Gap characteristics and gap regeneration in a subalpine coniferous forest on Mt Ontake,central Honshu,Japan

Gap characteristics and gap regeneration were studied in three mature stands belonging to different community types in a subalpine coniferous forest on Mt Ontake, central Honshu, Japan. Gap disturbance regimes were remarkably similar among stands studied; percentage gap area to surveyed area, gap density and mean gap size were 7.3–8.5%, 17.8–20.0 ha⁻¹ and 40.8–42.5 m², respectively. The gap size class distributions were also similar and showed a strong positive skewness with a few large and many small gaps; gaps <40m² were most frequent and gaps >200m² were rare. Forty-five to 66% of gaps were due to the death of single canopy trees. Canopy trees more often died leaving standing dead wood (40–5.7%) or broken trunks (43–49%). Shade tolerantAbies mariesii andAbies veitchii, frequently, andTsuga diversifolia, less frequently, regenerated in gaps, from advance regenerations recruited before gap formation.Picea jezoensis var.hondoensis may regenerate in gaps, from new individuals recruited after gap formation. The breakage of denseSasa coverage and the mineral soils exposed by the uprooted plants that form gaps might provide regeneration opportunities for shade intolerantBetula. Of the important species limited to the ridge site,Chamaecyparis obtusa, frequently, andThuja standishii, infrequently, regenerated from plants alreadyin situ. Regeneration ofPinus parviflora was not seen. Based on the gap characteristics and gap regeneration behaviour of each species described, stand dynamics in each stand are discussed.     

Gap characteristics and gap regeneration in subalpine old-growth coniferous forests,central Japan

Gap characteristics and gap regeneration were studied in three old-growth stands of subalpine coniferous forests in the northern Yatsugatake and the northern Akaishi mountains, central Japan. With the results of the present study and those of a previous study conducted in another locality, general features of gap characteristics and gap regeneration behavior of major tree species in subalpine coniferous forests of central Japan were summarized and discussed. Of the total 237 gaps investigated in the 14.48 ha of forested area, the percentage gap area to surveyed area, gap density and mean gap size were 7.3%, 17.2 ha⁻¹, and 43.3 m², respectively. The gap size distributions were similar among stands and showed a strong positive skewness with a few large and many small gaps; gaps <40m² were most frequent and those >200 m² were rare. Gaps due to the death of multiple canopy trees comprised 44.7% of the total ones. Canopy trees died in various states; standing dead (42.6%) or trunk broken (43.7%) were common and uprooted (12.2%) was an uncommon type of death of canopy trees. These figures indicate that general features of gap characteristics in this forest type are the low proportion of gap area and the high proportions of small gap size and multiple-tree gap formation. In general, shade-tolerantAbies frequently, andTsuga, infrequently, regenerate in gaps from advance regenerations recruited before gap formation, whilePicea and shade-intolerantBetula possibly regenerate in gaps from new individuals recruited after gap formation. Gap successors of conifers occurred in a wide range of gap size and did not show the clear preference to species specific gap size. In old-growth stands without large-scale disturbance (≥0.1 ha in area) of subalpine coniferous forests of central Japan, major tree species may coexist with their different gap-regeneration behaviors and, probably, different life history traits.     

Forest dynamics following eastern hemlock mortality in the southern Appalachians

Understanding changes in community composition caused by invasive species is critical for predicting effects on ecosystem function, particularly when the invasive threatens a foundation species. Here we focus on dynamics of forest structure, composition and microclimate, and how these interact in southern Appalachian riparian forests following invasion by hemlock woolly adelgid, HWA, Adelges tsugae. We measured and quantified changes in microclimate; canopy mortality; canopy and shrub growth; understory species composition; and the cover and diversity in riparian forests dominated by eastern hemlock Tsuga canadensis over a period of seven years. Treatments manipulated hemlock mortality either through invasion (HWA infested stands) or girdling (GDL) hemlock trees. Mortality was rapid, with 50% hemlock tree mortality occurring after six years of invasion, in contrast to more than 50% mortality in two years following girdling. Although 50% of hemlock trees were still alive five years after infestation, leaf area lost was similar to that of girdled trees. As such, overall responses over time (changes in light transmittance, growth, soil moisture) were identical to girdled stands with 100% mortality. Our results showed different growth responses of the canopy species, shrubs and ground layer, with the latter being substantially influenced by presence of the evergreen shrub, rhododendron Rhododendron maximum. Although ground layer richness in the infested and girdled stands increased by threefold, they did not approach levels recorded in hardwood forests without rhododendron. Increased growth of co‐occurring canopy trees occurred in the first few years following hemlock decline, with similar responses in both treatments. In contrast, growth of rhododendron continued to increase over time. By the end of the study it had a 2.6‐fold higher growth rate than expected, likely taking advantage of increased light available during leaf‐off periods of the deciduous species. Increased growth and dominance of rhododendron may be a major determinant of future responses in southern Appalachian ecosystems; however, our results suggest hemlock will be replaced by a mix of Acer, Betula, Fagus and Quercus canopy genera where establishment is not limited by rhododendron.     

Vegetation changes around the last glacial maximum and effects of the Aira-Tn ash,at the Itai-Teragatani site,central Japan

On the basis of fossil pollen, plant macrofossils and fossil wood, vegetation existing around the Itai-Teragatani Site between 26000 and 20000 yr ago was reconstructed. On the upland, there were mixed forests of conifers and deciduous broad-leaved trees throughout the studied period. The forests consisted mainly ofPinus subgen.Haploxylon andQuercus subgen.Lepidobalanus, together withTsuga, Cryptomeria andUlmus-Zelkova. In the wetland, there were stands ofBetula andAlnus in grassland consisting of Cyperaceae, Gramineae,Lysichiton, Thalictrum, Sangusiorba, Umbelliferae and Compositae. The vegetation changed three times during the studied period. The first change, which occurred prior to the fall of the Aira-Tn ash, was characterized by an increase inAbies, Pinus subgen.Haploxylon and hygrophytes, and a decrease inCryptomeria andAlnus. This is considered to reflect a cooler climate, and expansion of grassland. The second change was caused by the fall of the Aira-Tn ash, which caused serious damage to the vegetation. Many taxa, except forAlnus, Cyperaceae andLysichiton, decreased or vanished after the ash fall. The third change was the process of recovery after the damage. The regenerated forests differed from those before the ash fall. With the regeneration conifers andArtemisia increased. This seemed to reflect the changes in climate and ground conditions caused by the accumulation of the ash.     

The Importance of Large-Diameter Trees to Forest Structural Heterogeneity

Large-diameter trees dominate the structure, dynamics and function of many temperate and tropical forests. However, their attendant contributions to forest heterogeneity are rarely addressed. We established the Wind River Forest Dynamics Plot, a 25.6 ha permanent plot within which we tagged and mapped all 30,973 woody stems ≥1 cm dbh, all 1,966 snags ≥10 cm dbh, and all shrub patches ≥2 m². Basal area of the 26 woody species was 62.18 m²/ha, of which 61.60 m²/ha was trees and 0.58 m²/ha was tall shrubs. Large-diameter trees (≥100 cm dbh) comprised 1.5% of stems, 31.8% of basal area, and 17.6% of the heterogeneity of basal area, with basal area dominated by Tsuga heterophylla and Pseudotsuga menziesii. Small-diameter subpopulations of Pseudotsuga menziesii, Tsuga heterophylla and Thuja plicata, as well as all tree species combined, exhibited significant aggregation relative to the null model of complete spatial randomness (CSR) up to 9 m (P≤0.001). Patterns of large-diameter trees were either not different from CSR (Tsuga heterophylla), or exhibited slight aggregation (Pseudotsuga menziesii and Thuja plicata). Significant spatial repulsion between large-diameter and small-diameter Tsuga heterophylla suggests that large-diameter Tsuga heterophylla function as organizers of tree demography over decadal timescales through competitive interactions. Comparison among two forest dynamics plots suggests that forest structural diversity responds to intermediate-scale environmental heterogeneity and disturbances, similar to hypotheses about patterns of species richness, and richness- ecosystem function. Large mapped plots with detailed within-plot environmental spatial covariates will be required to test these hypotheses.     

Regeneration process of coniferous forests in northern Hokkaido I.Abies sachalinensis forest andPicea glehnii forest

Regeneration of natural forests was studied in the Nakagawa Experiment Forest of Hokkaido University using age distribution surveys made by the clear felling method. In Plot 1 (30 m × 65 m),Abies sachalinensis dominated the canopy layer but there were also a fewBetula ermanii trees.Sasa senanensis densely covered the forest floor. Most of the canopy trees were from 122 to 195 years old. Seedlings younger than 50 years old ofA. sachalinensis were found on fallen logs and root bases. There were, however, few trees from 50 to 120 years old. The present canopy trees seemed to have regenerated after competitive pressure from old canopy andSasa disappeared 180 years ago. Plot 2 (50 m × 100 m) on serpentinite soil was dominated byPicea glehnii. Sasa kulirensis covered the floor but not as densely asS. senanensis in Plot 1. The ages ofP. glehnii ranged from 1 to 586 years old, and the age distribution ofA. sachalinensis was L-shaped. A small gap in the canopy formed about 290 years ago, and it gradually extended. Conifers regenerated continuously in the extending gap butB. ermanii did not. One hundred thirty years ago, part of Plot 2 was again destroyed andA. sachalinensis andB. ermanii regenerated. Thus, two types of regeneration were found. One regenerated both conifers andBetula after a sudden disturbance of canopy layer or death ofSasa, and the other, under an extending gap, regenerated only conifers.     

Formation and closure of canopy gaps in the rain forest at Nouragues, French Guiana

Formation and closure of canopy gaps was monitored for three years in 12 ha of primary rain forest at Nouragues, French Guiana. At the first inventory, in April 1991, 74 openings in the canopy > 4 m² (sensu Brokaw 1982a) were located; 60 of these gaps were formed before January 1990. Between January 1990 and December 1993, 5 to 15 gaps were annually formed, opening 0.64–1.33% of the forest canopy each year. Of all gaps, 41% were created by a falling, snapped tree, 34% by a falling, uprooted tree, 22% by a falling branch, and 3% by a falling dead stem. A refined nearest neighbour analysis showed that gaps formed after January 1990 were clustered: uprooting of trees seemed to be related to shallow soils, and relatively many other trees fell when a tree uprooted, independent of the dbh of the uprooted tree. In 37 gaps, canopy openness in the gap centre (determined by hemispherical photographs) was monitored over three years. In 54% of the gaps, canopy openness increased in two successive years. It is reasoned that edges of especially large gaps may frequently be re-disturbed by falling trees or branches. Results suggest that gaps have closed after around 15 years. More data are needed to verify this.     

Stand structure and regeneration in a Kamchatka mixed boreal forest

Abstract. A 1‐ha plot was established in a Betula platyphylla‐Picea ajanensis mixed boreal forest in the central Kamchatka peninsula in Russia to investigate stand structure and regeneration. This forest was relatively sparse; total density and stand basal area were 1071/ha and 25.8 m²/ha, respectively, for trees > 2.0 cm in trunk diameter at breast height (DBH). 25% of Betula regenerated by sprouting, and its frequency distribution of DBH had a reverse J‐shaped pattern. In contrast, Picea had a bimodal distribution. The growth rates of both species were high, reaching 20 m in ca. 120 yr. The two species had clumped distributions, especially for saplings. Betula saplings were not distributed in canopy gaps. Small Picea saplings were distributed irrespective of the presence/absence of gaps, while larger saplings aggregated in gaps. At the examined spatial scales (6.25–400 m²) the spatial distribution of Betula saplings was positively correlated with living Betula canopy trees and negatively with dead Picea canopy trees. This suggests that Betula saplings regenerated under the crowns of Betula canopy trees and did not invade the gaps created by Picea canopy trees. The spatial distribution of Picea saplings was negatively correlated with living and dead Betula canopy trees and positively with dead Picea canopy trees. Most small Picea seedlings were distributed under the crowns of Picea trees but not under the crowns of Betula trees or in gaps. This suggests that Picea seedlings establish under the crowns of Picea canopy trees and can grow to large sizes after the death of overhead Picea canopy trees. Evidence of competitive exclusion between the two species was not found. At a 20 m × 20 m scale both skewness and the coefficient of variation of DBH frequency distribution of Picea decreased with an increase in total basal area of Picea while those of Betula were unchanged irrespective of the increase in total basal area of Betula. This indicates that the size structure of Picea is more variable with stand development than that of Betula on a small scale. This study suggests that Betula regenerates continuously by sprouting and Picea regenerates discontinuously after gap formation and that the species do not exclude each other.     

Tree population studies in low-diversity forests,Guyana. I. Floristic composition and stand structure

Studies were undertaken on the floristic composition and stand structure of four 1 hectare plots in the lowland forests of Kurupukari, Guyana. A total of 3897 trees, covering 153 species and 31 plant families were recorded at greater than 5 cm diameter at breast height (dbh). The number of species per hectare ranged from 61 to 84 (>5.0 cm dbh) and 50–71 (>10.0 cm dbh). The total number of trees per hectare varied two-fold between study plots, with 45–50% of the trees within the 5–10 cm size-class. Mean total basal area varied from 32.39–34.63 m² per 100 m². The four most dominant plant families represented 43.8% of the total number of trees, while representing only 11.2% of the species. No one plant family dominated in more than one of the four study plots, and all four plots held at least one plant family with more than 20% of the total number of trees. Although 14 tree species were common to all four plots, only 26%–35% of the species were represented by a single tree. Between three and seven species represented 50% of the trees within all size-classes, with species dominance occurring within the highest density plot.These tropical forest types of central Guyana may represent some of the lowest diversity forests in the neotropics, whereby the total number of tree species is relatively limited, typically with six dominant canopy species, but the relative abundance of these species is highly variable between the forest types. Mechanisms influencing the competitive interactions associated with species dominance are discussed in relation to the importance of mycorrhizae and the persistence of species dominance.     

Vegetation history since the last glacial period in the Mikata lowland,the Sea of Japan area,western Japan

Local and regional vegetation since the last glacial period was reconstructed on the basis of a palynological study of sediment at Iwaya, in the Sea of Japan area, western Japan. During the interstade (before about 30 000 years BP), forests were composed predominantly ofCryptomeria japonica withTsuga sieboldii and cool-temperate deciduous broad-leaved trees. In the pre-full-glacial, the full-glacial and the early late-glacial (30 000-12 000 years BP), forests were dominated by temperate (montane) and boreal (subalpine) Pinaceae andBetula. During the early full-glacial, the pinaceous forests were mixed with cool-temperate trees such asFagus crenata. In the late full-glacial (18 000-16 000 years BP), the maximum development of pinaceous conifer forests was recognized. Cool-temperate broad-leaved forests composed mainly ofF. crenata andQuercus (Lepidobalanus) replaced the pinaceous forests at about 12 000 years BP and were maintained to the early postglacial.Cryptomeria japonica was distributed around the Mikata lowland during the last glacial.Cryptomeria japonica, which began to increase at 16 000 years BP, increased abruptly in the early postglacial and spread throughout the postglacial in the lowlands. After 6300 years BP, lucidophyllous forests composed mainly ofQuercus (Cyclobalanopsis) andCastanopsis were established in the Mikata district; this was later than in the inland and the Pacific Ocean areas in the Kinki region, western Japan. In historic times (afterca 2000 years BP), secondary forest ofPinus densiflora, which can grow as a pioneer in disturbed habitats, spread.     

Differentiation and maintenance of topo-community patterns with reference to regeneration dynamics in mixed cool temperate forests in the Chichibu Mountains,central Japan

Topo-community structure and dynamics were studied in mixed cool temperate forests, using the regeneration dynamics, to clarify the maintenance mechanisms of community patterns along a microtopographic gradient. The 76 stands studied were classified into two groups (i.e. convex slope and concave slope stands). The soil surface was more eroded on the concave slope than on the convex slope, while water potential was not significantly different between topographies. On the convex slope, even-aged patches alternated between young phase patches dominated by shade intolerant species, such asAcer rufinerve andBetula grossa, and mature phase patches, withTsuga sieboldii andFagus crenata. A slower lateral growth rate ofTsuga canopy trees and the absence of suppressed saplings in the mature phase may prolong the gap phase, which provides a favorable situation to shade-intolerant species. On the concave slope, patch structure was less clear, and process of replacement of canopy species by previously suppressed individuals of the same species was seen in the mature phase, which was mainly composed ofF. crenata, Fagus japonica, Acer sieboldianum andStewartia pseudo-camellia. Gaps on the concave slope were formed frequently but were generally closed within 10 years by lateral growth of deciduous canopy trees and by upgrowth of suppressed trees, and thus some individuals underwent recurrent periods of suppression until they reached the canopy. We concluded that soil surface stability and gap encroachment pattern are critical to the maintenance of the community pattern along a microtopographic gradient.     

Patterns of tree replacement: canopy effects on understory pattern in hemlock-northern hardwood forests

The effect of canopy trees on understory seedling and sapling distribution is examined in near-climax hemlock-northern hardwood forests in order to predict tree replacement patterns and assess compositional stability. Canopy trees and saplings were mapped in 65 0.1-ha plots in 16 tracts of old-growth forests dominated by Tsuga canadensis, Acer saccharum, Fagus grandifolia, Tilia americana, and Betula lutea in the northeastern United States. Seedlings were tallied in sub-plots. Canopy influence on individual saplings and sub-plots was calculated, using several indices for canopy species individually and in total. For each species sapling and seedling distributions were compared to those distributions expected if saplings were located independently of canopy influence. Non-random distributions indicated that sapling and seedling establishment or mortality were related to the species of nearby canopy trees. Hemlock canopy trees discriminate against beech and maple saplings while sugar maple canopy favors beech saplings relative to other species. Basswood canopy discourages growth of saplings of other species, but produces basal sprouts. Yellow birch saplings were rarely seen beneath intact canopy. Since trees in these forests are usually replaced by suppressed seedlings or saplings, canopy-understory interactions should influence replacement probabilities and, ultimately, stand composition. I suggest that hemlock and basswood tend to be self-replacing, maple and beech tend to replace each other, and birch survives as a fugitive by occupying occasional suitable gaps. This suggests that these species may co-exist within stands for long periods with little likelihood of successional elimination of any species. There is some suggestion of geographical variation in these patterns.     

Long‐term mortality patterns of the deep‐rooted Acacia erioloba: The middle class shall die!

Question: Is there a relationship between size and death in the long‐lived, deep‐rooted tree, Acacia erioloba, in a semi‐arid savanna? What is the size‐class distribution of A. erioloba mortality? Does the mortality distribution differ from total tree size distribution? Does A. erioloba mortality distribution match the mortality distributions recorded thus far in other environments? Location: Dronfield Ranch, near Kimberley, Kalahari, South Africa. Methods: A combination of aerial photographs and a satellite image covering 61 year was used to provide long‐term spatial data on mortality. We used aerial photographs of the study area from 1940, 1964, 1984, 1993 and a satellite image from 2001 to follow three plots covering 510 ha. We were able to identify and individually follow ca. 3000 individual trees from 1940 till 2001. Results: The total number of trees increased over time. No relationship between total number of trees and mean tree size was detected. There were no trends over time in total number of deaths per plot or in size distributions of dead trees. Kolmogorov‐Smirnov tests showed no differences in size class distributions for living trees through time. The size distribution of dead trees was significantly different from the size distribution of all trees present on the plots. Overall, the number of dead trees was low in small size classes, reached a peak value when canopy area was 20 ‐ 30 m², and declined in larger size‐classes. Mortality as a ratio of dead vs. total trees peaked at intermediate canopy sizes too. Conclusion: A. erioloba mortality was size‐dependent, peaking at intermediate sizes. The mortality distribution differs from all other tree mortality distributions recorded thus far. We suggest that a possible mechanism for this unusual mortality distribution is intraspecific competition for water in this semi‐arid environment.     

SEEDLING ESTABLISHMENT IN FORESTS AFFECTED BY TEPHRA FROM MOUNT ST. HELENS

Seedling density on permanent plots at five sites was monitored for the first four summers following the deposition of 4.5 to 15 cm of tephra from the 1980 eruption of Mount St. Helens. Because the old-growth forests at the sites were not destroyed by the volcanic eruption, the plots were under a normal tree canopy. Almost no seedlings established in 1980. By 1983 tree seedling density exceeded 35 m^−-2 at all sites. Tsuga heterophylla seedlings were most common, with Abies amabilis seedlings next in abundance. A dense layer of small trees may develop on the tephra and long-term forest dynamics could be affected. There was no successful invasion by species not already present in the stands. Seedlings of some forest herbs, especially Tiarella trifoliata, were common, but many other common forest species produced none. It is still problematical if or when these species will return to areas of the forest from which they were extirpated.     

Dendrochronological analysis of the canopy history of two Ohio old-growth forests

This study examined the temporal patterns of establishment, suppression, and release of major tree species in two old-growth Ohio forest remnants as a means to determine the past disturbance history of these forests. Increment cores were taken from a total of 154 trees from two well-drained, upland plots and two poorly-drained, bottomland plots in each of the two forested areas. Acer saccharum and Fagus grandifolia exhibited multiple episodes of suppression and release prior to becoming canopy trees, and could tolerate suppressions as long as 84 years. In contrast, Quercus macrocarpa, Q. muehlenbergii, Prunus serotina, and Acer saccharinum rerely exhibited any tolerance to suppression and appeared to have entered the canopy after single disturbances had opened large areas of canopy. There was clear synchrony in the temporal pattern of establishment and final release from suppression among trees from bottomland plots scattered throughout the stands, indicating that relatively large disturbances were important in these poorly-drained areas. In contrast, there was little synchrony among trees from well-drained upland plots, except in a single instance where selective cutting of Quercus trees opened the canopy. Thus, the canopy of upland site was likely subjected only to small disturbances resulting from the death of one or a few trees. At the whole of forest level, there was evidence of episodic recruitment of canopy trees in both forests. Establishment of Fraxinus spp. and Quercus spp. were particularly episodic, and few Fraxinus or Quercus trees alive today established during the last century. These data suggest that large disturbances have affected canopy dynamics of both upland and bottomland areas prior to 1900 and in bottomland forests through this century. In contrast, disturbances in upland areas during this century have been restricted to small, treefall-generated canopy gaps.     

How do trees die? Mode of death in northern Amazonia

Question: How do trees die in high‐mortality and low‐mortality Amazonian forest regions? Why do trees die in different ways? Location: Humid, lowland forests in Amazonian Peru and Venezuela. Methods: Patterns of multiple treefall and mode of death (standing, broken or uprooted) were recorded for trees ≥10 cm in diameter in permanent plots. Logistic regression was used to relate mode of death to tree diameter, relative growth rate and wood density. Results: Frequency of multiple death events was higher in high‐mortality northwestern (NW) than in low‐mortality northeastern (NE) Amazonia, but these events were small, averaging two trees killed per multiple death event. Breakage was the dominant known mode of death (51±8%) in the NW, with half of fatal breakages caused by other treefalls or breakages. Small and slow‐growing trees were more prone to breaking than uprooting. In NE Amazonia, the dominant known mode of death was standing (48±10%); these trees tended to be relatively large and slow growing. Broken trees in NE forests have a lower wood density than uprooted trees. Conclusions: The major mortality mechanisms differ in the two regions. In the NW it involves an interaction between physiological failure and mechanical failure (small size, slow growth and broken mode). In the NE it is mainly driven by physiological failure (large size, slow growth and standing mode). We propose that by creating different‐sized gaps the different dominant modes of death would favour species from different functional groups and so help to maintain the contrasting functional composition and mortality rates of the two regions.     

Reactions of the herb and moss layer,tree saplings and the shrub layer to tree deaths in forests of the Wielkopolska National Park (Western Poland)

This study examines the impact of canopy and canopy gaps on the development of lower forest layers in five protected phytocoenoses of oak-hornbeam habitats (natural and regeneration stands) and oak-pine habitats in the Wielkopolska National Park (WPN). In the studied forests the most common form of dead trees are those which are uprooted (45–59%), while the most frequently dying tree is pine (40–88%). The total area of gaps in relation to the studied forest area ranges from 329 to 2356 m²/ha.     

Causes of spatial patterns of dead trees in forest fragments in Illinois

Natural disturbances introduce spatial heterogeneity into forests by causing non-random mortality of trees. We examined whether wind was the primary cause of spatial patterns of dead trees at fragment- and individual tree-levels in three fragments of temperate deciduous forests in Illinois, USA. Dead trees and wind-caused types of mortality were expected to be higher at forest edges, on windward aspects, in poorly-drained soils, and adjacent to existing canopy gaps. The extent of wind-related mortality was determined by comparing spatial and temporal patterns of dead trees, as well as characteristics of trees downed by single windstorms versus all dead trees. At the fragment-level, we used randomly located quadrats of 25×25 m to sample edge and interior areas of Trelease Woods, Brownfield Woods, and Hart Woods in 1995-1996 and again in 1999-2000. We noted type of mortality (standing dead, snapped-off, or uprooted trees), and measured DBH ( 10 cm) and direction of fall of each dead tree. The same measures were made for trees felled by two single storms in 1994. At the individual tree-level, domino effects were evaluated by comparing openness surrounding target treefalls vs. an equal sample size of living trees. The study provided limited evidence that wind caused spatial patterns of dead trees. Instead, spatial patterns of dead trees in the fragments accumulated from domino effects at the individual tree-level in two of the three fragments. Dead trees were more associated with preexisting gaps. Contrary to our predictions at the fragment-level, the frequency of dead trees was not greater at edges, on windward aspects, or in poorly drained soils. This study demonstrated the complexity of spatial patterns of dead trees in forest fragments. The significant domino effects indicated that the occurrence of dead trees was not random, but determined by previous disturbances.     

Composition and distribution of vascular epiphytes in a tropical semideciduous forest,Ghana

The composition and distribution of vascular epiphytes were studied in two 1‐ha plots in the KNUST Botanic garden, Ghana. One‐hectare plot each was randomly set up in secondary and cultivated forests for the identification and enumeration of trees and shrubs (≥10 cm dbh), and epiphytes. Each tree was carefully examined, noting the presence, positions and life‐forms of all epiphytes. Twenty‐nine epiphyte (29) species belonging to fourteen genera and eleven families were identified in the study. These were hosted by 48 tree species and occurred in three life‐forms: hemi‐epiphytes (45%), casual epiphytes (45%) and true epiphytes (10%). The vascular epiphyte species made up 25.7% of all the identified plant species (excluding herbs and climbers) encountered. Host species (P < 0.001), habitat (P = 0.001) and their interaction (P < 0.001) had strong effects on epiphyte composition in the forests. Moraceae was the most dominant family (44.8%), while Nephrolepis undulata J. Sm. and N. biserrata (Sw.) Scott. were the commonest species of epiphytes. In terms of vertical distribution, most epiphytes were located on the trunk, while a few occurred in the canopy.     

Gap-scale disturbance processes in secondary hardwood stands on the Cumberland Plateau,Tennessee, USA

Disturbance regimes in many temperate, old growth forests are characterized by gap-scale events. However, prior to a complex stage of development, canopy gaps may still serve as mechanisms for canopy tree replacement and stand structural changes associated with older forests. We investigated 40 canopy gaps in secondary hardwood stands on the Cumberland Plateau in Tennessee to analyze gap-scale disturbance processes in developing forests. Gap origin, age, land fraction, size, shape, orientation, and gap maker characteristics were documented to investigate gap formation mechanisms and physical gap attributes. We also quantified density and diversity within gaps, gap closure, and gap-phase replacement to examine the influence of localized disturbances on forest development. The majority of canopy gaps were single-treefall events caused by uprooted or snapped stems. The fraction of the forest in canopy gaps was within the range reported from old growth remnants throughout the region. However, gap size was smaller in the developing stands, indicating that secondary forests contain a higher density of smaller gaps. The majority of canopy gaps were projected to close by lateral crown expansion rather than height growth of subcanopy individuals. However, canopy gaps still provided a means for understory trees to recruit to larger size classes. This process may allow overtopped trees to reach intermediate positions, and eventually the canopy, after future disturbance events. Over half of the trees located in true gaps with intermediate crown classifications were Acer saccharum, A. rubrum, or Liriodendron tulipifera. Because the gaps were relatively small and close by lateral branch growth of perimeter trees, the most shade-tolerant A. saccharum has the greatest probability of becoming dominant in the canopy under the current disturbance regime. Half of the gap maker trees removed from the canopy were Quercus; however, Acer species are the most probable replacement trees. These data indicate that canopy gaps are important drivers of forest change prior to a complex stage of development. Even in relatively young forests, gaps provide the mechanisms for stands to develop a complex structure, and may be used to explain patterns of shifting species composition in secondary forests of eastern North America.