Floristic composition, structure, and dynamics of an upper montane rain forest in Southern Ecuador

Two 1 ha plots of undisturbed upper montane rain forest in southern Ecuador were sampled for all trees with a dbh ≥ 5 cm. An extraordinarily high α-diversity for a forest near treeline is described. The "non-ridge forest" plot at 2900 m elevation has 75 species and 28 families ha^-1 and is believed to represent an advanced stage of succession whereas, the "ridge forest" plot at 2700 m elevation with 90 species and 38 families ha^-1 has a more rapid turn-over rate probably due to a more unstable environment. Downslope forces (soil creep) are discussed as a possible cause for 32–44% of all trees being inclined more than 30° in the moderately steep terrain (average slope 20° in both study plots). The plot of "non-ridge forest" is characterized by a much greater biomass whether expressed as basal area (44 m² versus 15 m²) or stem volume (214 m³ versus 52 m³), while the density is equally high (2310 versus 2090 trees ha^-1) in both plots. Families with Family Importance Value >25 / ha^-1 are Clusiaceae, Cunoniaceae, Melastomataceae, Myrsinaceae, and Ternstroemiaceae.     

Species composition and vegetation structure of an upper montane forest at the summit of Mt. Doi Inthanon, Thailand

Upper montane forest (UMF) within Doi Inthanon National Park, Northern Thailand, was investigated by means of fifty, 40 × 40 m stratified random plots situated between 2080 and 2565 m altitude. The aim was to address a number of community ecological questions concerning woody species composition and structural heterogeneity of the forest. A total of 7474 individuals of trees and woody climbers ≥ 15 cm gbh (girth at breast height) were included in the study and these were identified to 47 species, 39 genera and 26 families. The average density was 934 individuals/ha and the average stem basal area was 71.8 m²/ha. The most important species were: Quercus eumorpha, Sjzygium angkae, Litsea martabanica, Helicia nilagirica, Lindera caudata, Schima wallichii, Osmanthus fiagrans, Eurya acuminata, Myrsine semiserrata and Ilex umbellulata and the most important families were Fagaceae, Lauraceae, Theaceae and Myrtaceae. Altitude was the most important environmental variable explaining species composition and vegetation structure. Most of the calculated vegetation variables showed significant correlation with altitude: species richness, family richness, diversity, density and crown cover declined with altitude, average tree height was uncorrelated with altitude and basal area increased with altitude. An analysis of size class distributions indicated good forest conditions and reverse-J-shaped age class distribution of most species.     

Tree population dynamics in Kibale National Park, Uganda 1975–1998

Changes in species composition, stem abundance, and basal area of trees taller than or equal to 10 m in a medium altitude tropical rain forest at the Ngogo study area, Kibale National Park, Uganda are described for the period between 1975 and 1998 ( n  = 23 years). Two enumeration episodes were conducted in 263 plots of 5 m by 50 m during 1975–80 and 1997–98. During this period, species richness decreased by 3% (from 92 to 89). Species diversity (H') also declined slightly from 2.97 to 2.86. The number of trees recorded in the sample plots decreased by 8% (from 2545 to 2329), while basal area decreased from 49.48 m² ha⁻¹ to 48.68 m² ha^−1. However, stem abundance and basal area increased for some species.     

The forest of Vorsø, Denmark: Succession towards a natural, decidous boreal forest influenced by breeding cormorants

The forest on the strictly protected island Vorsø in Horsens Fjord has since 1952 been examined every ten years to discover the number, basal area, wood volume and aboveground biomass of the species. This paper describes the results from 1982 and the development up to that. The dominating species are Fraxinus excelsior, Ulmus glabra and Fagus sylvatica. The two forests, "Vesterskov" (6.09 ha) and "Østerskov" (2.48 ha) respectively, were, in 1982, still immature. Their number of stems per ha were: 756 and 509; basal area: 38 m² and 45 m². volume: 544 m³ and 615 m³; biomass 207 t and 268 t. The annual increase 1972–1982 was respectively 8.8 m³/ha and 0.3 m³/ha. The impact of the cormorant ( Phalacrocorax carbo sinensis ) is especially in "Østerskov" increasing, since the colony here was established in 1976. The succession of the forest is discussed in relation to the disturbance by the cormorants.     

Soil seed bank dynamics of fire-prone wooded grassland, woodland and dry forest ecosystems in Ethiopia

In order to reveal the role of soil seed banks in vegetation recovery after fire in savanna, the spatial distribution and temporal changes in the soil seed banks of regularly burning savanna in Gambella, western Ethiopia, was studied. The seedling emergence technique was employed to determine the species composition and density of the soil seed bank of six sites ranging in fire severity from wooded grassland with frequent fires over woodland with intermediate fire frequency to forest with absence of fires. Species composition and density of seeds in the soil were compared between seasons, depths and sites with different types of standing vegetation. Fourteen plant species were recorded in the soil seed bank from the grassland and woodland sites and 6 from the dry forests; 60 % of the taxa in the soil seed bank were annuals and 40 % were perennials. The soil seed banks were largely dominated by graminoids and 48–97 % of the soil seed bank in the grasslands and woodlands was of a single grass species, Hyparrhenia confinis , which was absent from the dry forests. The soil seed pools ranged from less than 100 to 4700 seeds per m² depending upon the season. The soil seed bank of graminoids was nearly empty after the onset of the rainy season whereas seeds of broadleaved herbs and woody species able to germinate were still found after this time. Floristic composition, representation of life forms and density of seeds in soil did not correspond closely with that of the standing vegetation, but within graminoids there was a strong similarity between the soil seed bank and the standing vegetation. The current fire regime of Ethiopian savanna woodlands appears to maintain the dominance of graminoids over broadleaved herbs and woody plants both as seeds in the soil and in the standing vegetation.     

The important role of scattered trees on the herbaceous diversity of a grazed Mediterranean dehesa

Scattered trees are considered keystone structures and play an important role in Mediterranean sylvopastoral systems. Such systems are associated with high biodiversity and provide important natural resources and ecosystem services. In this study, we measured the contribution of scattered trees and different grazing management (cattle, sheep and wildlife only) to the diversity of the grassland sward in a dehesa (open holm oak woodland) located in Central Spain. We analyzed alpha and beta diversity through measurement of species richness, Shannon-Wiener, and Whittaker indices, respectively; and the floristic composition of the herb layer using subplots within two adjacent plots (trees present vs. trees absent) under three different grazing management regimes, including wildlife only, during a year. We found a 20–30% increment in the alpha diversity of wooded plots, compared to those without trees, regardless of grazing management. All beta indices calculated showed more than 60% species turnover. Wooded plots were occupied by different herbaceous species in different heterogeneous microsites (under the canopy, in the ecotone or on open land) created by the trees. Livestock grazing modified species composition (e.g. more nitrophilous species) compared to wildlife only plots. In addition to all their other benefits, trees are important to maintaining grassland diversity in Mediterranean dehesas.     

The influence of past land-use on understory plant distributions in a near-natural deciduous forest in Denmark

In regions with a long cultural history, past land-use may strongly determine floristic composition in seemingly natural vegetation. Here we examine if past land-use affects understory plant distribution in near-natural forest in a historically and environmentally heterogeneous setting, namely the 17 ha freshwater island of Borgnr (Denmark). In 107 randomly located 100 m² plots we recorded species composition and various environmental variables. The plots were classified into three groups according to past land-use: (1) Disturbed sites close to 12^th and 20^th century ruins and associated open areas; (2) formerly grazed meadow areas; and (3) forested sites without indications of strong anthropogenic influence. Past land-use, environmental variables and species traits were used to interpret floristic gradients. Unconstrained and constrained ordinations clearly separated plots with different past land-use, and showed that the main floristic gradients on Borgø correlated with hydrological-edaphic conditions and past land-use. Partial constrained ordination using the environmental parameters as covariables showed that past land-use had a unique influence on floristic composition.     

Effects of experimental small-scale grassland fragmentation on spatial distribution, density, and persistence of ant nests

Abstract.  1. Grassland fragmentation is expected to influence the abundance of different invertebrate species to a different extent. Fragmentation-related effects are of particular importance in species that interact with many other species.
2. The density and spatial distribution of nests of 15 ant species in experimentally fragmented calcareous grasslands at three sites in the Northern Swiss Jura mountains were examined. Fragments of different size (0.25 m², 2.25 m², and 20.25 m²) were isolated by a 5-m wide strip of frequently mown vegetation. Control plots of corresponding size were situated in adjacent undisturbed grassland.
3. Three years after initiation of the experiment, ant nest density did not differ between fragments and control plots. Six years after initiation of the experiment, however, ant nest density and forager abundance were higher in large fragments than in large control plots. Ant nests tended to occur more frequently along the edge of fragments than in the core area. Persistence time of nests of the most abundant species, Lasius paralienus , tended to be shorter in fragments than in control plots. Furthermore, persistence time was longer in nests situated close to the fragment edge than in nests in the core area.
4. Effects on nest density, edge effects on the spatial distribution of nests, and the relationships between nest density and environmental factors were more pronounced when only nests of L. paralienus were considered. The implications of these findings for plant and other invertebrate species are discussed.     

Bird community composition in an actively managed savanna reserve, importance of vegetation structure and vegetation composition

The mosaic of trees, shrubs and open grassland in mesic African savannas is highly dynamic and strongly influenced by mammal herbivory and fire. We investigated the bird fauna in four different savanna habitats to help assess the impacts of vegetation change on this component of faunal diversity. Birds were censused, plant species were identified and vegetation structure was measured in four different vegetation types (Acacia nilotica woodland, Acacia nigrescens woodland, broadleaf thicket and open grassland) in the Hluhluwe-Umfolozi Park in northern KwaZulu Natal, South Africa. Multivariate ordination analyses were used to determine the relative importance of vegetation structure and floristic composition in defining bird assemblages. The bird communities of the grasslands, the acacia woodlands, and the broadleaf woodlands were clearly separated on the first axis of the detrended canonical correspondence analysis (DCCA). Canopy cover and foliage height diversity (FHD) were strongly correlated with the first axis of DCCA, possibly reflecting a secondary successional series from grassland to woodland, known as bush encroachment. Floristic composition (based on presence–absence data only) seemed to be less important for bird community composition than vegetation structure. The results indicate that changes in vegetation structure, caused by bush encroachment, could cause concomitant changes in bird community composition.     

Fruit and seed ecology of wild robusta coffee (Coffea canephora Froehner) in Kibale National Park, Uganda

Coffee fruit production, frugivorous activities and seed longevity were investigated during two seasons. The average densities of fruiting coffee trees during the first sampling period ranged from four to seven per 25 m² and during the second sampling period, four fruiting trees per 25 m² were estimated. Fruiting coffee trees were significantly less than the non-fruiting trees ( P < 0.05, t -test). Periods of maximum coffee fruit fall coincided with those of lowest rainfall. Coffee fruit yield per tree from ground collections was 7–40 m⁻² in 1992 and 16–38 m⁻² in 1993/1994. Split coffee berries ranged between 71.7% and 83.6% of all fruits collected from the forest floor in the 1993/94 sampling period. Coffee seeds collected from the forest floor were mostly undamaged. Black and white colobus and redtail monkeys were observed to feed on ripe coffee fruits but did not crush the seeds. Coffee seed viability declined rapidly during storage. Seeds left on the forest floor survived longest, those stored under laboratory conditions lost viability fastest, and those in cold storage showed intermediate longevity.     

Composition and structure of a species rich Amazonian rain forest obtained by two different sample methods

Inventory data for trees ≥ 10 cm DBH from a hectare plot are compared to data obtained by the Point-Centered Quarter Method along a line transect from the same locality in Anangu, Amazonian Ecuador. The one-hectare quadrat plot of 100 × 100 m had 734 individuals, 153 species, 46 families, a total basal area of 22.2 m², and an estimated above ground tree volume of 240.5 m³. The line transect had a calculated density of 728 individuals per hectare, which included 239 species, 51 families, a total basal area of 34.1 m², and an estimated above ground tree volume of 409.6 m³. Of the 20 species with the highest IVI, only four were shared by the two samples. The most important species were Quararibea ochrocalyx on the hectare plot and Iriartea deltoidea on the line transect, constituting 26.6 and 13.3% of the individuals, respectively. The five families with the highest FIV on the hectare plot (Bombacaceae, Arecaceae, Moraceae, Caesalpinaceae, and Lauraceae) and on the line transect (Arecaceae, Moraceae, Meliaceae, Mimosaceae and Caesalpinaceae) constitute 40.4% and 35.4% of the Family Importance Values of the samples, respectively. The Point-Centered Quarter Method used along a line transect reflects maximum diversity and provides average values of density and tree size in the area. The quadrat plot reflects the local structure and composition of the forest within the plot.     

Restoration of wooded meadows ‐ a comparative analysis along a chronosequence on Öland (Sweden)

Abstract. Wooded meadows on the Baltic Island of Öland result from traditional agricultural management over centuries which has led to a species‐rich vegetation with high species diversity. Today, nearly all of these meadows have been abandoned and became rapidly overgrown by deciduous shrub and tree species forming a closed canopy which resulted in a rapid and strong decrease in species numbers of the herb layer. Recent efforts aim to restore overgrown wooded meadows by cutting single shrubs and trees to open the canopy. However, the effects of abandonment as well as of any restoration management in wooded meadows have rarely been documented until now. Mechanisms driving succession after restoration such as the dispersal potential of the respective species over time and space have not been analysed yet. Therefore, a chronosequence was studied which included a traditionally managed wooded meadow, an overgrown meadow which has been abandoned for more than 100 yr and a meadow which was restored 36 yr ago by cutting and is now grazed. We analysed the soil seed bank of the 3 meadows in comparison with the established vegetation and endozoochorous seed dispersal by cattle and sheep. After abandonment 87% of the typical grassland species vanished from the established vegetation and were replaced by species characteristic of woodland and disturbed grassland communities. The mean number of species decreased from 52 species per plot (4 m²) to 18 species. Mean species number and number of seeds in the seed bank declined significantly from the traditionally managed to the overgrown meadow. Most of the grassland species were assigned to a transient seed bank type while only 1/3 could be classified as having a short‐term persistent seed bank. Thus, restoration of wooded meadows cannot rely on the soil seed bank. Endozoochorous seed dispersal by cattle and sheep was shown for 15% of the species with seed densities per 100 g air dried dung of 737 and 767, respectively. Movement of animals between ancient and restored wooded meadows is recommended since many of the species only occurred in low densities and therefore, will probably not be found in the dung samples.     

Diel abundance, migration and feeding of fish larvae (Eleotridae) in a floodplain billabong

Eleotrid larvae (2.1–16 mm) were collected from surface waters of a billabong in south-eastern Australia. Estimates of larval density in plankton net samples at night averaged 148.3 larvae per m³ and 16.6 larvae per m³ during the day. In contrast, pump samples provided density estimates of 8.3 larvae per m³ at night and 0.9 larvae per m³ during the day. Larval densities did not differ between open water, snag (fallen tree) and Typha habitats, but Typha habitats yielded larger larvae than other habitats. 32.9% of larvae in pump samples were damaged and unmeasurable, creating a bias favouring larger larvae. The modal length of larvae in net samples at night was 5–6 mm, compared with 3–4 mm during the day, reflecting both greater net avoidance by larger larvae during the daytime and dispersal of smaller larvae from the surface at night. Dispersion patterns of larvae suggest that classes of larvae smaller than, and larger than 5.0 mm exhibit reciprocal diel vertical migration behaviour linked to ontogenetic changes in diet. Larvae less than 5 mm fed only during the day and preyed exclusively on rotifers, whereas larger larvae continued to feed at night and consumed mostly planktonic crustaceans.     

A MIDDLE JURASSIC FOSSIL FOREST FROM NEW ZEALAND

Abstract:  A Middle Jurassic fossil forest, thought to have been growing at high southern palaeolatitudes on the Gondwana margin, is described from New Zealand. Fossil stump horizons are exposed in stratigraphic section within the Urawitiki Measures Formation, Murihiku Supergroup. Tree bases were preserved by silica permineralisation in a sandy braided river setting. Fossil wood is referred to 'abietinean-type' and Agathoxylon . A total of 238 stumps were mapped with measured diameters allowing tree height estimates of 9–40 m. A two-dimensional area (102 m²) allows an interpretation of tree density (2353 trees/ha), biomass (579 t/ha) and stand basal area (146 m²). This horizon is interpreted as a highly productive, young, dense tree stand probably actively regenerating on a flood-cleared surface. Annual growth rings suggest a clearly defined growing season with a rapid onset of favourable temperature, light and water supply conditions at the start and a similarly rapid deterioration at the end. Mean ring width is 1·10 mm and the maximum 9·67 mm. Mean Sensitivity values suggest both 'complacent' and 'sensitive' growth patterns. Broad palaeoclimate interpretations suggest a humid, warm regional climate throughout the growing season.     

Relationship between seed and clonal growth in the reproduction of Carex rugulosa Kük. in riverside meadows

Carex rugulosa Kük. develops riverside vegetation in brackish marshes along the lower parts of rivers. Because the rhizomatous ramets grow densely (590–950/m²) in its developed population, it is expected that seedlings cannot get enough light to grow if seeds germinate in crowded ramets. We studied the reproductive contribution of seed in a marshy meadow of the Ohashi River, Japan. The number of seeds produced was 6900–14 200/m² in 2002. The number of buried seeds in the following germination season differed among sites from 1190 to 2690/m². No seedlings were observed at plots where the ramets from rhizomes grew densely. In contrast, seedlings appeared in plots where all rhizomatous ramets were artificially cut. The number of seedlings corresponded to 17.5–39.5% of that of buried seeds. All these seedlings were submerged and died when it rained heavily. In the genotype analysis using allozyme detection, one of the phosphoglucoisomerase loci ( Pgi-2 ) indicated a pair of alleles. Among eight plots (each plot was 50 or 60-cm square), four showed an excess of heterozygotes and two showed homozygote domination in the Pgi-2 genotype. These results suggest that: (i) seed germination is suppressed in crowded ramets; (ii) seedling survival is severely reduced by inundation; and (iii) the population is usually maintained by clonal growth. As germination is induced in open areas, it is hypothesized that the main role of seed is recovery after vegetation decline caused by environmental stresses or colonization to other areas without dense vegetation.     

Dissecting components of population-level variation in seed production and the evolution of masting behavior

Theory predicts that at higher trophic levels species are especially vulnerable to habitat fragmentation due to small population size and dependence on particular prey species. Using aphids as model organism, we tested the hypothesis that herbivore abundance increases in fragmented habitats as a result of reduced predator and parasitoid densities. In a 3 year-study, we examined the population dynamics of aphids with respect to host plant abundance and ant nest density in experimentally fragmented calcareous grasslands at two sites in the northern Jura mountains. Fragments of different size (area: 20.25 m², 2.25 m² and 0.25 m²) were isolated by a 5 m wide strip of frequently mown vegetation and corresponding control plots were situated in the adjacent undisturbed grassland. Aphid density was higher in fragments than in control plots. This was a combined result of a higher frequency of aphid-infested plants and larger aphid colonies in fragments than in control plots. Furthermore, a larger proportion of aphid colonies was ant-attended in fragments than in control plots. Aphid colonies were also more frequently visited by ants in fragments than in control plots in one of the 3 years. Parasitoid pressure on aphids was not influenced by the experimental fragmentation. Neither were aphid species richness and diversity affected by the fragmentation. Our study shows that even small-scale habitat fragmentation can have profound effects on the abundance of herbivorous insects. The effect on aphid density was consistent over 3 years and two sites with slightly different aphid communities.     

Control of Karoo paralysis ticks through vegetation management

Abstract. Karoo paralysis, caused by feeding Ixodes rubicundus females, is a major disease of small stock in South Africa. Control methods currently practised are almost exclusively chemical based. To limit overdependance on chemicals, vegetation management was investigated as a possible method for control, to be incorporated in an integrated tick management system. Laboratory and field experiments were conducted to determine, firstly, the extent of vertical migration and survival of ticks on long and short copper rods which simulated grasses as questing substrates; secondly, the infestation burdens of sheep exposed to similar tick challenges in pens with long and short grass; and thirdly, the effect of trimming the lower crown line of wild olive trees, simulating the browsing effect of goats, on tick density in the immediate environment of the trees. When ticks were exposed to optimal (>45 cm) and sub-optimal (<10 cm) length rods on which to quest, the extent of vertical migration over extended periods of time (up to 87 days) was significantly higher ( P < 0.001) for the ticks exposed to long rods. Also, almost 3 times as many ticks exposed to long rods survived compared to those exposed to short rods. Sheep exposed to long grass were infested by twice as many ticks compared to those exposed to short grass. Tick density at modified wild olive trees ( Olea europaea africana ) (0.027 ticks/m²) differed significantly ( P < 0.05) from that at control trees (0.088 ticks/m²). It is recommended that coarse grazers such as cattle and horses should be used to graze down long grasses before sheep are introduced into camps known to be infested with I. rubicundus. Similarly, domestic goats can, through their browsing effect on shrubs and trees, modify the vegetation and as such play an important role in an integrated tick management system.     

Spatial Distribution of Aboveground Carbon Stock of the Arboreal Vegetation in Brazilian Biomes of Savanna,Atlantic Forest and Semi-Arid Woodland

The objective of this study was to map the spatial distribution of aboveground carbon stock (using Regression-kriging) of arboreal plants in the Atlantic Forest, Semi-arid woodland, and Savanna Biomes in Minas Gerais State, southeastern Brazil. The database used in this study was obtained from 163 forest fragments, totaling 4,146 plots of 1,000 m² distributed in these Biomes. A geographical model for carbon stock estimation was parameterized as a function of Biome, latitude and altitude. This model was applied over the samples and the residuals generated were mapped based on geostatistical procedures, selecting the exponential semivariogram theoretical model for conducting ordinary Kriging. The aboveground carbon stock was found to have a greater concentration in the north of the State, where the largest contingent of native vegetation is located, mainly the Savanna Biome, with Wooded Savanna and Shrub Savanna phytophysiognomes. The largest weighted averages of carbon stock per hectare were found in the south-center region (48.6 Mg/ha) and in the southern part of the eastern region (48.4 Mg/ha) of Minas Gerais State, due to the greatest predominance of Atlantic Forest Biome forest fragments. The smallest weighted averages per hectare were found in the central (21.2 Mg/ha), northern (20.4 Mg/ha), and northwestern (20.7 Mg/ha) regions of Minas Gerais State, where Savanna Biome fragments are predominant, in the phytophysiognomes Wooded Savanna and Shrub Savanna.     

Long‐term changes in the vegetation after the cessation of livestock grazing in Eucalyptus marginata (jarrah) woodland remnants

This paper documents changes in the floristic composition of Eucalyptus marginata Donn (jarrah) woodlands over 7 years of recovery from continual, intensive livestock grazing. In remnants of native woodland left after agricultural clearing, which have been subjected to livestock grazing, comparisons were made between the floristics of fenced exclosure plots and open plots that continued to be grazed. The vegetation in nearby remnants, which had not been subjected to livestock grazing, was also surveyed. An initial increase in annual exotic pasture species after grazing relief was only temporary and highly influenced by fluctuations in annual climatic patterns, particularly rainfall distribution and abundance. Subsequent years saw a decrease in exotic annuals in exclosure plots and an increase in native perennials, in a trend towards becoming more floristically similar to the ungrazed sites. Germination of overstorey species was observed in the exclosure plots, however, development of seedlings and saplings was sparse. Results indicate that for jarrah woodland in southwestern Australia, natural regeneration is possible after the removal of livestock, with the return (within 6 years) of native species richness to levels similar to those found in ungrazed vegetation. Re‐establishment of cover, however, appears to take longer. The floristic dynamics are described in terms of a nonequilibrium model. Two vegetation states exist, degraded remnants with an understorey dominated by annual species, and ungrazed vegetation with an understorey dominated by perennial shrubs and herbs. The former state is maintained by continual heavy grazing by livestock. Upon relief from grazing, the vegetation undergoes a transition towards floristic similarity to ungrazed vegetation. After 6 years, vegetation change in the exclosure plots appears to be continuing and therefore it is still in transition.     

Influence of habitat characteristics on small mammals in a Mexican high-altitude grassland

Small mammals in a high-altitude grassland area close to Mexico City were studied. Populations of 10 species were censused using live traps in 48 sample quadrats. Within each quadrat, vegetation characterization, including complete floristic listings, cover values for species and layers and values of habitat modification, were assessed. Habitats were described according to plant communities identified using ordination and classification methods. Nine different plant communities were obtained. Densities and abundance of all small mammal species were calculated for each of the habitats classified. Peromyscus alsloni was the most abundant species in all habitats, reaching maximum densities of 55 ha⁻¹ in pine forest with dense ground and herb layer. Peromyscus melanotis also occurred in all habitats but at lower densities (maximum 29 ha⁻¹). Reithrodontomys megalotis was found in all habitats except in tall dense grassland. Densities for this species were generally low (1-9 ha⁻¹) but reached 19 ha⁻¹ in short dense grassland. All other species were largely absent from 4–8 habitats and showed very low densities (0.75–4 ha⁻¹). The densities of the more abundant species were largely correlated with more open habitats and higher indices of habitat modification. Lower altitude grassland habitats have a greater abundance of small mammals and a higher species richness than the medium and higher altitude, physiognomically more complex habitats. Species richness was highest in tall pine-alder forest with a species-rich, dense herb layer and lowest in pine forest with dense ground and herb layers. Species richness was positively correlated with overall small mammal density.