Vole population dynamics: factors affecting peak densities and amplitudes of annual population fluctuations of<Emphasis Type="Italic">Microtus pennsylvanicus</Emphasis>

We studied factors affecting peak densities and amplitudes of fluctuation during 20 annual population fluctuations ofMicrotus pennsylvanicus Ord, 1815 in alfalfa and bluegrass habitats over a 25-year period. Survival was correlated with population density over the 25 years and was the most consistent variable associated with stoppage of population growth. Although not correlated with population density over the 25 years, a decline in the proportion of reproductively active adult females contributed to cessation of growth of population fluctuations that peaked in late autumn-winter, and to cessation of growth of eight of eleven population fluctuations that peaked during summer-early autumn. We conclude variation in survival to be the primary factor affecting peak densities and amplitudes of population fluctuation ofM. pennsylvanicus.     

Intraspecific genetic polymorphism of Russian sturgeon <Emphasis Type="Italic">Acipenser gueldenstaedtii</Emphasis>

Three populations (Azov, Caspian, and Black Sea) of Russian sturgeon Acipenser queldenstaedtii were tested for polymorphism at nuclear (RAPD and microsatellites) and mitochondrial (PCR identification of two mitotypes) markers. In addition, morphometric analysis of he representatives of Azov population was carried out. According to the morphological characters, the Black Sea population occupied an intermediate position between the Caspian and Azov populations, reflecting the phylogeography of this species. In agreement with the morphometric data, genetic distances (the data of STR analysis) also placed the Black Sea population between the Caspian and Azov populations (F _ST = 0.058 and 0.043). The genetic distance between the Azov and Caspian population was somewhat higher (F _ST = 0.070). The highest allelic polymorphism at four microsatellite loci was found observed in Caspian population, while the lowest polymorphism was in the Sea of Azov. RAPD analysis distinguished high polymorphism within the populations, although it was not feasible for interpopulation analysis. Using the method differentiating the “baerii-like” and typical “gueldenstaedtii” mitotypes, the absence of the “baerii-like” marker in the Black Sea population was demonstrated. The frequency of this marker in Caspian and Azov populations constituted 31.1 and 1.8%, respectively. Possible evolutionary reasons for the interpopulation differences observed are discussed.     

Landscape pattern and genetic structure of a yellow-necked mouse <Emphasis Type="Italic">Apodemus flavicollis</Emphasis> population in north-eastern Poland

Reduced connectivity among local populations inhabiting a spatially heterogeneous landscape may restrict gene flow and thus contribute to diminished genetic variation within a population. The aim of this study was to determine the role of geographic distance and habitat barriers in developing genetic structure of a yellow-necked mouse Apodemus flavicollis (Melchior, 1834) population, taking into consideration the spatial organization of the landscape. A field study was carried out in two plots located in NE Poland that differed considerably in terms of the scale of habitat fragmentation: (1) a continuous forest complex, and (2) a mosaic of smaller forest habitats. The plots were separated by a water barrier comprised of a chain of lakes. DNA samples from a total of 654 individuals were examined by microsatellite analysis (5 loci). The results showed that the yellow-necked mouse population was characterized by a poorly pronounced genetic structure throughout the study area, although the statistical significance of F _ST for most location pairs indicated that gene flow in the area was not free. The division of the mouse population into three genetically distinct groups clearly demonstrated the significant role of water bodies as a natural barrier effectively hindering free movement of animals and thus gene flow. Analysis of the genetic structure of the mouse population throughout the study area and also within the distinguished groups indicated that the entire study population may be considered as a single metapopulation. Our results suggest that geographic distance alone is not the predominant factor affecting the genetic structure of population, but in the mosaic landscape the relative isolation of individual forest fragments, and barriers hindering movements of individuals and limiting gene flow among local populations played a much more important role.     

No evident spatial genetic structuring in the rapidly declining Black-tailed Godwit <Emphasis Type="Italic">Limosa limosa</Emphasis><Emphasis Type="Italic">limosa</Emphasis> in The Netherlands

With 40% of the European Black-tailed Godwit population breeding in The Netherlands, this country harbours internationally significant numbers of this species. However, ongoing agricultural intensification has resulted in the fragmentation of the population and drastic population declines since 1967. By establishing genetic diversity, genetic differentiation and gene flow on the basis of 12 microsatellites, we investigated whether the population genetic structure of the Dutch Black-tailed Godwit bears the marks of these changes. Genetic diversity appeared to be moderate, and Bayesian model-based analysis of individual genotypes revealed no clustering in the Dutch populations. This was supported by pairwise F_ST values and AMOVA, which indicated no differentiation among the nine breeding areas. Gene flow estimates were larger than “one migrant per generation” between sample locations, and no isolation by distance was demonstrated. Our results indicate the maintenance of moderate levels of genetic diversity and genetic connectivity between breeding sites throughout the Dutch Black-tailed Godwit breeding population. We suggest that the Dutch Black-tailed Godwit breeding areas should be managed as a single panmictic unit, much as it is presently done.     

Estimation of seasonal dynamics of number and age structure of South Baikal natural population of <Emphasis Type="Italic">Epischura baicalensis</Emphasis> sars using ANOVA

The seasonal dynamics of number and age structure of natural population of Epischura baicalensis Sars, 1900 (Copepoda, Calanoida) from South Baikal was studied using the materials of ten-day sampling of zooplankton samples in the water layer of 0–250 min 2001–2004. Using ANOVA we determined the statistical significance of seasonal changes in number and ratio of nauplius larvae, copepodid larvae, and adults. The method of sampling reduction was used to identify the three phases of population cycle: growth, peak, and depression of number, corresponding to three biological seasons. The importance of the obtained data is discussed considering the regulating mechanisms for numbers of population of E. baicalensis     

Detecting and managing an overgrazing-drought synergism in the threatened <Emphasis Type="Italic">Echeveria longissima</Emphasis> (Crassulaceae): the role of retrospective demographic analysis

Echeveria longissima, a threatened herb whose habitat has been severely overgrazed and eroded, was studied for three years in a currently grazed and a fenced area. Matrix population models were used to assess if livestock elimination provides a proper management strategy. The merits of retrospective perturbation analyses in terms of management planning have been debated. Nevertheless, they may prove useful when applied in combination with exclosures because they may detect the effects of anthropogenic disturbance on population dynamics. Thus, the results of retrospective and prospective methods were compared. A rapid decrease in population size was projected in both areas, even though it was faster in the exposed one. The demographic processes that were favourable or detrimental in a given year were magnified outside of the fence, but buffered in the exclosure, showing a strong drought-disturbance synergism. Thus, the largest difference in the population growth rate λ between areas was observed in the driest year. Higher nurse-plant density inside the fence seems to alleviate drought effects. The use of prospective analysis alone may lead to erroneous management decisions, since the highest elasticities corresponded to transitions that were favoured by human activities. While allowing for an increased λ in the short term, intervention aimed at increasing these transitions further without attending others that are lessened by disturbance may introduce large changes in the population dynamics, with negative long-term consequences. Retrospective methods can detect which processes have been altered by disturbance and its synergisms, so we may more efficiently restore healthy population dynamics.     

Demographic and genetic status of an isolated population of bog turtles (<Emphasis Type="Italic">Glyptemys muhlenbergii</Emphasis>): implications for managing small populations of long-lived animals

In this study, we sought to determine the population stability and genetic diversity of one isolated population of the federally-threatened bog turtle (Glyptemys muhlenbergii) in North Carolina. Using capture–recapture data, we estimated adult survival and population growth rate from 1992 to 2007. We found that the population decreased from an estimated 36 adult turtles in 1994 to approximately 11 adult turtles in 2007. We found a constant adult survival of 0.893 (SE = 0.018, 95% confidence interval, 0.853–0.924) between 1992 and 2007. Using 18 microsatellite markers, we compared the genetic status of this population with five other bog turtle populations. The target population displayed allelic richness (4.8 ± 0.5) and observed heterozygosity (0.619 ± 0.064) within the range of the other bog turtle populations. Coalescent analysis of population growth rate, effective population size, and timing of population structuring event also indicated the genetics of the target population were comparable to the other populations studied. Estimates of effective population size were a proportion of the census size in all populations except the target population, in which the effective population size was larger than the census size (30 turtles vs. 11 turtles). We attribute the high genetic diversity in the target population to the presence of multiple generations of old turtles. This study illustrates that the demographic status of populations of long-lived species may not be reflected genetically if a decline occurred recently. Consequently, the genetic integrity of populations of long-lived animals experiencing rapid demographic bottlenecks may be preserved through conservation efforts effective in addressing demographic problems.     

Low genetic variability of <Emphasis Type="Italic">Rhinolophus mehelyi</Emphasis> (Mehely's horseshoe bat) in Romania

Rhinolophus mehelyi (Mehely's horseshoe bat) is a vulnerable species with an increasingly fragmented distribution. In Romania, populations of R. mehelyi have experienced a dramatic decline over the past 50 years, and the current population size is estimated at only 100 adult individuals inhabiting almost exclusively the Limanu cave. In the present study, we investigated the genetic consequences of population decline for the viability of the remaining population of R. mehelyi in Romania. We sequenced and analyzed a 359-bp fragment of the mitochondrial control region from the only known Romanian population and compared it with two geographically close colonies from Bulgaria. A single haplotype was found in the Romanian population compared to 10 in the Bulgarian population, suggesting genetic isolation.     

Genetic monitoring and effects of stocking practices on small <Emphasis Type="Italic">Cyprinus carpio</Emphasis> populations

The ability to detect genetic differences both in space and time is crucial for conserving genetic variation. It can reveal genetic diversity and genetic composition changes of declining native populations that are supported through stocking with captive bred individuals. The present study was designed to analyse the temporal stability of a declining common carp (Cyprinus carpio) population from Lake Volvi (North Greece). Polymorphism was evaluated using seven microsatellite loci at two sampling time points (separated by 12 years). The genetic variability of four additional populations (from two rivers and two lakes) in Northern Greece was also investigated for comparison. Heterozygosity values (0.692–0.868) and allelic richness (8.530–11.148) were high for all studied populations and comparable to other European populations. However, the analysis of temporal common carp samples from Lake Volvi revealed a significant change in their genetic composition and admixture analysis demonstrated significant introgression of stocked individuals into the native population. Both temporal and point estimate methods revealed low effective size (Ne = 61–171.3) for this population, possibly a result of an ancient genetic bottleneck that led to population decline and/or recent anthropogenic interventions. This low Ne has rendered the native population vulnerable to alteration of its genetic composition. Our study demonstrates that enhancement programs should be applied cautiously, especially for small populations. Moreover, it underlines the need for temporal analyses, which may contribute to the evaluation of previous management policies and to future decision making.     

The state of populations of <Emphasis Type="Italic">Lernaea cyprinacea</Emphasis> L. (copepoda: lernaeidae) and crucian carp <Emphasis Type="Italic">Carassius carassius</Emphasis> L. in Dlinnoye Lake

The state of populations of the crucian carp Carassius carassius L. and the copepod parasite Lernaea cyprinacea L. was studied after 10 years of a sharp decrease in their abundance. Only one generation of Lernaea cyprinacea species was recorded each year. The alteration of the parasite generation occurred in the second half of July. Changes in the age structure of the copepod population did not affect the distribution pattern of its abundance in the host population, but the values of the K exponent changed in accordance with alterations of the parasite generations. By 2007 the sex ratio in the population became equal, males and females differed in their lengths and weights, and three dimensional groups were found in the catches. It is thought that males and females of the crucian carp have different roles in the maintenance of the L. cyprinacea abundance. The dominating fish sizes and the level of the host infection by copepods remained the same as in 1997–2003.     

Long-term monitoring of a Bengal tiger (<Emphasis Type="Italic">Panthera tigris tigris</Emphasis>) population in a human-dominated landscape of Central India

Long-term monitoring of tiger population is essential in any human-dominated landscape as globally their population is showing a declining trend due to unrelenting stress or stochastic events. In view of that, Bengal tiger population was monitored in Pench Tiger Reserve (Pench), Madhya Pradesh of Central India between May 2006 and April 2013. The population, age-sex ratios, survival rate, growth rate, and recruitment pattern of adult (≥36 month), subadult (≥18 to 36 month), and cubs (≤15 month) were studied using camera trap and radio-telemetry techniques. Overall, tiger density ± SE/100 km² using Maximum Likelihood Spatial Explicit Capture Recapture method was 5.5 ± 1.6 in first trapping period or year and 3.7 ± 0.8 in last or seventh trapping year. The estimated survival rate of all tiger (n = 66) was 0.66 (±0.04), whereas mean annual growth rate (±SE) was 1.15 (±0.11) i.e. 15%. The sex ratio (male/female) found to be female biased and more than 80% of overall recruitment was contributed by female tiger. The dispersal of subadults in the tiger population contributed gradual fluctuation in survival rate and annual growth rate, as resident adult tiger population remained almost stable over the study period. As the overall tigers are surviving in a small population size and operating at carrying capacity, the important corridors connecting with neighboring population of Kanha Tiger Reserve and Satpura Tiger Reserve may ensure constant emigration and immigration to reduce the chances of genetic drift or inbreeding in the Pench tiger population.     

Tolerance of a ruminant ciliate <Emphasis Type="Italic">Entodinium caudatum</Emphasis> against mercury,copper and chromium

The effects of three heavy metal cations, mercury (II), copper (II), and chromium (VI), on the growth of the rumen ciliate Entodinium caudatum in vitro culture was studied. The E. caudatum culture was challenged by HgCl₂, CuCl₂, and K₂Cr₂O₇ for a period of 4 days. The tested concentrations of mercury (II) and copper (II) were 1, 5, 10, 20, 50 mg/L and 2, 10, 20, 40 mg/L for chromium (VI) at single dose with either untreated or inhibited bacterial co-culture population. Effective metal concentrations required to reduce ciliate growth by 50% (EC₅₀) for mercury (II), copper (II), and chromium (VI) either with untreated or inhibited bacterial co-culture population after 24 h of metal application were 24, 20, and 21 or 15, 20, and 19 mg/L, respectively. After 4 days of metal application, corresponding EC₅₀ values for mercury (II), copper (II), and chromium (VI) were 16, 20, and 17 (with untreated bacterial population) or not determinable, 20, and 15 mg/L, respectively (with inhibited bacterial population). Increased sensitivity of E. caudatum to tested heavy metals with inhibited bacterial co-culture population indicate that the ciliate resistance to the heavy metal tested depends on detoxification abilities of rumen bacterial population.     

Rainfall and population dynamics of Grey Pileated Finch <Emphasis Type="Italic">Coryphospingus pileatus</Emphasis> (Aves: Passeriformes) in a Neotropical dry forest

In tropical dry environments rainfall periodicity may affect demographic parameters, resulting in fluctuations in bird abundance. We used capture–recapture data for the Grey Pileated Finch from a Neotropical dry forest to evaluate the hypothesis that intra- and inter-annual survival, individuals entrance and population abundance, are related to local rainfall. Sampling occurred across 3 years, with individuals captured, tagged and evaluated for age and presence of brood patch every 14 days. Using the POPAN formulation, we generated demographic models to evaluate study population temporal dynamics. Best-fit models indicated a low apparent annual survival in the first year (16%) compared to other years (between 47 and 62%), with this low value associated with an extreme drought. The abundance of juveniles at each capture occasion was significantly dependent on the accumulated precipitation in the previous 14 days, and the juvenile covariate was a strong predictor of the intra-annual entrance probability (natality). Individuals entrance during the reproductive period corresponded to 53, 52 and 75% of total ingress for each year, respectively. The trend in sampled population size indicated positive exponential growth (N_initial = 50, N_last = 600), with intra-annual fluctuations becoming progressively more intense. Low survival was relevant during population decline at study onset, while at study end intense Individuals entrance promoted rapid population growth. Thus, the indirect effects of rainfall and the combined effect of two demographic rates operated synergistically on the immediate population abundance of Grey Pileated Finch, an abundant bird in a Neotropical dry forest.     

Geographic variation in learning of escape behavior in the little brown skink (<Emphasis Type="Italic">Scincella lateralis</Emphasis>)

A previous study showed that little brown skinks, Scincella lateralis, from a population in Louisiana, USA were poor at learning to escape to a specific retreat in a series of laboratory trials. The study was repeated on little brown skinks from a population in Oklahoma (650 km from the Louisiana population). The Oklahoma lizards were significantly better at learning to escape to a specific retreat than their Louisiana counterparts. This result demonstrates that geographic variation in the ability to learn an escape behavior exists in this species.     

Multiplex PCR amplification of 13 microsatellite loci for <Emphasis Type="Italic">Aquila chrysaetos</Emphasis> in forensic applications

The golden eagle (Aquila chrysaetos) is an endangered raptor, which is threatened mainly by illegal egg and nestling robbery. Here we describe a fluorescently labeled, multiplex PCR method using 13 microsatellite markers, which provides a powerful tool for the individual identification and parentage testing of the Golden eagle. This test should be applicable to both forensic analysis and population studies. Fifteen polymorphic loci from A. chrysaetos were cross-amplified. Subsequent PCR condition optimization led to the successful co-amplification of 13 different loci in a single PCR reaction. Fifty samples from wild-living individuals and 89 samples from captive-bred individuals were examined. The results indicated that both populations have similar levels of moderate inbreeding, unsurprising in a small population. This probability of excluding a random individual in parentage analysis was 0.9912 for the wild population and 0.9932 in the captive-bred one in the case that both the individual and its mother were examined together. The probability of identity was estimated to be 3 × 10⁻⁸ for the wild and 4 × 10⁻⁸ for the captive-bred populations. Given the size of the Slovak golden eagle population, this test should therefore be sufficient to reliably identify individual raptors and assess parentage in both conservation studies and forensic analysis.     

A demographic analysis of population responses to the manipulation of adult males in <Emphasis Type="Italic">Calomys venustus</Emphasis> (Rodentia,Sigmodontinae)

In many species of mammals, adults play an important role in influencing the survival and/or reproduction of juveniles. Adult males could have a negative effect on population density when their absence becomes a limiting factor in female fertilization. We tested the hypotheses that the absence of overwintering males (adult males) reduces the population growth rate through a delay in the onset of reproductive activity of Cohort 1 females in Calomys venustus populations. The study was carried out in two control and two experimental enclosures (0.25 ha). Adult males were removed after their offspring were born. Weekly trapping sessions were carried out from spring to autumn. To estimate population growth rates (λ), apparent survival (ϕ) and seniority probability (γ) were estimated using capture–mark-recapture models. Models were constructed with these two parameters and recapture probability (p) constrained to vary as a function of time, enclosure and/or treatment. We derived estimates of population growth rates through the estimates of ϕ and γ. The best models for ϕ and γ did not show a treatment effect. Variability between the four enclosures was greater than between control and experimental enclosures. Enclosures had different growth rates at the beginning of the study but were equaled at the end. Temporal variation in population growth rates was a result of temporal variation of γ. The two controls showed the highest growth rates earlier in time. The results did not support the hypothesis tested in this study. It seems that the number of overwintering males do not affect the population growth rate.     

Genetic diversity of oil palm (<Emphasis Type="Italic">Elaeis guineensis</Emphasis> Jacq.) germplasm collections from Africa: implications for improvement and conservation of genetic resources

A total of 723 accessions of oil palm (Elaeis guineensis Jacq.) from 26 populations representing ten countries in Africa and one Deli dura family were screened for allelic variation at seven enzyme loci from six enzyme systems using starch gel electrophoresis. On average, 54.5% of the loci were polymorphic (0.99 criterion). The average and effective number of alleles per locus was 1.80 and 1.35, respectively. Mean expected heterozygosity was 0.184, with values ranging from 0.109 (population 8, Senegal) to 0.261 (population 29, Cameroon). The genetic differentiation among populations was high (F_ST=0.301), indicating high genetic divergence. The calculation of F_ST by geographic zones revealed that the high F_ST was largely due to F_ST among populations in West Africa, suggesting diversifying selection in this region. The mean genetic distance across populations was 0.113. The lowest genetic distance (D) was observed between population 5 from Tanzania and population 7 from the Democratic Republic of the Congo (0.000) and the highest was found between population 4 from Madagascar and population 13 from Sierra Leone (0.568). The total gene flow across oil palm populations was low, with an Nm of 0.576, enhancing genetic structuring, as evident from the high F_ST values. UPGMA cluster analysis revealed three main clusters; the western outlying populations from Senegal and Sierra Leone were in one cluster but separated into two distinct sub-clusters; the eastern outlying populations from Madagascar were in one cluster; the populations from Angola, Cameroon, The Democratic Republic of the Congo, Ghana, Tanzania, Nigeria and Guinea were in one cluster. The Deli dura family seems to be closely related to population 6 from Guinea. Oil palm populations with high genetic diversity—i.e. all of the populations from Nigeria, Cameroon and Sierra Leone, population 6 of Guinea, population 1 of Madagascar and population 2 of Senegal should be used in improvement programmes, whereas for conservation purposes, oil palm populations with high allelic diversity (A_e), which include populations 22 and 29 from Cameroon, populations 39 and 45 from Nigeria, population 6 from Guinea, populations 5 and 13 from Sierra Leone and population 1 from Madagascar should be selected for capturing as much genetic variation as possible.Communicated by D.B. Neale     

Significant genetic admixture after reintroduction of peregrine falcon (<Emphasis Type="Italic">Falco peregrinus</Emphasis>) in Southern Scandinavia

The peregrine falcon (Falco peregrinus) population in southern Scandinavia was almost extinct in the 1970’s. A successful reintroduction project was launched in 1974, using captive breeding birds of northern and southern Scandinavian, Finnish and Scottish origin. We examined the genetic structure in the pre-bottleneck population using eleven microsatellite markers and compared the data with the previously genotyped captive breeding population and contemporary wild population. Museum specimens between 53 and 130 years old were analyzed. Despite an apparent loss of historical genetic diversity, the contemporary population shows a relatively high level of genetic variation. Considerable gene introgression from captive breeding stock used to repopulate the former range of southern Scandinavian peregrines may have altered the genetic composition of this population. Both the historical and contemporary northern and southern Scandinavian populations are genetically differentiated. The reintroduction project implemented in the region and the use of non-native genetic stock likely prevented the southern Scandinavian population from extinction and thus helped maintain the level of genetic diversity and prevent inbreeding depression. The population is rapidly increasing in numbers and range and shows no indication of reduced fitness or adaptive capabilities in the wake of the severe bottleneck and the reintroduction.     

Genetic variation and population structure of a threatened timber tree <Emphasis Type="Italic">Dalbergia cochinchinensis</Emphasis> in Cambodia

Dalbergia cochinchinensis Pierre ex Laness. (Fabaceae) is a commercially important tree in Southeast Asia. Although this species is under legal protections, illegal logging and disorderly developments have reduced its populations, and the conservation of this species is currently of much concern. In this study, we determined nucleotide sequences at six chloroplasts and ten nuclear loci in four populations of D. cochinchinensis in Cambodia, followed by population genetic analyses. The average silent nucleotide diversity over the nuclear loci, excluding one with an exceptionally high value, was 0.0057 in the entire population, and the mean F _ST across the nuclear loci between each population pair was between 0.135 and 0.467. Thus, the nucleotide diversity in the studied populations was not low compared with that in other tree species, and the level of population differentiation was high. Neutrality test statistics indicated a recent reduction of population size and a subdivision of the population within this species. The divergence times and migration rates were estimated with a likelihood-based method assuming the isolation with migration model. Based on the results, the three populations split 68,000–138,000 years ago, possibly corresponding to the start of the last glacial period, and the level of gene flow among the populations was very low thereafter. Moreover, after the split, population sizes were reduced considerably. Notably, the nucleotide diversity in an insertion sequence in a noncoding region of nuclear C4H was much higher than the mean nucleotide diversity in silent sites across other nuclear genes, indicating that the region was affected by selection.     

Physiological responses of <Emphasis Type="Italic">Abies faxoniana</Emphasis> populations from different elevations to increased CO<Subscript>2</Subscript> and N application

The altitude-related responses to the increased application of CO₂, N, and their combination were investigated in two Abies faxoniana populations, which originated from a subalpine coniferous forest at elevations of 2,580 and 3,200 m using closed-top chambers. The two contrasting populations were subjected to two CO₂ regimes (350 and 700 µmol mol^?1) and two N levels (0 and 5 g N m^?2 year^?1). Their net photosynthetic rate, non-structural carbohydrate concentration, and photosynthetic N use efficiency (PNUE) increased under elevated CO₂. However, the increases detected in the high-elevation (HE) population were significantly greater than those found in the low-elevation (LE) population. Under elevated CO₂ and N application, the maximal carboxylation rate (V _cmax) increased in HE population, whereas no effects were found on V _cmax in LE population. The C to N ratio decreased under N application in both populations. N application also induced the HE population to show greater increases in free amino acids, soluble proteins, N concentration, and PNUE than LE population. These results suggested that the population from HE was more sensitive to elevated CO₂ and (or) N application than LE population. Results of this study provided valuable knowledge for predicting forest development under increased atmospheric CO₂ concentration and (or) N deposition.