Differential time allocation of foraging workers in the subterranean termite

Background

Worker reproduction has an important influence on the social cohesion and efficiency of social insect colonies, but its role in the success of invasive ants has been neglected. We used observations of 233 captive colonies, laboratory experiments, and genetic analyses to investigate the conditions for worker reproduction in the invasive Anoplolepis gracilipes (yellow crazy ant) and its potential cost on interspecific defence. We determined the prevalence of worker production of males and whether it is triggered by queen absence; whether physogastric workers with enlarged abdomens are more likely to be reproductive, how normal workers and physogastric workers compare in their contributions to foraging and defence; and whether worker-produced males and males that could have been queen- or worker-produced differ in their size and heterozygosity.

Results

Sixty-six of our 233 captive colonies produced males, and in 25 of these, some males could only have been produced by workers. Colonies with more workers were more likely to produce males, especially for queenless colonies. The average number of days between the first appearance of eggs and adult males in our colonies was 54.1 ± 10.2 (mean ± SD, n = 20). In our laboratory experiment, queen removal triggered an increase in the proportion of physogastric workers. Physogastric workers were more likely to have yolky oocytes (37–54.9%) than normal workers (2–25.6%), which is an indicator of fertile or trophic egg production. Physogastric workers were less aggressive during interspecific aggression tests and foraged less than normal workers. The head width and wing length of worker-produced males were on average 4.0 and 4.3% greater respectively than those of males of undetermined source. Our microsatellite DNA analyses indicate that 5.5% of worker-produced males and 14.3% of males of undetermined source were heterozygous, which suggests the presence of diploid males and/or genetic mosaics in A. gracilipes.

Conclusions

Our experimental work provides crucial information on worker reproduction in A. gracilipes and its potential cost to colony defence. The ability of A. gracilipes workers to produce males in the absence of queens may also contribute to its success as an invasive species if intranidal mating can take place between virgin queens and worker-produced males.

     

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.     

Body size variation in the sexually dimorphic scaphopod Rhabdus rectius (Carpenter, 1864) (Dentaliida: Rhabdidae)

ABSTRACT

Male-biased sexual size dimorphism typically evolves via sexual selection for larger males that are favoured by choosy females or are more successful in mate competition with other males. Among marine invertebrates that broadcast their gametes into the ocean for fertilisation, this form of sexual size dimorphism is rare because such species lack direct interactions among males or between the sexes. However, the broadcast-spawning tusk shell Rhabdus rectius was recently reported to show strong male-biased sexual size dimorphism. That pattern might imply interesting and undiscovered sexual selection in this species. We found instead that the distribution of body size variation (weight, shell length) was similar between males and females of R. rectius, and mean sizes were not different between the sexes. However, we noted a male-biased sex ratio (～1:1.3) in our large sample of individuals. Many live scaphopods (and several dead shells) showed partial or complete boreholes drilled by predatory gastropods. Boreholes were observed on males and females in similar proportions. We collected scaphopods along with multiple individuals of one likely scaphopod predator, the small moon snail Euspira pallida, and in the lab we observed successful attacks by moon snails on tusk shells.     

Taxonomy, distribution and nesting biology of species of the genus Dolichovespula (Hymenoptera, Vespidae)

The genus Dolichovespula consists provisionally of 18 species. Aerial nest site characteristics have been established for seven species but most of these species show flexibility in their choice of nest sites and two species often nest in shallow subterranean situations. Colony length is usually just over 3 months to approximately 4 months, but at lower latitudes may extend to more than 5 months. Mature colony size is usually approximately 1000 cells with more large cells than small cells. Parasitized colonies have a shorter colony cycle and smaller colonies. During the production of the sexual brood the larva/worker ratio reaches its lowest value of approximately 2.5 larvae per worker. Colonies often have upper mixed combs of small and large cells. Total adult production is usually less than 2000 adults. Colonies of D. arenaria and D. norwegica may specialize in mainly producing males or queens. Variations in mature colony size and production of queens is probably a consequence of the number of workers present, particularly early in the colony cycle.     

Arbuscular mycorrhiza and water and nutrient supply differently impact seedling performance of dry woodland species with different acquisition strategies

Background: We recently demonstrated that dwarf male plants are much more common among mosses than hitherto thought. Dwarf plants, producing functional male sexual organs occur in 10–20% of the moss species worldwide.

Aims: We investigated how an inadvertent omission of dwarf males affects estimates of male sex expression rates and sex ratios.

Methods: We recorded frequencies of specimens with female-expressing plants, male-expressing normal-sized and/or dwarf plants, and sporophytes in herbarium material of five model species from Macaronesia. We analysed how frequency estimates of male occurrence and sex ratios were affected by excluding dwarf males.

Results: Occurrence of dwarf males was positively associated with sporophyte formation. When dwarf males were excluded, male mate availability was missed in 51–61% of specimens (in 74–76% of sporophyte-containing specimens). Specimen-level sex ratio was balanced in material with sporophytes in the three species exhibiting male dwarfism, but strongly female biased if dwarf males were omitted.

Conclusions: We provide the first quantitative evidence that sex-expressing dwarf males are essential for sexual reproduction and sporophyte production in mosses that form such males. Considering its abundance, male dwarfism therefore deserves more attention among plant biologists. Innovative approaches are required to further study the minute male plants.     

Reproductive features and gonad development cycle of the soft bottom-gravel gorgonian Leptogorgia sarmentosa (Esper, 1791) in the NW Mediterranean Sea

Abstract

Leptogorgia sarmentosa (Cnidaria: Octocorallia) is a common soft bottom-gravel gorgonian in the northwestern Mediterranean Sea. This study examined the annual reproductive cycle of a L. sarmentosa. population at a depth of 30 m, and further determined the sex ratio, the intra-colonial gonadal output variability and the age at first reproduction. L. sarmentosa was gonochoric and the population studied was characterized by a sex ratio slightly biased towards female colonies (1 male:1.6 female). Secondary branches had higher polyp fertility (number of gonads per polyp) compared to primary branches, but because of the number of polyps on the former, most of the sexual effort was actually contributed by the primary branches, as in other gorgonian species. Male colonies became fertile above a height of 21 cm, but female colonies had gonads even in the 1–10 cm height class. Spawning occurred between late August and early September, although there was no direct observation of broadcast spawning or surface brooder strategy. The maximum gonad size was 550 μm in males and 500 μm in females, with a mean diameter of 270 μm for spermaries, and 190 μm for eggs. The number of gonads per polyp prior to spawning reached 4–5 spermaries polyp!1 in male colonies, and 3–4 eggs polyp!1 in female colonies. Spawning did not appear to be synchronized with the full moon. L. sarmentosa appeared to have a different reproductive strategy to other Mediterranean gorgonians (living on hard bottom substrata), probably because of its different habitat preferences and biological (morphology, diet, prey capture rates, growth, biochemical levels) features.     

Delayed maturation in the colonial coralGoniastrea aspera (Scleractinia): whole-colony mortality, colony growth and polyp egg production

The present study examines (1) the cost of reproduction on colony growth, and (2) relationships among sexual maturity, whole-colony mortality rate and colony growth rate inGoniastrea aspera free from external influences by macrobenthos. Survival of colonies in permanent plots was followed for two years. Egg production by polyps in colonies collected just before the first spawning of a year was estimated by dissecting the polyps. Growth of the colonies (increase in number of polyps) was followed over one annual reproductive cycle. The cost of egg production on colony growth was apparent through colony ontogeny: (1) immature colonies had a greater annual growth rate than mature colonies, but produced almost no eggs; (2) in mature colonies, growth rate was negatively correlated with NE/PV (number of eggs per polyp volume mm^-3). Annual whole-colony mortality was high in colonies with fewer than11 polyps in initial colony size, while mortality was extremely low once a colony grew beyond this size. This critical size for low whole-colony mortality was much smaller than the colony size (40 polyps) which would attain maturity one year later. Age at maturity was estimated as six years. While survival to maturity may be a selective force for the evolution of delayed maturation, the present data suggest that high colony fecundity, achieved after a long growth period as an immature colony, and an abrupt decrease of colony growth rate after maturation are the crucial forces.     

A simulation model of biological control of social wasps (Vespinae) using mermithid nematodes

Abstract

The nematode (Pheromermis spp.) is a potential biocontrol agent for wasps (Vespula spp.) in countries where invasive populations of wasps cause serious economic, social, and conservation problems. Using a simulation model previously developed for hornets, which belong to a genus with a similar biology to Vespula, we investigated the possibility of using nematodes as a biological control agent. The model wasp colony was exposed to different simulated levels of nematode infection during colony development, and the final number of wasp sexuals produced recorded. The model predicted that early and high levels of wasp infection had the greatest effect on reducing sexual production. However, even colonies with high (80%) levels of infection were still able to produce some sexuals, indicating that wasp colonies are resilient to infections. The model identified several key areas needing further research, including the effects of nematodes on the behaviour and physiology of wasps, of lengthening the infective period, and of increasing infection levels in both the wasps and intermediate transport hosts.     

Mating structure and male production in Vespa analis and Vespa simillima (Hymenoptera: Vespidae)

We estimated queen mating frequency, genetic relatedness between workers and worker reproduction in the hornets Vespa analis and Vespa simillima using microsatellite DNA genotyping. The 20 V. analis colonies studied each contained a queen inseminated by a single male. Of the 15 V. simillima colonies studied, nine had a queen inseminated by a single male, four had a queen inseminated by two males, and two had a queen inseminated by three males. The estimated effective number of matings was 1.33 ± 0.74 (mean ± SD), with 75–85% of the offspring of the six multiply mated queens sired by single males. The values for genetic relatedness between the workers of V. analis and V. simillima were 0.739 ± 0.004 and 0.698 ± 0.013 (mean ± SD), respectively. We conclude that V. analis and V. simillima colonies are genetically monogynous and monandrous. When high relatedness between the workers occurs within colonies, kin selection theory predicts a potential conflict between queens and workers over male production. To determine whether males were derived from queens or workers, males from V. analis and V. simillima colonies were genotyped at four microsatellite loci and the level of ovary activation in workers was determined. None of the 787 V. analis workers and only 15 of 3520 V. simillima workers had developed ovaries. Furthermore, the genotyping identified no worker‐produced males in any colony. The presence of reproductive workers correlated positively with the number of workers within the colony. These results suggest that eusocial colonies with an annual life cycle tend to break down socially when they become large and are close to dying.     

Polymorphic microsatellite loci in the ponerine ant,Hypoponera opacior (Hymenoptera,Formicidae)

The ant genus Hypoponera, with its high diversity of alternative reproductive tactics, is of particular interest in studies on sexual selection. In the species Hypoponera opacior, winged and wingless males and queens co‐occur and molecular markers are essential to study the reproductive success of these sexual tactics. Primers were developed for five polymorphic microsatellite loci isolated from H. opacior. Their variability was tested on 34 colonies from a population in the United States. Nine to 21 alleles per locus were found with observed heterozygosities between 0.1 and 0.7. A significantly positive F_IS value suggests inbreeding in this ant with predominant intracolonial matings.     

Sexual reproduction in the phyllodioicous bryophyte Homalothecium lutescens (Hedw.) H.Rob. in relation to habitat age,growth conditions and genetic variation

Sporophyte production and female fertility were investigated in seventeen calcareous grassland demes of the moss Homalothecium lutescens (Hedw.) H.Rob. on the Baltic island of Öland, with the aim of understanding the relationships between sexual reproduction, habitat age, genetic variation and factors related to growth conditions. The overall proportion of fertile female shoots (with perichaetia) was 35%. Fertility status at the level of individual shoots was positively associated with shoot length and density, while within deme fertility was positively associated with bush cover. There was no association between female fertility and habitat age, genetic diversity (H_S) or allelic richness. Out of 1344 investigated shoots, only two were normal-sized fertile males. Dwarf males were also extremely rare, and found almost exclusively on shoots with sporophytes. Few sporophytes were observed (in the two demes with highest fertility and bush cover). No relationship between genetic variation and the frequency of sporophytes and males was found. The lack of a relationship between sexual reproduction and genetic variation suggests that sexual reproduction may not occur in the same grassland fragments as the recruitment of new clones (from spores or vegetative fragments). The majority of the dry, open grassland habitats, where H. lutescens is typically found in the study area, appear to be suboptimal for both dwarf males and fertilization. Sexual reproduction is more likely to occur in shaded (although grazed) grassland patches, where moisture levels are likely to be higher and the moss colonies are generally more vigorous.     

Behavior and physiology of coexistence in two species of Dascyllus (Pomacentridae,Teleostei)

Synopsis In the Gulf of Aqaba Dascyllus aruanus and D. marginatus, form distinct populations organised into harems of single males and groups of females, living permanently in Stylophora and Acropora corals. Each harem consists of 4 sexual casts: dominating males, adult females, sex-changing fish and juveniles. The sexual performance of each adult fish in such social units is controlled by the presence or absence of a dominating male. In corals occupied by mixed groups of both species, only those females spawn that belong to the same species as the dominating male. Direct observation in the sea shows that such situations are produced by indiscriminative settling mechanisms of groups of juvenile fish. Evidence is also presented, demonstrating that dispersion of coral colonies suitable for occupancy and predation by piscivorous fish can modulate such complicated reproduction.     

Status,conservation and ecology of Pallavicinia lyellii (Hook.) Carruth. in a key region of England

ABSTRACT

Introduction. Pallavicinia lyellii is recognised as being of principal importance for conservation in England and is threatened with extinction in Europe. The aims of this study were to investigate the current status, conservation and ecology of the species in S Hampshire, a key region for the plant.

Methods. All recent and historic locations for Pallavicinia lyellii in S Hampshire were searched and information collected on distribution, abundance and ecology.

Results. Pallavicinia lyellii was re-found at three sites. It is almost certainly extinct at two sites and may have been lost from a third. Whilst few colonies were sterile, most comprised a single sex. Colonies with both male and female plants were confined to Cadnam Bog, which supported 49 (91%) of the colonies found. Overall, female shoots occurred at 41 (76%) colonies and males at 26 (48%), producing a female sex bias (1.6:1). All colonies occur in boggy woodland, mostly on the sides of low mounds of damp, acidic, peaty soil developed, due to poaching by ponies and cattle, at the base of trees, usually I. aquifolium. The community mainly comprises a species-poor mix of acidophilous mosses and liverworts, most frequently Mnium hornum, Pseudotaxiphyllum elegans and Leucobryum juniperoideum.

Conclusions. Cadnam Bog appears to support the largest sexually reproducing population of Pallavicinia lyellii in England. The site is specially protected, actively managed with consideration to P. lyellii and seemingly in favourable condition for the plant. Research is needed to quantify canopy shade favoured by the plant and assess implications for woodland management. Future monitoring should include re-survey of populations every 3?yr.     

Domestication-related changes in sexual performance of Queensland fruit fly

In Sterile Insect Technique (SIT) programs, massive numbers of insects are reared, sterilized, and released in the field to impede reproduction of pest populations. The domestication and rearing processes used to produce insects for SIT programs may have significant evolutionary impacts on life history and reproductive biology. We assessed the effects of domestication on sexual performance of laboratory reared Queensland fruit fly, Bactrocera tryoni, by comparing an old (49 generations) and a young colony (5 generations). We evaluated mating propensity, mating latency, copula duration, sperm transfer, and ability to induce sexual inhibition in mates. Overall, both males and females from the old colony had greater mating propensity than those from the young colony. Copula duration was longer when females were from the old colony. There was no evidence of sexual isolation between the colonies as males and females from the two colonies had similar propensity to mate with flies from either colony. Males from the old colony transferred more sperm regardless of which colony their mate was from. Finally, males from both colonies were similarly able to induce sexual inhibition in their mates and were also similarly able to secure copulations with already-mated females. Positive effects of domestication on sperm transfer, coupled with maintained ability to induce sexual inhibition in mates and to secure copulations with previously mated females, highlights that domestication may have little effect, or even positive effects, on some aspects of sexual performance that may advantage mass-reared B. tryoni in SIT programs.     

Lowered female reproductive effort as an indicator for increased male production and sexuality in Daphnia (Crustacea: Cladocera)

Summary

The relationship between environmental factors, sex ratio, mode of reproduction, and female reproductive effort was examined in the cyclically parthenogenetic Daphnia magna. Female reproductive effort was found to decline before a significant production of males and sexual eggs takes place. Unlike factors such as crowding, poor nourishment, low (sometimes high) temperature and short or long day length, lowered reproductive effort is not a cue but rather an indicator of the selective advantage that would accrue from a changed sex ratio and mode of reproduction. The mechanism of the complex system of environmental sex determination in Daphnia is reviewed, and the adaptivity of ESD is discussed.     

No worker reproduction in the Australian stingless bee Trigona carbonaria Smith (Hymenoptera, Apidae)

Brood of the Australian stingless bee Trigona carbonaria was investigated using microsatellites to determine the origin of males. Genotypes of over 1800 males sampled from ten unrelated colonies were consistent with the hypothesis that the resident queen was the sole mother of the males and that workers, either natal or non-natal (parasitic), did not contribute to the production of males. Dissections of 300 workers from a total of four colonies showed that ovaries are present but not activated. Received 6 June 2007; revised 20 August 2007; accepted 12 September 2007.     

Production of Sexuals and Mating Frequency in the Stingless Bee Tetragonisca angustula (Latreille) (Hymenoptera, Apidae)

Queen, worker, and male production was studied for 1 year in three queenright colonies of Tetragonisca angustula (Latreille). We sampled brood combs monthly and noticed that the number of brood cells and production of individuals were similar among colonies. Although the production of queens did not vary significantly, the frequencies of workers, males, and the number of cells among the combs varied over time. The production of males was highly seasonal, occurring mostly from February to April, coinciding with the period of intense brood cells production, when colonies produced more males and less workers, resulting in a negative correlation. Although the frequency of queens has not varied in time, the seasonal availability of males affected the mating frequency and the time spent since emergence until fertilization of queens. In the T. angustula colonies studied, the population dynamics was highly seasonal and the mating success depended of male production, according to the season.     

Reproductive conflicts in polyandrous and polygynous ant Formica sanguinea

The occurrence of multiple reproductives within an ant colony changes the balance between indirect fitness benefits and reproductive competition. We test whether the number of matings by an ant queen (polyandry) correlates negatively with the number of reproductive queens in the colony (polygyny), whether the patrilines and matrilines differ in their contribution to the sexual and worker progeny and whether there is an overall reproductive skew. For these aims, we genotyped both worker and sexual offspring from colonies of the ant Formica sanguinea in three populations. Most colonies were monogynous, but eight (11%) were polygynous with closely related queens. Most queens in the monogynous colonies (86%) had mated with multiple males. The effective paternity was lower than the actual number of mates, and the paternity skew was significant. Furthermore, in some monogynous colonies, the patrilines were differently represented in the worker pupae and sexual pupae produced at the same time. Likewise, the matrilines in polygynous colonies were differently present in worker pupae and male offspring. The effective number of matings by a queen was significantly lower in polygynous colonies (mean m_e = 1.68) than in monogynous colonies (means 2.06–2.61). The results give support to the hypotheses that polyandry and polygyny are alternative breeding strategies and that reproductive competition can lead to different representation of patrilines and matrilines among the sexual and worker broods.     

Aging and Loss of Fertility in Male and Female Brachionus plicatilis (Rotifera)

Summary

The decline with age of mictic female susceptibility to fertilization and male capacity for fertilization is characterized for the rotifer Brachionus plicatilis. All mictic females were susceptible to fertilization until age 4 hr. Susceptibility then declined non-linearly according to the quadratic equation Y = 140.6—14.3X + 0.36X ². By age 24 hr, sexual females no longer could be fertilized. Only 83% of newborn males were capable of fertilization. This level of fertility held until age 8 hr, then declined linearly. The age when 50% of individuals were no longer fertile was termed the length of fertilizability ₅₀(LF₅₀) and is 7.9 hr (16.7% of lifespan) and 18.8 hr (26.1% of lifespan) for females and males, respectively. Newborn males had an average of 30.1 ± 1.40 motile sperm. Males transferred a mean of 2.3 motile sperm into the pseudocoelom of females on each insemination. Sperm inseminated per copulation closely corresponds to the mean number of resting cysts produced by fertilized females. It is not likely that resting cyst production is limited by sperm availability.     

Reproductive modes in the epixylic hepatic Anastrophyllum hellerianum

The frequency and dynamics of sexual and asexual reproduction were investigated in a dioecious epixylic hepatic, Anastrophyllum hellerianum, which has declined in recent decades in Finland as a consequence of forestry practices. In our investigation asexual reproduction by gemmae was the dominant mode of reproduction and specialised gemmiparous shoots were present in all colonies studied. The proportions of dead shoots were considerably higher among sex-expressing than among non-sexexpressing shoots. Our results suggest that lower reproductive investment is required for asexual than for sexual reproduction. For instance, no trade-off is detected between asexual reproduction and survival of the gemmiparous shoots in A. hellerianum. Sexual reproduction occurred only in 12% of the colonies and it was promoted by the following factors: medium shoot density, high proportion of sex-expressing shoots, an even sex ratio and very short distances between individuals representing opposite sexes. The ratio of dead males to dead females was significantly female-biased, which suggests higher mortality among female shoots. At the level of individual shoots, more spores than gemmae were produced. However, as a consequence of the low frequency of sporophyte-bearing shoots, gemma production highly exceeded spore production at the colony level. Furthermore, cultivation tests of the propagules showed that gemmae germinate faster than spores.