Ant species distribution in a sandy coastal plain

Abstract.

• 1 Three species of ants, Lasius niger, Myrmica rubra and Myrmica scabrinodis, have been recorded in a 20-year-old coastal plain on the Dutch Wadden Zee island, Schiermonnikoog.
• 2 The habitats of these species appeared to be clearly segregated, both horizontally (different parts of the plain) and vertically (along dune slope gradients in the plain).
• 3 Lasius niger is distributed throughout the plain in sparsely vegetated sand-dunes and in mounds covered with turf of Festuca rubra ssp. litoralis, but always in areas that are only waterlogged or inundated in winter.
• 4 Myrmica rubra is limited to Festuca-turf, occurring in a distinct zone along the bases of small dunes and on isolated mounds.
• 5 Myrmica scabrinodis only occurs in mounds covered with Festuca-tutf and rarely in the small dunes.
• 6 Analysis of soil, vegetation, inundation and salinity of the ground-water demonstrated that Myrmica scabrinodis is dominant in areas with more silt and salt in the soil, characterized by a vegetation with more typical halophytes, whereas Myrmica rubra occurs most frequently in sandy habitats without much salt stress. The populations of both Myrmica species and Lasius niger do not seem to be affected by inundation during winter.
• 7 The distribution of the ant species is discussed in relation to data about ants in more stable habitats. In particular, attention is given to the absence, in this coastal plain, of Lasius flavus which is a dominant species in the sand-dunes and neighbouring parts of the salt-marsh on Schiermonnikoog.

     

Edge effects on ant community structure and species richness in an agricultural landscape

The effect of sharp edges between three different types of land use on the species richness and structure of ant communities was examined in an agricultural landscape within Central Hesse, Germany. Species richness and nest densities of ants at the centres and the edges of meadows, crop fields, and fallow land were recorded by hand sampling during 1997 and 1998. Edges between different land-use types did not increase ant species richness at the landscape scale, nor were they unique habitats for a specialised ant fauna. Nonetheless, most species shared ecotonal effects in the way that their relative abundance either decreased (e.g. Myrmica scabrinodis) or increased (e.g. Lasius niger, Lasius flavus) at the edges, resulting in different community structure between edges and centres of the land-use types. This was influenced by two major factors: (i) the boundary contrast between the neighbouring habitats (i.e. in terms of disturbance caused by agricultural practices), and (ii) the response of different species to changing abiotic conditions. High nest densities of aggressive species with large colonies occurred along edges. We hypothesise that this can significantly reduce edge permeability for surface-dwelling arthropods.     

Specific patterns of excavation in isolated ants

Summary The tunnels dug by isolated worker ants [Myrmica ruginodis Nylander,M. scabrinodis Nylander,Formica lemani Bondroit,F. fusca L. andLasius niger (L.)] have been studied photographically. Although there was much variation within species, certain interspecific differences in tunnel shape appeared. The tunnels ofL. niger are more spreading, longer and more convoluted than those of the other species, and often branch beneath the surface. The tunnels ofM. scabrinodis show the same features to a smaller degree. The tunnels ofF. fusca andF. lemani are straight, unbranched and vertical.

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Résumé Nous avons étudié par un moyen photographique les tunnels que creusent les ouvrières isolées deMyrmica ruginodis Nyl.,M. scabrinodis Nyl.,Formica fusca L.,F. lemani Bondroit etLasius niger (L.). Quoiqu'il y ait une variation intraspécifique très importante, nous avons constaté plusieurs différences entre ces espèces. Les tunnels deL. niger sont plus longs, plus ramifiés et moins rectilignes que chez les autres espèces. Les tunnels deM. scabrinodis montrent les mêmes caractéristiques, mais d'une façon moins prononcée. Les tunnels deFormica fusca etF. lemani sont plus on moins verticaux et rectilignes et ne se divisent pas.

     

Loss of mating flight and shift in the pattern of carbohydrate storage in sexuals of ants (Hymenoptera; Formicidae)

Summary In ants, energy for flying is derived from carbohydrates (glycogen and free sugars). The amount of these substrates was compared in sexuals participating or not participating in mating flights. Results show that in participating females (Lasius niger, L. flavus, Myrmica scabrinodis, Formica rufa, F. polyctena, F. lugubris), the amount of carbohydrates, especially glycogen, was higher than in non-participating females (Cataglyphis cursor, Iridomyrmex humilis). Similarly, male C. cursor and I. humilis which fly, exhibit a much higher carbohydrate content than do the non-flying females of these species. Furthermore, the quantity of carbohydrates stored was generally higher in males than in females for each species. These results are discussed with regard to the loss of the nuptial flight by some species of ants.     

Effects of inundation and salt on the survival of ants in a sandy coastal plain

Abstract. 1. Survival of four species of ants, Myrmica rubra, Myrmica scabrinodis, Lasius niger and Lasius flavus , exposed to prolonged inundation and the drinking of brackish water, was experimentally determined.
2. In most of the experiments, survival of Lasius flavus workers was much worse than either Lasius niger or Myrmica scabrinodis .
3. After inundation with brackish water, and drinking of brackish water for more than 3 weeks, survival of workers of Myrmica rubra was also more affected than that of Lasius niger and Myrmica scabrinodis .
4. As a rule, survival of dealated queens after inundation appeared to be better in Lasius flavus and Lasius niger , but worse in Myrmica rubra , compared with worker survival.
5. After surviving inundation, the capacity to produce eggs and workers was only slightly affected in queens of both Lusius species.
6. The conclusions based on the experimental mortality rates seem to be consistent both with ant species distribution and with frequency of inundation and salt stress in different parts of the coastal plain and surrounding sand-dunes on the Dutch Wadden island Schiermonnikoog.     

Distribution,host specificity,and the potential for cryptic speciation in hoverfly Microdon myrmicae (Diptera: Syrphidae), a social parasite of Myrmica ants

1. In 2002 Microdon myrmicae, a social parasite of Myrmica ants, was taxonomically separated from Microdon mutabilis. The original study in the U.K. found Microdon myrmicae to be specific to one ant species, Myrmica scabrinodis, yet it became apparent that the range of Microdon myrmicae includes at least the western Palaearctic. 2. Current knowledge of the European distributions of both Microdon myrmicae and Microdon mutabilis in Europe is reviewed. Also, in detailed studies of two Polish populations, Microdon myrmicae was found to survive equally well with two Myrmica ant species. We examine, however, the possibility that this reflects the presence of two separate Microdon species, each connected to one species of Myrmica. 3. Forty populations of Microdon myrmicae and 37 populations of Microdon mutabilis are currently known in Europe. All the populations in central and southern Europe that were visited after the separation of the two species were identified as Microdon myrmicae, while Microdon mutabilis’ recognised range is now restricted to the British Isles and Scandinavia. Myrmica scabrinodis was found to host Microdon myrmicae in 26 out of 31 populations investigated. Four other Myrmica species were identified to the host Microdon myrmicae: Myrmica gallienii (eight populations), Myrmica rubra (four), Myrmica vandeli (one), and Myrmica sabuleti (one). Microdon myrmicae occurs in waterlogged grassland habitats, mainly of the ‘Molinietum’ type, resulting in a patchy distribution relative to its host ants. 4. In two populations Myrmica scabrinodis and Myrmica gallienii are both abundant and rear Microdon myrmicae in equal proportions. Microdon myrmicae pupae from Myrmica gallienii nests were heavier and the anterior respiratory organs were of significantly different shape. In contrast, the comparisons of Microdon myrmicae pupae among all other populations showed no significant differences, suggesting only one species throughout the European range.     

How ants find each other; temporal and spatial patterns in nuptial flights

Reproduction is a key factor in understanding population ecology and therefore species occurrence. However, patterns in reproductive behaviour for distinct ant species remain insufficiently known. In this paper strategies in mate finding are studied for six ant species (Lasius niger, Lasius umbratus, Temnothorax nylanderi, Myrmica rubra, Myrmica ruginodis, Stenamma debile) in a forest – forest edge – agricultural field gradient. Using window traps, we studied whether these species had a restricted nuptial flight season, displayed swarming behaviour, and whether the alates aggregated at the forest edge. The flight season was limited to one month or less for L. niger, T. nylanderi, M. rubra, M. ruginodis and S. debile. Swarming behaviour occurred in all but one (L. umbratus) species. Although none of the six species seemed to have highest nest density at the forest edge, three of them, M. rubra, M. ruginodis and S. debile, showed male aggregations there, indicating this to be the main reproduction site. This last finding could be due to a more suitable micro-climate, but most likely, edges are conspicuous land marks which are used by ants to meet mates. The behavioural patterns of ant sexuals at the forest edge can influence dispersal possibilities in fragmented landscapes, reproductive success and nest densities. Received 27 November 2007; revised 27 March; accepted 5 April 2008.     

Isolation and characterization of microsatellite loci in the ant Myrmica scabrinodis

Five polymorphic microsatellite loci were developed for the ant Myrmica scabrinodis using a magnetic bead hybridization selection protocol. The number of alleles per locus varied between three and six. Cross‐species amplification of four of the loci yielded positive amplification products in four Myrmica species, suggesting their general suitability for microsatellite analysis within this taxonomic group.     

Larval niche selection and evening exposure enhance adoption of a predacious social parasite, Maculinea arion (large blue butterfly), by Myrmica ants

Larvae of the butterfly genus Maculinea feed briefly on a foodplant before being adopted as social parasites into Myrmica ant nests. Each Maculinea species typically survives only with a single Myrmica species, yet the eggs are laid across the overlapping territories of 3-5 Myrmica species and several other ants. The ability of Maculinea arion - a 'predatory' species of Maculinea - to influence its adoption into host Myrmica colonies was studied for the first time in the field. Some earlier reports, involving captive non-host ants, suggested that larvae of the predatory Maculinea follow ant trails into host nests or wander some distance from their foodplant before being discovered and (after a long interaction) carried away by Myrmica foragers. No dispersal from foodplants occurred in wild Maculinea arion larvae. Instead, they increased by >100-fold their probability of being discovered and adopted by Myrmica spp., rather than by ants of other genera, by exposing themselves in the micro-niche occupied by Myrmica foragers at their time of day of peak foraging. Despite a complex, hour-long interaction with Myrmica workers before being carried to the nest, Maculinea arion did not enhance its adoption by host species of Myrmica. Eggs were laid without bias in Myrmica sabuleti (host) and Myrmica scabrinodis (non-host) territories; larval survival on Thymus was the same in both ants' territories; larvae waited to be found beneath their foodplant rather than seek their host; Myrmica sabuleti and Myrmica scabrinodis foraged in the same vertical and temporal niches, and had the same probability of discovering larvae; both ants behaved identically after finding larvae and took the same time to adopt them; and the ratio of wild larvae taken into Myrmica sabuleti or Myrmica scabrinodis nests was the same as the distribution between these ants of Thymus, eggs and pre-adoption larvae.     

Vegetation of ant-hills in a mountain grassland:effects of mound history and of dominant ant species

Vegetation in grasslands with well-developed long-lastingant-hills in the Slovenské Rudohorie Mts., Slovakia, was studiedin relation to (i) position on the mound, (ii) ant speciesforming the mound, and (iii) history of the mound. Permanent plotrecordings of mound size and dominant ant species started fifteen years priorthe study began provided information on the history of individual mounds.The mound vegetation bears a striking similarity to vegetation insimilar habitats across a large part of Europe due to presence of species suchas Agrostis capillaris, Dianthusdeltoides, Polytrichum commune agg.,Thymus pulegioides, and Veronicaofficinalis. Out of the three major ant species-groups presentat the site (Lasius flavus, Tetramoriumcaespitum and Formica spp.), L.flavus had the most pronounced and the most lasting effect on themound vegetation. The dominance of the plant species listed above increased withthe time span over which the mound was inhabited by L.flavus. The effects of other species on vegetation composition,though discernible from short-term observation, disappeared over severalyears. The mounds proper did not differ from the undisturbed grassland in theproportion of myrmecochorous plants or plants with specific seed size ordormancy type. However, there was a highly significant concentration ofmyrmecochorous plants in the grassland patches immediately neighbouring themounds; this is likely to be due to seeds deposited there by the workers fromthe nest after the elaiosomes had been consumed. The mound vegetation wascomposed mainly of species with long stolons or rhizomes; however, there was nosignificant difference in formation type or length of stolons/rhizomesbetween mounds and the rest of the grassland or among mounds formed by differentant species.     

Ants benefit from attending facultatively myrmecophilous Lycaenidae caterpillars: evidence from a survival study

Workers of three ant species (Lasius niger, Lasius flavus, Myrmica rubra) were caged in the laboratory together with caterpillars and pupae of five species of lycaenid butterflies. Mortality of ants was 3–5 times higher when the ants were confined with larvae lacking a dorsal nectar organ (Lycaena phlaeas, Lycaena tityrus) rather than with caterpillars which possess a nectar gland (Aricia agestis, Polyommatus bellargus, P. icarus). For all five species, ant survival was always lower at the pupal stage (where a nectar organ is always absent) than at the caterpillar stage and was largely equivalent for the butterfly species tested. The experimental data confirm earlier estimates that ants can derive nutritive benefits from tending facultatively myrmecophilous lycaenid caterpillars, even though these caterpillars produce nectarlike secretions at low rates.     

Multiple host-ant use by the predatory social parasite <Emphasis Type="Italic">Phengaris</Emphasis> (=<Emphasis Type="Italic">Maculinea</Emphasis>) <Emphasis Type="Italic">arion</Emphasis> (Lepidoptera,Lycaenidae)

Phengaris (=Maculinea) arion is an endangered social parasite of Myrmica ants, and for a very long time was considered as specific to Myrmica sabuleti. Previous studies carried out in Poland suggested some discrepancies within this assumption, and therefore a much more intensive survey was undertaken. The host ant use of P. arion was studied at five sites in different types of biotopes in Poland, i.e. xerothermal grasslands where Thymus pulegioides was used as a larval food plant by the butterfly, and more or less sandy biotopes with Thymus serpyllum. Altogether nine Myrmica species were recorded, and considerable variation in species composition and density of nests was recorded. At four localities M. sabuleti proved to be the most common ant. A total of 529 Myrmica nests were examined, and only 20 of them contained larvae and pupae of P. arion. Host ants belonged to five different species, i.e. M. sabuleti, Myrmica scabrinodis, Myrmica schencki, Myrmica lobicornis and Myrmica hellenica. Only at one site (NE Poland) was a significant heterogeneity in parasitation rates among Myrmica species detected. M. lobicornis was the most often infested ant there, which may suggest local specialisation of the butterfly. Overall low parasitism rates may explain the vulnerability of P. arion in Central Europe but further studies are also necessary.     

Frequency of multiple paternity in Myrmica scabrinodis from southern Poland

In haplodiploid insects that create social systems, more than one male may contribute to the progeny of a single female (polyandry). Little is known about the possible evolutionary benefits of multiple mating or the occurrence of this phenomenon across different species of Hymenoptera. Here we present our observations of polyandry in a Myrmica scabrinodis Nylander population from southern Poland. The estimated proportion of multiply mated queens was 0.71 and the average pedigree‐effective mate number was 1.45.     

Not only the butterflies: managing ants on road verges to benefit Phengaris (Maculinea) butterflies

Obligate myrmecophilic butterfly species, such as Phengaris (Maculinea) teleius and P. nausithous, have narrow habitat requirements. Living as a caterpillar in the nests of the ant species Myrmica scabrinodis and M. rubra, respectively, they can only survive on sites with both host ants and the host plant Great Burnet Sanguisorba officinalis. After having been reintroduced into a nature reserve in the Netherlands in 1990, both butterfly species expanded their distribution to linear landscape elements such as road verges and ditch edges outside this reserve. As additional habitat of both butterfly species, vegetation management of these landscape elements became important. Our results show that a management beneficial for Phengaris butterflies should aim to increase the nest density of Myrmica species, at the same time reducing the density of nests of the competitor Lasius niger or at least keeping them at a low density. Unfavourable grassland management under which L. niger thrives, includes not mowing or flail-cutting the grass, or depositing dredgings along the side of the ditch. Management favourable for the two Myrmica species differs, demanding some flexibility if both species are to benefit. M. scabrinodis is best supported with early mowing of the road verge vegetation or late mowing in the nature reserve, both of which result in an open vegetation and warm microclimate. In contrast, the nest sites of M. rubra should be left undisturbed during the summer, and mown in late autumn. Mowing of butterfly habitat should be avoided between mid-June and mid-September as this would remove the flowerheads of the Sanguisorba plants, on which the butterflies lay their eggs.     

Local host ant specificity of Phengaris (Maculinea) teleius butterfly,an obligatory social parasite of Myrmica ants

1. Phengaris butterflies are obligatory social parasites of Myrmica ants. Early research suggested that there is a different Myrmica host species for each of the five European Phengaris social parasites, but more recent studies have shown that this was an oversimplification. 2. The pattern of host ant specificity within a Phengaris teleius metapopulation from southern Poland is reported. A combination of studying the frequency distribution of Phengaris occurrence and morphometrics on adult butterflies were used to test whether use of different host species is reflected in larval development. 3. Phengaris teleius larvae were found to survive in colonies of four Myrmica species: M. scabrinodis, M. rubra, M. ruginodis, and M. rugulosa. Myrmica scabrinodis was the most abundant species under the host plant but the percentage of infested nests was similar to other host ant species at two sites and lower in comparison to nests of M. rubra and M. ruginodis at the other two sites. Morphometric measurements of adult butterflies reared by wild colonies of M. scabrinodis and M. ruginodis showed that wing size and number of wing spots were slightly greater for adults eclosing from nests of M. ruginodis. 4. Our results suggest that P. teleius in the populations studied is less specialised than previously suggested. The results are consistent with the hypothesis that P. teleius is expected to be the least specific of the European Phengaris species, as it has the largest and best defended fourth‐instar caterpillars and, as a predatory species, it spends less time in the central larval chambers of the host colonies. The fact that individuals reared by M. ruginodis had wider hind wings may suggest that P. teleius had better access to resources in M. ruginodis than in M. scabrinodis colonies.     

Host specificity revisited: New data on Myrmica host ants of the lycaenid butterfly Maculinea rebeli

Larvae of Maculinea rebeli, one of the most endangered European butterflies, are obligatory social parasites of Myrmica ants. At present, this relationship is thought to be highly specific, with Myrmica schencki being regarded as the primary host. Here we present data on six populations from Poland and Austria, including the first record of Myrmica specioides as a host, together with published data from other central European countries, which severely questions the inference that M. schencki is the exclusive host of M. rebeli. Our results indicate that Myrmica sabuleti is the most frequently used host ant in central Europe, whereas M. scabrinodis, M. sulcinodis, M. specioides and M. schencki are used as secondary hosts. Possible explanations for this highly variable host use include (1) regional differences in semiochemicals, behaviour or social structure of the potential Myrmica host species and (2) the existence of different ecological subspecies or cryptic species of M. rebeli. Finally, we emphasize the importance of identifying local host ant species prior to further conservation strategies in order to avoid failure of management programs or even damage to populations on the edge of extinction.     

Habitat requirements and niche selection of Maculinea nausithous and M. teleius (Lepidoptera: Lycaenidae) within a large sympatric metapopulation

Identifying the factors determining the distribution and abundance of species is a fundamental question in population ecology. Based on habitat characteristics, we sought to identify factors affecting the abundance of two vulnerable large blue butterflies, Maculinea teleius and Maculinea nausithous. Both species often occur in sympatry and thus seem to have very similar requirements with regard to their wetland habitats. Even though, M. nausithous is usually far more widespread and abundant than M. teleius in Central Europe. For understanding this difference despite a similar ecology, more data is needed on the niche selection of both species. We have recorded Maculinea numbers, vegetation structure, food-plant abundance and ant community for 120 patches within a large sympatric metapopulation of both species in the Westerwald area (Germany). Unexpectedly (based on previous data), both species were equally common in the study area, reaching high densities of up to 61 (M. teleius) and 58 (M. nausithous) individuals per 50 m2. In 116 out of 120 patches both species occurred sympatrically. The preferred habitat was extensively managed grassland, being mown or grazed twice a year. The abundance of both Maculinea species was positively related to vegetation height, food-plant and Myrmica rubra abundance. There was no association between M. teleius and its putative main host-ant Myrmica scabrinodis, indicating that M. rubra may serve as an important host in the Westerwald area. The positive effect of food-plant abundance was more pronounced for M. teleius, coinciding with a higher food-plant abundance at sites dominated by M. teleius compared to those dominated by M. nausithous. Thus, M. teleius seems to be slightly more demanding than M. nausithous in terms of food-plant availability.     

Colony founding and social parasitism in <Emphasis Type="Italic">Lasius</Emphasis> (<Emphasis Type="Italic">Acanthomyops)</Emphasis>

We investigated colony foundation behavior of three species of the Lasius claviger group, L. latipes, L. interjectus and L. claviger, using field observations and laboratory experiments. Laboratory studies included observing gyne behavior prior to interaction with host colonies (e.g. grouping, overwintering, feeding) and experiments in which gynes were introduced to putative host colonies. Observations of gyne parasitic behavior immediately after mating flights in the field and the discovery of mixed colonies both confirm the parasitic behavior of these species and point toward a wide host species range including two cases of hyperparasitism. L. interjectus and L. latipes enter the host colony immediately after the nuptial flight, while L. claviger may hibernate and enter in the spring. Laboratory experiments showed that L. interjectus gynes prefer to group and suffer lower mortality when in a group, which is consistent with field observations of L. interjectus gynes entering host colonies in large numbers. Details on behavior of L. latipes alpha and beta form gynes are also given.     

The spatial distribution and environmental triggers of ant mating flights: using citizen‐science data to reveal national patterns

Many ant species produce winged reproductive males and females that embark on mating flights. Previous research has shown substantial synchrony in flights between colonies and that weather influences phenology but these studies have been limited by sample size and spatiotemporal scale. Using citizen science, we gathered the largest ever dataset (> 13 000 observations) on the location and timing of winged ant sightings over a three‐year period across a broad spatial scale (the United Kingdom). In total, 88.5% of winged ants sampled were Lasius niger. Observations occurred from June to September with 97% occurring in July/August but exact temporal patterns differed substantially between years. As expected, observations within each year showed a small but significant northward/westward trend as summer progressed. However, the predicted spatiotemporal synchrony was far less apparent; observations were not significantly spatially clustered at national, regional or local scales. Nests in urban (vs rural) areas and those associated with heat‐retaining structures produced winged ants earlier. Local weather conditions rather than broad geographical or seasonal factors were shown to be critical in the timing of winged ant activity, presumably to optimize mate finding and to minimize energy consumption and predation. Temperature and wind speed, but not barometric pressure, were significant predictors of observations (positively and negatively, respectively); winged ants were only observed at temperatures > 13°C and wind‐speeds < 6.3 m^–1. All days with a mean daily temperature > 25°C had observations. Intriguingly, changes in temperature and wind speed from the day before flight peaks were also significant. We conclude that: 1) spatiotemporal synchrony in flights is lower than previously thought for L. niger, 2) local temperature and wind are key predictors of flight phenology; and 3) ants appear able to determine, at least in a limited way, if weather is improving or deteriorating and adjust their behaviour accordingly.