Evidence for altitudinal migration of forest birds between montane Eastern Arc and lowland forests in East Africa

Burgess, N.D. & Mlingwa, C.O.F. 2000. Evidence for altitudinal migration of forest birds between montane Eastern Arc and lowland forests in East Africa. Ostrich 71 (1 & 2): 184–190.

In this paper we assess the evidence for altitudinal movements of forest birds from the montane forests of the Eastern Arc mountains of East Africa to nearby lowland forest patches. For 34 montane species, including all the Eastern Arc endemics except Banded Green Sunbird Anthreptes rubritorques there is no evidence that they undertake seasonal movements to lower altitudes. An additional 26 montane species, of somewhat wider distribution, have been recorded at low (<500 m) altitudes during the cold/dry season (June to September). Most records of these montane birds at lower altitudes are from sites adjoining montane forest areas, although a few records are from lowland coastal forests at 100–240 km distance from montane areas. Only five of the 26 species (White-chested Alethe Alethe fulleborni, White-starred Forest Robin Pogonocichla stellata, Orange Ground Thrush Zoothera gurneyi, Evergreen Forest Warbler Bradypterus mariae and Barred Long-tailed Cuckoo Cercococcyx montanus) are regularly and commonly reported in the lowlands. They are also found in the lowlands in small numbers during the warm/wet season (October to February), when they may breed. The abundance of at least four, and probably more, of the forest birds with a more widespread distribution in the lowland and montane forests of East Africa declines greatly at high altitudes from the onset of the cold/wet season (February) and only increases again at the start of the warm/wet season (September). It is not known how far these species move as they cannot be easily separated from resident populations in lowland forests, and there are no ringing recoveries in different forests. Altitudinal migration of a proportion of the Eastern Arc avifauna is the most likely explanation for available data, although source-sink metapopulation theories may be helpful to explain the distributions of some species. As the movement of forest birds from the Eastern Arc to the lowland forests does not involve the rare endemics, they are of lower conservation concern, but the presence of montane and lowland forest may be important for the long-term survival of some more widely distributed forest species.     

Birds and biogeography of Mount Mecula in Mozambique’s Niassa National Reserve

The montane forests of northern Mozambique’s isolated massifs are inhabited by numerous range-restricted and threatened bird species, but until recently were extremely little-known. We report on a first avifaunal survey of the isolated montane habitats of Mt Mecula (1 442 m), Niassa National Reserve, notable as the only currently protected montane area in northern Mozambique. Mount Mecula’s moist forest is small (approximately 136 ha in total) and patchy, and although known botanically to have some montane affinities, was found to support an avifauna more typical of riparian forests of medium to low altitude. The only montane forest species recorded was Lemon Dove Aplopelia larvata. Other montane elements included Vincent’s Bunting Emberiza (capensis) vincenti, one of six species recorded new to the Niassa National Reserve list. Overall, it appears that despite its intermediate location, Mt Mecula does not represent a biogeographical ‘stepping stone’ for montane forest bird species. This probably owes to its remoteness from the Eastern Arc Mountains of Tanzania to the north and the massifs of other parts of northern Mozambique, to the south and west.     

Structural comparison of tropical montane rain forests along latitudinal and altitudinal gradients in south and east Asia

Geographical patterns of altitudinal zonation, floristic composition, and structural features of tropical montane rain forests were examined along latitudinal gradients in south and east Asia. On equatorial mountains, the tropical montane rain forests occur above 1000 m. Toward middle latitudes, they come farther down and reach sea level at c. 35° N. Thus, the forests are equivalent to the subtropical rain forests of the latitudinal, horizontal zonation series. They exhibit gradual changes in floristic composition and structure along both altitudinal and latitudinal gradients. On equatorial mountains, they are divided into three types, i.e. tropical lower montane, upper montane, and subalpine forests. The three tree regeneration types, having emergent, sporadic and inverse-J type stem-diameter class frequency distributions, coexist in the lower montane forests, but the upper and subalpine forests display only the inverse-J type species with a few species of the sporadic type. Toward the northern latitudinal limit, the distinction between the three tropical montane forest zones in equatorial mountains becomes less clear. This can be explained by temperature conditions: on equatorial mountains, a temperature sum of 85° C months which controls the upper limit of the lower montane forests, and a coldest month mean temperature of-1° C which controls the evergreen broad-leaved trees, appear at c. 2500 and c. 4000 m respectively. The altitudinal range between 2500 m and 3800 m, which is the upper forest limit, is covered by upper montane and subalpine forests. On the other hand, at the latitudinal northern limit, the tropical upper montane and subalpine forests cannot exist because the above mentioned two temperature conditions occur at nearly the same point. Thus, at the northern latitudinal limit of the tropical montane forests, the three zones of equatorial mountains amalgamate into a single subtropical lowland forest community. This is due to the seasonal temperature climate in middle latitudes in, e.g., central Japan and central China.A part of this paper was presented as an oral presentation at the Vth International Congress of Ecology, Yokohama 23–30.8.1990.     

High diversity and regional endemism in land snails of eastern Tanzania

In February/March 1995 we collected land snails (including slugs) at 12 stations in eastern Tanzania. A total of 571 person-hours yielded 9174 snails assigned to 159 morpho-species. The richest two sites each (<4ha of uniform forest) had 50 species (Amboni Cave) and 48 species (near Amani, Usambaras), nearly as great as the most species-rich sites known in the world; sieving of litter and soil would probably yield more species. In lowland (coastal) forests, both diversity and endemism seemed to decrease from north to south. Most snail species were found within only one of four coastal or one montane geographic regions, indicating substantial regional endemism. Only one species (Achatina fulica) appeared in all five regions, and 84% of all other species were found in only one (61%) or two regions (23%). The predatory streptaxids comprised about half the species and a third of the individuals at the Usambara site, an extremely high ratio of carnivores. Small snails (< 5mm greatest adult shell dimension) – many of which are probably undescribed species – comprise a substantial proportion of Tanzanian molluscan diversity; more surveys are needed, especially because of human pressures on the few forest patches remaining.     

Altitudinal distribution and habitat preferences of forest birds on Halmahera and Buru, Indonesia: implications for conservation of Moluccan avifaunas

The BOU Indonesia Expedition, with BirdLife International, carried out surveys on the Moluccan Islands of Buru and Halmahera in 1994-96 as part of a process of protected area identification. Data on resident forest birds were collected during ten-minute sampling periods at points along altitudinal gradients, in primary and modified forests and in rainforests on different geology. While all species ever reported from Halmahera have been recorded down to near sea level, many of the forest birds of Buru, including a number of restricted-range and endemic species, are largely confined to montane forest, and only common above 800–900 m altitude. The existence of a distinct montane bird community on Buru, but not on Halmahera, is attributed to the relatively large, continuous area of forest at high altitude on this island. Forest on Halmahera at such altitudes is limited and fragmented. While most species survive in modified forests, some were significantly rarer in this habitat and may depend on adjacent areas of primary forest. Mangroves supported impoverished forest bird communities, typically of more adaptable species. On Halmahera, rainforest on ultrabasic rock appears to be impoverished when compared with rainforest on other rock formations, in particular supporting very low numbers of two species of threatened parrot. Implications for conservation concern lowland forests on Halmahera. Despite the surveys, a number of species of conservation concern that occur on Halmahera and Buru still remain largely unknown.     

The distribution of soil macrofauna on the west ridge of Gunung (Mount) Mulu,Sarawak

Summary The macrofauna of the soils on the west ridge of Gunung Mulu in 4th Division, Sarawak, were sampled during February and March 1978. The eleven sampling sites ranged from near the base of the mountain at 130 m a.s.1. to the summit at 2.376 m.Altitudinal changes from lowland rain forest (mixed dipterocarp forest) to lower montane and upper montane rain forests are concomitant with changes in soil from red yellow podzolics and regosols to peaty gley podzolics and organic peats.The abundance of the total macrofauna declined from 2,579 individuals m^-2 at 130 m to 145 m^-2 at 2,376 m. Declining population densities of ants and termites correlated significantly with increasing altitude but the effect on other groups was variable. Changes in total biomass were erratic and varied from 4.1–6.2 g m^-2 (alc. w.w.) in the dipterocarp forest soils to 5.8 g m^-2 in the lower montane, 9.3–20.2 g m^-2 in the upper montane (tall facies) and 1.9–9.5 g m^-2 in the upper montane (short and summit facies). Only the decline in the biomass of termites and ants correlated significantly with altitude. Other groups remained fairly constant, varied erratically or increased in the middle altitudes.The dipterocarp forest soil macrofaunal biomass was dominated by termites, beetles and earthworms (Megascolecidae and Moniligastridae), with ants the dominant predators. The lower montane forest was a transitional and ill-defined zone on the mountain and the soil macrofauna was also transitional to some extent. Termite biomass fell substantially and earthworms replaced them as the dominant detritivores, with beetles in a secondary role. Formicidae remained as the major predators. With the inception of peats in the upper montane forest (tall facies), the macrofauna was dominated by Coleoptera with earthworms, Diptera larvae and Blattodea in lesser roles. With increasing exposure in the upper montane forests (short and summit facies), several major groups disappeared altogether. The soils were dominated by Blattodea with Coleoptera and Megascolecidae of lesser importance. Chilopoda and Arachnida replaced Formicidae as the dominant predators.     

Environmental correlates of gastrointestinal parasitism in montane and lowland baboons in Natal,South Africa

We compared the gastrointestinal parasites of the baboon,Papio cynocephalus ursinus, living in montane (altitude, >1800 m) and coastal lowland (altitude, 100–200 m) habitats in Natal, South Africa, using fecal analysis. While the montane animals harbored a smaller number of species, helminth egg-output rates were higher in them than in the lowland animals. The decrease in parasite diversity with increasing altitude was expected, but the difference in helminth egg output was not. It may be due to a combination of food shortage, which characterizes the montane environment, especially at the end of winter, and the high proportion of soil-contaminated items in the diets of the montane animals.     

The effect of altitude,patch size and disturbance on species richness and density of lianas in montane forest patches

The species richness and density of lianas (woody vines) in tropical forests is determined by various abiotic and biotic factors. Factors such as altitude, forest patch size and the degree of forest disturbance are known to exert strong influences on liana species richness and density. We investigated how liana species richness and density were concurrently influenced by altitude (1700–2360 m), forest patch size, forest patch location (edge or interior) and disturbance intensity in the tropical montane evergreen forests, of the Nilgiri and Palni hills, Western Ghats, southern India. All woody lianas (≥1 cm dbh) were enumerated in plots of 30 × 30 m in small, medium and large forest patches, which were located along an altitudinal gradient ranging from 1700 to 2360 m. A total of 1980 individual lianas were recorded, belonging to 45 species, 32 genera and 21 families, from a total sampling area of 13.86 ha (across 154 plots). Liana species richness and density decreased significantly with increasing altitude and increased with increasing forest patch size. Within forest patches, the proportion of forest edge or interior habitat influenced liana distribution and succession especially when compared across the patch size categories. Liana species richness and density also varied along the altitudinal gradient when examined using eco-physiological guilds (i.e. shade tolerance, dispersal mode and climbing mechanism). The species richness and density of lianas within these ecological guilds responded negatively to increasing altitude and positively to increasing patch size and additionally displayed differing sensitivities to forest disturbance. Importantly, the degree of forest disturbance significantly altered the relationship between liana species richness and density to increasing altitude and patches size, and as such is likely the primary influence on liana response to montane forest succession. Our findings suggest that managing forest disturbance in the examined montane forests would assist in conserving local liana diversity across the examined altitudinal range.     

An altitudinal transect study of the vegetation on Mount Kinabalu,Borneo

A quantitative transect analysis of altitudinal sequences of forest canopy species from 600 to 3400 m asl on Mt. Kinabalu (4101 m), Borneo, resulted in four discrete altitudinal vegetation zones. These were made up of mutually exclusive species groups for lowland (<1200 m asl), lower montane (1200 to 2000–2350 m asl), upper montane (2000–2350 to 2800 m asl), and subalpine (2800 to the forest line, 3400 m asl) zones. Zonal soil types were correlated with the vegetation zones. In upslope sequence, these were: lowland Oxisols, montane Histosol/Spodosol complex, and subalpine Inceptisols. The highest contents of organic carbon, extractable phosphorus, and exchangeable magnesium and potassium were recorded in the lower and upper montane zones. The upper boundaries of the lowland, upper montane and subalpine zones coincided with thermal thresholds of latitudinal bioclimatic zones: 18°C TMIN (Köppen's tropical), WI 85 (Kira's warm temperate), and WI 45 (Kira's cool temperate), respectively. The upper limit of the lower montane zone was correlated with an abrupt increase of water surplus estimated from the annual rainfall minus annual potential evaporation. These climatic characteristics appear to define ecological altitudinal turnover points, so called critical altitudes, where groups of associated species are displaced by other groups.Abbreviations asl = above sea level - DBH = diameter at breast height - PHQ = Park headquarters - TMAX = Mean daily maximum air temperature - TMIN = Mean daily minimum air temperature - TWINSPAN = Two-way indicator species analysis - WI = Warmth index     

海南霸王岭不同森林类型附生兰科植物的多样性和分布

附生兰科植物是热带林附生植物的主要类群之一,对于维持热带林生态系统的物种多样性及生态功能具有重要的作用。以海南岛霸王岭国家级自然保护区内的6种热带原始林类型(热带季雨林、低地雨林、热带针叶林、山地雨林、山地常绿林及山顶矮林)中的附生兰科植物为研究对象,通过样带调查(每个森林类型设置12个10m×50m的样带,记录每个样带内胸径(DBH)≥5cm的树木及藤本上附生兰科植物的物种名称、株数及附生位置)分析了附生兰科植物的物种多样性、附生位置及其在不同森林类型中的分布规律。结果表明:1)3.6hm2森林调查样带内共记录到附生兰科植物9634株,分属于26属60种;2)除趋势对应分析(DCA)结果表明,6种森林类型中的附生兰科植物可分成5组(其中,山地常绿林与山顶矮林内的附生兰科植物归为一组);3)分布海拔范围相临近的森林类型的附生兰科植物具有较高的相似性,山地常绿林和山顶矮林附生兰科植物的相似性最高(88.9%);4)6种森林类型中,较高海拔的3种森林类型(山地雨林、山地常绿林和山顶矮林)中,附生兰科植物的丰富度和多度均显著高于其在较低海拔的3种森林类型(热带季雨林、低地雨林和热带针叶林),其中,附生兰科植物在山地常绿林内的丰富度和多度均最高;5)热带季雨林、低地雨林、热带针叶林及山地雨林内,宿主冠区附生兰科植物的多度均高于干区;山地常绿林内两者之间无显著差异;而山顶矮林干区的附生兰科植物的多度高于冠区;6)调查木上附生兰科植物的发生率在高海拔森林类型均高于其在低海拔森林类型,各森林类型内附生兰科植物的多度及物种丰富度与宿主胸径均存在显著正相关关系。     

Consistent Effects of Disturbance and Forest Edges on the Invasion of a Continental Rain Forest by Alien Plants

Continental tropical forests are thought to be resistant to alien plant invasion due to a lack of disturbance, or low propagule pressure from introduced species. We assessed the importance of disturbance and edge effects by surveying areas of submontane and lowland forest of Amani Nature Reserve in the East Usambara mountains, Tanzania. These areas are in the vicinity of Amani Botanic Garden (ABG)—a propagule source for many alien plant species. We surveyed three edges in the vicinity of the ABG plantations, using plots interspersed along multiple 250 m transects. Survey plots were either in secondary or seminatural forest, representing a difference in past disturbance). Alien plant species richness and abundance declined with increasing distance from forest edges, indicating that edge effects were important. In addition, the effect of distance on richness and abundance of alien species as adults was much smaller in seminatural than secondary forest, emphasizing that invasion of seminatural forest is less likely to occur. Abundance and occurrence of individual species showed broadly similar declines with increasing distance from the forest edge, and lower abundance in seminatural compared to secondary forest. Alien species were dominant in 15 percent of plots surveyed. As 28 percent of the Amani nature reserve forest is within 250 m of an edge, the importance of disturbance and edges could make a potentially large proportion of the forest vulnerable to alien species invasion.     

Diversity and endemism of woody plant species in the Equatorial Pacific seasonally dry forests

The biodiversity hotspot of the Equatorial Pacific region in western Ecuador and northwestern Peru comprises the most extensive seasonally dry forest formations west of the Andes. Based on a recently assembled checklist of the woody plants occurring in this region, we analysed their geographical and altitudinal distribution patterns. The montane seasonally dry forest region (at an altitude between 1,000 and 1,100 m, and the smallest in terms of area) was outstanding in terms of total species richness and number of endemics. The extensive seasonally dry forest formations in the Ecuadorean and Peruvian lowlands and hills (i.e., forests below 500 m altitude) were comparatively much more species poor. It is remarkable though, that there were so many fewer collections in the Peruvian departments and Ecuadorean provinces with substantial mountainous areas, such as Cajamarca and Loja, respectively, indicating that these places have a potentially higher number of species. We estimate that some form of protected area (at country, state or private level) is currently conserving only 5% of the approximately 55,000 km² of remaining SDF in the region, and many of these areas protect vegetation at altitudes below 500 m altitude. In contrast, the more diverse seasonally dry forests in mountainous areas remain little protected.     

Amphibian Diversity in East African Biodiversity Hotspots: Altitudinal and latitudinal Patterns

The Eastern Arc mountain chain and adjoining coastal forests of Tanzania and Kenya have been listed as world biodiversity hotspots. We report on an ongoing attempt to estimate amphibian diversity on the three best studied mountains of the Eastern Arc, the East Usambara, Uluguru and Udzungwa mountains of Tanzania, complemented by an estimate of diversity on the adjoining coastal lowland. This proves to be a complex task, which introduces a note of caution into evaluating global biodiversity estimates. Most amphibian species in eastern Tanzania occur on the coastal lowlands and are widely distributed, extending at least north or south of Tanzania and, to a variable extent, westwards to the elevated interior. Diversity patterns along the length of the lowlands are complex, with the presence of a Sahelian element in the extreme north. On the three Eastern Arc mountains studied, species turnover associated with rising altitude is greater than turnover associated with latitudinal distance between the mountain blocks, leading to greater altitudinal than latitudinal diversity in this equatorial region. A long-standing divergence is indicated between montane and lowland endemics. Although forest-associated species are not the largest contributor to the eastern Tanzanian total species diversity (some 48%), the uniqueness of these species both in lowland and montane forests, combined with their evident vulnerability to disturbance, makes them a subject for particular conservation concern, and justifies hotspot status for both montane and lowland forests.     

Species and structural diversity of church forests in a fragmented Ethiopian Highland landscape

Question: Thousands of small isolated forest fragments remain around churches (“church forests”) in the almost completely deforested Ethiopian Highlands. We questioned how the forest structure and composition varied with altitude, forest area and human influence. Location: South Gondar, Amhara National Regional State, Northern Ethiopia. Methods: The structure and species composition was assessed for 810 plots in 28 church forests. All woody plants were inventoried, identified and measured (stem diameter) in seven to 56 10 m x 10‐m plots per forest. Results: In total, 168 woody species were recorded, of which 160 were indigeneous. The basal area decreased with tree harvest intensity; understorey and middle‐storey density (<5 cm DBH trees) decreased with grazing; overstorey density (>5 cm DBH trees) increased with altitude. The dominance of a small set of species increased with altitude and grazing intensity. Species richness decreased with altitude, mainly due to variation in the richness of the overstorey community. Moreover, species richness in the understorey decreased with grazing intensity. Conclusions: We show how tree harvesting intensity, grazing intensity and altitude contribute to observed variations in forest structure, composition and species richness. Species richness was, however, not related to forest area. Our study emphasizes the significant role played by the remaining church forests for conservation of woody plant species in North Ethiopian Highlands, and the need to protect these forests for plant species conservation purposes.     

Species composition of termites of the Nyika plateau forests, northern Malawi, over an altitudinal gradient

Termites were surveyed at three altitudes (Brachystegia woodland at 1676 m and 1905 m, and Juniper woodland at 2210 m) in forests within the Nyika Plateau, northern Malawi. Sampling was by a standardized 100 m transect protocol. Termite diversity was highest in the mid‐altitude site and lowest in the Juniper forest. The assemblages were dominated by soil‐feeding termites in the Termitidae subfamilies Apicotermitinae and Termitinae, and included one new soldierless Apicotermitinae genus. The structure of the assemblages was clearly due to a mixture of altitudinal and site history factors. This was especially true of the lowest altitude forest where burning and other anthropogenic disturbance factors appear to have reduced termite diversity relative to the mid‐altitude site. The Nyika plateau shows a much higher diversity at mid‐altitudes than similar SE Asian sites, probably due to the larger area of highland in Africa than in SE Asia. In addition, the clade composition of the Nyika assemblages differs completely from that found at similar altitudes in SE Asia. This preliminary study supports the hypothesis that mid‐ to high‐ altitude assemblages in both SE Asia and Africa appear to be derived from depauperated random subsets of the lowland fauna rather than from clades specifically adapted to higher altitudes.     

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

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

Air temperature and soil phosphorus availability correlate with trait differences between two types of tropical cloud forests

Tropical cloud forests are characterized by lower air temperature and high frequency of fog condensation at canopy level, as compared with forests at lower altitudes. Few studies have been conducted to understand differences of plant functional traits in relation to the environment in this kind of forest. In this paper, we explored the community-level differences of specific leaf area (SLA) and height of plants in relation to major environmental conditions between two adjacent tropical cloud forests on Hainan Island, South China. We measured the two functional traits for all individual plants within twenty-nine and thirty-two 10 m × 10 m plots located in a low altitude tropical montane evergreen forest (TMEF) and a high altitude tropical dwarf forest (TDF), respectively.The results showed that both mean SLA and height decreased from TMEF to TDF, while phenotypic plasticity for the two functional traits increased from TMEF to TDF. Correlation analysis and multiple regression analysis showed that the mean SLA and its plasticity were significantly correlated with both air temperature and soil phosphorus. The mean height was only significantly correlated with air temperature, but its plasticity was significantly correlated with both air temperature and soil phosphorus.Our results suggest that plants in dwarf tropical cloud forests have decreased SLA and height, correlated with less favorable soil and atmospheric conditions, with a higher plasticity of these traits, as compared with the tropical montane evergreen forest. Community-level differences in SLA and plant height thus can be taken as indicators characterizing plant distribution to different types of tropical cloud forests.     

Elemental concentrations in a tropical montane rain forest in Sri Lanka

As a preliminary to studies on biogeochemistry of a tropical montane rain forest at about 1800 m altitude in Sri Lanka, the concentrations of 17 chemical elements in different forest compartments, i.e. in leaves of six floristically dominant tree species, undergrowth, litter, and soil at 0–10 cm and 11–60 cm depth were determined. Foliar chemical differences between species were large. Mean leaf water content, mean leaf area per leaf, and specific leaf weight of each species were also determined. Calcium seems to be used in relatively large quantities by these plants. Phosphorus concentration in all compartments was consistently low. A few species investigated appeared to concentrate aluminium and silicon at high amounts. Concentration differences between compartments were significant only for certain elements. Location effect on the concentrations in soil was considerable for most of the elements studied. When studying plant-soil chemical interactions in these forests, species-level concentrations must be taken into account as the species-specific chemical differences are obscured when treated as canopy leaves.     

Forest/environment relationships in Yosemite National Park,California, USA

Forest compositional patterns in Yosemite National Park, California, were related to environmental factors through numerical classification of forest types, arrangement of forest types along elevational and topographic gradients, and development of regression models relating basal area of common tree species to environmental variables. The eight forest types are differentiated primarily by elevation zone and secondarily by topographic setting. Lower montane forests (1200–1900 m) were divided into the Abies concolor/Calocedrus type occurring primarily on mesic sites and the Pinus ponderosa/Calocedrus type predominantly on xeric sites. Upper montane forests (1900–2500 m) included the Abies concolor/Abies magnifica type on mesic sites, the Abies magnifica/Pinus type on somewhat more xeric sites, and Juniperus occidentalis/Pinus jeffreyi woodlands on granitic domes. Subalpine forests (2500–3300 m) embraced three types: Tsuga mertensiana/Pinus forests on mesic sites, monotypic Pinus contorta forests on drier sites, and Pinus albicaulis/Pinus contorta groves at treeline. Regression models consistently included elevation and soil magnesium content as explanatory variables of species basal area totals. The two Abies spp. were negatively correlated with soil magnesium levels, whereas other montane species (e.g. Calocedrus decurrens, Pinus lambertiana, and Pinus ponderosa) exhibited positive correlation with soil magnesium. Topography and soil physical properties were only infrequently incorporated into species regression models.Abbreviations DBH= diameter at breast height (1.4 m) - DCA= detrended correspondence analysis - TWINSPAN= two-way indicator species analysis