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.     

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.

     

Testing the enemies hypothesis in forest stands: the important role of tree species composition

Numerous studies conducted in agro-ecosystems support the enemies hypothesis, which states that predators and parasites are more efficient in controlling pest densities in polycultures than in monocultures. Few similar studies, however, have been conducted in forest ecosystems, and we do not yet have evidence as to whether the enemies hypothesis holds true in forests. In a 2-year study, we investigated whether the survival of autumnal moth (Epirrita autumnata) larvae and pupae differs between silver birch monocultures and two-species mixtures of birch with black alder, Norway spruce and Scots pine. We placed young larvae on birch saplings and monitored their survival until the end of the larval period, when we checked whether they had been parasitized. After the larvae had pupated, pupal survival was tested in a field trial. In 2002, the larvae disappeared earlier and their overall survival was lower in birch–pine mixtures than in other stand types. In 2003, survival probability was lowest in birch–pine stands only during the first week and there were no differences between stands in overall survival. Larval parasitism was not affected by tree species composition. Pupal weight and pupal survival were likewise not affected by stand type. Among the predators, wood ants were more abundant on birches growing in birch–pine mixtures than in other stand types probably because colonies of myrmecophilic aphids were common on pines. In contrast, spider numbers did not differ between stand types. Ant exclusion by means of a glue ring around the birch trunk increased larval survival, indicating that ants are important predators of the autumnal moth larvae; differences in larval survival between stands are probably due to differential ant predation. Our results provide only partial support for the enemies hypothesis, and suggest that it is both tree species composition and species diversity which affect herbivore survival and predation.     

CO2 efflux from subterranean nests of ant communities in a seasonal tropical forest,Thailand

Many ant species construct subterranean nests. The presence of their nests may explain soil respiration “hot spots”, an important factor in the high CO₂ efflux from tropical forests. However, no studies have directly measured CO₂ efflux from ant nests. We established 61 experimental plots containing 13 subterranean ant species to evaluate the CO₂ efflux from subterranean ant nests in a tropical seasonal forest, Thailand. We examined differences in nest CO₂ efflux among ant species. We determined the effects of environmental factors on nest CO₂ efflux and calculated an index of nest structure. The mean CO₂ efflux from nests was significantly higher than those from the surrounding soil in the wet and dry seasons. The CO₂ efflux was species‐specific, showing significant differences among the 13 ant species. The soil moisture content significantly affected nest CO₂ efflux, but there was no clear relationship between nest CO₂ efflux and nest soil temperature. The diameter of the nest entrance hole affected CO₂ efflux. However, there was no significant difference in CO₂ efflux rates between single‐hole and multiple‐hole nests. Our results suggest that in a tropical forest ecosystem the increase in CO₂ efflux from subterranean ant nests is caused by species‐specific activity of ants, the nest soil environment, and nest structure.     

The effect of nest moisture on daily temperature regime in the nests of Formica polyctena wood ants

Summary: Relations between nest moisture and daily temperature regime were studied. Two extreme situations were distinguished among the variable patterns of daily temperature regime found. The first one was characterized by increasing temperature during the day and decreasing temperature during the night and was typical for dry nests exposed to sun. The second one was characterized by a lower temperature during the day that increased rapidly during the night. This pattern occurred in moist shaded nests and was less frequent than the first one.¶Patterns of surface nest temperature, which closely correspond with thermal loss, differ between dry and wet nests as well. In dry nests, the temperature decreased during the night on the whole surface in a similar way. In moist nests, the top parts constantly had a high temperature. The average surface temperature during the night was significantly higher in moist than in dry nests, which implies a higher thermal loss in moist nests. Microbial respiration of nest material strongly correlated with nest moisture, which implies a higher microbial heat production in moist nests.¶The results indicated two mechanisms for the maintenance of internal nest temperature. The first one, used in dry nests, is based on a combination of ant metabolic heat production and the isolation properties of a dry nest. The second one, used in moist nests, is based on the metabolic heat production provided by both the ants and the microorganisms in the nest material. These two strategies differ in the pattern of daily temperature fluctuation.     

Ecological aspects of development of testacean complexes in nests of Formica lugubris and Formica exsecta (Hymenoptera, Formicidae)

Testacean population in the nests of Formica lugubris and F. exsecta was studied in a spruce forest and in a mixed birch forest. Samples of the nest material (spruce litter and grass fragments) were taken from the surface layers, the inner parts of the nests, and from underlying soil. In all, 33 species of testaceans were identified. The highest species diversity and abundance were observed in the Formica lugubris nests, in the upper layer of the F. exsecta nest, and in the spruce litter. In all the samples, 10 widespread aerophilic and edaphic species (Centropyxis aerophila, C. sylvatica, C. orbicularis, Cyclopyxis kahli, Trinema lineare, and others) were the most common, resulting in a uniform species composition in all the habitats studied. In the presence of the relevant substrate, this group is supplemented by species of appropriate ecological groups (bryophilic, acidophilic, and inhabitants of coarse humus). In the inner part of the Formica lugubris nest, the testacean population is characterized by a very high abundance of Plagiopyxis penardi, whereas the surface layer of the F. exsecta nest is characterized by abundant Cyclopyxis eurystoma and very low species diversity in the inner layers. The initial composition and decree of decomposition of plant remains due to ant activity are considered as the main factors responsible for the testacean species diversity, as well as the availability of substrates suitable for the development of different ecological groups of testaceans.     

The simulation of soil organic matter and nitrogen accumulation in Scots pine plantations on bare parent material using the combined forest model EFIMOD

The individual-based combined forest model EFIMOD including the soil-sub model SOMM has been used for the simulation of Scots pine stand growth and soil organic matter (SOM) accumulation on a humus-free bare mineral surface. The growth of Scots pine plantation, with an initial density of 10 000 trees ha⁻¹ and average tree biomass of 0.01 kg was simulated for 50 yr under Central European climatic conditions (i) with varying atmospheric nitrogen inputs and (ii) different rates of initial application of raw undecomposed organic material or compost, on humus-free parent material. The accumulation of typical raw humus was simulated in all cases. The accumulation was most intensive in the simulation of high atmospheric nitrogen input. The humus pool in the mineral topsoil was small but achieved its maximum value with compost application. SOM nitrogen accumulation was scant in all cases, except the compost applications with low atmospheric nitrogen input. No statistically significant differences of SOM and stand parameters were found between variants without organic matter and those with low input of organic manure. However, the maximum relative rate of SOM and nitrogen accumulation was found in the scenario without organic manure, under slowly growing unstable Scots pine plantation. This revised version was published online in June 2006 with corrections to the Cover Date. This revised version was published online in June 2006 with corrections to the Cover Date. This revised version was published online in June 2006 with corrections to the Cover Date.     

Ant colonization and coarse woody debris decomposition in temperate forests

Ants are ubiquitous, abundant and have widespread impacts on ecological communities and ecosystem processes. However, ant effects on coarse woody debris decomposition are unexplored. Several ant species colonize coarse woody debris for nesting, and this puts them in contact with fauna and microbes that utilize coarse woody debris as habitat and food, potentially influencing nutrient cycling and, ultimately, forest productivity. We report results from a field experiment employing 138 artificial ant nests (routed pine blocks) across five locations in southeastern US deciduous forests. We examine the correspondence between ant, termite and wood-eating fungi colonization and variation in coarse woody debris decomposition. After 1 year, nests colonized by ants had 5% more mass than those not colonized. Ant colonization corresponded with significantly less termite- and fungal-mediated decomposition of the nests. Without ants, termites removed 11.5% and fungi removed 4% more wood biomass. Ants, termites and wood-eating fungi all colonized pine nests where temperatures were highest, and ants also preferred higher soil moisture whereas termites and fungi responded negatively to high soil moisture when temperatures were higher. Ants reduce termite colonies through predation, and may inhibit fungi through the secretion of antimicrobial compounds. Our results indicate that interactions between forest understory ants, termites and fungi may influence the rate of coarse woody debris decomposition—biotic interactions that potentially influence forest structure and function.     

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.

     

Thiosulfate sulfurtransferase (rhodanese) in forest soils

Concentrations of thiosulfate sulfurtransferase (rhodanese; EC 2.8.1.1) in soils of five types of forest stands (spruce and pine stands without ground vegetation, and mountain ash, birch and pine stands with grass cover) were followed. Soils from grassed stands contained much higher concentrations of the enzyme than soils without ground vegetation. On grassed locations higher enzyme concentrations were found in fermentation horizons, whereas on locations without the ground vegetation they were detected in humus horizons. Urea fertilization and SO₂ pollution did not exhibit any significant effect.     

Changes in diversity and storage function of ectomycorrhiza and soil organoprofile dynamics after introduction of beech into Scots pine forests

Diversity and storage function of mycorrhiza as well as soil organoprofile formation were investigated in a chronosequence of a pure Scots pine (Pinus sylvestris L.) stand, of Scots pine stands that were underplanted with beech (Fagus sylvatica L.) and in three pure beech stands of different age. Mycorrhiza diversity was higher in the pure beech stands compared to the pure pine stand. Beech and pine trees in the mixed stands had similar dominant mycorrhiza morphotypes. However, trees in two of the three pure beech stands were mycorrhized with other types. Mycorrhizal abundance and nutrient amounts of mycorrhizae associated with beech trees were higher in the mixed and in the pure beech stands compared to pine mycorrhizae indicating that nutrient uptake was higher in older beech than in older pine trees. Humus quality varied from pine to beech stands. Plant litter storage in the humus layer was highest in the youngest mixed stand and lowest in the oldest beech stand. Humus forms changed from moder grass-type in the pure Scots pine stand to mor-like moder and moder rich in fine humus with increasing age of beeches in the mixed stands. The older beech stands were characterised by oligomull and mull-like moder as the dominating humus forms. The ecologically favourable humus forms, i.e., nutrient rich humus forms in the older beech stands correlate well with the higher mycorrhizal diversity and abundance as well as the higher nutrient storage of their mycorrhizae in these stands. The results are also discussed with regard to the 'base-pump effect' of beech trees.     

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     

Soil CO2 efflux in a mixed pine-oak forest in Valsaín (central Spain)

Soil-surface CO2 efflux and its spatial and temporal variation were investigated in a southern Mediterranean, mixed pine-oak forest ecosystem on the northern slopes of the Sierra de Guadarrama in Spain from February 2006 to July 2006. Measurements of soil CO2 efflux, soil temperatures, and moisture were conducted in nine 1963-m2 sampling plots distributed in a gradient around the ecotone between Pinus sylvestris L. and Quercus pyrenaica Lam. forest stands. Total soil organic matter, Walkey-Black C, particulate organic matter, organic matter fraction below 53 microm, total soil nitrogen content, total soil organic carbon content, and pH were also measured under three representative mature oak, pine, and mixed pine-oak forest stands. Soil respiration showed a typical seasonal pattern with minimums in winter and summer, and maximums in spring, more pronounced in oak and oak-pine stands. Soil respiration values were highest in pine stands during winter and in oak stands during spring and summer. Soil respiration was highly correlated with soil temperatures in oak and pine-oak stands when soil moisture was above a drought threshold of 15%. Below this threshold value, soil moisture was a good predictor of soil respiration in pine stands. Greater soil organic matter, particulate organic matter, Walkey-Black C, total organic C, and total N content in pine compared to oak sites potentially contributed to the greater total soil CO2 efflux in these stands during the winter. Furthermore, opposing trends in the organic matter fraction below 53 microm and soil respiration between plots suggest that in oak stands, the C forms are less affected by possible changes in use. The effects of soil properties on soil respiration were masked by differences in soil temperature and moisture during the rest of the year. Understanding the spatial and temporal variation even within small geographic areas is essential to assess C budgets at ecosystem level accurately. Thus, this study bears important implications for the study of large-scale ecosystem dynamics, particularly in response to climatic change.     

An Analysis of Vegetation Restoration on Opencast Oil Shale Mines in Estonia

We compared four types of 30‐year‐old forest stands growing on spoil of opencast oil shale mines in Estonia. The stand types were: (1) natural stands formed by spontaneous succession, and plantations of (2) Pinus sylvestris (Scots pine), (3) Betula pendula (silver birch), and (4) Alnus glutinosa (European black alder). In all stands we measured properties of the tree layer (species richness, stand density, and volume of growing stock), understory (density and species richness of shrubs and tree saplings), and ground vegetation (aboveground biomass, species richness, and species diversity). The tree layer was most diverse though sparse in the natural stands. Understory species richness per 100‐m² plot was highest in the natural stand, but total stand richness was equal in the natural and alder stands, which were higher than the birch and pine stands. The understory sapling density was lower than 50 saplings/100 m² in the plantations, while it varied between 50 and 180 saplings/100 m² in the natural stands. Growing stock volume was the least in natural stands and greatest in birch stands. The aboveground biomass of ground vegetation was highest in alder stands and lowest in the pine stands. We can conclude that spontaneous succession promotes establishment of diverse vegetation. In plantations the establishment of diverse ground vegetation depends on planted tree species.     

Predation on artificial nests in a forest dominated landscape – the effects of nest type, patch size and edge structure

We studied the effects of forest patch size and forest edge structure on nest predation in a boreal coniferous forest landscape. The following predictions were tested. Nest predation should be higher in small than in large stands, in edges than in interior areas of forest stands, and in barren forest/clear–cut edges created by forestry than in natural forest/open marsh edges. Four types of artificial above ground nests (total of 261) were used; open cup nests with reindeer Rangifer t. tarandus hair, open cup nests with domestic hen Gallus domesticus feathers, and unlined open cup and nest–box nests. Nests were baited with one Japanese quail Coturnix coturnix japonica egg. Nest–boxes were depredated significantly less than open cup nests of all types. No edge- or stand size–related nest predation was found. The predation rate, regardless of the nest type, did not differ relative to the edge type and vegetation characteristics. However, better horizontal visibility of open cup nests due to more open vegetation structure increased predation risk in man–made edges compared to inherent edges. The results suggest that edge–related nest predation is absent or weak in forest dominated landscapes. This may be due to predator types present in the landscape and/or predators habitat use in forest dominated areas. Therefore, it might be that findings documented in other areas, such as in agricultural dominated landscapes, cannot be directly applied to managed forest landscapes.     

Diel pattern driven by free convection controls leaf-cutter ant nest ventilation and greenhouse gas emissions in a Neotropical rain forest

Leaf-cutter ant nests are biogeochemical hot spots where ants live and import vegetation to grow fungus. Metabolic activity and (in wet tropical forests) soil gas flux to the nest may result in high nest CO2 concentrations if not adequately ventilated. Wind-driven ventilation mitigates high CO2 concentrations in grasslands, but little is known about exchange for forest species faced with prolonged windless conditions. We studied Atta cephalotes nests located under dense canopy (leaf area index > 5) in a wet tropical rainforest in Costa Rica, where wind events are infrequent. We instrumented nests with thermocouples and flow-through CO2 sensing chambers. The results showed that CO2 concentrations exiting leaf-cutter ant nests follow a diel pattern with higher values at night. We developed an efflux model based on pressure differences that evaluated the observed CO2 diel pattern in terms of ventilation by (1) free convection (warm, less dense air rises out the nest more prominently at night) and (2) episodic wind-forced convection events providing occasional supplemental ventilation during daytime. Average greenhouse gas emissions were estimated through nest vents at about 78 kg CO2eq nest−1 year−1. At the ecosystem level, leaf-cutter ant nest vents accounted for 0.2% to 1% of total rainforest soil emissions. In wet, clayey tropical soils, leaf-cutter ant nests act as free convection-driven conduits for exporting CO2 and other greenhouse gases produced within the nest (fungus and ant respiration, refuse decay), and by roots and soil microbes surrounding the nest. This allows A. cephalotes nests to be ventilated without reliable wind conditions.     

Degradability of dissolved soil organic carbon and nitrogen in relation to tree species

The degradability and chemical characteristics of water-extractable dissolved organic carbon (DOC) and nitrogen (DON) from the humus layer of silver birch (Betula pendula Roth), Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) stands were compared in short-term incubation of soil solutions. For all extracts the degradation of DOC and DON was low (12-17% loss) and increased in the order: birch, spruce and pine. In the humus layer under pine a relatively larger pool of rapidly degrading dissolved soil organic matter (DOM) was indicated by the [3H]thymidine incorporation technique, which measures the availability of DOM to bacteria. The degradation of DOC was explained by a decrease in the hydrophilic fraction. For DON, however, both the hydrophilic and hydrophobic fractions tended to decrease during incubation. No major differences in concentrations of hydrophilic and hydrophobic fractions were detected between tree species. Molecular size distribution of DOC and DON, however, revealed slight initial differences between birch and conifers as well as a change in birch extract during incubation. The depletion of very rapidly degrading fractions (e.g., root exudates and compounds from the litter) may explain the low degradability of DOM in the humus layer under birch.     

土壤温度和水分对长白山不同森林类型土壤呼吸的影响

在实验室条件下,将不同含水量的3种森林类型的土柱分别置于0、5、15、25和35℃条件下,进行土壤呼吸测定．结果表明,在0～35℃范围内。土壤呼吸速率与温度呈正相关．在一定含水量范围内(0．21～0．37kg·kg^-1),土壤呼吸随含水量的增加而升高,当含水量超出该范围,土壤呼吸速率则随含水量的变化而降低．土壤温度和水分对土壤呼吸作用存在明显的交互作用．不同森林类型土壤呼吸作用强弱存在显著差异,大小顺序为阔叶红松林>岳桦林>云冷杉暗针叶林．阔叶红松林土壤呼吸作用的最佳条件是土壤温度35℃、含水量0．37kg·kg^-1;云冷杉暗针叶林下的山地棕色针叶林土壤呼吸作用的最佳条件是25℃、0．21kg·kg^-1;岳桦林土壤呼吸作用的最佳条件是35℃、含水量0．37kg·kg^-1。但是．由于长白山阔叶红松林、云冷杉林和岳桦林处在不同的海拔带上,同期不同森林类型土壤温度各不相同,相差4～5℃,所以野外所测的同期山地棕色针叶林土呼吸速率应低于暗棕色森林土呼吸速率,山地生草森林土呼吸速率应高于山地棕色针叶林土的呼吸速率．     

小兴安岭5种林型土壤呼吸时空变异

原始阔叶红松林、谷地云冷杉林、阔叶红松择伐林、次生白桦林、人工落叶松林是小兴安岭乃至东北地区的重要森林类型。采用红外气体分析法比较测定了这几种森林类型的土壤呼吸及其相关环境因子,分析探讨了这几种森林类型土壤呼吸的时空变异。结果表明:各林型土壤呼吸与5 cm深土壤温度(T5)呈显著的指数相关,并且土壤呼吸与土壤温度、土壤湿度及其相互作用的回归模型可以解释各林型土壤呼吸约71%的季节变异。生长季平均土壤呼吸速率为次生白桦林(3.59μmolCO.2m-.2s-1)>谷地云冷杉林(3.52μmolCO.2m-.2s-1)>阔叶红松择伐林(3.44μmolCO.2m-.2s-1)>原始阔叶红松林(2.58μmolCO.2m-.2s-1)>人工落叶松林(2.29μmolCO.2m-.2s-1),说明土壤呼吸对原始阔叶红松林人为干扰的响应是不同的。各林型Q10值介于1.84(人工落叶松林)—2.32(次生白桦林)之间。在整个生长季,各林型之间土壤呼吸的变异系数变化幅度为19.74%—37.39%,而各林型内土壤环间其变化幅度为32.13%—60.20%,显著大于样地间的变化幅度14.28%—35.70%(P<0.001),说明土壤呼吸在细微尺度上的差异更大。土壤湿度可以解释各林型(阔叶红松林除外)内部土壤呼吸15.8%—33.5%的空间异质性。     

Habitat heterogeneity affects predation of European pine sawfly cocoons

Habitat heterogeneity is thought to affect top‐down control of herbivorous insects and contribute to population stability by providing a more attractive microhabitat for natural enemies, potentially leading to reduced population fluctuations. Identifying the parameters that contribute to habitat heterogeneity promoting top‐down control of herbivorous insects by natural enemies could facilitate appropriate management decisions, resulting in a decreased risk of pest insect outbreaks because of a higher level of predation. In our study, we measured the top‐down pressure exerted by small mammals on the cocoons of a notorious pest insect in pine forests, the European pine sawfly (Neodiprion sertifer), which is known to be regulated by small mammal predation. The forest stands used differed in heterogeneity measured in terms of differences in tree diversity and density, understory vegetation height, presence/absence, and density of dead wood. We found higher predation in more dense spots within forest stands. Further, the effect of dead wood on sawfly cocoon predation depended on the pine proportion in forest stands. The addition of dead wood in a manipulation experiment had a slight positive effect on cocoon predation, while dead wood removal caused a clear decrease in predation rate, and the decrease was more pronounced when the proportion of pine increased. Our results show that habitat heterogeneity affects predation by generalist predators on herbivorous insects. This knowledge could be applied to reduce the risk of insect outbreaks by applying management methods that increase heterogeneity in perennial systems such as forests and orchards, thus decreasing the levels of insect damage.