Respiratory patterns in nurses of the red wood ant Formica polyctena (Hymenoptera, Formicidae)

Summary Cycles of discontinuous gas exchange (DGCs) and abdominal ventilatory movements were studied in nurses of red wood ant, Formica polyctena, using an electrolytic respirometer and an infra-red (IR) gas analyser or flow-through respirometry. Both respirometry systems were combined with an IR actographic device based on IR-emitting and IR-sensor diodes. After recovering from handling and apparatus stress, lasting 1–3 h, completely motionless intact ants displayed regular DGC. After decapitation the ants displayed DGC whose frequency was somewhat lower than that of the intact individuals (7.17 ± 0.79 mHz and 10.43 ± 01.12 mHz, respectively). In headless ants, there occurred continuous slow movements of legs. Bursts of carbon dioxide in the intact and in the headless ants always coincided with a bout of telescoping movements (contractions) of abdominal segments, which was interpreted as active ventilation. During interburst periods, the headless ants exhibited telescoping movements characterised by rapid protraction, lasting 0.07–0.09 s, followed by a slow retraction of segments, suggesting passive ventilation. The intact ants were very sensitive to the flowing air and tended to be continuously active during flow-through respirometry. The decapitated ants, on the contrary, were insensitive to the air current.Received 26 March 2003; revised 30 July 2003; accepted 6 August 2003.     

Feinstruktur des Komplexauges der Roten Waldameise,Formica polyctena (Hymenoptera,Formicidae)

Zusammenfassung Die Analyse der Feinstruktur des Komplexauges der Ameise Formica polyctena ergab, daß der dioptrische Apparat der insgesamt 750 Ommatidien aus einer lamellierten Cornealinse und einem euconen Kristallkegel besteht. Zwei Hauptpigmentzellen umgeben schalenförmig den Kristallkegel, 8 Nebenpigmentzellen schirmen das Ommatidium in seiner ganzen Länge von der Cornea bis zur Basalmembran ab. Jedes Ommatidium besteht in seinem distalen Teil aus 8 Retinulazellen, 2 gegenüberliegenden schmalen und je 3 gegenüberliegenden großen Sehzellen. Weiter proximal tritt eine 9. Retinulazelle hinzu. Die Mikrovillisäume der Sehzellen verschmelzen zu einem zentralen Rhabdom. Im distalen Teil sind die Mikrovilli in 3 Richtungen angeordnet. Es wird im besonderen die Orientierung der Mikrovilli zur Augenlängsachse und zur Ommatidien-Symmetrieachse untersucht. Auch die 9. Sehzelle bildet Mikrovilli. Das Rhabdom endet an 4 basalen Pigmentzellen.Auf den Mikrovillisaum folgt im Querschnitt des dunkel adaptierten Ommatidiums ein Kranz von großen intrazellulären Vakuolen. Die anschließende cytoplasmatische Zone der Retinulazellen enthält viele Pigmentgranula und Mitochondrien; multivesikuläre und multilamelläre Körper sowie Golgiapparate treten nur selten auf. Die funktionelle Bedeutung des Ommatidienaufbaues und die Verteilung der Organellen bei Dunkeladaptation werden diskutiert.

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The fine structure of the complex eye of the ant Formica polyctena (Hymenoptera, Formicidae)

Summary The fine structure of the compound eye of the ant Formica polyctena was investigated. The eye consists of a total of 750 ommatidia, each containing a dioptric apparatus of a lamellated cornea lens, a eucone-type crystalline cone, and 8 long pigment cells which surround the ommatidium for its total length, i.e. from the cornea to the basal membrane. Each ommatidium has in its distal portion 8 retinula cells—6 large plus 2 small ones. The retinula cells are arranged in such a way that 3 pairs of large cells, and the one pair of the small cells lie opposite each other. Further proximally, a 9th retinula cell is encountered. The fused, centrally located rhabdom is built up by the microvilli of all nine retinula cells. The rhabdom ends at 4 basal pigment cells. In dark adapted ommatidia, a ring of large intracellular vacuoles is to be seen immediately peripherally to the rhabdom. The outer, cytoplasmic zone of the retinula cells contains many pigment granules and mitochondria; multivesicular bodies, onion bodies and Golgi apparatus are of relatively rare occurrence. The functional significance of the ommatidial structure and the arrangement of the cell organelles in the dark adapted condition are discussed.

Für technische Hilfe danke ich Frau Langer und Frl. Müller, Herrn Prof. H. Markl für die Durchsicht des Manuskripts.     

Immunocytochemical localization of a vacuolar-type ATPase in Malpighian tubules of the ant Formica polyctena

The presence of a vacuolar-type ATPase in Malpighian tubules of the ant Formica polyctena was investigated immunocytochemically, using antibodies to vacuolar ATPases of Manduca sexta midgut and bovine kidney. Specific labelling was observed at the brush border of the epithelium extending along the entire length of the tubules. These findings agree with the current view that a vacuolar ATPase is situated at the apical membrane of Malpighian tubule cells and other insect epithelial cells, being the energizing element of an electrogenic potassium pump. When antibodies were tested on tubules in different secretion conditions prior to fixation, no differences were observed in the distribution of the vacuolar ATPase.This work was supported by grants from the European Community (SCI-CT90-0480), from the Ministerio de Educación y Ciencia DGICYT, Spain (CE 91-0002), and from the Deutsche Forschungsgemeinschaft (Wi 698-3).     

Effects of seed aggregation on the removal rates of elaiosome-bearing Chelidonium majus and Viola odourata seeds carried by Formica polyctena ants

In a field dominated by Formica polyctena Foerst. ants, we examined the effect of seed aggregation on the seed-removal rates of two plant species: a large-seeded obligate myrmecochore Viola odorata L. and a small-seeded diplochore Chelidonium majus L., which was autochorous as well as myrmecochorous. The effect was statistically non-significant in V. odorata but significant in C. majus, with more closely aggregated seeds having higher removal rates. The large seeds of the obligate myrmecochore were more quickly discovered and repeatedly removed by ant workers than were the small seeds of the diplochore.     

Impact of vegetation removal on the temperature and moisture content of red wood ant nests

We studied the influence of vegetation growing on red wood ant, Formica polyctena, hills and its removal, by cutting, on the temperature and moisture content of the nests. Vegetation was removed from half of nests (n = 10) in June and August. Generally, ant hills were small (0.1–1.1 m³) and their daily temperature fluctuations and moisture (16–38%) were low. The centre temperature of ant hills was positively correlated with ambient air temperature. Daily temperatures peaked at 13:00 and slowly decreased until 09:00 then increased again until 13:00. Nest moisture content was not related to nest volume. All the ant hills were covered with Reed Grass (Calamagrostis arundinacea) that grows through nests and reaches about 185–1085 g/dcm² of dry matter. Dry vegetation matter (g/dcm² of anthill surface) was the same on cut and uncut nests as well because all ant hills were under long-term management (vegetation removal) for at least 10 years. Although vegetation removal did not result in an immediate increase in the temperature of denuded ant hills removal did results in higher daily and night temperatures during the July-September period. The moisture content of nests with undisturbed and intact vegetation was the same. The dry matter content of vegetation strongly affected their moisture content. However, temperature at the inside centre of nests decreased as vegetation dry matter increased, i.e. there was a statistically negative correlation between temperature and moisture. The results support removing grass as a suitable management tool to facilitate the survival of wood ants. Received 30 December 2007; revised 22 April 2008; accepted 10 June 2008.     

Nest structure and colony cycle of the Allegheny mound ant,Formica exsectoides Forel (Hymenoptera: Formicidae)

Summary The nest structure and colony cycle of a population of Allegheny mound ants,Formica exsectoides, were examined in central Michigan. The dispersion pattern of mounds was random. Nest structure and presence of brood were primarily determined by excavation of twenty-three nests over three intervals from June through September. Additional excavations of five nests in 1990 and ten nests in 1991 provided further details on nest structure and colony cycle. Most galleries occurred within the mound and upper 30 cm of soil, but some activity reached depths of 100 to 270 cm. Depth of nests showed little correlation with external measurements of height and diameter. Immature stages were recovered from two strata: the upper 20 cm of nest and mound and the lowest nest depths. Alate sexual forms were found in or near the mound in July, and numerous dealate queens were collected in September from peripheral galleries near the soil surface.     

Effect of an invasive ground cover plant on the abundance and diversity of a forest floor spider assemblage

Ground dwelling spiders are important predators in the detrital food web, which plays important roles in nutrient cycling and energy flow in forest ecosystems. The cursorial spider assemblage in a Beech-Maple forest in southwestern Michigan at sites where and invasive plant, Vinca minor, has invaded was compared to a native site within the same forest and to the forest prior to invasion by the plant. Pitfall traps were used to sample cursorial spiders over the course of a summer. Vinca minor substantially altered the forest floor spider assemblage. The invasive plant reduced the total activity-abundance of spiders by nearly 49% and depressed species diversity and evenness; in contrast, species richness was not affected. We found that V. minor changed the guild and family structure with wolf spiders being common at sites where the plant had invaded. Vinca minor reduced the abundance of vagrant web building and crab spiders. Similarity indices revealed that the spider communities between the two sites were quite dissimilar (Bray-Curtis = 0.506; Jaccard’s = 0.520). Importantly, comparison to a study conducted in the same forest 28 years earlier showed that the cursorial spider assemblage in the forest prior to Vinca invasion was very different than it was after Vinca invaded but was similar to the current native site in species and guild composition. We conclude that invasion by Vinca has caused the striking changes we observed in community organization of this important group of forest floor predators. We suggest that changes in the physical structure of the litter/soil microhabitat with the invasion of V. minor are likely the cause of the substantial impacts of the plant on the spider assemblage.     

Effect of past and present behavioural specialization on brain levels of biogenic amines in workers of the red wood ant Formica polyctena

Social insect workers usually participate first in intranidal tasks (i.e. act as nurses within the nest) and then switch to extranidal tasks and become foragers. However, foragers sometimes switch back again to brood care and become reverted nurses. Behavioural and physiological correlates of the transition nurse–forager (behavioural maturation) and forager–reverted nurse (behavioural reversion) are relatively well known in the honeybee, although they are less explored in ants. To understand better the role of biogenic amines in ant behavioural maturation and behavioural reversion, the levels of octopamine (OA), dopamine (DA) and serotonin (5‐HT) are examined in the brains of nurses, foragers and reverted nurses of the red wood ant Formica polyctena Först. (Hymenoptera: Formicidae). Brain OA levels and the ratios OA : DA and OA : 5‐HT are higher in nurses than in foragers and reverted nurses. Reverted nurses and foragers do not differ significantly with respect to brain biogenic amine levels and amine ratios. Biogenic amine levels in brains of workers of F. polyctena are thus maturation‐related rather than task‐related. This is one of the first studies of neurochemical correlates of ant behavioural maturation and the first attempt to identify neurobiological correlates of ant behavioural reversion. The data obtained provide further evidence that neurobiological processes underlying honeybee and ant behavioural maturation and behavioural reversion reveal important differences.     

The soil fauna of a beech forest on limestone: trophic structure and energy budget

Summary The soil fauna of a mull beech forest on lime-stone in southern Lower Saxony (West Germany) was sampled quantitatively. Biomass estimates, trophic characteristics, and measurement and calculation of the energetic parameters of the constituent animal populations were used to construct an energy budget of the total heterotrophic subsystem of the forest. Mean annual zoomass amounted to about 15 g d wt m^–2; earthworms (about 10 g d wt m^–2) and other groups of the macrofauna were dominant. Protozoa constituted about 1.5 g d wt m^–2. Relative distribution of zoomass among the trophic categories was 50% macrosaprophages, 30% microsaprophages, 12% microphytophages, and 4% zoophages. Total annual consumption rate of the saprophagous and microphytophagous soil fauna (6328 and 4096 kJ m^–2 yr^–1, respectively) was of the same order of magnitude as annual litter fall (canopy leaves 6124 kJ m^–2 yr^–1, flowers and fruits 944 kJ m^–2 yr^–1, herbs 1839 kJ m^–2 yr^–1, fine woody material 870 kJ m^–2 yr^–1, tree roots 3404 kJ m^–2 yr^–1, without coarse woody litter). Primary decomposers (macrosaprophages) were the key group for litter comminution and translocation onto and into the soil, thus contributing to the high decomposition rate (k=0.8) for leaf litter. Consumption rates of the other trophic groups were (values as kJ m^–2 yr^–1): bacteriophages 2954, micromycophages 416, zoophages 153. Grazing pressure of macrophytophages (including rhizophages) was low. Faeces input from the canopy layer was not significant. Grazing pressure on soil microflora almost equalled microbial biomass; hence, a large fraction of microbial production is channelled into the animal component. Predator pressure on soil animals is high, as a comparison between consumption rates by zoophages and production by potential prey — mainly microsaprophages, microphytophages and zoophages — demonstrated. Soil animals contributed only about 11% to heterotrophic respiration. However, there is evidence that animals are important driving variables for matter and energy transfer: key processes are the transformation of dead organic material and grazing on the microflora. It is hypothesized that the soil macrosaprophages are donor-limited.     

Responses of Two Potential Host Species (Formica gnava and Formica occulta) to Pupae of the Obligatory Slave-Making Ant, Polyergus breviceps (Hymenoptera: Formicidae)

Queens of the slave-making ant, Polyergus breviceps, take over nests of adult Formica workers when establishing new colonies. Although naïve to slave-maker brood, the usurped Formica rear Polyergus offspring and nests containing both host and parasite species forms. Host worker acceptance of parasite brood has been attributed to the similarity of brood tending signals between these closely related taxa and/or the presence of an attractive pheromone in the slave-maker brood. By presenting single-species groups of Formica occulta and Formica gnava (two host species of P. breviceps) with a choice of Formica pupae of both species or with a choice of P. breviceps pupae from both types of mixed-species nests, it seems that neither close phylogenetic relatedness nor an attractive brood odor alone can account for the propensity of host workers to adopt slave-maker pupae. Significantly greater numbers of P. breviceps pupae were adopted by enslaved workers than by free-living workers, and within the enslaved groups and the free-living F. gnava group, greater numbers of P. breviceps pupae were adopted if they were from nests where the host species was conspecific to workers used in tests. When presented with F. gnava and F. occulta pupae, Formica workers adopted conspecific pupae almost exclusively and ignored or consumed pupae of the other host species. Taken together, these results imply that P. breviceps pupae have both a species-specific odor and a general brood-tending pheromone, upon which a host odor may be imposed. The disparate requirements of immatures at different stages of development for cue specificity or generality in maintaining nest exclusivity and maximizing inclusive fitness are discussed.     

Investigation on catechin as a beech wood decay biomarker

A high-performance liquid chromatography (HPLC) method based on the evolution of wood extractives was developed to follow the first stages of fungal degradation of beech wood exposed to Coriolus versicolor. The nature and the quantity of the extracts initially present in wood depended on the extraction conditions and also on the wood-drying conditions (time and temperature). The most interesting fraction was soxhlet extracted with acetone at 56 °C for 6 h. The best conditions to avoid extractives degradation consisted of a moderate drying at 55 °C for 48 h allowing identification of catechin as potential tracer. After 2 weeks of wood blocks exposure to C. versicolor, analysis of their acetonic extractives showed that catechin signal initially detected in beech wood, had totally disappeared. Treatment of wood with an appropriate fungicide such as propiconazole before exposure to C. versicolor, prevents the catechin amount from any variation. The comparison of these results with the classical weight loss (WL) measurements obtained after long-time experiments on treated and untreated wood blocks shows that the catechin amount evolution, monitored during 2 weeks, correlates with the wood resistance evaluated during 16 weeks, allowing the use of this flavonoid as a valuable biomarker of wood decay.     

Myrmecotopical transformation of grass stand structure on a plot of fragmented forest

The authors have ecologically and cenotically characterized the grass stand around anthills of Formica polyctena Foerst. on a plot of fragmented forest and have determined species composition of myrmecotopic grass assembly. The study revealed prairification and synanthropization of the grass stand as well as similarity of cenotic and morphologic changes in grass covers around anthills of F. aquilonia Yarr. and F. polyctena.     

Limited phylogeographical structure across Eurasia in two red wood ant species Formica pratensis and F. lugubris (Hymenoptera, Formicidae)

The phylogeography and demographic history of two closely related species of the red wood ant (Formica pratensis and F. lugubris) were examined across Eurasia. The phylogeny based on a 1.5-kilobase mitochondrial DNA fragment, including the cytochrome b gene and part of the ND6 gene, showed one phylogeographical division in F. pratensis. This division (0.7% of nucleotide divergence) suggests postglacial colonization of western Europe and of a wide area ranging from Sweden on the west to Lake Baikal on the east from separate forest refugia. In two localities, mitochondrial DNA has been transferred from F. lugubris to F. pratensis and all the individuals of F. pratensis sampled from the Pyrenees had haplotypes clustering in the lugubris clade. No phylogeographical divisions were detected in F. lugubris. Comparison of species-wide phylogeography between the two sympatrically distributed species of ant demonstrates a difference in phylogeographical structure that implies different vicariant histories. However, over most of the species' distribution ranges, similar signs of demographic expansion predating the last glaciation and the lack of phylogeographical structure were found in both the eastern phylogroup of F. pratensis and F. lugubris. This finding is highly consistent with the results reported for all other boreal forest animal species studied to date in Eurasia. Contraction of the distribution range of each species to a single refugial area at different times during the late Pleistocene and a subsequent population expansion seem to be an explanation for the lack of phylogeographical structure across most of Eurasia in species that are ecologically associated with the boreal forest.     

Conservation genetics of the wood ant, Formica lugubris, in a fragmented landscape

Various intrinsic factors connected to the special features of sociality influence the persistence of social insect populations, including low effective population size, reduced amount of genetic variation easily leading to inbreeding depression, and spatially structured populations. In this work, we studied an isolated, small and fragmented population system of the red wood ant Formica lugubris, and evaluated the impact of social and genetic population structure on the persistence and conservation of the populations. The effective population size was large in our study population because all nests were polygynous. As a result, and despite the apparent isolation, the amount of nuclear genetic variability was similar to that in a nonisolated population system. Lack of inbreeding, as well as a high level of variability, indirectly suggests that this population does not suffer from inbreeding depression. The spatial distribution of genetic variation between local populations suggests intensive, but strongly male-biased, nuclear gene flow. Thus, the persistence of this population system does not seem to be threatened by any immediate social or genetic factor, but colonization of new habitat patches may be difficult because of restricted female dispersal.     

On the spider genus Arboricaria with the description of a new species (Araneae,Gnaphosidae)

The spider genus Arboricaria Bosmans, 2000 is redefined and an updated diagnosis given. The differences between Arboricaria and Micaria Westring, 1851 are discussed in detail. A key to all five species of the genus is provided. One new species, Arboricaria zonsteini sp. n. (♂♀), is described based on specimens from Kyrgyzstan and Azerbaijan. One new synonym is proposed: Arboricaria koeni Bosmans in Bosmans & Blick, 2000, syn. n. is assigned to Arboricaria sociabilis Kulczyński in Chyzer & Kulczyński, 1897. Data on the distribution of Arboricaria in Russia and adjacent countries are presented with references to the papers on local spider faunas.     

The effect of red wood ant abundance on feeding damage by the pine weevil Hylobius abietis

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Influence of the wood ant, Formica polyctena, on soil nutrient and the spruce tree growth

The effect of Formica polyctena ant nests on the distribution of soil nutrients, soil pH and the growth of Norway spruce trees was studied in the southern part of the Czech Republic. Soil nutrient content (exchangeable P, N, K and pH) and growth of mature spruce trees were measured at four distances from the nearest ant hill (0–1, 3–5, 10–50 and >200 m). Trees at all distances were visited by ants, except for those >200 m from the nearest nest. Soil pH and of P, K and NO₃ concentrations were higher near ant nests (<1 m), and pH and K at distances of 3–5 m, when compared with distances of 10–50 and >200 m from the nests, where no significant differences in these variables were detected. In contrast, tree ring analyses (1974–2004) showed that trees >200 m from the ant nests grew significantly faster than trees at other distances, followed by trees within 1 m of the nests. No growth differences were found between the growth of trees at 3–5 and 10–50 m from ant nests. We postulate that nutrient and carbohydrate removal of honeydew collected by ant‐tended aphids are slowing growth of tree. However, trees may partly compensate for this depletion by having access to a larger supply of soil nutrients near ant nests.     

Raiding and Foraging Behavior of the Blood-Red Ant,Formica sanguinea Latr. (Hymenoptera,Formicidae)

In this paper we report the results of a detailed study on the behavioral ecology of slave raiding and foraging activity in the European blood-red ant, Formica sanguinea Latr. The field study was conducted over an unbroken period of 78 days, during which the activity of two dulotic colonies of this facultative slave-maker was observed for 10 h each day. It was possible to observe 26 raids distributed over 23 days, among which 18 were followed by the sacking of nests belonging to the species F. cunicularia, F. fusca, and Lasius emarginatus, whereas 8 failed. Simple, continuous, and simultaneous raids occurred. We recorded the timing, frequency, distance, and direction of slave raids, including the number of participants and the type of booty. Particular attention was devoted to the scouting behavior and raiding organization. Moreover, every day, we observed foraging and predatory behavior, during which adult insects (mainly ants), seeds, and berries were retrieved to the dulotic colonies. On the basis of our observations F. sanguinea seems to be a very efficient slave-maker and predatory species of the Raptiformica subgenus. Moreover, its dulotic behavior may be regarded as a continuation and an expansion of its foraging and predatory behavior, as predicted by Darwin's hypothesis for the origin and evolution of slavery in ants.