Roosting Requirements of Two Frugivorous Bats (Sturnira lilium and Arbiteus intermedius) in Fragmented Neotropical Forest1

As tropical forest fragmentation accelerates, scientists are concerned with the loss of species, particularly those that play important ecological roles. Because bats play a vital role as the primary seed dispersers in cleared areas, maintaining healthy bat populations is critical to natural forest regeneration. Observations of foraging bats suggest that many Neotropical fruit‐eating species have fairly general habitat requirements and can forage in many different kinds of disturbed vegetation; however, their roosting requirements may be quite different. To test whether or not general foraging requirements are matched by equally broad roosting requirements, we used radiotelemetry to locate roost sites of two common frugivorous bat species (Sturnira lilium and Artibeus intermedius) in a fragmented forest in southeastern Mexico. Sturnira lilium roosted inside tree cavities and selected large‐diameter roost trees in remnant patches of mature forest. Fewer than 2 percent of trees surveyed had a mean diameter equal to or greater than roost trees used by . S. lilium, Artibeus intermedius roosted externally on branches and vines and under palm leaves and selected roost trees of much smaller diameter. Compared to random trees, roost trees chosen by A. intermedius were closer to neighboring taller trees and also closer in height to these trees. Such trees likely provide cryptic roosts beneath multiple overlapping crowns, with sufficient shelter from predators and the elements. While males of A. intermedius generally roosted alone in small trees within secondary forest, females roosted in small groups in larger trees within mature forest and commuted more than three times farther than males to reach their roost sites. Loss of mature forest could impair the ability of frugivorous bats to locate suitable roost sites. This could have a negative impact on bat populations, which in turn could decrease forest regeneration in impacted areas.     

Structure and dynamics of subalpine forests in the Wang Lang Natural Reserve, Sichuan, China

Population structure (size, age, spatial patterns) and radial growth patterns are used to analyze regeneration patterns of Abies faxoniana, Betula albosinensis, Betula utilis, Larix potaninii, Picea purpurea, and Sabina saltuaria and reconstruct disturbance history in 8 subalpine forest stands in Wang Lang Natural Reserve, Sichuan, China. In old-growth stands tree regeneration occurs in tree-fall gaps whereby A. faxoniana, Betula sp., P. purpurea, and S. saltuaria persist at stand scales by gap-phase regeneration. Clump sizes of young populations are similar to canopy gap sizes but clumps sizes vary among species. Young Betula patches are larger than those of A. faxoniana suggesting that gap-partitioning by size contributes to species coexistence in mixed stands. Picea purpurea and S. saltuaria are longer lived than A. faxoniana which may compensate for lower recruitment and prevent their replacement by A. faxoniana. Tree regeneration and community structure are also influenced by the understory bamboo Fargesia denudata. Seedlings, saplings, and shrub density all decline with an increase in bamboo cover. Species that regenerate in old-growth forest also regenerate after flooding as do species that establish only on bare substrates (i.e. Larix potaninii, Prunus sp.). Structural and compositional patterns in Wang Lang forests are a reflection of disturbance history, canopy species life history attributes such as dispersal ability, shade tolerance, growth rates, and longevity, and competition of trees and shrubs with understory bamboos.     

The limited role of canopy gaps in the successional dynamics of a mature mixed Quercus forest remnant

Question: Are canopy gap dynamics responsible for driving the structural and compositional changes that have occurred over a 26‐year period in a mature Quercus forest remnant? Location: Dobbs Natural Area, an unlogged 3.6‐ha forest preserve in west‐central Indiana, USA. Methods: We analyzed mapped permanent plot data for a site that illustrates a trend common in Quercus‐dominated forests in eastern North America, where recruitment of new stems is dominated by mesophytic, shade‐tolerant species such as Acer saccharum, rather than Quercus. We developed a GIS database from stand census measurements taken in 1974 and 2000, employing it to conduct tree‐by‐tree comparisons that allow direct determination of ingrowth, mortality and survivorship, and to relate the spatial patterns of subcanopy dynamics to canopy gap occurrence. Results: The re‐census shows modest changes in canopy composition, but much greater turnover in the subcanopy. Nearly half of all individuals originally present died; much of this mortality resulted from a major decline in subcanopy Ulmus americana. While overall density remained fairly constant, the subcanopy experienced substantial ingrowth of shade‐tolerant Acer saccharum, Fagus grandifolia, and Tilia americana. Canopy gaps, although forming at rates in the upper range of regional averages, did not significantly benefit subcanopy populations of Quercus spp. or most other taxa with limited shade tolerance. Conclusions: Canopy gaps play a minor role in driving the recent demographic trends of this stand. The spatial and temporal scales of light availability in gaps do not support regeneration of most shade‐intolerant species. Compositional change parallels a historical shift in light regimes.     

Use of soil transfer for reforestation on abandoned mined lands in Alaska

Soil transfers from an intermediate successional site and a mature forest site were applied to Populus balsamifera L. cuttings and Alnus crispa (Ait.) Pursh seedlings placed on an abandoned mined site in south central Alaska to improve plant establishment. Mycorrhizal fungi in the soil transfers from the two successional stages were hypothesized to have different effects on plant species that colonize disturbed sites at different times or on different substrates. The site consisted of coarse, dry, low-nutrient spoils and was naturally colonized by scattered P. balsamifera but not A. crispa, although seed sources for both were adjacent to the site. Physical dimensions of the transplanted seedlings and cuttings were measured at the beginning and end of each growing season. Selected plants were harvested at the end of the 2-year study and examined for mycorrhizal formation, current growth, and leaf tissue nutrient concentrations. Both plant species were taller when treated with the soil transfers from the mature forest than with soils from the intermediate site although the increase for A. crispa was greater. Physical dimensions, current growth, and nutrient concentrations were greater when A. crispa was treated with the mature soil transfer compared with the intermediate soil transfer. Mycorrhizae which infected Alnus were predominantly a brown woody type, while other types accounted for greater relative mycorrhizal infection percentage on Populus. Insufficient quantities of mycorrhizal inoculum of suitable species, as well as low moisture and low nutrient conditions, may be factors limiting A. crispa colonization on primary disturbed sites in south central Alaska.     

Establishment and growth of mangrove seedling in mangrove forests of southern Thailand

The demography and growth of mangrove seedlings in southern Thailand were studied for an 18-month period. Plots for measurement of seedlings were established using gradients of light and surface soil texture. The number of established propagules peaked from April to August, the peak forRhizophora being earlier than that forBruguiera. Under the mangrove forest canopy, the survivorship of seedlings established during May to December 1982 was 33% in June 1983. Light conditions and soil texture did not strongly affect initial seedling establishment, and the relationship between species seed size and water depth at a particular site influenced that were able to become established there. The elongation growth was large during the rainy season and in open sites. The growth ofR. apiculata Blume,B. parviflora Wight & Arn. andB. cylindrica Blume in open sites was more than ten times that of seedlings established in the shade. It has noted that light conditions affected the growth of young trees (>1 year old) to a greater degree than that of current-year seedlings.     

Structural characterization of a Podocarpus parlatorei and Juglans australis forest in Salta, Argentina

Stands of Subtropical Montane Cloud Forest were studied in areas under different land use regimes near Los Toldos (NW Argentina). Circular plots were used to calculate density and basal area of trees with dbh > 10 cm; and density of trees with dbh < 10 cm. The stands were classified and grouped as a function of basal area. Five structures were recognized, with different proportions of Juglans australis, Podocarpus parlatorei and shade-tolerant species like Blepharocalyx salicifolius, Allophylus edulis and Myrcianthes sp. Less disturbed area stands had a higher basal area and greater dbh for all species, and shade-tolerant species in the canopy. In the most disturbed area, all canopy species were shade intolerant and regeneration was dominated by shade-tolerant species. The differences in composition, basal area, stem diameter class distribution and regeneration indicate that the structure types corresponded to different stages of the successional process, and the regeneration of the most disturbed areas suggest a tendency towards the composition of mature forest.     

Arrested succession in logging gaps: is tree seedling growth and survival limiting?

Thirty years after selective timber harvest in the Kibale National Park, Uganda, many abandoned logging gaps are dominated by Acanthus pubescens, and show little forest recovery. To examine if this arrested successional state was caused by limited tree seedling growth and survival, we planted seedlings of four forest tree species (Albizia grandibracteata, Mimusops bagshawei, Prunus africana and Uvariopsis congensis) in A. pubescens‐dominated logging gaps and in control areas of adjacent forest. To assess if clearing A. pubescens facilitates forest regeneration, we planted seedlings of two species (A. grandibracteata and U. congensis) in small clearings cut within the logging gaps. We examined mortality, growth, herbivory and site characteristics among the treatments. Finally, we described the physical attributes of the A. pubescens‐dominated gaps. Seedlings of all the four species survived and grew equally well in A. pubescens and forest treatments, and most site characteristics were also similar. Seedlings planted in clearings grew more than in either forest or A. pubescens sites. Very few established trees were found in A. pubescens sites, and most of these were near the forest edges. We also discussed the role of elephants (Loxodonta africana) and collapsing A. pubescens canopies in the maintenance of an arrested successional state in these logging gaps.     

Regeneration dynamics and life history differences in southern Chilean Nothofagus forests: a synthesis

Large, infrequent natural disturbances have been proposed as a key component in determining the distribution and abundance of Nothofagus in southern Andean temperate forests. In this study, a comparison of influences of small-to-intermediate-sized natural disturbances and effects of selective logging on the establishment and growth of Nothofagus forests in south-central Chile are synthesized with the results of other forest studies to develop general hypotheses on the regeneration dynamics of southern Chilean Nothofagus alpina forests. A synthesis is given for regeneration pattern in the abundance of 18 temperate rain forests in the Andean Range, Chile. The study aimed to determine the role of life history differences in promoting coexistence of the five main tree species (N. alpina, N. dombeyi, N. pumilio, N. macrocarpa and Laurelia philippiana) in N. alpina-dominated forests. Age data reported for N. alpina, a shade mid-tolerant emergent tree in the temperate rain forests of southern Chile, indicate maximum lifespans > 650 years, figures unprecedented for N. alpina. In low elevation stands, N. alpina coexists with broad-leaved evergreen tree species, such as L. philippiana, L. sempervirens, Persea lingue, and in these stands an intermittent establishment of Nothofagus occurred and appeared to be most dependent on small-to-intermediate disturbances. In high-elevation stands, in contrast, Nothofagus establishment was less dependent on disturbance, regeneration being much more continuous even in the absence of canopy openings. The forests studied provide another example of the general pattern of increasing dependence of Nothofagus on disturbance towards the more productive end of the environmental gradients. As a long-lived pioneer and despite its dependence on disturbance in lowland sites, N. alpina has been subject to selection for complementary growth, adult survivorship, and mid-tolerance to shade. Thus, interspecific differences in juvenile and adult life history characteristics of N. alpina and its competitors may be sufficient to maintain its persistence in the landscape. In conclusion, this study of population structures and replacement patterns provides a comprehensive picture on our understanding of regeneration dynamics and trait differentiation in southern temperate forests by recognizing both the influences of environmental gradients (i.e., altitude) on competing species and the disturbance regimes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Losses of seeds of temperate trees to soil fungi: effects of habitat and host ecology

Evidence suggests that impacts of fungal pathogens on tree recruitment tend to be greater in the forest understory than in openings, and that shade-tolerant trees are less vulnerable than shade-intolerant species. To investigate the role that harmful soil fungi may have in reducing regeneration of temperate trees, we applied fungicide to buried seeds of matched pairs of species differing in their relative shade tolerance and/or successional status (Acer negundo versus Acer saccharum, Prunus virginiana versus Prunus serotina, and Pinus strobus versus Tsuga canadensis), in three habitats that differed in their degree of openness (old field, forest gap, and forest understory). Our results indicated that soil fungi reduced germination of A. negundo, P. virginiana, P. serotina. and T. canadensis, and reduced viability of ungerminated seeds of P. strobus; no significant effects of fungi on seeds of A. saccharum were detected. However, we found seeds were not less likely to survive following burial in forest understory than in gaps. As well, results for only one species pair (A. negundo versus A. saccharum) were consistent with the prediction that shade-intolerant or successional species should be more susceptible to fungal attack than mature forest species. These results contrast with other studies of temperate and especially tropical forest trees.     

Quercus rugosa seedling dynamics in relation to its re-introduction in a disturbed Mexican landscape

Abstract. In order to define the ‘regeneration niche’ and then promote the re-introduction of Quercus rugosa into a disturbed area we studied seed predation, germination, seedling survival and growth of this species at three different sites: the forest interior, the forest border and a disturbed site, along a disturbance gradient. Acorn removal on the ground was high at the sites, with higher removal rates from 25-seed clusters than from 5- and 1-seed clusters. More seeds were removed at the forest border than at the other two sites, although in all cases some seeds remained in the soil for periods long enough for them to germinate and produce seedlings. Seed germination and seedling establishment were more successful and similar in the forest interior and the forest border, and much less successful at the disturbed site, due to acorn desiccation. Seedling survival was strongly affected by site, with the highest survival at the forest border and the lowest survival at the disturbed site after a year. The subsequent dry season, which was particularly harsh, caused a drop in survival both at the forest border and the disturbed site. In the former, a closer analysis revealed that partially shaded microsites enhanced survival, while in the latter overall survival was very low, with approximately 5 % of the seedlings surviving, compared to 50 % final survival at the other sites. There was a positive correlation between seed size and seedling survival at the forest border. The three size variables measured (seedling height, basal diameter and crown area) did not increase noticeably during the 20-month study period at the forest interior, while growth still occurred at the forest border and the disturbed site. However, during the two dry seasons there were large drops in mean seedling height and crown area at the disturbed site, due to more frequent dieback, causing large variation in seedling size. These results show that extensive re-introduction of this species to the disturbed area will probably fail under present conditions, and that careful selection or duplication of microsites that mimic the partial shade of the forest border is a sound method for improving oak establishment at this site.     

Comparing the Importance of Seedbed and Canopy Type in the Restoration of Upland Thuja occidentalis Forests of Northeastern Minnesota

In cold‐temperate ecosystems of the upper Great Lakes Region, evergreen conifer‐dominated forests were once common. As a result of past management practices, early‐successional deciduous forests now dominate the landscape. Embedded in this matrix are stands of shade‐tolerant conifers, including Thuja occidentalis. For the past several decades, large‐scale T. occidentalis regeneration in remnant T. occidentalis and adjacent Betula papyrifera forests has not occurred. Using a combination of restoration experiments and field surveys at three study sites on the Lake Superior Highlands, Minnesota, U.S.A., we examined safe sites for T. occidentalis regeneration under both Thuja and Betula canopy types. This study focused on the colonization and establishment phases of regeneration, differentiating among safe‐site components for T. occidentalis. Seedbed type determined colonization success, with higher rates on conifer logs than on B. papyrifera logs, mounds, or pits. Mortality rates for seedlings on natural decayed wood seedbeds were higher under Thuja canopy than under Betula canopy, but the reverse was true for seedlings on manipulated seedbeds, suggesting that seedbed type was more important than the dominant canopy type. Growth rates for seedlings on moved log segments were greater under the Betula than the Thuja canopy type, but seedlings on natural decayed wood seedbeds did not exhibit this difference. Results indicated that T. occidentalis regeneration was more limited in Betula forest by seedbed availability, while in Thuja forest canopy conditions were more limiting.     

Community structure of a species-rich temperate forest,Ogawa Forest Reserve,central Japan

Species assembly and niche differentiation were studied, and future species composition was predicted by simple Markov models, in an old-growth deciduous forest at the Ogawa Forest Reserve in central Japan. The dominant species in our 6ha study site are Quercus serrata, Fagus japonica, and F. crenata.An ordination by population parameters revealed four different combination 3 of life forms and regeneration niches. Cluster analysis based on interspecific spatial correlation revealed three groups of species. The species in cluster A, such as F. japonica, occurred at the bottom of the valley, while those in cluster B, such as Q. serrata, occurred along ridges. Species in cluster C, such as F. crenata, did not show any particular habitat preference. Clusters B and C were further divided into three smaller clusters (a-c). Both clusters Ba and Bb included shade intolerant species. Species in cluster Ba had large clump sizes (>1500 m²), reflecting regeneration following large-scale disturbances. Species in cluster Bb had smaller clump sizes (<400 m²) reflecting regeneration following local disturbances. Clusters Ca and Cb mainly included shade tolerant species and shade intolerant species, respectively. Markov models predicted that shade intolerant species, particularly those in cluster Ba, would be eliminated. Thus, species coexist by differentiation of both habitats and regeneration niche in this forest community. Some species such as Quercus serrata, however, regenerate following large-scale disturbances or human activity.     

The role of canopy gaps in the regeneration of coastal dune forest

In regenerating coastal dune forest, the canopy consists almost exclusively of a single species, Acacia karroo. When these trees die, they create large canopy gaps. If this promotes the persistence of pioneer species to the detriment of other forest species, then the end goal of a restored coastal dune forest may be unobtainable. We wished to ascertain whether tree species composition and richness differed significantly between canopy gaps and intact canopy, and across a gradient of gap sizes. In three known‐age regenerating coastal dune forest sites, we measured 146 gaps, the species responsible for gap creation, the species most likely to reach the canopy and the composition of adults, seedlings and saplings. We paired each gap with an adjacent plot of the same area that was entirely under intact canopy and sampled in the same way. Most species (15 of 23) had higher abundance in canopy gaps. The probability of self‐replacement was low for A. karroo even in the largest gaps. Despite this predominance of shade‐intolerant species, regenerating dune forest appears to be in the first phase of succession with ‘forest pioneers’ replacing the dominant canopy species. The nature of these species should lead to successful regeneration of dune forest.     

Anthropogenic disturbances and plant biodiversity in forests of Uttaranchal,central Himalaya

Eight forest types varying in disturbance frequencies were identified along an elevational gradient in Uttaranchal, central Himalaya. Low elevation forests were close to human habitation and had high disturbance frequency, while high elevation forests were situated far from the human habitation and had low disturbance. The dominant tree species at low elevation were Pinus roxburghii and Quercus leucotrichophora, while Q. floribunda and Q. semecarpifolia dominated the high elevation forests. Pyracantha crenulata was the shrub present in all the forests except in Q. semecarpifolia forest and Anaphalis contorta, a herb species, was present in all the forests. Disturbance decreased the dominance of single species and increased the plant biodiversity by mixing species of different successional status. Species richness and diversity for all the vegetation layers were higher in low elevation–high disturbance forests. Mean tree density decreased from high to moderate and increased in low disturbance. The shrub density decreased from high to low disturbance while the reverse occured for herbs. High proportion of early successional species in disturbed forests indicated that disturbance induces succession. The mean number of young individuals increasing from high to low disturbance indicates that disturbance adversely affects regeneration. But, however, the high number of young individuals of Coriaria nepalensis, a small non-leguminous nitrogen fixing tree, in disturbed forests shows that the forest is regenerating. This species could be helpful in the re-establishment of original vegetation through triggering the regeneration of these forests. High elevation–low disturbed forests separated from low elevation–high disturbed forests. Forest type and elevation may have more influence on tree richness while shrub and herb richness may be more sensitive to disturbance and forest types.     

Defining species guilds in the Central Hardwood Forest,USA

Tree regeneration outcomes are challenging to generalize and difficult to predict. Many tree species can establish new propagules in a variety of post-disturbance environments and many different reproductive mechanisms may be used. In order to develop conceptual models that accurately reflect reproductive potential, we need a better understanding of the similarities in regeneration ecology among species. We used information from the forest ecology literature to evaluate the reproductive attributes of sixty-two tree species in the central hardwood region of the eastern United States. Each species was classified categorically for features such as flowering, seed production and dispersal, seed dormancy, germination requirements, seedling characteristics, and vegetative reproduction. Cluster analysis (Jaccard's similarity coefficient, complete linkage method) and ordination (homogeneity analysis) were used to separate nine groups (guilds) of species that had similar reproductive attributes. Individual attributes that had high variance in the first and second dimensions included: seed banking, seed dispersal, seedling shade tolerance, and seedbed requirements. Members of each guild had similar levels of reproductive specialization and guilds were either pioneer-like, opportunistic, or persistent. Pioneer guilds included: short-lived or fugitive species that colonize sites rapidly and are too shade intolerant to replace themselves; shade-tolerant species that colonize frequently disturbed sites; and stress-tolerant pioneers that survive on dry or nutrient-poor sites. Opportunistic guilds contained species that are remarkably versatile in their reproductive effort. The most flexible opportunists can colonize new sites, maintain seed in a seed bank, sprout from existing stems and persist as a seedling or sapling bank. Persistent guilds contain species that develop and maintain advance regeneration. These include: species with moderate understory tolerance that regenerate via cycles of dieback and resprouting; and more tolerant species that maintain seedling or sapling banks. Our regeneration guilds may provide a useful approach for more realistically representing large and diverse sets of tree species in forest ecosystem models.     

Characteristics of the Psidium cattleianum invasion of secondary rainforests

Strawberry guava (Psidium cattleianum) is a shade‐tolerant shrub or small tree invader in tropical and subtropical regions and is considered among the world's top 100 worst invasive species. Studies from affected regions report deleterious effects of strawberry guava invasion on native vegetation. Here we examine the life history demographics and environmental determinants of strawberry guava invasions to inform effective weed management in affected rainforest regions. We surveyed the vegetation of 8 mature rainforest and 33 successional sites at various stages of regeneration in the Australian Wet Tropics and found that strawberry guava invasion was largely restricted to successional forests. Strawberry guava exhibited high stem and seedling densities, represented approximately 8% of all individual stems recorded and 20% of all seedlings recorded. The species also had the highest basal area among all the non‐native woody species measured. We compared environmental and community level effects between strawberry guava‐invaded and non‐invaded sites, and modelled how the species basal area and recruitment patterns respond to these effects. Invaded sites differed from non‐invaded sites in several environmental features such as aspect, distance from intact forest blocks, as well as supported higher grass and herb stem densities. Our analysis showed that invasion is currently ongoing in secondary forests, and also that strawberry guava is able to establish and persist under closed canopies. If left unchecked, strawberry guava invasion will have deleterious consequences for native regenerating forest in the Australian Wet Tropics.     

Climbing asparagus,Asparagus scandens Thunb.: a South African in your forest patch

Climbing asparagus, Asparagus scandens Thunb., currently has a patchy distribution in New Zealand, mainly in disturbed lowland forest remnants near towns and the coast, predominantly in the North Island but found as far south as Banks Peninsula. Here it prevents regeneration of native seedlings and smothers or strangles understorey shrubs and saplings. Based on its current distribution, it tolerates open sunny sites to deep shade and damp or dry forest. Left unchecked and aided by birds and humans, climbing asparagus is likely to spread considerably in the next decade, consolidating its current distribution in New Zealand. Several characteristics of the species make it difficult to control: the small, fine seedlings are hard to find, adult plants entwine with native plants, much of the plant’s biomass is underground and the apical meristem will resprout if left in the ground after control.     

A long‐term record of Nothofagus dominance in the southern Andes,Chile

Abstract The general model of regeneration dynamics in Nothofagus forests of southern South America could have value in community ecology if predictive relationships between disturbance history, functional traits and site attributes could be identified. Examined here is the proposal that on favourable sites shade‐intolerant Nothofagus are likely not to survive in competition with shade‐tolerant, broad‐leaved evergreen taxa of temperate rain forests, and persistence, thus, is dependent on periodic coarse‐scale disturbance. Comparison of stand dynamics of three old‐growth Nothofagus forests at different elevations in the southern Andes, Chile where deciduous Nothofagus alpina dominates the upper canopy, and examination of the life history trade‐offs of this variation were made. Stem density of all stems ≥5.0 cm d.b.h. was 233–303 stems per hectare, and basal area was 123.9–171.0 m²ha^?1. Maximum lifespan of N. alpina was found to be greater than ca 640 years, exceeding all previously reported ages for this species in the region. Forests had a stable canopy composition for this long‐term, but some appeared to lack effective regeneration of N. alpina in recent years. Regeneration of N. alpina was generally greater in disturbed stands and higher elevation than in undisturbed stands and at lower elevation. Recruitment emerged to be strongly affected by competitive over‐ and understorey associates. There was a gradient of increasing dependence of N. alpina on disturbance towards the more productive end of the environment gradients, and hence less dependence of N. alpina on disturbance for its regeneration towards higher elevation. The study confirms that changes in forest composition may be explained by processes occurring in accordance with the predictions of the existing model of Nothofagus regeneration dynamics, providing stronger evidence specifically directed at mid‐tolerant N. alpina, and by factoring out regeneration dynamics on favourable sites. Thus, for N. alpina, trait differences probably contribute to the competitive advantage over its associates in productive habitats, and may be linked to small‐to‐intermediate‐sized disturbances which inevitably occur as older trees die, enabling N. alpina to persist in forests and therefore maintain species coexistence for the long‐term.     

Evidence for the widespread occurrence of ancient forests on cliffs

Aim The objective of this work was to determine if the existence of ancient forests on cliffs was specific to the Niagara Escarpment, Canada, or part of a globally widespread pattern. Location Sixty‐five cliff sites were visited in five countries in the temperate climatic zone, and trees were sampled for age and growth rate on forty‐six of these. Methods Two hundred and twenty‐four core samples or cross‐sections were taken from trees on cliffs that varied in height, aspect, rock‐type, and exposure. General observations were also made of regeneration of the tree species forming the mature canopy, and other habitat conditions. Results The evidence shows that ancient slow‐growing forest occurs on most cliffs. Age and growth rate distributions were similar at all treed sites. Small‐statured Thuja, Juniperus, or Taxus stems with age estimates in excess of 1000 years were found in the United States, the United Kingdom and France, and small Pinus and Quercus stems nearly 400 years in Germany. There was a high rate of recurrence of plants in the genera Polypodium, Asplenium, Cystopteris, Campanula, Rosa, Prunus, Hedera, and Sorbus. Most of the sites appear to be habitats of completely natural origin. Conclusions We conclude that ancient natural forest is a normal feature of cliffs, at least in the temperate zone.