Sources of variation in the incidence of ant–aphid mutualism in boreal forests

• 1 The mutualism between wood ants of the Formica rufa group and aphids living in the canopy of trees is a widespread phenomenon in boreal forests, and it can affect tree growth. However, not all trees in the forest are involved in this interaction.
• 2 To assess the incidence of host trees involved in this ant–aphid mutualism and its spatial distribution in boreal forests, we inventoried sample plots with a radius of 10–15 m around wood ant mounds in 12 forest stands of two age classes (5–12‐year‐old sapling stands and 30–45‐year‐old pole stands) and two dominant tree species (Scots pine and silver birch) in Eastern Finland from 2007 to 2009.
• 3 The proportion of trees visited by ants out of all trees on the individual study plots were in the range 4–62%, and 1.5–39% of the trees on the plots were consistently visited by ants during all 3 years. The percentage of host trees increased with the ant mound base area on the plots. Trees visited by ants were larger and closer to the mound than trees not visited by ants. Within the group of visited trees, more ants were found on bigger trees and on trees close to the ant mounds.
• 4 Extrapolated from plot to stand level, we estimated that 0.5–6.6% of the trees were host trees in at least one of the three study years, and that only 0.01–2.3% of all the trees were consistently visited by ants during all 3 years. It is concluded that ant–aphid mutualism is a minor occurrence at the stand level.

     

Distribution of ant species and mounds (Formica) in different-aged managed spruce stands in eastern Finland

Mound‐building ants (Formica spp.), as key species, have large impacts on organisms and ecosystem functions in boreal Eurasian forests. The density, sizes and locations of ant mounds determine the magnitude and the spatial distribution of ant activities in forest ecosystems. Clear‐cutting can destroy wood ant colonies, and the species, abundance, dimensions and locations of ant mounds may change as forest stand structure changes with stand age. We compared ant species composition, ant mound numbers and dimensions, and the spatial distribution of mounds in Norway spruce [Picea abies (L.) Karst.] stands of different age (5, 30, 60 and 100 years) in eastern Finland. The mound density of Formica aquilonia Yarr. was greater in the two oldest stand age classes, while most mounds of Formica rufa L., Formica polyctena Först., Formica lugubris Zett., Formica exsecta Nyl. and Formica pressilabris Nyl. were found in the two youngest age classes. The mean volume, the volume per area and height/diameter ratio of F. aquilonia mounds increased with stand age. In the oldest stand age class, mounds were slightly smaller in well‐lit locations than in shade and near stand edges than further from the edges indicating that new mounds are established in well‐lit locations. Similarly, the longest slopes of the mounds faced south, indicating the importance of exposure to the sun. F. aquilonia mounds were concentrated near stand edges, and the spatial distribution of the mounds was aggregated in some stands. At the ecosystem level, the aggregation of ant mounds near stand edges may increase the edge productivity, as mounds concentrate resources to the edges and release nutrients after abandonment.     

Nest mounds of red wood ants (Formicaaquilonia): hot spots for litter-dwelling earthworms

A previously undocumented association between earthworms and red wood ants (Formicaaquilonia Yarr.) was found during an investigation of the influence of wood ants on the distribution and abundance of soil animals in boreal forest soil. Ant nest mounds and the surrounding soil of the ant territories were sampled. The ant nest mound surface (the uppermost 5-cm layer) harboured a much more abundant earthworm community than the surrounding soil; the biomass of the earthworms was about 7 times higher in the nests than in the soil. Dendrodrilusrubidus dominated the earthworm community in the nests, while in soils Dendrobaenaoctaedra was more abundant. Favorable temperature, moisture and pH (Ca content), together with abundant food supply (microbes and decomposing litter) are likely to make a nest mound a preferred habitat for earthworms, provided that they are not preyed upon by the ants. We also conducted laboratory experiments to study antipredation mechanisms of earthworms against ants. The experiments showed that earthworms do not escape predation by avoiding contact with ants in their nests. The earthworm mucus repelled the ants, suggesting a chemical defence against predation. Earthworms probably prevent the nest mounds from becoming overgrown by moulds and fungi, indicating possible mutualistic relationships between the earthworms and the ants. Received: 21 November 1996 / Accepted: 3 April 1997     

Wood ants and a geometrid defoliator of birch: predation outweighs beneficial effects through the host plant

Soil amelioration by a wood-ant species and its consequences for the larval performance of autumnal moths feeding on mountain birch were studied at various distances from the nest mound. Soil nitrate and ammonium nitrogen did not show any clear relationship with distance. However, trees growing in the mound had over 20% more foliar nitrogen than more distant trees. When moth larvae were experimentally protected from predation, their survival rate and final weight tended to decrease with increasing distance. In a laboratory experiment with detached leaves, the relative growth rate of larvae was roughly 30% higher on leaves from trees located on the mound. Differences in larval performance refute the Plant Stress Hypothesis proposed by T.C.R. White and support P.W. Price's Plant Vigor Hypothesis. Predation by ants was examined along the same gradient in trees with and without a glue band that excluded ants from the canopy. Reduction in the daily survival rate of larvae attributable to ant predation was about 35% in trees growing in the mound and around 5% at a distance of 20 m. Other things being equal, about 25 times more larvae entering the penultimate instar would achieve the pupal stage outside the wood-ant territory than in the vicinity of the mound. While both the fertilizing and predatory influence of wood ants is clear, the domain of predation is much larger than the area where trees and their herbivores can exploit enhanced nutrient levels in and around ant mounds. The existence of undamaged green islands around ant mounds in otherwise totally defoliated mountain-birch forests cannot be explained by soil amelioration by wood ants but rather by their predatory activity. Received: 21 November 1996 / Accepted: 8 September 1997     

Impact of Formica exsecta Nyl. on seed bank and vegetation patterns in a subalpine grassland ecosystem

The mound building ant Formica exsecta Nyl. is widely distributed in grassland ecosystems of the Central European Alps. We studied the impact of these ants on seed bank and vegetation patterns in a 11 ha subalpine grassland, where we counted over 700 active ant mounds. The mounds showed a distinct spatial distribution with most of them being located in tall‐grass, which was rarely visited by ungulates (red deer; Cervus elaphus L.). Heavily grazed short‐grass, in contrast, seemed to be completely avoided by ants as only few mounds were found in this vegetation type. The species composition of the ant mound and grassland seed banks was quite similar, i.e. from 15 common plant species 12 were found in both seed bank types. We found the same proportions of myrmecochorous seeds in ant mound and grassland soil samples. In contrast, the number of seeds was 15 times higher in mound compared with the grassland soil samples. Also, the vegetation growing on ant mounds significantly differed from the vegetation outside the mounds: graminoids dominated on ant mounds, herbaceous and myrmecochorous species in the grassland vegetation. We found significant continuous changes in vegetation composition on gradients from the ant mound centre to 1 m away from the mound edge. Overall, F. exsecta was found to have a considerable impact on seed bank and vegetation patterns in the grassland ecosystem studied. These insects not only altered grassland characteristics in the close surrounding of their mounds, but also seem to affect the entire ecosystem including, for example, the spatial use of the grassland by red deer.     

Leaf‐cutting ants,seasonal burning and nutrient distribution in Cerrado vegetation

Abstract Fire and herbivory are known to modify plant community structure. Many studies have suggested that fire ashes may increase soil nutrients in dystrophic soils. Herbivores may also change plant community structure through direct effects of herbivory and affecting nutrient cycling. Leaf‐cutting ants were traditionally viewed as herbivores, although their role may be more complex, because their nests affect both chemical and physical soil properties, thus affecting plants indirectly. We investigated the effects of frequent burning and of leaf‐cutting ants on the nutrient status of an herbaceous and a shrub species occurring in the Brazilian Cerrado, a habitat that is characterized by natural burnings. The proximity of ant nests resulted in an increase of nutrients in the leaves of both vegetation strata, whereas burning sometimes resulted in a decrease of nutrients. Our results do not lead to a possible positive effect of fire on plant nutrient content. On the other hand, ant nests may represent an important source of nutrients for plants on the nutrient‐depleted Cerrado soils and may accelerate vegetation recovery after burning.     

The Role of Wood Ants (Formica rufa group) in Carbon and Nutrient Dynamics of a Boreal Norway Spruce Forest Ecosystem

Wood ants (Formica rufa group) are regarded as keystone species in boreal and mountain forests of Europe and Asia by their effect on ecosystem carbon (C) and nutrient pools and fluxes. To quantify the impact of their activity on boreal forest ecosystems, C, nitrogen (N), phosphorus (P), potassium (K) and calcium (Ca) pools and fluxes in wood ant nests (WAN), and soil were assessed along a 5-, 30-, 60-, and 100-year-old Norway spruce (Picea abies L. Karsten) dominated successional gradient in eastern Finland. Amounts of C and nutrients in WAN increased with stand age, but contained less than 1% of total C and nutrient pools in these stands. The CO₂-efflux from nests was also insignificant, as compared to CO₂-efflux from the forest floor. Annually, the amount of C brought by wood ants into their nests as honeydew, prey and nest-building materials ranged from 2.7 to 49.3 kg ha^?1 C, but this is only 0.1–0.7% of the combined net primary production of trees and understorey in boreal forests. The difference between wood ant nest C inputs and outputs was very small in the younger-aged stands, and increased in the older stands. Carbon accumulation rates in nests over a 100 year period are estimated to be less than 10 kg ha^?1 a^?1. In contrast to C, annual inputs of N, P, and K are larger compared to wood ant nest nutrient pool size, ranging from 3 to 6% of the annual tree stand and understorey uptake. This indicates a more rapid turnover and transport of N, P, and K out of WAN, and suggests that wood ants increase the cycling rate of these nutrients in boreal forests.     

Multivariate analyses of the factors affecting the distribution,abundance and social form of Louisiana fire ants, <Emphasis Type="Italic">Solenopsis invicta</Emphasis>

We surveyed 165 sites to determine the ecological factors influencing the distribution, abundance, and occurrence of polygyny in the red imported fire ant (Solenopsis invicta) in Louisiana. On average, sites had 220 nests/ha, 14% of mounds were polygyne, and 22% of sites had ≥ one polygyne mound. The density of nests and ants per site both increased with the proportion of mounds that were polygyne and the organic and phosphorous content of the soil but decreased with longitude, latitude, and the silt: clay, calcium and sodium content of the soil. Ant density also declined with ambient relative humidity. These multivariate models explained ~25% of the variation in nest and ant density per site. Mean mound size per site increased with the phosphorous content of the soil and the number of nests at the site suggesting that prospective queens may select sites that are conducive to produce large mounds. Mean nest size, however, decreased with the proportion of nests that were polygyne and soil potassium while mounds in forests were typically larger than those in residential areas. Overall, this model accounted for 29% of the variation in mean nest size per site. Polygyne sites were patchily distributed across Louisiana. The probability of a site being polygyne declined with mean monthly temperature for 1999 – 2003 and distance to the nearest commercial waterway suggesting that shipping activities may have played a role in the introduction of polygyne colonists to an area. Forested sites were also less likely to be polygyne than those in residential areas. Finally, the density of polygyne nests and ants increased with latitude whereas that of the monogyne form generally declined with latitude. The abundance of both social forms was also greater when they occurred alone. These data are consistent with the hypothesis that monogyne and polygyne S. invicta compete with one another. Received 28 July 2006; revised 2 March 2007; accepted 29 May 2007.     

Foraging activity and dietary spectrum of wood ants (Formica rufa group) and their role in nutrient fluxes in boreal forests

Abstract 1. We monitored three different‐sized wood ant (Formica aquilonia Yarrow) mounds over a 3‐year period in Finnish boreal forests dominated by Norway spruce (Picea abies Karst.), to assess the seasonal temperature dependency of ant activity. Additionally, we also monitored Norway spruce trees around the mounds for descending honeydew foragers. 2. The amount of collected honeydew and prey and its composition, as well as the carbon (C), nitrogen (N), and phosphorus (P) in honeydew and invertebrate prey was also investigated. 3. The number of warm days (average temperature above 20 °C) and the amount of precipitation differed among the years. Ant activity at the mounds (but not on the trees) was highly correlated with air temperature throughout the ant‐active season (May–September), but ant activity in spring and autumn was lower than in summer at similar temperatures. During all 3 years, honeydew played a major role in wood ant nutrition (78–92% of dry mass). Invertebrate prey was mainly Diptera (on average 26.2%), Coleoptera (12.5%), Aphidina (9.3%), and Arachnoida (8.5%). 4. The total amounts of C, N, and P input brought into the ant mounds in the form of food (both honeydew and prey) on the stand level were 12.6–39.0, 1.6–4.6 and 0.1–0.4 kg ha^?1 year^?1, respectively, which is equivalent to 2–6%, 12–33% and 27–58% of the fluxes in annual needle litterfall in typical boreal Norway spruce forests. Thus, wood ants can play a significant role in short term and local N and P cycling of boreal forest ecosystems.     

热带森林不同演替阶段蚂蚁巢穴的分布特征及其影响因素

为探明热带森林蚂蚁巢穴的分布特征及其影响因素, 采用样方法研究了西双版纳不同演替阶段热带森林定居巢穴蚂蚁的种类及其巢穴的密度、盖度和空间分布特征, 并分析了土壤理化环境与蚂蚁种类总数、巢穴密度及盖度的相关性。结果表明, 不同演替阶段热带森林蚂蚁种类总数、巢穴的密度及盖度大小顺序为: 小果野芭蕉 (Musa acuminata)群落>白背桐(Mallotus paniculatus)群落>思茅崖豆(Mellettia leptobotrya)群落, 并且热带森林的演替类型显著影响蚂蚁种类总数及巢穴密度, 而对巢穴盖度的影响未达到显著水平; 蚂蚁种类总数、巢穴密度与土壤总有机碳和水解氮显著正相关, 与土壤容重和土壤含水率显著负相关, 但所选择的土壤理化指标与巢穴盖度的相关性均未达到显著水平; 蚂蚁巢穴的空间分布呈随机分布格局。我们的数据表明, 不同演替阶段热带森林所形成的植被类型及土壤环境状况共同影响定居的蚂蚁种类总数与筑巢密度。     

Stand type is more important than red wood ant abundance for the structure of ground‐dwelling arthropod assemblages in managed boreal forests

• 1 The relationships between red wood ants (Formica rufa group) and other ground‐dwelling arthropods were studied in young managed forests stands in Eastern Finland. The main objectives were: (i) to test the influence of stand type (dominant tree species; age: sapling versus pole stage) and numbers of red wood ants on the occurrence of other ground‐dwelling arthropods and (ii) to study the occurrence of red wood ants versus other arthropods on a distance gradient from ant mounds. We used pitfall traps set in 5–14‐year‐old sapling stands and 30–45‐year‐old pole‐stage stands of Scots pine (Pinus sylvestris L.) and birch (Betula spp.) forests.
• 2 Pitfall trap catches of red wood ants did not vary significantly between the forest stand types, although some groups of other arthropods showed clear responses to stand type (e.g. catches of other Formicinae and Gnaphosidae were higher in sapling stands than in pole‐stage stands). The number of red wood ants clearly explained less of the variation in assemblages of other ground‐dwelling arthropods than the forest stand type.
• 3 Red wood ant numbers decreased significantly with distance from the mounds, but the other ground‐dwelling arthropods were insensitive to this gradient or even showed a preference for proximity to ant mounds and high ant activity.
• 4 The results obtained in the present study suggest that wood ants do not have strong effects on several other ground‐dwelling arthropod groups in young managed forests other than in the immediate vicinity of their mounds.

     

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.     

Interactive disturbance effects of two disparate ecosystem engineers in North American shortgrass steppe

Disturbances such as fire, grazing, and soil mixing by animals interact to shape vegetation in grassland ecosystems. Animal-generated disturbances are unique in that they arise from a suite of behaviors that are themselves subject to modification by external factors. The manner in which co-occurring animal taxa interact to alter vegetation is a function of their respective behaviors, which shape the characteristics (e.g., the magnitude or extent) of their disturbances. To determine whether prairie dogs (Cynomys ludovicianus) and harvester ants (Pogonomyrmex occidentalis) interactively alter vegetation structure and heterogeneity on the Colorado shortgrass steppe, we characterized the size, dispersion, and vegetation of prairie dog burrow mounds and ant nests (located on and off prairie dog colonies) and vegetation growing beyond mound and nest perimeters. Ants located on prairie dog colonies engineered significantly larger nests and disturbed nearly twice as much total soil area as their off-colony counterparts. Ant nests were overdispersed both on and off prairie dog colonies, while prairie dog mounds were randomly dispersed. Where harvester ants and prairie dogs co-occur, the overdispersed pattern of on-colony ant nests is in effect "overlaid" onto the random pattern of prairie dog mounds, resulting in a unique, aggregated pattern of soil disturbance. Ant nests on prairie dog colonies had significantly less vegetation and lower plant species diversity than did prairie dog mounds, while off-colony nests were similar to mounds. These results suggest that ant nests are more highly disturbed when located on prairie dog colonies. Beyond nests proper, ants did not appear to alter vegetation in a manner distinct from prairie dogs. As such, the interactive effects of prairie dogs and ants on vegetation arise mainly from the disturbance characteristics of mounds and nests proper.     

Influence of mound construction by red and hybrid imported fire ants on soil chemical properties and turfgrass in a sod production agroecosystem

Mound construction by imported fire ants (Solenopsis spp.) actively modify the biogeochemical and physical properties of soil; however, their influence on nutrient levels in surrounding vegetation is poorly understood. Aside from the reported persistence of elevated available P and K levels in clay-rich soils one year after mound abandonment, the relative stability of nutrient concentrations from one season to the next is largely unknown. Nutrient concentrations were concurrently analysed from ant mounds and undisturbed soils as well as plant samples collected from warm-season turfgrass in a commercial sod production agroecosystem. Initial collection of soil and turfgrass samples coincided with peak annual biomass (September 2006); the second soil sample collection occurred over twelve weeks later during turfgrass dormancy and ant brood minimum (December 2006). Total C, C/N ratios, organic matter (OM), and Zn²⁺ concentrations as well as pH of ant mound soils were significantly higher than control plot soils; these trends persisted across seasons. Turfgrass harvested from ant mound perimeters in September exhibited elevated N, P, Ca²⁺, S, Cu²⁺, Fe²⁺, and Na⁺ concentrations. Evaluation of the relative stability of soil parameters across seasons revealed a significant drop in ant nest pH from September to December 2006. Total N of mound soils was distinctively greater than control soil counterparts during September only. Soil P, K⁺, Ca²⁺, Mg²⁺ and S (all macronutrients), as well as Na⁺ concentrations from ant mound soils were substantially elevated during the late Autumn to Winter transition compared to control soil locations, whereas Fe²⁺ and Mn²⁺ levels (both micronutrients) were significantly lower in ant mound soils versus control soil environments. Continuous pedoturbation by imported fire ants as well as seasonal shifts in mound soil chemistry resulting from changes in assimilation/dissimilation among mound biota may influence the site-specific effectiveness of microfaunal pathogens (e.g., Thelohania solenopsae) or parasites (e.g., Orasema spp.) identified as classical biological control agents of non-native Solenopsis spp. Therefore, further study of the intrinsic complexities of soil ecosystem dynamics of imported fire ant mounds across several seasons is warranted. Received 10 September 2007; revised 23 April 2008; accepted 1 May 2008.     

Interactive effects of soil-dwelling ants, ant mounds and simulated grazing on local plant community composition

Interactions between aboveground vertebrate herbivores and subterranean yellow meadow ants (Lasius flavus) can drive plant community patterns in grassland ecosystems. Here, we study the relative importance of the presence of ants (L. flavus) and ant mounds under different simulated grazing regimes for biomass production and species composition in plant communities. We set up a greenhouse experiment using intact soil cores with their associated vegetation.We found that plant biomass production in the short term was affected by an interaction between simulated grazing (clipping) and ant mound presence. Clipping homogenized production on and off mounds, while in unclipped situations production was higher off than on mounds. During the experiment, these differences in unclipped situations disappeared, because production on unclipped mounds increased. Plant species richness was on average higher in clipped treatments and patterns did not change significantly over the experimental period. Plant community composition was mainly affected by clipping, which increased the cover of grazing-tolerant plant species. The actual presence of yellow meadow ants did not affect plant community composition and production.We conclude that the interaction between ant mounds and clipping determined plant community composition and biomass production, while the actual presence of ants themselves was not important. Moreover, clipping can overrule effects of ant mounds on biomass production. Only shortly after the cessation of clipping biomass production was affected by ant mound presence, suggesting that only under low intensity clipping ant mounds may become important determining plant production. Therefore, under low intensity grazing ant mounds may drive the formation of small-scale plant patches.     

Factors affecting distribution of the mound-building ant Lasius neoniger (Hymenoptera: Formicidae) and implications for management on golf course putting greens

Lasius neoniger (Emery), a cosmopolitan ant species, can be a serious pest when its mound-building activities occur on golf course putting greens and other closely mowed turfgrass sites. We mapped the distribution of 735 ant mounds on 30 sand-based putting greens of three golf courses. We then examined factors that might explain why >90% of the mounds on such greens were concentrated in a 2-m wide band just inside the perimeter. Root aphids (Homoptera: Aphididae) from which L. neoniger obtains honeydew were largely absent from high-sand root zone mix of greens but present in surrounding turfgrass on natural soil. Main ant nests, with brood, also were absent from sand-based greens but abundant in adjacent roughs. Although more root aphids were found within ant nests than away from nests, their numbers seem too low to be the main factor restricting the ants' distribution to edges of putting greens. In manipulative experiments, ants responded to low cut (scalped) turf and to sand-filled holes by increased mound building. We suggest that most ant mounds on sand-based greens are associated with subnests, used by foraging workers, which are connected to main nests located just outside the collar in natural soil. Encroachment of mounds into greens occurs when the polydomous colonies seasonally expand their foraging territories, accounting for mounds being concentrated around the perimeter. Control actions for L. neoniger on golf courses should focus on the perimeter of sand-based greens.     

不同形成时间高原鼢鼠鼠丘土壤养分分配规律

高原鼢鼠(Myospalax baileyi)作为青藏高原唯一营地下生活的鼠类,其造丘活动不仅能影响草地群落演替的方向和进程,而且对鼠丘土壤养分含量和分配造成空间异质性。关于不同形成时期的高原鼢鼠鼠丘养分含量及在垂直高度的分配规律研究报道尚少。基于此本研究以不同形成时期的鼢鼠鼠丘为研究对象,对鼠丘土壤养分含量及空间分配规律进行探讨。结果表明:随着鼠丘形成时间的推移,土壤养分总体表现为:1年鼠丘多年鼠丘对照区;与非鼠丘区相比,随着土壤土层增加,土壤养分未呈现规律性的变化。其中,土壤全氮和速效氮含量表现为1年鼠丘3年鼠丘5年鼠丘对照区;0—10cm土壤全磷和速效磷含量变化总体表现为多年鼠丘1年鼠丘对照区,10—30cm土层1年鼠丘土壤中全磷和速效磷含量最高。土壤速效钾和有机质含量随着鼠丘形成时间推移总体表现为:1年鼠丘5年鼠丘3年鼠丘;其中在10—30cm土层,1年鼠丘速效钾和有机质含量显著高于多年鼠丘和对照区。由此可见,高原鼢鼠造丘活动能显著改变土壤养分分配格局,短期内形成养分富集的肥力岛,为鼠丘土壤种子库中种子的萌发和植被生长提供必需的养分保证。     

Thermoregulatory brood transport in the fire ant, <Emphasis Type="Italic">Solenopsis invicta</Emphasis>

Nest structure in ants is often designed to optimize the colony’s ability to thermoregulate, and this specialization is most highly developed in mound-building ant species. Solenopsis invicta invest a large amount of energy in building mounds and transporting their brood up and down in their nests as a means of thermoregulation. Because few ant species build true mounds, we wanted to determine the effectiveness of these mounds in harvesting solar heat as well as to distinguish what factors (temperature vs. circadian rhythm) govern where fire ants place their brood in the mound and when they place it. We measured temperature patterns in the mound over several days at different depths and under different conditions (under direct sunlight or shade), and then conducted a series of field experiments to manipulate the orientation and time of heating. On cool mornings in spring or fall, surface temperatures of the mound rise at the fastest rate on the side receiving the most direct sunlight (usually the south side). This heating causes a temperature gradient through different depths in the mound, and shows little difference from outside ground temperature at a depth greater than ~40 cm inside the nest. In the morning, fire ants move their brood up into the mound on the side most directly heated, and when temperatures exceed optimal (~32°C) they move their brood down the temperature gradient to lower depths in the nest. In addition to this, mound temperature does not only increase due to direct sunlight, but temperature also increases higher than ground temperatures when the mound is in the shade due to its low specific heat. Experiments in which sunlight was mirrored to the normally shaded side of the mound, or when mounds were heated at night, revealed that S. invicta primarily track temperature patterns and do not rely on behavioral habits or circadian rhythms for the thermoregulatory transport of their brood. When mounds were shaded, S. invicta brood was evenly distributed directly under the surface of the mound rather than aggregating towards a specific side. The fire ant mound is important for thermoregulation because, compared to moundless subterranean nests, it absorbs heat more rapidly both in direct sunlight and shady conditions. Temperature tracking within the nest is key to understanding thermoregulatory placement of fire ant brood, as well as insight into the production of sexual brood and reproduction. Received 9 August 2007; revised 31 January 2008; accepted 7 February 2008.     

Interaction between the non-myrmecochorous herb Galium aparine and the ant Formica polyctena

Non-myrmecochorous plant species are able to colonize Formica polyctena mounds. Galium aparine has been chosen as a model to study this kind of ant-plant association. In order to understand benefits to ants and plants, we have studied removal rates of fruits by ant workers, and compared soil seed material, seedling pattern and development of plants of the nest versus forest.Ant workers remove fruits at very low rates; removal time for half of the sample of 10 seeds was on average 5 h. Removed fruits were usually lost within 20–30 cm distance of the ant route. The soil samples from the nests contained a higher number of fruits, than those from randomly chosen sites in the forest. Seeds start to germinate in late August; on the F. polyctena mounds significantly earlier than at surrounding places. The seedling density was higher within experimental quadrats located on mound sites especially in late September–early October. The highest density of seedlings was on the slope of the mound. Plants from mounds are significantly longer, have a higher number of nodes during all stages of development and a higher mortality rate than that at surrounding places.The interactions between G. aparine and F. polyctena ants are asymmetrical. In spite of higher mortality rate of seedlings on mounds, ants provide some benefits to the plant. Firstly, workers are additional dispersal agents for G. aparine fruits. Furthermore, the soil peculiarities of ant mounds are beneficial for seed germination and seedling growth. G. aparine does not provide ants with energetically valuable food like plants with elaiosome-bearing seeds. Moreover, the high density of seedlings and plants on the mounds shades the nest.