Habitat Assessment, Phytosociology and Conservation of the Tunbridge Filmy-fern Hymenophyllum tunbrigense (L) Sm. in its Isolated Locations in the Vosges Mountains

The Tunbridge Filmy-fern, Hymenophyllum tunbrigense, has been discovered within the last 10 years in several places in a 15 km long by 2 km wide area located in the western part of the Vosges Mountains. After the disappearance of the Saxony and Ardennes sites, these Vosgian locations have to be considered as one of the last continental areas for this fern in Europe, meriting the same level of interest as the well-known Luxembourg locations. In its Vosgian area, the fern occurs in Abies alba forests on small (less than 2.5 m high) and moist sandstone rocks, generally located near running water in wet valleys, at altitudes between 400 and 640 meters a.s.l. As assessed through the impact of the 26 December 1999 storm on one Hymenophyllum location, this fern appears to be sensitive to direct sunlight in the Vosges mountains. Therefore the main measure to ensure its conservation would be to maintain a dense and permanent forest canopy above the Hymenophyllum sites in order to avoid any direct sunlight.     

Impact of river dynamics on the genetic variation of Gypsophila repens (Caryophyllaceae): a comparison of heath forest and more dynamic gravel bank populations along an alpine river

• Alpine rivers are, despite anthropogenic water flow regulation, still often highly dynamic ecosystems. Plant species occurring along these rivers are subject to ecological disturbance, mainly caused by seasonal flooding. Gypsophila repens typically grows at higher altitudes in the Alps, but also occurs at lower altitudes on gravel banks directly along the river and in heath forests at larger distances from the river. Populations on gravel banks are considered non‐permanent and it is assumed that new individuals originate from seed periodically washed down from higher altitudes. Populations in heath forests are, in contrast, permanent and not regularly provided with seeds from higher altitudes through flooding. If the genetic structure of this plant species is strongly affected by gene flow via seed dispersal, then higher levels of genetic diversity in populations but less differentiation among populations on gravel banks than in heath forests can be expected.
• In this study, we analysed genetic diversity within and differentiation among 15 populations of G. repens from gravel banks and heath forests along the alpine River Isar using amplified fragment length polymorphisms (AFLP).
• Genetic diversity was, as assumed, slightly higher in gravel bank than in heath forest populations, but genetic differentiation was, in contrast to our expectations, comparable among populations in both habitat types.
• Our study provides evidence for increased genetic diversity under conditions of higher ecological disturbance and increased seed dispersal on gravel banks. Similar levels of genetic differentiation among populations in both habitat types can be attributed to the species' long lifetime, a permanent soil seed bank and gene flow by pollinators among different habitats/locations.

     

Response of ants to human‐altered habitats with reference to seed dispersal of the myrmecochore Corydalis giraldii Fedde (Papaveraceae)

Anthropogenic habitat disturbance has potential consequences for ant communities. However, there is limited information on the effects of ant responses on associated ecological processes such as seed dispersal. We investigated the effect of disturbance on the abundance, richness, and composition of ant communities and the resulting seed‐dispersal services for a herbaceous myrmecochore, Corydalis giraldii (Papaveraceae), in an undisturbed habitat (forest understory), moderately disturbed habitat (abandoned arable field), and highly disturbed habitat (road verge) on Qinling Mountains, China. In total, we recorded 13 ant species, and five out of these were observed to transport seeds. The community composition of dispersers was significantly different amongst habitats. The richness of the dispersers did not differ among the habitats, but their total abundance varied significantly across habitats and was 21% lower in the road verge than in the abandoned arable fields. The major seed‐dispersing ant species in both the forest understory and the abandoned arable field were large‐bodied (Myrmica sp. and Formica fusca, respectively), whereas the major seed‐dispersing ants found in the road verge were the small‐bodied Lasius alienus. This difference resulted in lower seed removal rates and dispersal distances in the road verge than in the other two habitats. The different dispersal patterns were attributed primarily to differences in dispersing ant abundance and identity, most likely in response to habitats with different degree of anthropogenic disturbance. The possible influence of disturbance on the ecological specialization of ant‐seed dispersal interaction is also discussed.     

Indirect effects of black spruce (Picea mariana) cover on community structure and function in sheep laurel (Kalmia angustifolia) dominated heath of eastern Canada

Recent studies on phenotypic plasticity of plant traits indicate that within-species variation in litter quality may be a significant factor that feeds back on litter decomposition and nutrient cycling rates at the stand level. These findings may be especially significant for understanding biodiversity-stability relationships in species-poor ecosystems that have little functional redundancy among primary producers. We tested the null hypothesis that black spruce and Kalmia were functional equivalents with respect to their structuring roles of subordinate vegetation and their influence on site biogeochemistry. The purpose of the study was to determine the degree to which forest cover exerts top-down control on community structure and function of Kalmia-black spruce communities. This community type dominates much of the forest understory and unforested heathlands in Atlantic Canada. We intensively studied a representative stand of Kalmia heath in Terra Nova National Park in eastern Newfoundland. Thirty-two 0.5 m × 0.5 m sample plots were randomly distributed among five transects bisecting gradients in dominance of black spruce and Kalmia. Light levels, species composition, vascular plant cover and soil respiration rate were determined for each plot. Tissue samples of litter, mature and current year leaves of Kalmia were collected and analyzed for nutrient status. Herbaceous species richness and cover peaked at intermediate light levels. Kalmia foliar N concentration and above-ground biomass increased with increasing shade. Soil respiration rates were strongly related to the light gradient and increased with increasing quality of Kalmia litter inputs. Our data indicate that Kalmias vigour and foliar nitrogen concentrations are greater under black spruce canopy as opposed to heath condition and that the shaded phenotype has relatively benign feedbacks on soil productivity compared to the open-habitat phenotype. In the absence of functional diversity at the species level in these species-poor habitats, phenotypic plasticity in Kalmia appears to be an important dimension of the biodiversity-stability relationship in these communities since our data suggest that this species has the potential either to inhibit or facilitate carbon cycling and the pathway is strongly linked to the presence or absence of overstory cover. The role of forest regeneration as an indirect control of forest soil processes such as carbon and nitrogen cycling in this ecosystem is discussed.     

Diversity of geometrid moths (Lepidoptera: Geometridae) along an Afrotropical elevational rainforest transect

Geometrid moths were investigated at 26 sites on 9 elevational levels along an elevational transect at Mt. Kilimanjaro (Tanzania), stretching from the fine‐grained mosaic of small agroforest plots with combined cultivation of trees, shrubs and crops at 1650 m through mountain rainforest to heathland at 3300 m. We sampled moths manually at light between 19 : 00 and 22 : 00 in the rainy seasons of March to May and October to January in the years 2000, 2001 and 2002. Along the transect, the composition of moth communities changed from a domination by Sterrhinae and Ennominae to a dominance of Larentiinae with increasing elevation. Overall, alpha diversity was very low compared to other tropical mountain regions. Fisher's alpha showed a maximum of 30 in the agroforest mosaic at 1650 m and decreased to values around 12 in the mountain rainforest. Communities of geometrid moths within the forest belt were significantly dissimilar from communities outside the forest. The diversity patterns on Mt. Kilimanjaro can be related to the young age, island‐like position and history of the mountain. These factors have led to the formation of a homogeneous upper mountain rainforest habitat which in turn houses homogeneous moth communities with a low diversity compared to habitats at lower elevations. Here, a heterogeneous habitat mosaic allowing the intrusion of savannah species into this former forest habitat may account for an increased diversity. In the heath zone above the forest, climatic conditions are very harsh, permitting only few specialists to thrive in this ericaceous woodland. Edge effects were discernible at the forest–heathland boundary where some moth species from heathland invaded the closed forest. At the boundary between agroforest and a forest mosaic of exotic Acacia and Eucalyptus forest plantations and natural mountain forest, diversity values remained low as the dominant species Chiasmia fuscataria accounted for far higher proportions than other dominant species in any of the other habitats.     

Cytotype distribution and phylogeography of Hieracium intybaceum (Asteraceae)

Using flow cytometry and amplified fragment length polymorphism (AFLP), we explored the cytogeography and phylogeography of Hieracium intybaceum, a silicicolous species distributed in the Alps and spatially isolated in the Vosges Mountains and the Schwarzwald Mountains. We detected two ploidies, diploid and tetraploid, but no triploid or mixed‐ploidy populations. Whereas diploids are sexual and distributed all across the Alps, tetraploids are apomictic and seem to be confined to the western Alps and the Vosges. We detected a low level of genetic variation. Bayesian clustering identified four clusters/genetic groups, which are partly congruent with the ploidal pattern. The first two groups consisting exclusively of diploids dominate the whole distribution range in the Alps and show east–west geographical separation with a diffuse borderline running from eastern Switzerland to the eastern part of North Tyrol. The third genetic group lacks a defined geographical range and includes diploid and tetraploid plants. The last genetic group comprises tetraploid plants in the French Alps and the Vosges. We suppose that diploids colonized the deglaciated areas from source populations most likely located mainly in the southern part of the recent distribution range and occasionally also in the western Alps. Gene flow and further differentiation likely took place. Apomictic tetraploids most likely originated in the western Alps or in the refugium at the south‐western foot of the Alps. Their rather limited geographical range (partly contrasting with the theory of geographical parthenogenesis) can be explained by their rather recent origin. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 487–498.     

Succession in the southern part of the Canadian boreal forest

Forest succession following fire in a forest mosaic of northwestern Quebec has been studied in order to: (1) describe the successional pathways using communities of different ages and (2) evaluate convergence of successional pathways and possible effect of fire suppression on the establishment of steady-state communities. As a first step, ordination and classification techniques were used in order to remove changes in forest composition which are related to abiotic conditions. Then, ordinations based on tree diameter distributions were used to study shifts in species composition in relation to time since the last fire.Even under similar abiotic conditions, successional pathways are numerous. However, regardless of forest composition after fire, most stands show convergence toward dominance of Thuja occidentalis and Picea mariana on xeric sites and dominance of Abies balsamea and Thuja occidentalis on more mesic sites. Stable communities of >300 yr occur on xeric sites while on mesic sites directional succession still occurs after 224 yr. Nearly all species involved in succession are present in the first 50 yr following fire. Only Abies balsamea and Thuja occidentalis increase significantly in frequency during succession. Following initial establishment, successional processes can generally be explained by species longevity and shade tolerance. Early successional species may be abundant in the canopy for more than 200 yr while the rapid decrease of Picea glauca, a late successional species could be related to spruce budworm outbreaks. Considering the short fire rotation observed (about 150 yr), a steady-state forest is unlikely to occur under natural conditions, though it may be possible if fire is controlled.     

Effects of patch characteristics and isolation on relative abundance of the scarce heath butterfly <Emphasis Type="Italic">Coenonympha hero</Emphasis> (Nymphalidae)

The scarce heath (Coenonympha hero) is an internationally threatened butterfly in Western Europe, where it occurs primarily on hay fields and abandoned arable land in a small-scale agricultural landscape of south-central Scandinavia. Due to afforestation, this habitat is becoming increasingly fragmented in Sweden, and it can be expected that the scarce heath will decline abruptly when threshold conditions for metapopulation persistence are no longer met. We used stepwise polychotomous logistic regression to compare habitat characteristics and isolation measures for patches that harbour large, small or no populations, respectively, in an area of south-western Sweden. We found that patch area, distance to the nearest large population and amount of Galium spp. explained a significant part of the variation in relative abundance among patches. Distance to nearest large population resulted in a better model to predict occupancy than both distance to the nearest inhabited patch and connectivity, which suggests that primarily large populations act as sources for small satellite populations. Today, sites of three of the eight larger populations in the study area have been planted with spruce or pine and will disappear within 20 years. We argue that the disappearance of these patches may very well lead to rapid extinction of the whole metapopulation system.     

Potential Effects of Ateline Extinction and Forest Fragmentation on Plant Diversity and Composition in the Western Orinoco Basin,Colombia

Cattle and agricultural farming in the western Orinoco Basin began in 1555, and since then fragmentation of continuous forest has occurred. We evaluated the effects of the disturbances and the absence of large primates on plant community composition, diversity, and regeneration patterns. Atelines (Lagothrix and Ateles) inhabited the lowlands close to the Andean mountains, but no longer live in fragmented habitats. Their absence may have negative effects on plant populations because atelines play important roles as seed dispersers in neotropical forests, especially for large-seeded plants, which are rarely swallowed by other seed dispersers. We compared 2 1-ha vegetation plots in forest fragments north of the La Macarena Mountains with 7 plots in continuous forest in Tinigua National Park. Both sites share the same climatic conditions and have similar geological origins. There is floristic affinity between forests with similar ecological characteristics; the fragmented forests are also less diverse than the continuous forests. As predicted, the forest fragments have fewer individuals with large seeds. The results suggest that forest fragmentation and local ateline extinctions affect plant communities, reducing diversity and affecting large-seeded plants.     

Habitat‐dependent population regulation in an irrupting population of long‐nosed bandicoots (Perameles nasuta)

We used isodars to analyse habitat‐dependent population regulation by long‐nosed bandicoots Perameles nasuta during an irruption and subsequent population crash in three habitats (heath, woodland and forest) at Booderee National Park, south‐eastern Australia. Specifically, we aimed to see whether patterns of habitat‐dependent population regulation matched a priori estimates of quantitative and qualitative differences between habitats. We also tested if habitat preference changed between the increasing and decreasing phase of the irruption as predicted by the reciprocating dispersal theory. Quantitative differences in habitat quality were indexed by the relative abundance of the main food of long‐nosed bandicoots (terrestrial invertebrates), while qualitative differences were indexed by the availability of refuge from predation (vegetation understorey density). One index of fitness, body weight, was highest in forest, and lowest in heath, suggesting an ideal despotic model of habitat selection. Over the entire course of the irruption, there was density‐dependent habitat selection with forest and woodland both quantitatively superior to heath. This reflected the overall abundance of invertebrates with highest abundance in woodland and forest and less in heath. Isodar analysis also revealed that although forest was quantitatively better than heath and equivalent to woodland it was qualitatively poorer than either habitat. Heath had a higher density of understorey than woodland and woodland having a higher density of understorey than forest giving crossover population regulation. When the increasing and declining phase of the irruption were analysed separately, no habitat was quantitatively superior to any other during either phase. The lack of switching in preference between habitats from the increasing to the declining phase of the irruption and the virtual absence of any dispersal by adults, does not support the reciprocating dispersal hypothesis.     

Occurrence and evolutionary origins of polyploids in the clubmoss genus Diphasiastrum (Lycopodiaceae)

Two polyploid taxa are commonly recognized in the genus Diphasiastrum, D. wightianum from Asia and D. zanclophyllum from South Africa and Madagascar. Here we present results from Feulgen DNA image densitometry analyses providing the first evidence for the polyploid origin of D. zanclophyllum. Also reported for the first time are data confirming that D. multispicatum and D. veitchii, representing the putative parent lineages of D. wightianum, are diploids. Phylogenetic analyses of the nuclear regions RPB2, LEAFY and LAMB4 reveal that putative tetraploid accessions are of allopolyploid origin. Diphasiastrum zanclophyllum shows close relationship to the North American taxon D. digitatum on the maternal side, but the paternal relationship is less clear. Two accessions from Asia, both inferred to be polyploid, have D. veitchii as the maternal parent, whereas the paternal paralogs show relationships to D. multispicatum and D. tristachyum, respectively. None of these parental combinations have previously been hypothesized.     

Changes of dung beetle communities from rainforests towards agroforestry systems and annual cultures in Sulawesi (Indonesia)

Little is known about how tropical land-use systems contribute to the conservation of functionally important insect groups, including dung beetles. In a study at the margin of Lore Lindu National Park (a biodiversity hotspot in Central Sulawesi, Indonesia) dung-beetle communities were sampled in natural forest, young secondary forest, agroforestry systems (cacao plantations with shade trees) and annual cultures (maize fields), each with four replicates (n = 16 sites). At each site we used 10 pitfall traps, baited with cattle dung, along a 100 m transect for six 3-day periods. The number of trapped specimens and species richness at the natural forest sites was higher than in all land-use systems, which did not significantly differ. Each land-use system contained, on average, 75% of the species richness of the natural forest, thereby indicating their importance for conservation. However, a two-dimensional scaling plot based on NESS indices (m = 6) indicated distinct dung beetle communities for both forest types, while agroforestry systems and annual cultures exhibited a pronounced overlap. Mean body size of dung beetles was not significantly influenced by land-use intensity. Five of the six most abundant dung beetle species were recorded in all habitats, whereas the abundance of five other species was significantly related to habitat type. Mean local abundance and number of occupied sites were closely correlated, further indicating little habitat specialisation. The low dung beetle diversity (total of 18 recorded species) may be due to the absence of larger mammals in Sulawesi during historical times, even though Sulawesi is the largest island of Wallacea. In conclusion, the dung beetle fauna of the lower montane forest zone in Central Sulawesi appears to be relatively robust to man-made habitat changes and the majority of species did not exhibit strong habitat preferences.     

Limiting factors for seedling emergence and establishment of the invasive non-native Rosa rugosa in a coastal dune system

The relative importance of seed, habitat and microhabitat limitation has rarely been investigated for invasive non-native species, although this is critical for their effective management and for predicting future range expansion. Rosa rugosa is an abundant non-native shrub in coastal habitats of NW Europe; it is common along the Baltic coast but more scarce in exposed dunes of the North Sea. To investigate whether invasion of exposed dunes is limited by seed, habitat and microhabitat limitation, seedling emergence and establishment were examined in a factorial sowing, transplant and disturbance experiment. Twenty plots were randomly placed in each of five dune habitats (white dune, Empetrum dune, grey dune, outer dune heath, inner dune heath), and studied over 2 years. Seedling emergence in control subplots was zero in all habitats, whereas 96% and 98% of the undisturbed and disturbed seeded subplots produced seedlings. Disturbance had a positive effect on emergence and subsequent survival in white dune, outer and inner dune heath. Seedling survival and growth, and growth of transplanted seedlings, were markedly lower in grey dune. These findings indicate that establishment of R. rugosa is seed-limited in coastal dune habitats, and that the species is able to establish in both active and fixed dunes once seeds have arrived. Although differential seedling emergence and growth indicate that habitats differ in their degree of invasion susceptibility by R. rugosa, the positive influence of small-scale disturbance suggests microhabitat limitation in some dune habitats as well. Dune management should aim to reduce seed production and dispersal of R. rugosa in near-natural sites, and anthropogenic changes of habitat dynamics should be prevented.     

Ecological genetics of Bromus tectorum

Summary By incorporating demographic analyses of fitness components (e.g., survival and reproduction) within a reciprocal sowing design, we tested for 3 consecutive years whether local adaptation has occurred in the alien grass Bromus tectorum (cheatgrass) within 7 habitats along an environmental gradient from arid steppe to subalpine forest in the Intermontain Region of western North America. Patterns of emergence and survival were strongly influenced by the local environment. In terms of survival, expression of significant local adaptation in Tsuga heterophylla habitat varied among years. In contrast, relative differences in flowering time among seed sources were stable across sites and years. Populations from the arid steppe were the earliest to flower; flowering was latest in populations from the mesic Tsuga heterophylla habitat. In terms of net reproductive rate, evidence for local adaptation in B. tectorum was obtained in populations from habitats representing environmental extremes: an arid, saline site dominated by the shrub Sarcobatus vermiculatus and clearings within the cool, mesic Tsuga heterophylla forest habitat. Unlike the plants introduced from other sites, members of the resident population at the Sarcobatus site flowered and produced seeds before soil water became limiting. In contrast, net reproductive rates in other habitats were sometimes the lowest for populations in their home site. This lack of an advantage for local populations within more environmentally moderate sites suggests that limited dispersal may restrict the rate at which superior genotypes are introduced into a particular site.     

Geranium kikianum sp. nov. (Geraniaceae) from the southern Peloponnese,Greece

Geranium kikianum Kit Tan & G. Vold sp. nov. (Geraniaceae) is illustrated and described as a new species endemic to Greece. It occurs beside streams and other wet places in open Pinus nigra forest at the foothills of Mt Taigetos in the southern Peloponnese. It is closely related to G. macrorrhizum, a species more widely distributed in southern and central Europe, ranging from the southern Alps to the Balkan Peninsula. It differs from the latter by its smaller and narrower deflexed petal limb, slender claw and occurrence in wet habitats. The genome sizes of G. kikianum and of Greek populations of G. macrorrhizum are presented here for the first time. The nuclear DNA content (2C values) of 2.84 pg for G. kikianum and 2.83 and 2.87 pg for two G. macrorrhizum populations from Greece probably corresponds to a 2n=92 cytotype while the smaller genome size of 1.45 pg in a population of G. macrorrhizum from Mt Smolikas, Greece may correspond to a 2n=46 cytotype.     

Forest snail faunas from Crimea (Ukraine), an isolated and incomplete Pleistocene refugium

The land snail faunas of 26 forest sites and two open rocky sites in the Crimean Mountains were sampled in 2011. Of the 40 species found within the forests (about half the known fauna of Crimea as a whole), 28 were species with wide western Palaearctic distributions, and only eight were endemic to Crimea. While there were significant differences in the faunas of different sampling areas, these seemed to be a consequence of ecological differences among them rather than a product of geographical isolation and differentiation. Endemic species were large, and not entirely restricted to forest; known endemics not found in these forests are mainly typical of more open habitats. There is no local radiation of small species living in damp forest litter, as with Leiostyla species in the Transcaucasian forest refugium, and families such as the Clausiliidae with many endemic forest species in both Transcaucasia and the Carpathians are sparsely represented. The one endemic clausiliid genus, Mentissa, occurs in open as well as in wooded habitats. The present faunas are rather poor considering the soil conditions and climate, and the forests hold widespread species often associated with open habitats elsewhere. While there is evidence that these mountains provided a refuge for many animals and plants during glacial episodes further north, the forest snail fauna suggests that full forest cover did not survive throughout the Pleistocene. Rather, the present fauna contains endemics that survived in other habitats and widespread species with good powers of passive dispersal. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 424–433.     

山地隔离和空间距离对高黎贡山灰腹地莺(Tesia cyaniventer)鸣唱特征的影响

鸟类鸣唱存在广泛的地理变异,研究鸟类鸣唱变异的模式及其影响因素可帮助解释自然界中广泛而复杂的鸣声变异现象。灰腹地莺（Tesia cyaniventer）是在高黎贡山海拔2000-2800 m分布的小型地栖性森林鸟类。高黎贡山南北走向的山脊海拔通常在3000m以上,这导致灰腹地莺东、西坡种群被山脊所隔离。该种小鸟鸣声洪亮易于鉴别,其鸣声地理变异可揭示山地对鸟类种群产生的隔离效应。在高黎贡山片马垭口和独龙江垭口的东西坡4个地点录制了灰腹地莺的鸣声（n=58）,基于声谱分析比较了种群间鸣唱的质量特征,发现种群间鸣唱型的共享程度极低,而音节型在4个种群间均有共享。进一步测量了11个鸣唱的数量特征参数,有6个参数在不同种群间有显著差异：最低频率、中心频率、频率宽度、起始音节频率、首二音节的时间间隔、句子平均音节数。种群间的两两比较表明,鸣声特征差异呈现"隔离-距离"共同作用的格局,但隔离的影响更大。研究表明山地系统中影响鸟类鸣声地理变异的因素较为复杂,山地隔离和空间距离均对灰腹地莺的鸣唱特征产生了影响。     

Habitat use and seed removal by invasive rats (Rattus rattus) in disturbed and undisturbed rain forest,Puerto Rico

Despite frequent occurrences of invasive rats (Rattus spp.) on islands, their known effects on forests are limited. Where invasive rats have been studied, they generally have significant negative impacts on native plants, birds, and other animals. This study aimed to determine invasive rat distribution and effects on native plant populations via short‐term seed removal trials in tropical rain forest habitats in the Luquillo Experimental Forest, Puerto Rico. To address the first objective, we used tracking tunnels (inked and baited cards inside tunnels enabling animal visitors’ footprints to be identified) placed on the ground and in the lower canopy within disturbed (treefall gaps, hurricane plots, stream edges) and undisturbed (continuous forest) habitats. We found that rats are present in all habitats tested. Secondly, we compared seed removal of four native tree species (Guarea guidonia, Buchenavia capitata, Tetragastris balsamifera, and Prestoea acuminata) between vertebrate‐excluded and free‐access treatments in the same disturbed and undisturbed habitats. Trail cameras were used to identify animals responsible for seed contact and removal. Black rats (Rattus rattus) were responsible for 65.1% of the interactions with seeds, of which 28.6% were confirmed seed removals. Two plant species had significantly more seeds removed in disturbed (gaps) than undisturbed forest. Prestoea acuminata had the lowest seed removal (9% in 10 days), whereas all other species had >30% removal. Black rats are likely influencing fates of seeds on the forest floor, and possibly forest community composition, through dispersal or predation. Further understanding of rat–plant interactions may be useful for formulating conservation strategies.     

The distribution of heath balds in the Great Smoky Mountains,North Carolina and Tennessee

Abstract. We used remote sensing and a geographic information system to model the distribution of evergreen shrub communities, called ‘heath balds’, in the Great Smoky Mountains, North Carolina and Tennessee, USA. The 421 heath balds averaged 1.8 ha in size and covered 0.3% of the landscape. They reached their greatest importance on upper slopes (92% had relative slope positions > 80), convex topography (82% occurred on sites with a curvature greater than 2.6), and elevations between 1100 and 1600 m (94% of the balds). Although heath balds were found in old‐growth watersheds, the two watersheds with the greatest number of balds burned extensively after logging in the early 1900s. Bald occurrence was positively correlated with burned sites, old growth condition, and a highly acidic rock type. Heath balds showed a striking geographic pattern, with 88.1% of the area of this community found in six watersheds comprising only 35.4% of the study area. Despite similar topography, geology, and history, the eleven other watersheds had only 11.9% of the bald area while comprising 64.4% of the study area. Multivariate models showed that this community occurs on only 0.4 to 9.0% of the seemingly appropriate sites. Once established, this shrub community, with its dense evergreen canopy and thick leaf litter, is resistant to tree invasion. Both forest and shrub communities are stable on sites that are seemingly ideal for heath bald occurrence.