Persistent effects of fire severity on early successional forests in interior Alaska

There has been a recent increase in the frequency and extent of wildfires in interior Alaska, and this trend is predicted to continue under a warming climate. Although less well documented, corresponding increases in fire severity are expected. Previous research from boreal forests in Alaska and western Canada indicate that severe fire promotes the recruitment of deciduous tree species and decreases the relative abundance of black spruce (Picea mariana) immediately after fire. Here we extend these observations by (1) examining changes in patterns of aspen and spruce density and biomass that occurred during the first two decades of post-fire succession, and (2) comparing patterns of tree composition in relation to variations in post-fire organic layer depth in four burned black spruce forests in interior Alaska after 10-20 years of succession. We found that initial effects of fire severity on recruitment and establishment of aspen and black spruce were maintained by subsequent effects of organic layer depth and initial plant biomass on plant growth during post-fire succession. The proportional contribution of aspen (Populus tremuloides) to total stand biomass remained above 90% during the first and second decades of succession in severely burned sites, while in lightly burned sites the proportional contribution of aspen was reduced due to a 40-fold increase in spruce biomass in these sites. Relationships between organic layer depth and stem density and biomass were consistently negative for aspen, and positive or neutral for black spruce in all four burns. Our results suggest that initial effects of post-fire organic layer depths on deciduous recruitment are likely to translate into a prolonged phase of deciduous dominance during post-fire succession in severely burned stands. This shift in vegetation distribution has important implications for climate-albedo feedbacks, future fire regime, wildlife habitat quality and natural resources for indigenous subsistence activities in interior Alaska.     

Effects of overstory retention and site preparation on growth of planted white spruce seedlings in deciduous and coniferous dominated boreal plains mixedwoods

Survival and growth of planted white spruce was assessed under partial harvest treatments and different site preparation techniques in mixedwood forests of two compositions prior to logging: deciduous dominated (d-dom) – primarily comprised of mature trembling aspen (Populus tremuloides Michx.) and coniferous dominated (c-dom) – primarily comprised of mature white spruce (Picea glauca (Moench) Voss). Levels of overstory retention were 0% (clearcut), 50% and 75% of original basal area, and site preparation techniques were inverted mounding, high speed mixing, scalping and control (no treatment). The survival and growth of white spruce were assessed seven years after planting. The experiment was established as a part of the Ecosystem Management Emulating Natural Disturbance (EMEND) experiment located in northern Alberta, Canada. In the c-dom, the 50% and 75% retention of overstory resulted in reduced growth and survival of white spruce seedlings compared to clearcuts. In contrast, in the d-dom, the seedlings performed best in sites that had 50% of the overstory retained. For the c-dom, the mounding and mixing treatments yielded the best growth of spruce seedlings, while scalping yielded the worst. In the d-dom, spruce growth was highest in sites with the mixing treatment. In the d-dom, growth and survival of the planted spruce was greater than in the c-dom. The natural regeneration of deciduous trees was suppressed by the retention of canopy regardless of original composition.     

Wildfire promotes broadleaves and species mixture in boreal forest

Postfire tree species compositions are predicted to be the same prior to fire according to the direct regeneration hypothesis (DRH). We studied 94 upland boreal forest stands between 5 and 18 years after fire in Ontario, Canada. Postfire species-specific regeneration density was positively related to prefire stand basal area for Pinus banksiana, Populus spp., Betula papyrifera and Picea mariana, but not for Picea glauca and Abies balsamea. In addition, seedling density of Populus spp., B. papyrifera, P. mariana, P. glauca and A. balsamea were positively affected by build up index and, except Populus spp., their density increased with age of burn. To facilitate testing the DRH, we introduced a term called compositional difference (CD) that is the difference in a species relative percentage between the postfire and prefire stand. The testable null hypothesis is CD = 0 for a given species. CD was not different from 0 for P. banksiana, was 19.8% for Populus spp., 10.4% for B. papyrifera, −17.9% for P. mariana, −14.6% for P. glauca, and −14.9% for A. balsamea, indicating fire increases broadleaves at the expenses of mid- and late-successional coniferous species. Compositional increases of Populus spp. and B. papyrifera in postfire stands occurred mostly where these species were a minor component prior to fire. In conclusion, the DRH was supported by the specific positive relationships between postfire regeneration densities and prefire basal area for P. banksiana, Populus spp., B. papyrifera and P. mariana. However, if the DRH is used for predicting postfire composition, P. banksiana is the only species that had the same composition between postfire and prefire stands. Nevertheless, CD for P. banksiana was negatively related to its prefire composition. Similarly, CD for other species was negatively related to their prefire compositions with varying effects of build up index and age of burn. Our results suggest, if fire occurrences increase with global change, the boreal landscape will be more dominated by hardwoods and mixtures of conifers and hardwoods.     

Successional development of silviculturally treated and untreated high-latitude Populus tremuloides clearcuts in northern Alberta,Canada

Seventy 1–28-year-old clearcuts were sampled to characterize post-harvest vegetation development and to determine the effect of mechanical site treatment and Picea glauca (Moench) Voss (white spruce) crop-seedling planting on regenerating boreal forest stands in the John D’Or—Wood Buffalo National Park area of northern Alberta in western Canada (58°35′N, 114°37′W). Natural Populus tremuloides/Rosa–Viburnum stands of wildfire origin (n = 25), widespread occurrence, and 52–91-year-old were sampled as a benchmark for comparison. Clearcut Populus-Picea and Picea stands reverted to early successional Populus tremuloides Michx. (trembling aspen)—dominated vegetation, with maximum sucker densities (mean 18 716, S.D. 13 239) within 4 years after stand initiation. Stem exclusion occurred most intensively 5–20 years after initiation, but was expected to continue until stands were >40–50-year-old. In untreated clearcuts, tree and understory shrub cover peaked near natural stand levels 18–20 years after harvesting, and graminoid cover remained constant (∼3%) but elevated compared to natural levels (<1%); whereas forb cover decreased linearly to natural stand levels by Year 28. The early composition of clearcuts was primarily composed of species that were common to the natural stands and also vegetatively reproduced. Mechanical site treatment and crop-seedling planting delayed attainment of maximum tree cover by 7 years, with total cover similar to natural stands. Site treatment reduced total shrub cover and prolonged the occurrence of elevated forb and graminoid cover values, probably in response to disruption of the pre-treatment ground vegetation. Calamagrostis canadensis L., a common crop-seedling competitor, was typically of minor importance on the sampled clearcuts compared to levels associated with more southerly boreal clearcuts. Detrended correspondence analysis ordinations based on species cover suggested untreated and treated clearcuts >13–16-year-old approximated the composition of natural stands. The data also suggested that silvicultural planting of P. glauca will accelerate stand development toward late-successional conifer-dominated vegetation relative to unplanted and natural stands.     

Determinants of floodplain forest development illustrated by the example of the floodplain forest in the District of Leipzig

This paper discusses determinants of the historical and current spatial extent of the floodplain forest in Leipzig as well as its tree species composition using a GIS-data based delineation model and historical forest inventories for the floodplain forest in the district of Leipzig in Germany from the 19th to the 20th century. We found that the spatial extent of the floodplain forest remained considerably stable in spite of an overall decline in the entire floodplain area from the period where the city first experienced industrialisation in the 19th century to now. However, with river regulations and the alteration of forest management from coppice-with-standards forest to high forest in the 19th century, major changes can be found in the tree species composition of the floodplain forest. Comparing these findings with references from other European floodplain forests we discuss the impact of historical and current forest management as well as the city location's influence on the extent and tree species composition of urban floodplain forests. For urban forest management in particular there is a great need to integrate biophysical, historical and forestry knowledge when predicting future developmental trends.     

Spatial impacts of soil disturbance and residual overstory on density and growth of regenerating aspen

We examined spatial aspects of harvesting impacts on aspen regeneration at 25 sites in northern Minnesota. These sites had been clearcut or partially harvested 4–11 years ago. At each site, residual overstory, which was composed of trees other than aspen, soil disturbance, and tree regeneration were determined along transects leading away from skid trails into the neighboring stand. We characterized spatial extent of soil disturbance as soil strength using an Eijkelkamp soil cone penetrometer. Soil disturbance dropped off very quickly at the edge of skid trails, suggesting that the impact of harvesting traffic on areas adjacent to skid trails is minor. On skid trails, disturbance levels were higher on sites harvested in summer than on sites harvested in winter. Even after adjustment for differences in soil disturbance, stands harvested in winter had higher regeneration densities and greater aspen height growth than stands harvested in summer, suggesting that aspen regeneration was more sensitive to a given level of soil disturbance on summer-harvested sites versus on winter-harvested sites. Soil disturbance and residual overstory interactively reduced aspen regeneration densities and height growth, indicating that avoidance of soil disturbance is even more critical in partially harvested stands. Predictions based in the spatial patterns of impact found in this study indicated that harvesting conditions may have a great impact in future productivity of a site.     

Effects of bamboo stands on seed rain and seed limitation in a rainforest

Bamboos often negatively affect tree recruitment, survival, and growth, leading to arrested tree regeneration in forested habitats. Studies so far have focused on the effects of bamboos on the performance of seedlings and saplings, but the influence of bamboos on forest dynamics may start very early in the forest regeneration process by altering seed rain patterns. We tested the prediction that the density and composition of the seed rain are altered and seed limitation is higher in stands of Guadua tagoara (B or bamboo stands), a large-sized woody bamboo native from the Brazilian Atlantic Forest, compared to forest patches without bamboos (NB or non-bamboo stands). Forty 1 m² seed traps were set in B and NB stands, and the seed rain was monitored monthly for 1 year. The seed rain was not greatly altered by the presence of bamboos: rarefied seed species richness was higher for B stands, patterns of dominance and density of seeds were similar between stands, and differences in overall composition were slight. Seed limitation, however, was greater at B stands, likely as a resulted of reduced tree density. Despite such reduced density, the presence of trees growing amidst and over the bamboos seems to play a key role in keeping the seeds falling in B stands because they serve as food sources for frugivores or simply as perches for them. The loss of such trees may lead to enhanced seed limitation, contributing ultimately to the self-perpetuating bamboo disturbance cycle.     

Root carbohydrates and aspen regeneration in relation to season of harvest and machine traffic

Season of harvest has often been suggested as a driver for the erratic success of aspen (Populus tremuloides) sucker regeneration, partially due to root carbohydrate reserves and soil conditions at the time of harvest. A field experiment in western Manitoba, Canada, assessed root suckering and root carbohydrates of aspen in response to season of harvest and machine traffic. Six sites (120 m × 120 m) were selected within two large mature aspen stands slated for summer harvest. Plots (50 m × 50 m) were hand-felled (without machine traffic) in mid-summer, late summer, winter, and one plot was left uncut as a control. Season of cut with no traffic had no effect on sucker density, height or leaf dry mass per sucker. During the dormant season, root starch reserves were highest in the winter cut plots, however, just prior to suckering, this difference in carbohydrate reserves among the three seasons of harvest disappeared and by the end of the first growing season root reserves in all three seasons of cut had recovered to near control levels. Adjacent plots that were conventionally harvested in the summer and impacted by logging traffic had similar sucker densities but had 19% less height growth of suckers and 29% less leaf dry mass per sucker compared to suckers in plots harvested at the same time without traffic. After one growing season, root carbohydrate levels were similar whether or not machine traffic was used; however, the reduction in leaf dry mass in plots with machine traffic could have negative implications for carbohydrate accumulation and growth. The study suggests that the phenological state of the mature aspen plays a very small role in aspen regeneration and that harvesting practices and site conditions are likely the main drivers of aspen regeneration success.     

The lateral spread of tree root systems in boreal forests: Estimates based on N uptake and distribution of sporocarps of ectomycorrhizal fungi

In nutrient poor environments, such as boreal forests, many of the most important interactions between plants take place belowground. Here, we report the results of two approaches to obtain estimates of the lateral spread of tree roots.     

Effects of non-native spruce plantations on small mammal communities in subarctic birch forests

Voles and shrews are key species in northern forest ecosystems. Thus, it is important to quantify to what extent new forestry practices such as planting of non-native tree species impact these small mammals. In northern Norway stands of coastal subarctic birch forests have increasingly been converted to non-native spruce stands during the last century. This leads to changes in the forest floor vegetation and soil conditions that can be expected to negatively impact the community of ground-dwelling small mammals. In this 10-year trapping study we contrasted seasonal small mammal population abundances in spruce plantations with four birch forest varieties. Six different small mammal species were trapped (in descending order of abundance; common shrew Sorex araneus, red vole Myodes rutilus, field vole Microtus agrestis, grey-sided vole M. rufocanus, pygmy shrew S. minutus and water shrew Neomys fodiens). None of the voles appeared to exhibit temporal dynamics resembling population cycles. The three most numerous species were clearly less abundant in the spruce plantations compared to the other forest types. Autumn abundances were most impacted by spruce plantations, indicating that growth rates in the reproductive season were more influenced than winter declines. Species associated with productive forest habitats (i.e. field vole and common shrew) were most impacted by tree species conversion. Still young spruce plantations inter-mixed with birch trees and the ecotone habitat, sustained small mammal abundances comparable to the native birch forests. This implies that managing spruce plantations to maintain a mix of different tree species and high spatial heterogeneity (i.e. more ecotones), will reduce the negative impacts on the small mammal community. On the contrary, if young spruce plantations, as they age become spruce monocultures covering larger parts of the landscapes than they do presently, the negative effects on small mammal communities may be larger than observed in the present study.     

Increased mortality can promote evolutionary adaptation of forest trees to climate change

Forecasts of rapid climate change raise the question how quickly species can evolutionarily adapt to future climates. The adaptability of forest trees to environmental changes is generally promoted by high levels of genetic diversity and gene flow, but it can also be slowed down by long generation times and low mortality of established trees. Here, we investigate the adaptation of Scots pine (Pinus sylvestris) and Silver birch (Betula pendula) to climate change induced prolongation of the thermal growing season. We use quantitative genetic individual-based simulations to disentangle the relative roles of mortality, dispersal ability and maturation age for the speed of adaptation. The simulations predict that after 100 years of climate change, the genotypic growth period length of both species will lag more than 50% behind the climatically determined optimum. This lag is reduced by increased mortality of established trees, whereas earlier maturation and higher dispersal ability had comparatively minor effects. The evolutionary lag behind environmental change shown in our simulations stresses the importance of accounting for evolutionary processes in forecasts of the future dynamics and productivity of forests. Sensitivity of the adaptation speed to mortality suggests that species experiencing high mortality rates as well as populations subject to regular disturbances such as storms or fires might be the quickest to adapt to a warming climate.     

Modeling heliotropic tree growth in hardwood tree species—A case study on Maesopsis eminii

There are many impacts during a tree's life that affect bole development. In Maesopsis eminii, a high-potential timber species in Uganda, studies have shown that the occurrence of strongly bent boles is the overwhelming reason that boles fail quality criteria. This observation is incongruous with the tree architecture model of Roux, which describes M. eminii as a tree with a strong apical dominance, meaning that it has a strong genetically based preference for vertical growth of the terminal sprout relative to side branches. As external causes for bent boles could be excluded, we demonstrate in this study the proof of heliotropic growth, i.e. an active bending towards light, for M. eminii beyond the sapling stage. We develop a model used to describe the effects of competition on bole quality using bole form parameters and basic information about the neighbouring trees, and without having to incorporate crown parameters. By means of calculated bole parameters and a mathematical equation to calculate the intensity and direction of competition, we prove the existence of a heliotropic growth reaction. However, we are not able to predict the intensity of this reaction. Finally, general silvicultural recommendations are discussed for tree species with strong heliotropic growth.     

Mycorrhizas and secondary succession in aspen-conifer forests: Light limitation differentially affects a dominant early and late successional species

Plant succession and mycorrhizal fungi both play crucial roles in shaping the development of forest ecosystems. However, despite the strong potential for interactions between them, few studies have examined how patterns of forest succession affect mycorrhizal associations that a majority of plant species depend on to alleviate soil resource constraints. Fire suppression in subalpine forests over the last century has changed successional patterns in ways that may have important implications for mycorrhizal associations of forest tree species. To better understand these relationships we conducted a field and greenhouse study in which we examined mycorrhizal infection along gradients of light intensity and soil nutrient availability that develop as aspen becomes seral to conifers under longer fire return intervals. We examined whether ectomycorrhizal associations of quaking aspen (Populus tremuloides), a shade intolerant, early succession species, were more sensitive to light and soil resource limitations than subalpine fir (Abies lasiocarpa), a shade tolerant, late succession species. In the field study, ectomycorrhizal infection of aspen roots was reduced by 50% in conifer dominated stands relative to aspen stands. In contrast, subalpine fir maintained its EM associations regardless of the successional status of the stand. The greenhouse results were consistent with field results and indicated that light limitation was the driving force behind reductions in EM infection of aspen roots in later stages of succession. These results suggest that nutrient limitations constraining early successional species may be exacerbated by losses in EM associations via light limitations created by late successional species. This is one potential mechanism by which climax forest species create a competitive advantage over early successional species and these results suggest that it is likely exacerbated by longer fire return intervals.     

Does the mutualism between wood ants (Formica rufa group) and Cinara aphids affect Norway spruce growth?

Ant–aphid mutualisms, in which ants tend aphids, which in turn provide honeydew to the ants, are widespread and have been shown to affect plant growth. In boreal forests the effect of ant–aphid mutualism on tree growth can vary with stand age, because forest clear-cutting harms the ecologically most dominant ant partner in such mutualisms, wood ants (Formica rufa group). We studied whether the mutualism between wood ants and Cinara aphids affects the growth of boreal Norway spruces (Picea abies L. Karst.) in stands of different ages. In boreal forests, conifers, unlike deciduous trees, have only few defoliating insects, and therefore we expected the growth loss of conifers due to sap sucking by aphids not to be compensated by reduced insect herbivory due to predation by wood ants. The study was conducted in medium-fertile spruce-dominated stands in eastern Finland. We used stands of four different age classes (5, 30, 60 and 100 years) and selected ten spruces heavily visited and ten spruces lightly visited by ants around five medium-sized ant mounds in each stand age class. The access of ants was blocked on half of the trees in both groups. In the 5-year-old stands, the mean annual height growth of individual heavily visited seedlings was 16.3% greater than in the ones where ant traffic was blocked, but this difference was not significant. In the 30-year-old stands, the mean annual radial growth of the heavily visited spruces was 7.3% smaller than in trees where ant traffic was blocked, and this difference was significant. The mutualism had no significant effect on the radial growth in the 60- and 100-year-old stands. In the 60-year-old stands, however, the spruces that were visited heavily prior to the beginning of the study grew significantly less relative to their past growth than the initially lightly visited trees during the study. This suggests that the ant–aphid mutualism may have long-term effects on tree growth. The ant–aphid mutualism had no significant effect on the growth at the stand level. The results indicate that ant–aphid mutualism can have a significant effect on the growth of individual spruces, but the effect is negligible at ecosystem level.     

Bird community assembly in Bornean industrial tree plantations: Effects of forest age and structure

Plantations of exotic trees for industrial and agricultural purposes are burgeoning in the tropics, and some of them offer the opportunity to study community ecology of animals in a simplified forest setting. We examined bird community assembly in different aged groves of the industrial tree mangium (Acacia mangium) at two plantations in Malaysian Borneo: Sabah Softwoods near Tawau, Sabah, and the Planted Forest Project, near Bintulu, Sarawak. Bird communities were compared among three age-groups of mangium (2-, 5-, and 7-years old) and logged native forest. Mangium rapidly developed into a secondary forest consisting of a wide diversity of understory trees and shrubs. The bird community correspondingly increased in species richness and diversity, and we were able to relate these increases specifically to canopy height, secondary canopy development, and shrub cover. Species of common, small bodied frugivores, nectarivores, and insectivores were diverse in older plantation groves, as were common mid-sized insectivores. However, large, specialized, and normally uncommon taxa (e.g., galliforms, pigeons, hornbills, barbets, midsized woodpeckers, muscicapine flycatchers, and wren babblers) were rare or nonexistent in the plantations. Because we lacked species-specific data on foraging, nesting, and other behaviors of most groups of birds, it was difficult to explain the precise causes of seral diversification in any group except woodpeckers, which have been well studied in Southeast Asia. Thus, in future, particular emphasis needs to be placed on obtaining such data. Industrial plantations, by virtue of their simple structure, variably aged groves, and bird community richness, are good places to gather such data.     

Effects of thinning on growth of six tree species in north-temperate forests of Lithuania

This paper summarises the results from 35 years-observed thinning experiments on 256 permanent sample plots in 10–60 year-old stands of ash, aspen, birch, oak, pine and spruce in Lithuania. Thinning enhanced crown projection area increment of residual trees. The largest effect was observed in stands of aspen and birch (growth increase by 200%), followed by ash and oak (over 100%), and spruce and pine (about 80%). Thinning also promoted dbh increment, especially in younger stands, and the increase of dbh increment was positively correlated with the thinning intensity. The strongest reaction was exhibited by oak and aspen, while ash, birch and conifers reacted to a lower extent. Low and moderate intensities of thinning stimulated volume production in younger stands while the opposite was observed in older stands with increasing removals. Spruce stands exhibited relatively strongest increase of volume increment and pine, –the weakest, while the effect on deciduous species was intermediate. The results demonstrate that significant increase in volume increment is achievable with thinning of only young forest stands, e.g. 10–20 year-old pine, birch and ash, or 10–30 year-old oak, aspen and spruce.     

Influence of woody and herbaceous competition on microclimate and growth of eastern white pine (Pinus strobus L.) seedlings planted in a central Ontario clearcut

The influence of woody and herbaceous plant competition, either alone or in combination, on microclimate and growth of planted eastern white pine (Pinus strobus L.) seedlings was examined over four consecutive growing seasons in a central Ontario clearcut. Treatments that manipulated the comparative abundance of these two plant functional groups significantly affected light availability, soil moisture, and air and soil temperature regimes. These microclimate alterations, coupled with the relative competitiveness of herbaceous and woody vegetation, corresponded to temporal changes in vegetation cover and dominance. The more rapid colonization and growth of the herbaceous plant community, dominated by bracken fern (Pteridium aquilinum) and ericaceous shrubs (Kalmia sp., Vaccinium sp.), resulted in this form of vegetation being a comparatively important early competitor for soil moisture. As the woody plant community, dominated by naturally regenerated trembling aspen (Populus tremuloides Michx.), grew in height and leaf area, it became a comparatively strong competitor for both light and soil moisture. For all vegetation treatments combined, white pine seedling growth responses were strongly correlated with total cover of competing vegetation and its relative influence on above- and belowground microclimatic variables. Higher total cover of competing vegetation was generally associated with lower light and soil moisture availability and cooler soil temperatures. Multiple regression analyses indicated that pine seedling relative height growth increased with soil moisture content and growing season soil heat sum, while seedling relative diameter and relative volume growth increased with light availability.     

Predicting productivity of trembling aspen in the Boreal Shield ecozone of Quebec using different sources of soil and site information

Site quality index (SQI) of trembling aspen (Populus tremuloides Michx) in the Boreal Shield of Quebec was predicted using two sources of information: (1) mappable permanent site variables derived from permanent sampling plots and other sources and (2) measured soil and site properties including both biological and permanent site variables. General mappable information did not produce reasonable relationships (R² < 0.25) with SQI while measured variables were able to explain much of trembling aspen SQI variability. For the two parent material types found in our study, i.e. fluvial and till, there was no difference between median SQI values between groups. However, different soil and site variables were better at predicting trembling aspen productivity for the individual parent material types. As much as 60% of the variability in trembling aspen productivity was explained when both biological and permanent site variables were considered in stepwise regression models. When treated individually, models developed for fluvial sites better explained trembling aspen productivity compared to models developed for till sites. Moreover, the ability of the model to predict trembling aspen productivity on till sites when using permanent site variables alone, e.g. soil texture, elemental chemistry and elevation was decreased (R² < 0.3). This indicates that the inclusion of biological site variables such as overstory species composition and forest floor properties provide a major contribution to SQI prediction and are necessary to yield high R². Overall, the data indicate that the traditional mapping of landscape attributes such as drainage and deposit as well as inferred soil geochemistry do not contribute much to explaining SQI. At present, field measurements are needed to predict SQI with a reasonable degree of precision within a forest management unit.     

Effect of species and spacing of fast-growing nurse trees on growth of an indigenous tree, Hopea odorata Roxb., in northeast Thailand

We tested the effects of species and spacing of nurse trees on the growth of Hopea odorata, a dipterocarp tree indigenous to Southeast Asia, in a two-storied forest management system in northeast Thailand. Eucalyptus camaldulensis, Acacia auriculiformis, and Senna siamea were planted as nurse trees in 1987 at spacings of 4 m × 8 m, 2 m × 8 m, 4 m × 4 m, and 2 m × 4 m in the Sakaerat Silvicultural Research Station of the Royal Forest Department, Thailand. Seedlings of H. odorata were planted in the nurse tree stands at a uniform spacing of 4 m × 4 m and in control plots (no nurse trees) in 1990. Stem numbers of some nurse trees were thinned by half in 1994. The stem diameter and height of all trees were measured annually until 1995 and again in 2007. The mean annual increment (MAI) in volume was estimated as 8.2–10.1 m³ ha⁻¹ year⁻¹ for E. camaldulensis and 0.9–1.2 m³ ha⁻¹ year⁻¹ for S. siamea, smaller than reported elsewhere. This suggests that the site properties were not suitable for them. The MAI of A. auriculiformis was 7.9–9.8 m³ ha⁻¹ year⁻¹, within the reported range. Survival rates of H. odorata in the S. siamea stands and the control plots decreased rapidly during the first 2 years but then stayed constant from 1992. In contrast, survival rates of H. odorata in the E. camaldulensis and A. auriculiformis stands were initially high (>70%), but then decreased after 1995. Stem diameter, tree height, and stand basal area of H. odorata were large in both the S. siamea stands and the control plots from then. The growth of H. odorata was largest in the 2 m × 8 m S. siamea stands. In contrast, it was restricted in the E. camaldulensis and A. auriculiformis stands owing to strong shading by their canopies. Thinning by 50% tended to facilitate the growth of H. odorata temporarily in the E. camaldulensis and A. auriculiformis stands. The stand basal areas of nurse trees and of H. odorata showed a trade-off. These results suggest that the growth of H. odorata was maximized in the S. siamea stands. We assume, however, that the growth of H. odorata could be improved even in the E. camaldulensis and A. auriculiformis stands by frequent or heavy thinning.     

Early seedling establishment of two tropical montane cloud forest tree species: The role of native and exotic grasses

Tropical montane cloud forest has been undergoing a drastic reduction because of its widespread conversion to pastures. Once these forests have been cleared exotic grasses are deliberately introduced for forage production. Exotic grass species commonly form monodominant stands and produce more biomass than native grass species, resulting in the inhibition of secondary succession and tree regeneration. The purpose of this study was to assess the effect of native vs. exotic grass species on the early establishment of two native tree seedlings (Mexican alder, Alnus acuminata and Jalapa oak, Quercus xalapensis) on an abandoned farm in central Veracruz, Mexico. Seedling survival and growth were monitored (over 46 weeks) in relation to grass cover and height, and available photosynthetic active radiation (PAR). More seedlings survived in the presence of the native grass Panicum glutinosum than those growing with the exotic grass Cynodon plectostachyus (92% vs. 48%). The causes of seedling mortality varied between species; Q. xalapensis was affected by herbivory by voles but mainly in the exotic grass-dominated stands, whereas A. acuminata seedlings died due to competition with the exotic grass. A. acuminata seedlings increased more in height in the exotic grass-dominated stands (102 ± 7.8 cm) compared to native grass-dominated stands (51 ± 4.7 cm). Grass layer height, cover and available PAR were correlated (Pearson; p < 0.05). In the exotic grass dominated plots, grass layer height was correlated with the relative height growth rates of Q. xalapensis (Pearson; p < 0.05). These results indicate that the exotic grass may be affecting tree regeneration directly (grass competition) and indirectly (higher herbivory). Passive restoration may occur once P. glutinosum dominated pastures are abandoned. However, when C. plectostachyus dominates, introduction of early and mid successional tree seedlings protected against vole damage is needed.