Abundance and distribution of organic mound-building ants of the Formica rufa group in Yellowstone National Park

Red wood ants (Formica rufa group) are ubiquitous in many conifer and mixed‐conifer forests of northern Europe and Asia. In contrast, relatively little is known about the abundance and distribution of the 24 North American F. rufa group species. As ants are important components of most soil invertebrate communities and are considered ecosystem engineers that alter the flow of energy and nutrients through terrestrial systems, it is important to gain information on their distribution and abundance. We conducted a survey for red wood ant mounds in Yellowstone National Park (YNP), Wyoming/Montana, USA, where human disturbance has been kept to a minimum for over 130 years. We found a total of 85 red wood ant mounds (0.11 to 0.17 mounds/ha) on 327 km of roads and 180 km of the hiking trails we surveyed. The occurrence of ant mounds was higher then expected by random distribution at elevations between 1600 and 2400 m, annual precipitation of 250 to 760 mm, middle and late successional lodgepole pine, late successional Douglas fir forest and non‐forested grassland/sagebrush prairie vegetation. Additionally, mounds were clustered in gently sloped not north‐exposed locations and in areas that had not recently burned. Most of the mounds detected were inhabited by Formica obscuripes Forel, which occupied 94% of the mounds sampled. Based on a multi‐criteria binary Geographic Information System model that we developed, we found that ant mounds were to be expected with a high probability in less then 1% of the YNP area. These results together with the detected low density and small size of the red wood ant mounds within the study area suggest that these insects have a much lower impact on invertebrate biodiversity and ecosystem processes, such as forest productivity and carbon and nutrient cycling on the ecosystem scale compared with their counterparts in European or Asian systems.     

RETRACTED ARTICLE: Interspecific competition among ants in the boreal forest: Testing predictions from a linear hierarchical competition model

Competitive interactions often play an important role in local community structure, and particularly so in ant communities. We test predictions derived from a competition model proposed for Scandinavian ant communities by comparing ant nest densities inside and outside fifty randomly-selected territories of competitively dominant wood ants within 30 km of Ume?, northern Sweden. As predicted by the model, competitively-intermediate encounter species, as well as other territorial species, showed complementary occurrences with both studied wood ants, Formica aquilonia Yarrow and F. lugubris Zett.. In contrast, complementary abundances, i.e. lower density of competitively-inferior submissive species in the presence of wood ants, as predicted by the model, was not supported for all submissive species. Of the two studied wood ant species, the nest density of submissive species was negatively correlated only with F. aquilonia. Submissives as a group, as well as Myrmica-species, showed complementary abundances with F. aquilonia, but one Leptothorax and two Serviformica species had higher nest densities in the presence of this competitively-dominant species. We propose that, for Leptothorax, these deviations from the model predictions may be because of limited niche overlap with dominant wood ants, a small worker force and a timid behaviour which does not elicit aggression in wood ants. For the two Serviformica species, we propose a combination of protection against social parasites, inter-specific social control performed by F. aquilonia, and dominance relationships between competitively-inferior submissive species as reasons for the higher nest density inside F. aquilonia territories. Monogyny, and thereby smaller nests, lower worker force, smaller territory (as shown by this study) in F. lugubris, as compared to the polygyny in F. aquilonia, may also help explain the differences in their effects on subordinate species. Our analyses indicate that the linear competition hierarchy model proposed for Scandinavian ants accurately predicts the outcome of interaction between and community composition for dominant territorial and encounter species, but that it needs refinement with respect to the relationship between territorial and submissive species and the resulting community composition. Further studies are needed, especially addressing the complex relationships between these latter groups, and the effects of different competitively-dominant wood ants, to determine the mechanisms determining the outcome of these relationships and to more accurately predict community composition. Received 11 June 2007; revised 11 September 2007; accepted 17 September 2007.     

A size-distance relation in Homoptera-tending thatch ants (Formica obscuripes,Formica planipilis)

Summary In 20 of 22 study colonies, Homoptera-tending workers of the thatch antsFormica obscuripes andF. planipilis exhibited a significant size-distance relation, with smallest tenders working closest to the primary nest. No such relation was observed in scavengers working over the same range of distances in any of the 22 colonies. This paper describes the size-distance relation in tenders quantitatively, examines its expression given variation in Homoptera distribution, and compares it to similar patterns observed in otherF. rufa-group species. Three hypotheses on the adaptive significance of the size-distance relation are discussed: that the relation increases energetic efficiency, improves colony defense, or decreases the rate of getting lost in smaller tenders.     

Rhizospheric and heterotrophic components of soil respiration in six Chinese temperate forests

Partitioning soil respiration (R_S) into heterotrophic (R_H) and rhizospheric (R_R) components is an important step for understanding and modeling carbon cycling in forest ecosystems, but few studies on R_R and R_H exist in Chinese temperate forests. In this study, we used a trenching plot approach to partition R_S in six temperate forests in northeastern China. Our specific objectives were to (1) examine seasonal patterns of soil surface CO₂ fluxes from trenched (R_T) and untrenched plots (R_UT) of these forests; (2) quantify annual fluxes of R_S components and their relative contributions in the forest ecosystems; and (3) examine effects of plot trenching on measurements of R_S and related environmental factors. The R_T maximized in early growing season, but the difference between R_UT and R_T peaked in later summer. The annual fluxes of R_H and R_R varied with forest types. The estimated values of R_H for the Korean pine (Pinus koraiensis Sieb. et Zucc.), Dahurian larch (Larix gmelinii Rupr.), aspen‐birch (Populous davidiana Dode and Betula platyphylla Suk.), hardwood (Fraxinus mandshurica Rupr., Juglans mandshurica Maxim. and Phellodendron amurense Rupr.), Mongolian oak (Quercus mongolica Fisch.) and mixed deciduous (no dominant tree species) forests averaged 89, 196, 187, 245, 261 and 301 g C m⁻² yr⁻¹, respectively; those of R_R averaged 424, 209, 628, 538, 524 and 483 g C m⁻² yr⁻¹, correspondingly; calculated contribution of R_R to R_S (RC) varied from 52% in the larch forest to 83% in the pine forest. The annual flux of R_R was strongly correlated to biomass of roots <0.5 cm in diameter, while that of R_H was weakly correlated to soil organic carbon concentration at A horizon. We concluded that vegetation type and associated carbon metabolisms of temperate forests should be considered in assessing and modeling R_S components. The significant impacts of changed soil physical environments and substrate availability by plot trenching should be appropriately tackled in analyzing and interpreting measurements of R_S components.     

Ideal phenotypes and mismatching haplotypes – errors of mtDNA treeing in ants (Hymenoptera: Formicidae) detected by standardized morphometry

A total of 401 nest samples of Formica lugubris Zetterstedt, F. pratensis Retzius, F. aquilonia Yarrow, F. rufa Linnaeus, and F. polyctena Förster, covering the entire Palaearctic range of these species and including 2100 individual workers, was phenotypically investigated by a system of standardized morphometry, pairwise removal of allometric variance, and canonical discriminant functions. A mitochondrial DNA fragment including the cytochrome b gene was sequenced in 148 samples from basically the same range. In the more difficult F. pratensis vs. F. lugubris case, the phenotypic system correctly determined 99.6% of all nest samples, and 95.1% with p<0.05. In all other pairwise species discriminations any nest sample was correctly determined with p<0.01, and three samples with hybrids F. rufa×lugubris were identified. At four localities in the Pyrenees and the Urals, 9 samples with F. pratensis phenotypes (7 of them ideal) but F. lugubris mtDNA haplotypes could be identified, resulting in 14.5% of phenotype/haplotype mismatches. A local dominance of this mismatch combination was observed at one Pyrenean and one Ural locality. There was no indication of an F. pratensis haplotype associated with an F. lugubris phenotype. One ideal F. polyctena phenotype was associated with an F. aquilonia haplotype in a sample from the Urals, and one ideal F. aquilonia phenotype was combined with an F. lugubris haplotype in a sample from central Siberia, resulting in overall phenotype/haplotype mismatch frequencies of 12.5% and 11.1%, respectively. We conclude that all these samples cannot represent actual F₁ hybrids but are the result of hybridizations in the past followed by unidirectional purging of the nuclear genome. Whether this process of purging worked very fast or over longer periods of population history, and whether or not it was complete or incomplete, cannot be assessed from the available information. These facts of hybridizing in two thirds of the W Palaearctic wood ant species, of extreme regional hybrid frequencies (up to 26%), of unidirectional purging of nDNA associated with mismatching mtDNA haplotypes, and of occasional achievement of local dominance of these mismatch combinations, may serve as urgent warning not to perform isolated mtDNA phylogenetic studies without a geographically and locally wide sampling basis and without control by nDNA information or reliable phenotypic determination. The latter two systems definitely have superior significance when conflicts with mtDNA indications arise.     

CO2 fertilization in temperate FACE experiments not representative of boreal and tropical forests

Results from free-air CO₂ enrichment (FACE) experiments in temperate climates indicate that the response of forest net primary productivity (NPP) to elevated CO₂ might be highly conserved across a broad range of productivities. In this study, we show that the LPJ-GUESS dynamic vegetation model reproduces the magnitude of the NPP enhancement at temperate forest FACE experiments. A global application of the model suggests that the response found in the experiments might also be representative of the average response of forests globally. However, the predicted NPP enhancement in tropical forests is more than twice as high as in boreal forests, suggesting that currently available FACE results are not applicable to these ecosystems. The modeled geographic pattern is to a large extent driven by the temperature dependence of the relative affinities of the primary assimilation enzyme (Rubisco) for CO₂ and O₂.     

Soil respiration in six temperate forests in China

Scaling soil respiration (R_S), the major CO₂ source to the atmosphere from terrestrial ecosystems, from chamber‐based measurements to ecosystems requires studies on variations and correlations of R_S from various biomes and across geographic regions. However, few studies on R_S are available for Chinese temperate forest despite the importance of this forest in the national and global carbon budgets. In this study, we conducted 18‐month R_S measurements during 2004–2005 in six temperate forest types, representing the typical secondary forest ecosystems across various site conditions in northeastern China: Mongolian oak (Quercus mongolica Fisch.), aspen‐birch (Populous davidiana Dode and Betula platyphylla Suk.), mixed deciduous (no dominant tree species), hardwood (dominated by Fraxinus mandshurica Rupr., Juglans mandshurica Maxim., and Phellodendron amurense Rupr.) forests, Korean pine (Pinus koraiensis Sieb. et Zucc.) and Dahurian larch (Larix gmelinii Rupr.) plantations. Our specific objectives were to: (1) explore relationships of R_S against soil temperature and water content for the six forest ecosystems, (2) quantify annual soil surface CO₂ flux and its relations to belowground carbon storage, (3) examine seasonal variations in R_S and related environmental factors, and (4) quantify among‐ and within‐ecosystem variations in R_S. The R_S was positively correlated to soil temperature in all forest types, and was significantly influenced by the interactions of soil temperature and water content in the pine, larch, and mixed deciduous forests. The sensitivity of R_S to soil temperature at 10 cm depth (Q₁₀) ranged from 2.61 in the oak forest to 3.75 in the aspen‐birch forests. The Q₁₀ tended to increase with soil water content until reaching a threshold, and then decline. The annual R_S for the larch, pine, hardwood, oak, mixed deciduous, and aspen‐birch forests averaged 403, 514, 781, 785, 786, and 813 g C m^?2 yr^?1, respectively. The annual R_S of the broadleaved forests was 72% greater than that of the coniferous forests. The annual R_S was positively correlated to soil organic carbon (SOC) concentration at O horizon (R²=0.868) and total biomass of roots <0.5 cm in diameter (R²=0.748). The coefficient of variation (CV) of R_S among forest types averaged 25% across the 18‐month measurements. The CV of R_S within plots varied from 20% to 27%, significantly (P<0.001) greater than those among plots (9–15%), indicating the importance of the fine‐scaled heterogeneity in R_S. This study emphasized that variations in soil respiration and potential sampling bias should be appropriately tackled for accurate soil CO₂ flux estimates.     

CO2 balance of boreal, temperate, and tropical forests derived from a global database

Terrestrial ecosystems sequester 2.1 Pg of atmospheric carbon annually. A large amount of the terrestrial sink is realized by forests. However, considerable uncertainties remain regarding the fate of this carbon over both short and long timescales. Relevant data to address these uncertainties are being collected at many sites around the world, but syntheses of these data are still sparse. To facilitate future synthesis activities, we have assembled a comprehensive global database for forest ecosystems, which includes carbon budget variables (fluxes and stocks), ecosystem traits (e.g. leaf area index, age), as well as ancillary site information such as management regime, climate, and soil characteristics. This publicly available database can be used to quantify global, regional or biome‐specific carbon budgets; to re‐examine established relationships; to test emerging hypotheses about ecosystem functioning [e.g. a constant net ecosystem production (NEP) to gross primary production (GPP) ratio]; and as benchmarks for model evaluations. In this paper, we present the first analysis of this database. We discuss the climatic influences on GPP, net primary production (NPP) and NEP and present the CO₂ balances for boreal, temperate, and tropical forest biomes based on micrometeorological, ecophysiological, and biometric flux and inventory estimates. Globally, GPP of forests benefited from higher temperatures and precipitation whereas NPP saturated above either a threshold of 1500 mm precipitation or a mean annual temperature of 10 °C. The global pattern in NEP was insensitive to climate and is hypothesized to be mainly determined by nonclimatic conditions such as successional stage, management, site history, and site disturbance. In all biomes, closing the CO₂ balance required the introduction of substantial biome‐specific closure terms. Nonclosure was taken as an indication that respiratory processes, advection, and non‐CO₂ carbon fluxes are not presently being adequately accounted for.     

Arboreal search in ants: Search on branches (Hymenoptera: Formicidae)

Ants (Formicidae) perform two distinct search behaviors for resources: on the ground they use irregular, almost random alternating looping, and on branches and leaves they resort to outline-tracing (arboreal systematic search), whereby the individual systematically turns to one side at bifurcations and to the opposite side when turning about at end points. Experiments with searching Formica pallidefulva and Crematogaster cerasion artificial stick mazes under seminatural conditions demonstrated that bifurcations and end points only trigger turn decisions, whereas an intrinsic mechanism specifies the handedness of such turns. Arboreal homing differs from arboreal searching by a much stronger tendency to rectify paths by counterturning. The theory is advanced that searching on branches by outline-tracing is evolutionarily derived from ranging search by superposing a sustained intrinsic turn bias and by suppressing random turns.     

The likely impact of elevated [CO2], nitrogen deposition, increased temperature and management on carbon sequestration in temperate and boreal forest ecosystems: a literature review

Temperate and boreal forest ecosystems contain a large part of the carbon stored on land, in the form of both biomass and soil organic matter. Increasing atmospheric [CO2], increasing temperature, elevated nitrogen deposition and intensified management will change this C store. Well documented single-factor responses of net primary production are: higher photosynthetic rate (the main [CO2] response); increasing length of growing season (the main temperature response); and higher leaf-area index (the main N deposition and partly [CO2] response). Soil organic matter will increase with increasing litter input, although priming may decrease the soil C stock initially, but litter quality effects should be minimal (response to [CO2], N deposition, and temperature); will decrease because of increasing temperature; and will increase because of retardation of decomposition with N deposition, although the rate of decomposition of high-quality litter can be increased and that of low-quality litter decreased. Single-factor responses can be misleading because of interactions between factors, in particular those between N and other factors, and indirect effects such as increased N availability from temperature-induced decomposition. In the long term the strength of feedbacks, for example the increasing demand for N from increased growth, will dominate over short-term responses to single factors. However, management has considerable potential for controlling the C store.     

Effect of soil water stress on soil respiration and its temperature sensitivity in an 18-year-old temperate Douglas-fir stand

We analyzed 17 months (August 2005 to December 2006) of continuous measurements of soil CO₂ efflux or soil respiration (R_S) in an 18‐year‐old west‐coast temperate Douglas‐fir stand that experienced somewhat greater than normal summertime water deficit. For soil water content at the 4 cm depth (θ) > 0.11 m³ m^?3 (corresponding to a soil water matric potential of ?2 MPa), R_S was positively correlated to soil temperature at the 2 cm depth (T_S). Below this value of θ, however, R_S was largely decoupled from T_S, and evapotranspiration, ecosystem respiration and gross primary productivity (GPP) began to decrease, dropping to about half of their maximum values when θ reached 0.07 m³ m^?3. Soil water deficit substantially reduced R_S sensitivity to temperature resulting in a Q₁₀ significantly < 2. The absolute temperature sensitivity of R_S (i.e. dR_S/dT_S) increased with θ up to 0.15 m³ m^?3, above which it slowly declined. The value of dR_S/dT_S was nearly 0 for θ < 0.08 m³ m^?3, thereby confirming that R_S was largely unaffected by temperature under soil water stress conditions. Despite the possible effects of seasonality of photosynthesis, root activity and litterfall on R_S, the observed decrease in its temperature sensitivity at low θ was consistent with the reduction in substrate availability due to a decrease in (a) microbial mobility, and diffusion of substrates and extracellular enzymes, and (b) the fraction of substrate that can react at high T_S, which is associated with low θ. We found that an exponential (van't Hoff type) model with Q₁₀ and R₁₀ dependent on only θ explained 92% of the variance in half‐hourly values of R_S, including the period with soil water stress conditions. We hypothesize that relating Q₁₀ and R₁₀ to θ not only accounted for the effects of T_S on R_S and its temperature sensitivity but also accounted for the seasonality of biotic (photosynthesis, root activity, and litterfall) and abiotic (soil moisture and temperature) controls and their interactions.     

Contrasting soil respiration in young and old-growth ponderosa pine forests

Three years of fully automated and manual measurements of soil CO₂ efflux, soil moisture and temperature were used to explore the diel, seasonal and inter‐annual patterns of soil efflux in an old‐growth (250‐year‐old, O site) and recently regenerating (14‐year‐old, Y site) ponderosa pine forest in central Oregon. The data were used in conjunction with empirical models to determine which variables could be used to predict soil efflux in forests of contrasting ages and disturbance histories. Both stands experienced similar meteorological conditions with moderately cold wet winters and hot dry summers. Soil CO₂ efflux at both sites showed large inter‐annual variability that could be attributed to soil moisture availability in the deeper soil horizons (O site) and the quantity of summer rainfall (Y site). Seasonal patterns of soil CO₂ efflux at the O site showed a strong positive correlation between diel mean soil CO₂ efflux and soil temperature at 64 cm depth whereas diel mean soil efflux at the Y site declined before maximum soil temperature occurred during summer drought. The use of diel mean soil temperature and soil water potential inferred from predawn foliage water potential measurements could account for 80% of the variance of diel mean soil efflux across 3 years at both sites, however, the functional shape of the soil water potential constraint was site‐specific. Based on the similarity of the decomposition rates of litter and fine roots between sites, but greater productivity and amount of fine litter detritus available for decomposition at the O site, we would expect higher rates of soil CO₂ efflux at the O site. However, annual rates were only higher at the O site in one of the 3 years (597 ± 45 vs. 427 ± 80 g C m^?2). Seasonal patterns of soil efflux at both sites showed influences of soil water limitations that were also reflected in patterns of canopy stomatal conductance, suggesting strong linkages between above and below ground processes.     

Selection harvest in temperate deciduous forests: impact on herb layer richness and composition

Herb layer richness and composition of four forest types and three different management treatments were investigated in 16 deciduous forest stands of northern Germany. Specifically, we compared the species richness and composition occurring in mature forest stands that were single-tree and group selection harvested to those in unmanaged reference stands. Mean species richness of all herb layer species increased significantly with increasing harvest severity. When analyzing plant groups separately, it became obvious that this overall pattern was not consistent. While a negative relationship was detected between vernal herb richness and harvest severity, group selection harvest significantly increased species richness of summer herbaceous forest species and generalists. Woody species richness was not related to harvest severity. Community composition of the spring aspect was not significantly affected by selection harvest, whereas herb layer species composition in the summer aspect differed significantly among the three harvest treatments. A dominance of highly competitive shrub and generalist species was confined to some parts of the most intensively harvested stands. Overall, our results indicate that the herb layer community was not severely adversely affected by selection harvest at the intensities used in the studied stands. It is suggested that selection harvest systems may be feasible tools with which to conserve local forest vascular plant diversity and at the same time to meet the demand for timber products. However, information about forest history and the implementation of the selection harvest system are basic requirements when interpreting the results of studies on understorey response to selection harvest.     

Organic carbon cycling in Taylor Valley, Antarctica: quantifying soil reservoirs and soil respiration

Organic carbon reservoirs and respiration rates in soils have been calculated for most major biomes on Earth revealing patterns related to temperature, precipitation, and location. Yet data from one of the Earth's coldest, driest, and most southerly soil ecosystems, that of the McMurdo Dry Valleys of Antarctica, are currently not a part of this global database. In this paper, we present the first regional calculations of the soil organic carbon reservoirs in a dry valley ecosystem (Taylor Valley) and report measurements of CO₂ efflux from Antarctic soils. Our analyses indicate that, despite the absence of visible accumulations of organic matter in most of Taylor Valley's arid soils, this soil environment contained a significant percentage (up to 72%) of the seasonally unfrozen organic carbon reservoir in the terrestrial ecosystem. Field measurements of soil CO₂‐efflux in Taylor Valley soils were used to evaluate biotic respiration and averaged 0.10 ± 0.08 μmol CO₂ m^?2 s^?1. Laboratory soil microcosms suggested that this respiration rate was sensitive to increases in temperature, moisture, and carbon addition. Finally, a steady‐state calculation of the mean residence time for organic carbon in Taylor Valley soils was 23 years. Because this value contradicts all that is currently known about carbon cycling rates in the dry valleys, we suggest that the dry valley soil carbon dynamics is not steady state. Instead, we suggest that the dynamic is complex, with at least two (short‐ and long‐term) organic carbon reservoirs. We also suggest that organic carbon in the dry valley soil environment may be more important, and play a more active role in long‐term ecosystem processes, than previously believed.     

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

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

Effects of ants on water and soil losses from organically-managed citrus orchards in eastern Spain

Ants can play a key role in the erosion processes on agriculture land by modifying soil properties and increasing macropore flow. Ants are abundant in organically-managed orchards in the Mediterranean region due to climate conditions, no-till practices, no pesticide use, and the resulting vegetation cover. In order to determine the effect of ants on soil and water losses from these orchards growing on moderately-sloped land (4–8%), forty 1.0 m² plots (20 with ants mounts and 20 without ants — controls) were established during the summer of 2007. A rainfall simulator was used to apply 78 mm of water to each plot over a one-hour period, equivalent to a 20-year return-period thunderstorm. Runoff was collected at 1-minute intervals and sediment concentration measured every 10 minutes. Sediment concentrations were 300% higher on plots with ant mounds, but runoff rates were similar to the plots without ants. Average soil erosion rates averaged 41 kg ha⁻¹ h⁻¹ on the ant plots and 13 kg ha⁻¹ h⁻¹ on the control plots. The low erosion rates are due to the effect of the vegetation and litter cover in this organically-managed soil, which were little impacted by ant activity at the pedon scale.     

A genetic model for disruptive selection on colony social organisation,reproduction, and ecotype distribution in wood ants inhabiting different woodland habitats

Summary Wood ants of theFormica rufa species complex are typically bound to woodland habitats and their eventual colonization of more open habitats depends on the presence of a sufficient number of woody plants. Computer simulations of the long-term development of large wood ant populations in woodland systems of differing structure were performed. A cellular, stochastic automation model simulated the alternatives of a compact, coherent woodland system and a fragmented, coarse-grained woodland system. The simulation of a compact woodland system gave evidence that disruptive selection alone may be sufficient to produce two distinct ectotypes, which are known asFormica rufa andFormica polyctena, even if there is a full fertility of the crosses. The simulation of a fragmented, coarse-grained woodland system has supported the view, that a high local frequency of nests with intermediate phenotypes might be explainable by a particular woodland pattern which favour mixed strategies. It was shown that queen dominance and intraspecific social parasitism are likely to be important factors in the dynamics of large wood ant populations.     

The importance of phenology for the evaluation of impact of climate change on growth of boreal, temperate and Mediterranean forests ecosystems: an overview

An overview is presented of the phenological models relevant for boreal coniferous, temperate-zone deciduous and Mediterranean coniferous forest ecosystems. The phenology of the boreal forests is mainly driven by temperature, affecting the timing of the start of the growing season and thereby its duration, and the level of frost hardiness and thereby the reduction of foliage area and photosynthetic capacity by severe frost events. The phenology of temperate-zone forests is also mainly driven by temperature. Since temperate-zone forests are mostly mixed-species deciduous forests, differences in phenological response may affect competition between tree species. The phenology of Mediterranean coniferous forests is mainly driven by water availability, affecting the development of leaf area, rather than the timing of phenological events. These phenological models were subsequently coupled to the process-based forest model FORGRO to evaluate the effect of different climate change scenarios on growth. The results indicate that the phenology of each of the forest types significantly affects the growth response to a given climate change scenario. The absolute responses presented in this study should, however, be used with caution as there are still uncertainties in the phenological models, the growth models, the parameter values obtained and the climate change scenarios used. Future research should attempt to reduce these uncertainties. It is recommended that phenological models that describe the mechanisms by which seasonality in climatic drivers affects the phenological aspects of trees should be developed and carefully tested. Only by using such models may we make an assessment of the impact of climate change on the functioning and productivity of different forest ecosystems. Received: 21 October 1999 / Revised: 10 May 2000 / Accepted: 10 May 2000