Changes in tree species diversity after successive clearcuts in the Southern Appalachians

A 16 ha watershed in the Coweeta Basin was clearcut in 1939 and 1962. Vegetation was inventoried in 1934 and at about 7-year intervals to 1991. After the first clearcut, tree diversity remained high until after the second cut. Diversity based on density and basal area decreased significantly 14 years after the second clearcut and remained low through 1991. Diversity was highest in the early establishment stage of stand development, then declined at the intermediate stage with canopy closure. Evenness based on basal area declined more than evenness based on density because basal area of Liriodendron tulipifera increased substantially from 1977 to 1991. Trends in diversity were due to changes in evenness rather than changes in species richness.     

Density-dependence and co-existence of conifer and broad-leaved trees in a Japanese northern mixed forest

Abstract. The growth and survival of coniferous trees (particularly Abies sachalinensis) and broad-leaved trees (particularly Quercus crispula) were followed over a 15-yr period in a 15.5-ha area in a northern mixed forest in Japan, and the coexistence of the two groups was simulated by a density-dependent projection matrix model. The density-dependent model assumes that the density effect of mother trees due to one-sided competition for light on smaller-sized tree regulates the demographic functions. The mother tree densities of conifers and broad-leaved trees have stronger negative effects on the recruitment and survival of seedlings of their own group than of the other group. These results support the idea of reciprocal replacement for conifer and broad-leaved trees. Simulations using the density-dependent model showed that the two groups will co-exist within a particular range of recruitment rates. However, the density of both groups did not affect the growth rate of any tree, and equilibrium DBH-distributions from density-dependent matrices were quite different from present distributions both for conifers and broad-leaved trees. On the other hand, equilibrium DBH-distributions of conifer and broad-leaved trees from density-independent matrices were quite distinct from each other, reflecting different survivorship curves of the two dominants. These results suggest that density-dependent processes are not so important for shaping population structures in this northern mixed forest.     

Patterns of growth,mortality and biomass change in a coastal Picea sitchensis - Tsuga heterophylla forest

Abstract. Ten years (1979-1989) of growth and mortality were determined in a 130-yr old stand on the Oregon coast based on periodic remeasurements in 441000 m² plots. Western hemlock (Tsuga heterophylla) constituted 90 % of the individuals and 57 % of the biomass. Wind is a major form of disturbance in this area, creating both small discrete and large diffuse disturbance patches; wind therefore has a direct effect on the location and extent of regeneration. Rates of tree mortality were high for this coastal stand (2.8 %/yr), especially compared to similar-aged stands in the western and eastern Cascade Ranges. Though low in absolute density, Sitka spruce (Picea sitchensis) persisted in competition with the more tolerant western hemlock. Net production of bole biomass (4.9 Mg ha^-1 yr^-1) did not equal mortality (8.7 Mg ha^-1 yr^-1), and total biomass declined over the 10-yr measurement period from 499 to 460 Mg/ha; this trend may have begun as early as the mid-1950's at a peak biomass of about 600 Mg/ha. The decline may have been due to a positive feedback in which new gaps and enlarging gap perimeters exposed more and more trees to potential wind damage.     

Effects of snow pressure on growth form and life history of tree species in Japanese beech forest

Abstract. In the cool temperate zone in Japan Fagus crenata (beech) is a highly dominating climax species, especially in the snow-rich regions at higher altitudes. The explanation for this dominance was studied with special reference to the tolerance of tree trunks to snow pressure. Traits of six tree species: trunk bend, trunk height decrease, trunk damage and basal sprouting rate were measured in Fagus forest in the Echigo Mountains, central Japan along a snow pressure gradient. The following general trends were recognized: (1) trunk bend and trunk height decrease; (2) snow pressure caused trunk injury; (3) when the trunk bend exceeded a critical angle, the number of sprouts increased remarkably. Although the tolerances were different among the six species, most of them were seriously damaged by snow pressure, and many trunks were reduced in size. On the other hand, many sprouts emerged from deformed trunks and contributed to the maintenance of the population under heavy snow pressure. F. crenata was the only species whose trunk form and maximum size were hardly influenced by snow pressure.     

Population dynamics of coniferous and broad-leaved trees in a Japanese temperate mixed forest

The growth and survival of coniferous and broad-leaved trees were followed over a 5-yr period in a temperate old-growth mixed forest in Japan, and dynamic features of the forest were studied in relation to the life history of the dominants, the coniferous Abies homolepis and the broad-leaved Fagus crenata. During this period, the gap formation rate was 31m² ha^?1yr^?1, the mortality of trees > 2m high was 1.7%/yr, and the rate of loss in basal area 1.4%/yr. These values were much higher than the recruitment, 0.3%/yr, and the total growth of surviving and new trees, 0.6%/yr, owing to the inhibition of regeneration by understorey dwarf bamboo (Sasa borealis). A transition matrix model based on DBH size classes predicts that the basal area of the forest will decrease by 14% in 50 yr, but that the DBH distribution of trees > 10 cm diameter will change little. Equilibrium DBH distributions assuming recruitment being equal to mortality, were quite different between broad-leaved and coniferous trees, reflecting different survivorship curves of the two dominants. The composition and structure of the forest may change depending on the pattern and frequency of disturbances, or episodic events, notably the synchronous death of Sasa borealis.     

Spatial patterning in the canopy gap regime of a subalpine Abies-Picea forest in the northeastern United States

Abstract. Spatial patterning in the disturbance regime of a forest affects the vegetation dynamics. Therefore, the distribution of canopy gaps was examined in detail for a subalpine Abies-Picea forest in the northeastern United States. Gaps were not randomly distributed. The fraction of forest area in gaps and the abundance of gaps varied significantly with topographic position, elevation, and slope percent. On average, 15 % of the forest was influenced by gaps, but the gap fraction was greater near ridges (23 %) and near streams (27 %) than on the backslope (13 %). Also, gaps were larger and more abundant near streams and ridges. Gap fraction varied with elevation as well: more of the forest was disturbed at lower and higher elevations than at mid-elevations. Significantly more of the forest on steep slopes (≥ 30 %) was under gaps. As a result of this patterning, some parts of the Abies-Picea forest were predictably more disturbed than others. A remaining question is whether this patterning is sufficient to influence the regeneration environment and thus forest composition.     

Dampening effects of long‐term experimental drought on growth and mortality rates of a Holm oak forest

Forests respond to increasing intensities and frequencies of drought by reducing growth and with higher tree mortality rates. Little is known, however, about the long‐term consequences of generally drier conditions and more frequent extreme droughts. A Holm oak forest was exposed to experimental rainfall manipulation for 13 years to study the effect of increasing drought on growth and mortality of the dominant species Quercus ilex, Phillyrea latifolia, and Arbutus unedo. The drought treatment reduced stem growth of A. unedo (?66.5%) and Q. ilex (?17.5%), whereas P. latifolia remained unaffected. Higher stem mortality rates were noticeable in Q. ilex (+42.3%), but not in the other two species. Stem growth was a function of the drought index of early spring in the three species. Stem mortality rates depended on the drought index of winter and spring for Q. ilex and in spring and summer for P. latifolia, but showed no relation to climate in A. unedo. Following a long and intense drought (2005–2006), stem growth of Q. ilex and P. latifolia increased, whereas it decreased in A. unedo. Q. ilex also enhanced its survival after this period. Furthermore, the effect of drought treatment on stem growth in Q. ilex and A. unedo was attenuated as the study progressed. These results highlight the different vulnerabilities of Mediterranean species to more frequent and intense droughts, which may lead to partial species substitution and changes in forest structure and thus in carbon uptake. The response to drought, however, changed over time. Decreased intra‐ and interspecific competition after extreme events with high mortality, together with probable morphological and physiological acclimation to drought during the study period, may, at least in the short term, buffer forests against drier conditions. The long‐term effects of drought consequently deserve more attention, because the ecosystemic responses are unlikely to be stable over time.Nontechnical summaryIn this study, we evaluate the effect of long‐term (13 years) experimental drought on growth and mortality rates of three forest Mediterranean species, and their response to the different intensities and durations of natural drought. We provide evidence for species‐specific responses to drought, what may eventually lead to a partial community shift favoring the more drought‐resistant species. However, we also report a dampening of the treatment effect on the two drought‐sensitive species, which may indicate a potential adaptation to drier conditions at the ecosystem or population level. These results are thus relevant to account for the stabilizing processes that would alter the initial response of ecosystem to drought through changes in plant physiology, morphology, and demography compensation.     

Fertilization effects on fineroot biomass, rhizosphere microbes and respiratory fluxes in hardwood forest soils

Fertilizer-induced reductions in CO(2) flux from soil ((F)CO(2)) in forests have previously been attributed to decreased carbon allocation to roots, and decreased decomposition as a result of nitrogen suppression of fungal activity. Here, we present evidence that decreased microbial respiration in the rhizosphere may also contribute to (F)CO(2) reductions in fertilized forest soils. Fertilization reduced (F)CO(2) by 16-19% in 65-yr-old plantations of northern red oak (Quercus rubra) and sugar maple (Acer saccharum), and in a natural 85-yr-old yellow birch (Betula allegheniensis) stand. In oak plots, fertilization had no effects on fine root biomass but reduced mycorrhizal colonization by 18% and microbial respiration by 43%. In maple plots, fertilization reduced root biomass, mycorrhizal colonization and microbial respiration by 22, 16 and 46%, respectively. In birch plots, fertilization reduced microbial respiration by 36%, but had variable effects on root biomass and mycorrhizal colonization. In plots of all three species, fertilization effects on microbial respiration were greater in rhizosphere than in bulk soil, possibly as a result of decreased rhizosphere carbon flux from these species in fertile soils. Because rhizosphere processes may influence nutrient availability and carbon storage in forest ecosystems, future research is needed to better quantify rhizo-microbial contributions to (F)CO(2).     

Impacts of a spring heat wave on canopy processes in a northern hardwood forest

Heat wave frequency, duration, and intensity are predicted to increase with global warming, but the potential impacts of short‐term high temperature events on forest functioning remain virtually unstudied. We examined canopy processes in a forest in Central Ontario following 3 days of record‐setting high temperatures (31–33 °C) that coincided with the peak in leaf expansion of dominant trees in late May 2010. Leaf area dynamics, leaf morphology, and leaf‐level gas‐exchange were compared to data from prior years of sampling (2002–2008) at the same site, focusing on Acer saccharum Marsh., the dominant tree in the region. Extensive shedding of partially expanded leaves was observed immediately following high temperature days, with A. saccharum losing ca. 25% of total leaf production but subsequently producing an unusual second flush of neoformed leaves. Both leaf losses and subsequent reflushing were highest in the upper canopy; however, retained preformed leaves and neoformed leaves showed reduced size, resulting in an overall decline in end‐of‐season leaf area index of 64% in A. saccharum, and 16% in the entire forest. Saplings showed lower leaf losses, but also a lower capacity to reflush relative to mature trees. Both surviving preformed and neoformed leaves had severely depressed photosynthetic capacity early in the summer of 2010, but largely regained photosynthetic competence by the end of the growing season. These results indicate that even short‐term heat waves can have severe impacts in northern forests, and suggest a particular vulnerability to high temperatures during the spring period of leaf expansion in temperate deciduous forests.     

Influence of micro-landforms on forest structure,tree death and recruitment in a Japanese temperate mixed forest

The micro-landform unit system offers an effective way of analyzing vegetation–geomorphology relationships at a 10-m scale in areas such as the hilly regions of Japan. We analyzed relationships between micro-landforms and tree population parameters over a 9-year interval to elucidate the influence of geomorphic processes on vegetation dynamics. A 2.16-ha permanent plot was established in a temperate mixed forest. Each 5m×5m quadrat within this plot was classified according to six types of micro-landform units: (i) crest slope (CS); (ii) upper sideslope (US); (iii) head hollow (HH); (iv) lower sideslope (LS); (v) foot slope (FS); and (vi) river bed (RB). All living trees larger than 10cm in diameter at breast height (d.b.h.) were identified, mapped and marked in 1989 and resurveyed in 1998. Almost all of the 23 common tree species persisted in their own core habitats (i.e. the micro-landforms) between the two surveys. The species distribution in both surveys showed that the six micro-landforms could be combined into two larger groups: upper and lower hillslope areas. The upper hillslope area had higher tree densities and larger basal areas than the lower hillslope area. It is possible that these differences result from the longer lifespans of trees on the upper hillslope area rather than from differences in mortality and recruitment rates. In addition, the different ways in which trees die in the different micro-landform units may affect the regeneration process in hilly regions through different gap formation. The effects of different geomorphic processes are reflected in the lifespans of the trees and may result in different forest structure and dynamics among micro-landform units.     

Small rodents as significant dispersers of tree seeds in a Neotropical forest

Abstract. Through seed dispersal and predation, terrestrial mammals should be an important component of the mechanisms that determine patterns of tree recruitment in tropical forests. Despite their great abundance and ubiquity in Neotropical forests, small rodents as seed predators and dispersers remain largely forgotten. To investigate the fates of seeds in a hunted primary forest in Belize, we tagged seeds of Astrocaryum mexicanum (Palmae), Ampelocera hottlei (Ulmaceae), and Pouteria sapota (Sapotaceae) and placed them into open plots, exclosures accessible only to small mammals, and exclosures accessible to medium-sized and small mammals. The exclosure experiments and fates of the seeds show that the spiny pocket mouse, Heteromys desmarestianus (Heteromyidae), was the dominant handler of seeds of the first two species and also removed a significant proportion of the very large-seeded Pouteria. Most of the seeds were killed immediately upon removal, but many of the seeds (3–18 %) of the first two species were scatterhoarded (dispersed and buried in the soil) by Heteromys. Some of the scatterhoarded seeds (29%) remain buried and therefore protected from predation by other animals. Agoutis (Dasyprocta punctata), a caviomorph rodent, buried 13 % of the seeds of Pouteria, and Heteromys consumed and dispersed but did not bury Pouteria seeds. Results of this study support predictions by some researchers that small rodents are dominant terrestrial granivores in Neotropical forests. The role of small rodents as seed dispersers, however, has never been fully appreciated.     

Hurricane disturbance in a temperate deciduous forest: patch dynamics, tree mortality, and coarse woody detritus

Patch dynamics, tree injury and mortality, and coarse woody detritus were quantified to examine the ecological impacts of Hurricane Fran on an oak-hickory-pine forest near Chapel Hill, NC. Data from long-term vegetation plots (1990–1997) and aerial photographs (1998) indicated that this 1996 storm caused patchy disturbance of intermediate severity (10–50% tree mortality; Woods, J Ecol 92:464–476, 2004). The area in large disturbance patches (>0.1 ha) increased from <1% to approximately 4% of the forested landscape. Of the forty-two 0.1-ha plots that were studied, 23 were damaged by the storm and lost 1–66% of their original live basal area. Although the remaining 19 plots gained basal area (1–15% increase), across all 42 stands basal area decreased by 17% because of storm impacts. Overall mortality of trees >10 cm dbh was 18%. The basal area of standing dead trees after the storm was 0.9 m²/ha, which was not substantially different from the original value of 0.7 m²/ha. In contrast, the volume and mass of fallen dead trees after the storm (129 m³/ha; 55 Mg/ha) were 6.1 and 7.9 times greater than the original levels (21 m³/ha; 7 Mg/ha), respectively. Uprooting was the most frequent type of damage, and it increased with tree size. However, two other forms of injury, severe canopy breakage and toppling by other trees, decreased with increasing tree size. Two dominant oak species of intermediate shade-tolerance suffered the largest losses in basal area (30–41% lost). Before the storm they comprised almost half of the total basal area in a forest of 13% shade-tolerant, 69% intermediate, and 18% shade-intolerant trees. Recovery is expected to differ with respect to vegetation (e.g., species composition and diversity) and ecosystem properties (e.g., biomass, detritus mass, and carbon balance). Vegetation may not revert to its former composition; however, reversion of biomass, detritus mass, and carbon balance to pre-storm conditions is projected to occur within a few decades. For example, the net change in ecosystem carbon balance may initially be negative from losses to decomposition, but it is expected to be positive within a decade after the storm. Repeated intermediate-disturbance events of this nature would likely have cumulative effects, particularly on vegetation properties.     

Influence of site conditions on the survival of Fagus sylvatica seedlings in an old‐growth beech forest

Abstract. Following a full mast production in autumn 1995 in the old‐growth beech forest of la Tillaie (France), cupules, live and dead 1‐yr old seedlings were counted in September 1996 in 40 plots, representing a wide range of ecological conditions. The influence of geomorphology and forest stage on mast production and on seedling establishment and survival was studied. Maximum seedling abundance did not occur in areas with the largest cupule abundance, but this result was not statistically significant. The presence of a shallow sandstone table under Fontainebleau sand, where beech is far from its ecological optimum, positively influenced the production of mast but negatively influenced seedling survival. Contrary to expectation, the early biostatic rather than the late biostatic phase seemed to be the best stage for beech regeneration. Gaps also provide good environmental conditions for seedling establishment. Humus and light conditions, water availability and competition between beech individuals may explain the results. Light was the main factor influencing mast production in mature stages, and drought was an important factor causing seedling mortality. This study points out the importance of seedling survival for regeneration patterns in beech forests.     

Dynamics of composition and structure in an old Sequoia sempervirens forest

Abstract. Dynamics of a Sequoia sempervirens forest in northern California were studied with long‐term plot data (1.44 ha) and recent transect data. The study was conducted in an old stand (> 1100 yr) on alluvial flats. Over three decades (1972–2001), changes in the composition and structure of the tree stratum were minor. Sequoia maintained a broad distribution of stem diameters throughout the period. Annual rates of Sequoia mortality (0.0029) and ingrowth (0.0029) were low, reflecting the great longevity of Sequoia and the slow canopy turnover of the study forest. Transect data also indicated a low frequency of canopy gap disturbance (≤ 0.4% of total land area per yr), but gap size was potentially large (> 0.1 ha) and the fraction of area in gaps (ca. 20%) was similar to other temperate forests. Regeneration quadrats sampled along transects, in gap centers, and on logs revealed that Sequoia regeneration is elevated at gap edges. The longevity of Sequoia and its response to gap disturbances ensure that it will remain a dominant species in the study forest.     

Quantifying the effects of drought on abrupt growth decreases of major tree species in Switzerland

Drought entails important effects on tree physiology, which may result in short‐ to long‐term radial growth decreases. While the majority of studies have focused on annual drought‐related variability of growth, relatively little is known about sustained growth decreases following drought years. We apply a statistical framework to identify climatic factors that induce abrupt growth decreases and may eventually result in tree mortality. We used tree‐ring data from almost 500 standing dead trees and 200 living trees in eight sites of the Swiss network of strict forest reserves, including four of the most important Central European tree species (Abies alba, Picea abies, Fagus sylvatica and Quercus spp.). First, to assess short‐term growth responses to drought under various climate and site conditions, we calculated correlations and linear mixed‐effects models between ring‐width indices (RWIs) and drought based on the Standardized Precipitation Evapotranspiration Index (SPEI). Second, to quantify drought effects on abrupt growth decreases, we applied distributed lag nonlinear models (DLNMs), which account for both delayed effects and the nonlinear relationship between the SPEI and the occurrence of abrupt growth decreases. Positive correlations between RWIs and the SPEI indicated short‐term growth responses of all species, particularly at arid sites. Results of the DLNMs revealed species‐specific growth responses to drought. For Quercus spp., abrupt growth decreases were more likely to occur several years following severe drought, whereas for P. abies, A. alba, and F. sylvatica abrupt growth decreases started frequently immediately in the drought year. We conclude that the statistical framework allows for quantifying the effects of drought intensity on the probability of abrupt growth decreases, which ultimately contributes to an improved understanding of climate impacts on forest community dynamics.     

The role of anthropogenic influences in a mixed-conifer forest mortality episode

Abstract. Anthropogenic influences on recent tree mortality in mid-montane mixed conifer forests of southern California, USA, and northern Baja California, Mexico, were investigated. The Pinus jeffreyi-Abies concolor phase of the mixed-conifer montane community was sampled at three sites, characterized by: (1) low levels of air pollution and long-term fire suppression (Holcomb Valley in the San Bernardino Mountains, California); (2) chronic, severe air pollution and long-term fire suppression (Barton Flats in the San Bernardino Mountains, California); and (3) little air pollution and no fire suppression (La Corona Arriba in the Sierra San Pedro Martir in Baja California, Mexico). Similar percentages of dead standing trees, around 14 %, were found at both San Bernardino sites, but a comparatively low level, 4 %, was found in the Mexican forest. Anthropogenic effects, in particular fire suppression, may play an important role in enhancing the impact of natural stresses on the dynamics of mixed conifer forests.     

Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.