Population Biology of Maculinea rebeli (Lepidoptera: Lycaenidae) on the Chalk Grasslands of Eastern Westphalia (Germany) and Implications for Conservation

The population biology of Maculinea rebeli Hirschke, 1904, on dry grassland biotypes in different successional stages was investigated in the summers of 1990 and 1991 in the district of Höxter (North Rhine–Westphalia, Germany). The population characteristics of M. rebeli were studied by mark-recapture and eggshell counts on the larval host plant Gentiana cruciata. Pitfall traps, ant baits, excavating and counting of ant nests were used to examine the Formicidae community and the nest density of Myrmica sabuleti. In 1990, the total population of M. rebeli was estimated as 528 individuals, with subpopulations varying from 2 to 273 individuals. Populations existed in areas differing in size and vegetation structure. However, the limiting factor for population size was the density of the host ant population. The differences in population size did not depend on the size of the habitat or the density of the larval food plant. The butterflies can switch their host. In the absence of M. schenki, the colonies of M. sabuleti can maintain M. rebeli populations. Adult males and females have a maximum life expectancy of 13 days. The females prefer to oviposit on G. cruciata plants of 10–30cm height. Only plants which stuck out of the surrounding vegetation were oviposited on. The average number of eggs deposited per female was 100–150. Low density populations are probably most threatened. Therefore, future protective measures should work towards further maintaining and optimizing the habitats of the small populations, as well as creating the prerequisites so that dense stable populations of M. rebeli can develop.     

Social parasitism in fire ants (Solenopsis spp.): a potential mechanism for interspecies transfer of Wolbachia

One possible mechanism for interspecific transfer of Wolbachia is through the intimate contact between parasites and their hosts. We surveyed 10 species of fly parasitoids (Pseudacteon spp.) and one inquiline social parasite, Solenopsis daguerrei, for the presence and sequence identity (wsp gene) of Wolbachia. Two Wolbachia variants infecting S. daguerrei were identical to known variants infecting the two common ant host species, Solenopsis invicta and Solenopsis richteri, suggesting possible transfers of Wolbachia between this parasite and their hosts have occurred. Our data also revealed an unexpectedly high diversity of Wolbachia variants within S. daguerrei: up to eight variants were found within each individual, which, to our knowledge, is the highest reported number of Wolbachia variants infecting a single individual of any host species.     

Species delimitation in native South American fire ants

The taxonomy of fire ants has been plagued by difficulties in recognizing species on the basis of morphological characters. We surveyed allozyme markers and sequences of the mtDNA COI gene in several closely related nominal species from two areas of sympatry in the native ranges to learn whether the morphology-based delimitation of these species is supported by genetic data. We found that Solenopsis invicta and Solenopsis richteri, pest species whose distinctiveness has been debated, appear to be fully reproductively isolated at both study sites. This isolation contrasts with the extensive hybridization occurring between them in the USA, where both have been introduced. We also found strong genetic differentiation consistent with barriers to gene flow between Solenopsis quinquecuspis and the other two species. However, several lines of evidence suggest that nuclear and mitochondrial genes of S. invicta and S. richteri are introgressing into S. quinquecuspis. The latter apparently is a recently derived member of the clade that includes all three species, suggesting that there has been insufficient time for its full development of intrinsic isolating mechanisms. Finally, our discovery of genetically distinct populations within both S. invicta and S. richteri suggests the presence of previously unrecognized (cryptic) species. Their existence, together with the difficulties in developing diagnostic morphological characters for described species, imply that the group is actively radiating species and that morphological divergence generally does not keep pace with the development of reproductive isolation and neutral genetic divergence in this process.     

Catching ants with honey: an experimental test of distraction and satiation as alternative modes of escape from flower-damaging ants

According to the distraction hypothesis, extrafloral nectaries (EFN) evolved under selection to entice ants away from floral nectaries, reducing ant-mediated damage to flowers and/or interference with pollinators. Predator-satiation, through production of nectar in either surplus flowers or EFN, provides an alternative mechanism for reducing the impact of ants as flower visitors. I tested these two hypotheses by experimentally adding EFN to flowering plants of the alpine wildflower, Polemonium viscosum, and by surveying the relationship between ant visitation and nectary number in nature. Plants of P. viscosum lack EFN and experience flower damage by ants of Formica neorufibarbus gelida. Ant behavior was compared on plants with five flowers and three experimental EFN and on controls with equal floral display, but no EFN. Addition of EFN increased flower visitation by ants. The effect of EFN on flower visitation did not depend on proximity of EFN to flowers or attractiveness of EFN to ants. Findings suggest that ants perceived patch quality on a whole plant basis, rather than responding to EFN and flowers as distinct nectar patches. Ant visitation did not keep pace with nectary number in nature. The relationship between ant visitation and nectary number per plant was weak and shallow as predicted under satiation. Ant foraging choices on experimental inflorescences showed that ants bypass flowers avoided by earlier ants, enhancing probability of escape via satiation. Results do not support the idea that EFN evolve to reduce flower visitation by ants, but show instead that nectar in surplus flowers can satiate ants and reduce their negative impacts on flower function and integrity.     

The coevolutionary dynamics of obligate ant social parasite systems--between prudence and antagonism

In this synthesis we apply coevolutionary models to the interactions between socially parasitic ants and their hosts. Obligate social parasite systems are ideal models for coevolution, because the close phylogenetic relationship between these parasites and their hosts results in similar evolutionary potentials, thus making mutual adaptations in a stepwise fashion especially likely to occur. The evolutionary dynamics of host-parasite interactions are influenced by a number of parameters, for example the parasite's transmission mode and rate, the genetic structure of host and parasite populations, the antagonists' migration rates, and the degree of mutual specialisation. For the three types of obligate ant social parasites, queen-tolerant and queen-intolerant inquilines and slavemakers, several of these parameters, and thus the evolutionary trajectory, are likely to differ. Because of the fundamental differences in lifestyle between these social parasite systems, coevolution should further select for different traits in the parasites and their hosts. Queen-tolerant inquilines are true parasites that exert a low selection pressure on their host, because of their rarity and the fact that they do not conduct slave raids to replenish their labour force. Due to their high degree of specialisation and the potential for vertical transmission, coevolutionary theory would predict interactions between these workerless parasites and their hosts to become even more benign over time. Queen-intolerant inquilines that kill the host queen during colony take-over are best described as parasitoids, and their reproductive success is limited by the existing worker force of the invaded host nest. These parasites should therefore evolve strategies to best exploit this fixed resource. Slavemaking ants, by contrast, act as parasites only during colony foundation, while their frequent slave raids follow a predator prey dynamic. They often exploit a number of host species at a given site, and theory predicts that their associations are best described in terms of a highly antagonistic coevolutionary arms race.     

Taxonomic level, trophic biology and the regulation of local abundance

• 1 Taxocenes — monophyletic ecological assemblages — are a key focus of macroecology. Abundance (individuals per area) is a basic property of taxocenes but has received less attention than diversity, although the two are probably related. Abundance reflects a taxocene’s ability to harvest and sequester available energy and divide it among individuals. This paper explores how two properties of all taxocenes — trophic makeup and taxonomic level (e.g. genus, tribe, subfamily, family … ) — may contribute to patterns of local abundance at geographical scales.
• 2 Forty‐nine ground ant taxocenes, in habitats ranging from New World deserts to rain forests, were surveyed along a three‐orders of magnitude productivity gradient using transects of 30 1‐m² quadrats at each site. Abundance — the number of nests per transect — varied over two orders of magnitude.
• 3 Over 80% of the genera collected were omnivores. However, herbivore, omnivore, and predator taxa were added to ant taxocenes in roughly 1 order of magnitude steps up the productivity gradient. Specialist detritivores were added last.
• 4 Net primary productivity and mean monthly temperature both consistently entered regression models predicting abundance. However, while productivity was the dominant predictor of abundance for higher taxa (families, subfamilies), temperature was the dominant predictor of abundance for lower taxa (tribes, genera). The answer to the question ‘What regulates the abundance of a taxocene?’ is thus sensitive to the taxonomic level of analysis.
• 5 These data support the following scenario. Lower taxa are abiotic specialists given the insufficient number of genomes and generations required for the exploration of the entire abiotic envelope. Higher taxa, in contrast, consist of suites of abiotic specialists arrayed along the entire productivity gradient, with access to productivity everywhere the taxon occurs. If this scenario is true, individual species may respond to global changes in temperature; the higher taxa they belong to may most respond to global changes in productivity.

     

Mating behaviour of Rhytidoponera sp. 12 ants inferred from microsatellite analysis

In the queenless ponerine ant Rhytidoponera sp. 12, all workers have a spermatheca and functional ovaries and are potentially able to mate and reproduce. Within a colony gamergates may either be full sisters to each other (Type 1 colony), or they may not be full sisters but still be significantly related to each other (Type 2 colony) due to daughter gamergates reproducing in their natal colonies after mating. Despite many studies the mating behaviour of R. sp. 12 has been poorly understood. In this study, we used microsatellite markers to investigate intracolony relatednesses of male mates to the gamergates (bmq) and between male mates (bmm), and mating frequencies and mating patterns, using gamergate DNA and sperm DNA isolated from the spermathecae of gamergates from five colonies. Average bmm and bmq estimates for all five colonies studied were not significantly different from zero, suggesting that on average, within colonies, mating males were unrelated both to each other and to the gamergates. A low frequency (3%) of multiple mating by gamergates was detected. Multiple mating by individual males with sister gamergates within Type 1 colonies was also detected at 3% and could give rise to half-sister nestmate workers. Polygamy in R. sp. 12 might indicate local female-biased operational sex ratios despite the expectation of overall male biases. Our results concur with previous reports that gamergates mate within the colony or nearby, but indicate more diversity in mating patterns than previously indicated for this polygynous ponerine ant species.     

Hunting strategy of a generalist ant species proposed as a biological control agent against termites

We studied the hunting behaviour of Myrmicaria opaciventris (Hymenoptera: Formicidae) in order to evaluate if it can be used as a biological control agent against the termites that damage sugarcane plantations. Hunting workers foraged in groups and recruited nestmates at short-range when they encountered large termite soldiers or groups of small termite workers. Differences in prey capture concerned the: (1) means of detection (from a distance or by contact); (2) termite body part seized (small termites seized by the body; large termites by an appendage); (3) percentages of prey abandoned; and (4) use of venom. The sting of the workers is spatulated implying a topical application of the venom on the prey. Large termites were stretched by several workers whose adherence to the substrate is facilitated by well-developed arolia and claws on the legs while others spread venom on the body and carved it up. An adaptation to termite capture was noted with a distribution of tasks between the workers which subdued prey, and those which transported it. In the former case, the workers easily eliminated termite soldiers, successively attacked several termite workers and even captured new individuals while holding the first ones captured between their mandibles before retrieving them all at once. The remaining individuals were retrieved by the transporting workers. Given this particularly effective predatory strategy, we concluded that, under certain conditions, M. opaciventris can be used as a biological control agent against termites.     

Brood sex ratio determination by flow cytometry in ants

Sex ratio variations during brood development have important implications for the study of sex allocation in haplodiploid insects. So far, few studies have addressed this question because of the difficulty to determine the sex of the brood. We used flow cytometry to differentiate haploid males from diploid females in the ant Linepithema humile. Our data show that flow cytometry can be used successfully to distinguish between male and female brood on the basis of their DNA content, from the very first larval stage. Moreover, we show that flow cytometry allows sex brood determination in other ant species, as well as in nonsocial Hymenoptera.     

Parallel evolution of thermophilia: daily and seasonal foraging patterns of heat‐adapted desert ants: Cataglyphis and Ocymyrmex species*

In the deserts of northern and southern Africa, respectively, ants of the genera Cataglyphisf oerster (Formicinae) and Ocymyrmexe mery (Myrmicinae) occupy the same ecological niche, which comprises that of a strictly diurnal thermophilic scavenger. Their daily foraging activities exhibit a bimodal pattern in summer and unimodality or complete inactivity in winter. The present study investigates whether these overall patterns are a result of endogenous annual activity rhythms of the colony or are triggered directly by the prevailing ambient temperatures. By exploiting various seasonal temperature regimes and, in particular, by creating near‐nest winter conditions experimentally in summer, it is shown that the latter hypothesis is generally true. However, there are daily and annual variations in the temperature set points at which foraging activities start and finish. These temperatures are lower in the winter than in the summer months and, in summer, they are lower in the morning than in the afternoon. The level of foraging activity in the afternoon reaches maximum values at surface temperatures of 60–63 °C. This means that, in summer months, these thermophilic ants concentrate their foraging activities into a period of almost lethal temperature regimes, during which they have to devote a substantial portion of their time outside the nest to respite (i.e. cooling‐off) behaviour.