defense.     

Social parasitism of <Emphasis Type="Italic">Polistes dominulus</Emphasis> by <Emphasis Type="Italic">Polistes nimphus</Emphasis> (Hymenoptera,Vespidae)

Summary Interspecific facultative social parasitism is well known in ants and in bumble-bees, but it is rarer in wasps. This form of parasitism is traditionally considered to be an intermediate stage in the evolution of obligate interspecific parasitism, where the parasites are no longer able to nest alone. We report field and experimental observations of a newly discovered facultative parasitic relationship between two closely related free-living Polistes species: P. nimphus and P. dominulus. P. nimphus foundresses sometimes usurp the nests of the larger P. dominulus before worker emergence. The invading queen takes over the nest with abundant abdomen stroking on the nest surface and is accepted by workers if they emerge 6 or more days after usurpation. Morphometric comparisons show that the usurper species, though smaller than its victims, has morphological adaptations consisting of larger heads, mandibles and front femora relative to their body size that may give it an advantage during nest invasion. This strategy is likely to be taken only after the foundress loses her original nest because invading P. nimphus queens have lower reproductive success than they would have had on their own nest. Overall, we found that P. nimphus usurpers use strategies of invasion similar to those of two obligate parasites, suggesting that this may be an example of one of the pathways by which social parasitism evolved.Received 4 April 2003; revised 8 August 2003; accepted 14 September 2003.     

Potential and realized reproduction by different worker castes in queen-less and queen-right colonies of <Emphasis Type="Italic">Pogonomyrmex badius</Emphasis>

Workers of the Florida harvester ant (Pogonomyrmex badius), the only North American Pogonomyrmex with a polymorphic worker caste, produce males when colonies are orphaned. In this study,we assessed the reproductive potential of workers of each caste group, minors and majors, in the presence and absence of the queen, and tested whether males produced in natural queen-right colonies are derived from workers. Worker size was positively correlated with ovariole number such that major workers had approximately double the number of ovarioles as minor workers. The number of vitellogenic oocytes, a measure of reproductive potential, was greater in major compared to minor workers and increased in both worker castes when queens were removed. Major workers have greater reproductive potential than minors although they represent a minority within the colony (~5% of workers are majors). Worker produced eggs were visible in colonies 28 – 35 days after queen removal. This time lag, from queen removal to egg production, is similar to other ants and bees. Though workers are capable of producing viable eggs, we found no evidence that they do so in queen-right colonies, suggesting that worker reproduction is controlled via some social mechanism (self restraint, policing, or inhibition). This result supports predictions of kin selection theory – that due to multiple mating by the queen workers are more related to queen-produced males than most worker-produced males and should thus favor reproduction by the queen and inhibit reproduction by other workers. Received 25 January 2007; revised 1 May 2007; accepted 21 May 2007.     

Breeding behaviour, relatedness and sex-investment ratios inLeptothorax tuberum Fabricius

The ant speciesLeptothorax tuberum was shown to be predominantly monogynous. Queens usually mate once only but some nests may have a multiply-mated queen or are partially or serially polygynous. As expected from these results, within nest relatedness between workers and between workers and alate queens was found to be high. Almost fifty percent of nests had no nest queen which may indicate high queen mortality, queens leaving to found new nests or nest fragmentation. Observed female investment frequencies (IF _o ) were not significantly different from those expected on the basis of worker control of sexual production and the relatedness estimates of workers to alate queens and workers to males calculated from isozyme data (IF _E ). These values were not consistent with queen control. There was no evidence for lower IF _o s in queenless nests nor for higher IF _o s in larger nests classified by worker number. When nests were classified by sexual productivity, however, there was a strong rank correlation between productivity and female bias. This is the first study of an ant species to test observed IF against expected IF calculated without inferring between caste relatedness from worker data or pooling of data from different castes.     

Queen–worker caste ratio depends on colony size in the pharaoh ant (<Emphasis Type="Italic">Monomorium pharaonis</Emphasis>)

The success of an ant colony depends on the simultaneous presence of reproducing queens and non-reproducing workers in a ratio that will maximize colony growth and reproduction. Despite its presumably crucial role, queen–worker caste ratios (the ratio of adult queens to workers) and the factors affecting this variable remain scarcely studied. Maintaining polygynous pharaoh ant (Monomorium pharaonis) colonies in the laboratory has provided us with the opportunity to experimentally manipulate colony size, one of the key factors that can be expected to affect colony level queen–worker caste ratios and body size of eclosing workers, gynes and males. We found that smaller colonies produced more new queens relative to workers, and that these queens and workers both tended to be larger. However, colony size had no effect on the size of males or on the sex ratio of the individuals reared. Furthermore, for the first time in a social insect, we confirmed the general life history prediction by Smith and Fretwell (Am Nat 108:499–506, 1974) that offspring number varies more than offspring size. Our findings document a high level of plasticity in energy allocation toward female castes and suggest that polygynous species with budding colonies may adaptively adjust caste ratios to ensure rapid growth.     

Male Behavior and Regulation of Worker Mating in a Ponerine Ant, Pachycondyla (Bothroponera) sp. (Hymenoptera: Formicidae)

Male behavior and regulation of worker mating were observed in Indonesian Pachycondyla sp. The colonies had some mated workers but only one gamergate (mated and egg-laying workers). Males frequently tried to mate with workers in their natal nests irrespective of worker age and dominance rank, however, workers never accepted mounting of nestmate males. Worker mating occurred only under the gamergateless condition with alien males.     

Kin structure and colony male reproduction in the hornet<Emphasis Type="Italic"> Vespa crabro</Emphasis> (Hymenoptera: Vespidae)

We estimated queen mating frequency, genetic relatedness among workers, and worker reproduction in Vespa crabro flavofasciata using microsatellite DNA markers. Of 20 colonies examined, 15 contained queens inseminated by a single male, 3 colonies contained queens inseminated by two males, and 2 colonies contained queens inseminated by three males. The genetic relatedness among workers was estimated to be 0.73±0.003 (mean±SE). For this high relatedness, kin selection theory predicts a potential conflict between queens and workers over male production. To verify whether males are derived from queens or workers, 260 males from 13 colonies were genotyped at four microsatellite loci. We found that all of the males were derived from the queens. This finding was further supported by the fact that only 33 of 2,990 workers dissected had developed ovaries. These workers belonged to 2 of the 20 colonies. There was no relationship between queen mating frequency and worker reproduction, and no workers produced male offspring in any of the colonies. These results suggest that male production dominated by queens in V. crabro flavofasciata is possibly due to worker policing.     

Non-lethal sampling of DNA from bumble bees for conservation genetics

Summary Non-lethal sampling of DNA from individuals in wild populations will often be required for studies of the conservation genetics of social insects, since it avoids destroying members of scarce or declining species. We investigated the effectiveness and consequences of methods of non-lethal sampling of DNA from bumble bee workers. In an experiment with two captive and confined Bombus terrestris colonies, we found that, unlike sampling haemolymph, sampling the terminal portion of the tarsus of a mid-leg of a worker reliably yielded amplifiable microsatellite DNA and did not significantly reduce worker survivorship. In a further experiment with four B. terrestris colonies whose workers were allowed to forage freely at flowers in the external environment, tarsal sampling of either a mid-leg or a hind-leg had no significant effects on worker survivorship, the mean body mass of foraging workers, the frequency or duration of foraging trips, mass of pollen loads or mass of nectar loads. We therefore suggest that tarsal sampling of either a mid-leg or a hind-leg is an effective and acceptable means of non-lethally sampling DNA from workers in wild populations of bumble bees, because effects on individual and colony performance are likely to be absent or minimal.Received 2 December 2002; revised 10 April 2003; accepted 25 April 2003.     

Bioecology of the fiddler crab <Emphasis Type="Italic">Uca uruguayensis</Emphasis> and the burrowing crab <Emphasis Type="Italic">Chasmagnathus granulatus</Emphasis> (Decapoda,Brachyura) in the Refugio de Vida Silvestre Bahía Samborombón,Argentina

The aim of the present work conducted at the Refugio de Vida Silvestre Bahía Samborombón, Argentina, is to analyse the most relevant aspects of the life cycles of Chasmagnathus granulatus and Uca uruguayensis: their abundance, size distribution and sex ratio in order to be used for make management purposes. A total of 1200 individuals of U. uruguayensis (megalopae, juveniles and adults) and 957 individuals of C. granulatus (juveniles and adults) were collected from March 2001 to February 2003. U.␣uruguayensis had a maximun density of 42 ind./m² in February 2003, while the maximun density of C.␣granulatus was 52 ind./m² in June 2001. Minimun density was zero individual for both species in July 2002. Each environmental factor sampled (temperature of water, salinity and pH) was correlated with the abundance of each species by Pearson’s linear correlation analysis. Sex ratio did not significantly differ from the expected Mendelian ratio, except for U. uruguayensis in December 2002 (male-biassed) and C.␣granulatus in June and December 2001, April 2002 and February 2003 (female-biassed). Ovigerous females of U. uruguayensis and C. granulatus were found during the summer.     

Nesting biology of the fungus growing ants <Emphasis Type="Italic">Mycetarotes</Emphasis> Emery (Attini,Formicidae)

Summary. Fungus-growing ants of the genus Mycetarotes are among the least studied in the tribe Attini. This report documents nest architecture and worker population numbers for 19 nests of M. parallelus and 5 nests of M. acutus, including the first such report for M. acutus. This new information is integrated with the scant biological information reported on Mycetarotes to date. The resulting picture of Mycetarotes life history, as well as the relative ease with which large numbers of nests can be collected and observed in the field, suggest that Mycetarotes (particularly M. parallelus) is an ideal model system for the study of coevolution of lower-attine ants and their cultivated fungi.Received 30 June 2003; revised 17 December 2003; accepted 23 March 2004.     

Temporal variation in the effects of habitat fragmentation on reproduction of the Mediterranean shrub <Emphasis Type="Italic">Colutea hispanica</Emphasis>

Habitat fragmentation poses a major threat to the viability of plant populations. However, the intensity of fragmentation effects may vary among years. We studied two possible effects of habitat fragmentation (patch size and isolation) on the reproduction and proportion of damaged fruits in 24 patches of the self-compatible shrub Colutea hispanica for three consecutive years with different climate conditions. We also studied the effect of fragmentation on the incidence of two main pre-dispersal seed predators, the butterflies Iolana iolas and Lampides boeticus. High between-year variability was found in number of viable seeds per fruit, number of fruits per plant, total number of viable seeds per plant and proportion of damaged fruits. In 2003, small, isolated patches had a higher fruit set and number of fruits per plant. The proportion of damaged fruits was significantly lower in isolated populations in 2003, while it was very high in all patches in 2004 and 2005. High between-year variability was also found in the proportion of fruits per plant with I. iolas eggs. In 2003 isolated patches had a lower proportion of fruits with I. iolas eggs, but no significant effect of patch size and isolation was found in 2004 or 2005. The proportion of fruits with L. boeticus eggs was similar in the three years of study, although it was slightly higher in large, non-isolated patches in 2003. Thus, the effects of fragmentation on plant reproduction cannot be generalized from one single-year survey. In contrast to the generally accepted idea that fragmentation reduces plant reproduction, plant fitness may increase in isolated patches in years with high fruit production and low seed predation.     

Colony pollen reserves affect body size,sperm production and sexual development in males of the stingless bee <Emphasis Type="Italic">Melipona beecheii</Emphasis>

The production of male sexual offspring by social insect colonies is often strongly seasonal or resource-dependent. In stingless bees, males are produced in smaller numbers under conditions of low colony food reserves; whether such males are negatively affected in traits related to reproductive success is not known. We compared body size, sperm production and sexual maturity in Melipona beecheii males reared with experimentally supplemented or reduced pollen reserves, but with otherwise equal numbers of workers and equal quantities of honey reserves. We also studied the same traits in males collected from non-manipulated colonies with pollen reserves intermediate between the supplemented or reduced groups but with more workers and honey reserves. Males reared under experimentally reduced pollen reserves had significantly smaller bodies and lower sperm counts compared to those reared in colonies with experimentally supplemented pollen reserves. There was also a significantly positive relationship between the number of sperm and body size in males across all colony treatments. The maximum number of sperm in seminal vesicles was recorded 2 days later in males from colonies with reduced pollen compared to males from colonies with supplementary pollen. Males from non-manipulated colonies were intermediate in size, sperm count and speed of maturation. Our study documents for the first time the existence of large size variation in males of stingless bees that is related with the amount of pollen reserves in their natal colony. We conclude that a colony’s pollen reserves have a major impact on male body size, sperm production and speed of sexual maturity in this stingless bee, which may be the case in other social insects. Stingless bees are a good model system to study the balance between colony-level selection and individual-level selection on male sexually selected traits such as body size.     

Testosterone treatment can increase circulating carotenoids but does not affect yellow carotenoid‐based plumage colour in blue tits Cyanistes caeruleus

A number of mechanisms are responsible for producing the variation in natural colours, and these need not act in isolation. A recent hypothesis states that carotenoid‐based coloration, in addition to carotenoid availability, is also enhanced by elevated levels of circulating testosterone (T). This has only been tested for carotenoid‐coloured bare parts in birds. We performed an experimental manipulation of T levels and examined the effects on the yellow carotenoid‐based breast plumage in captive yearling blue tits Cyanistes caeruleus, of which half received a diet supplemented with carotenoids. T treatment resulted in elevated plasma T compared to controls and carotenoid supplementation strongly increased plasma carotenoid levels. T treatment resulted in an additional increase in plasma carotenoid levels but only in the carotenoid‐supplemented males. Carotenoid supplementation resulted in more intense breast colour (carotenoid chroma), as expected. However, there was no effect of testosterone on plumage coloration at either dietary carotenoid level. Our results suggest that T can cause an increase in plasma carotenoid concentration, but that this does not necessarily lead to improved carotenoid‐based plumage coloration.     

Consequences of chronic infections with three different avian malaria lineages on reproductive performance of Lesser Kestrels (Falco naumanni)

We studied the consequences of chronic infections by three different lineages of avian malaria, two Plasmodium (RTSR1, LK6) and one Haemoproteus (LK2), on reproductive performance of Lesser Kestrels (Falco naumanni). Malaria infections in male and female parents had no effect on clutch size, hatching success or nesting success. However, when only successful nests were considered, we found that males parasitized by LK6 raised a lower number of fledglings, suggesting that the level of parental effort by males may be limited by this particular lineage of Plasmodium. This effect was not evident in females, probably due to the higher investment of males during the chick rearing period in this species. Overall, we have found that chronic stages of specific malaria lineages have certain negative consequences on host reproductive performance, highlighting the importance of considering genetic differences among malaria parasites to study their consequences on natural bird populations.     

Nutritional supplementation improves ovulation and pregnancy rates in female goats managed under natural grazing conditions and exposed to the male effect

Two experiments were conducted to determine if nutritional supplementation improved ovulation and pregnancy rates in female goats managed under grazing conditions and submitted to the male effect. In Experiment 1, one group of does did not receive nutritional supplementation, while the other group was supplemented daily for 7 days starting at the time when the males were introduced to the females. The ovulation rate at the second male-induced ovulation was greater (P < 0.05) in supplemented (2.0 ± 0.1) than in non-supplemented (1.6 ± 0.1) does. For Experiment 2, female goats were supplemented for 0, 7, 14 or 28 days, starting 9 days following buck introduction. The proportion of does that were pregnant in the group supplemented for 28 days was greater (P < 0.05) than in the non-supplemented group, but did not differ from 14-day and the 7-day supplemented groups. The proportion of pregnant does was greater (P < 0.05) in the group supplemented for 14 days compared to the group supplemented for 7 days and the non-supplemented group. These latter two groups did not differ (P > 0.05). In conclusion, feed supplementation for 7 days, starting at the time when males were introduced increased ovulation rate and feed supplementation for 14 or 28 days starting 9 days after males were introduced improved pregnancy rates in goats managed under grazing conditions and exposed to males.     

Colony size affects collective decision-making in the ant Temnothorax albipennis

Social insects are well-known for their ability to achieve robust collective behaviours even when individuals have limited information. It is often assumed that such behaviours rely on very large group sizes, but many insect colonies start out with only a few workers. Here we investigate the influence of colony size on collective decision-making in the house-hunting of the ant Temnothorax albipennis. In experiments where colony size was manipulated by splitting colonies, we show that worker number has an influence on the speed with which colonies discover new nest sites, but not on the time needed to make a decision (achieve a quorum threshold) or total emigration time. This occurred because split colonies adopted a lower quorum threshold, in fact they adopted the same threshold in proportion to their size as full-size colonies. This indicates that ants may be measuring relative quorum, i.e. population in the new nest relative to that of the old nest, rather than the absolute number. Experimentally reduced colonies also seemed to gain more from experience through repeated emigrations, as they could then reduce nest discovery times to those of larger colonies. In colonies of different sizes collected from the field, total emigration time was also not correlated with colony size. However, quorum threshold was not correlated with colony size, meaning that individuals in larger colonies adopted relatively lower quorum thresholds. Since this is a different result to that from size-manipulated colonies, it strongly suggests that the differences between natural small and large colonies were not caused by worker number alone. Individual ants may have adjusted their behaviour to their colony’s size, or other factors may correlate with colony size in the field. Our study thus shows the importance of experimentally manipulating colony size if the effect of worker number on the emergence of collective behaviour is to be studied. Received 13 December 2005; revised 9 May 2006; accepted 15 May 2006.