Influence of tree age, tree size and crown structure on lichen communities in mature Alpine spruce forests

Testing the relations between tree parameters and the richness and composition of lichen communities in near-natural stands could be a first step to gather information for forest managers interested in conservation and in biodiversity assessment and monitoring. This work aims at evaluating the influence of tree age and age-related parameters on tree-level richness and community composition of lichens on spruce in an Alpine forest. The lichen survey was carried out in four sites used for long-term monitoring. In each site, tree age, diameter at breast height, tree height, the first branch height, and crown projection area were measured for each tree. Trees were stratified into three age classes: (1) <100 years old, immature trees usually not suitable for felling, (2) 100–200 years old, mature trees suitable for felling, and (3) >200 years old, over-mature trees normally rare or absent in managed stands. In each site, seven trees in each age class were selected randomly. Tree age and related parameters proved to influence both tree-level species richness and composition of lichen communities. Species richness increased with tree age and related parameters indicative of tree size. This relation could be interpreted as the result of different joint effects of age per se and tree size with its area-effect. Species turnover is also suspected to improve species richness on over-mature trees. Similarly to species richness, tree-level species composition can be partially explained by tree-related parameters. Species composition changed from young to old trees, several lichens being associated with over-mature trees. This pool of species, including nationally rare lichens, represents a community which is probably poorly developed in managed forests. In accordance to the general aims of near-to-nature forestry, the presence of over-mature trees should be enhanced in the future forest landscape of the Alps especially in protected areas and Natura 2,000 sites, where conservation purposes are explicitly included in the management guidelines.     

Relationships between soil microbial properties and aboveground stand characteristics of conifer forests in Oregon

Eight forest sites representing a large range of climate, vegetation, and productivity were sampled in a transect across Oregon to study the relationships between aboveground stand characteristics and soil microbial properties. These sites had a range in leaf area index of 0.6 to 16 m² m^–2 and net primary productivity of 0.3 to 14 Mg ha^–1 yr^–1.Measurements of soil and forest floor inorganic N concentrations and in situ net N mineralization, nitrification, denitrification, and soil respiration were made monthly for one year. Microbial biomass C and anaerobic N mineralization, an index of N availability, were also measured. Annual mean concentrations of NH ₄ ⁺ ranged from 37 to 96 mg N kg^–1 in the forest floor and from 1.7 to 10.7 mg N kg^–1 in the mineral soil. Concentrations of NO ₃ ^– were low ( < 1 mg N kg^–1) at all sites. Net N mineralization and nitrification, as measured by the buried bag technique, were low on most sites and denitrification was not detected at any site. Available N varied from 17 to 101 mg N kg^–1, microbial biomass C ranged from 190 to 1230 mg Ckg^–1, and soil respiration rates varied from 1.3 to 49 mg C kg^–1 day^–1 across these sites. Seasonal peaks in NH ₄ ⁺ concentrations and soil respiration rates were usually observed in the spring and fall.The soils data were positively correlated with several aboveground variables, including leaf area index and net primary productivity, and the near infrared-to-red reflectance ratio obtained from the airborne simulator of the Thematic Mapper satellite. The data suggest that close relationships between aboveground productivity and soil microbial processes exist in forests approaching semi-equilibrium conditions.Abbreviations IR infrared - LAI leaf area index - k _c proportion of microbial biomass C mineralized to CO₂ - NPP net primary productivity - TM Thematic Mapper     

Soil properties under Norway spruce differ in spruce dominated and mixed broadleaf forests of the Southern Taiga

In natural forest, disturbance changes tree species composition which in turn affects soil properties. Two areas in the Central Forest State Biosphere Reserve, in the Russian Southern Taiga Zone, differed in the intensity of disturbance: Norway spruce was the dominant species at one site, while at the other spruce was mixed with broadleaves. The presence of broadleaves was due to large gaps in the canopy having been formed, which have triggered vegetation succession. At both sites, five plots were selected to evaluate how the presence of broadleaves influences the properties of the soils under spruce. Soil samples were taken close to spruce trees and the O, A and E horizons were analysed. A difference in the distribution of organic matter in the soil horizons was evident, with a higher concentration in the O and A horizons at the spruce dominated site, while a more homogeneous distribution was found under spruce at the site where broadleaves were abundant. The organic matter did not only differ in quantity, but also in quality as estimated by the C/N ratio, and therefore affected the CEC and element relative availability. No differences at the two sites were found for water-extractable and exchangeable elements, but the ratio between the exchangeable and the acid-extractable forms were different, suggesting a higher relative availability of the elements at the spruce dominated site, and thus potentially higher leaching. Both theoretical and empirical studies have suggested that podzolisation and accumulation of organic matter in the O horizon are related to stagnation of ecosystem processes and ecosystem decline. Our data suggest that the presence to windthrow sites and the inclusion of broadleaf species acts to slow or even reverse podzolisation even in spruce dominated sites.     

Effects of stand age, wildfire and clearcut harvesting on forest floor in boreal mixedwood forests

Carbon (C) in the forest floor (FF) of the boreal region is an important reservoir of terrestrial C. We examined the effects of stand age and disturbance type (clearcutting vs. wildfire) on quantity and quality of organic C of FF in a boreal mixedwood forest of central Canada. Forest floor samples were collected from 6 post-fire (2- to 203-year-old) and 3 post-harvest age classes (2- to 28-year-old) on mesic sites, each randomly replicated three times. Samples were analyzed to determine the physical and chemical properties and the C quality was assessed by quantifying C fractions as easily labile, moderately labile and recalcitrant. Bulk density, total organic C concentration, N concentration and the cation exchange capacity increased with stand age and peaked at 85-year-old sites. Soil pH and concentration of P and K decreased with stand age. In post-fire stands, the depth of FF, total organic C, and labile C fractions increased with stand age in the 2- to 85-year-old stands, while recalcitrant C was lower in 2-year-old stands than older stands. In stands ≤28 years old, post-harvest sites had significantly higher concentration of total organic C and the three C fractions than post-fire sites in 2-year-old stands. No or marginal difference occurred between the two stand origins in 10- and 28-year-old stands. The relative proportions of C fractions did not differ with stand age or stand origin. Our results showed that the quantity of organic C in FF of boreal mixedwoods increased with stand development till 85 years and then slightly decreased in older stands, and post-harvest stands had a higher amount of organic C than post-fire stands immediately after disturbance, but the effect of two disturbances on C in FF converged shortly (within 10 years). The quality of organic C remains the same through stand development and between the two studied stand origins.     

The importance of host tree age, size and growth rate as determinants of epiphytic lichen diversity in boreal spruce forests

The amount of large and old trees has decreased in the boreal forests during the last centuries of forestry. Such trees are important habitats for epiphytic lichens and there is a growing concern for lichen species that are associated with large and old trees. However, only little is known about the relative importance of tree size versus age as determinants of lichen diversity. Here we have determined the size, age and growth rate of 157 Norway spruce trees and recorded the occurrence of epiphytic lichen species on their branches and lower stems. The study includes crustose lichens and was done in two old-growth forests in SE Norway. Tree age and tree size were the parameters that explained the largest part of epiphytic lichen diversity. Only the growth rate of the most recent time period, i.e. 1984–2004, showed a statistically significant relationship to diversity. There was no indication of a stabilising species number with increasing tree age. Slow-growing and old trees were, however, mainly of importance to the lichen species growing on stems, and this set of species were in general adversely affected by a large amount of branches. The opposite was the case for the species that were confined to branches as their diversity increased when the amount of branches increased. Our study adds empirical data to support the importance of large and old trees as bearers of biodiversity in boreal forests. Site preservation and patch retention of groups of old and large trees is recommended as measures to maintain epiphytic lichen diversity.     

Site identity and moss species as determinants of soil microbial community structure in Norway spruce forests across three vegetation zones

Soil microbial community structure was investigated by PLFA-analysis in four spruce forests in Norway. The maximum latitudinal distance between the sites was approximately 350 km. Bilberry Vaccinium myrtillus dominated the forest floor vegetation in the study sites, which were selected because of the vegetation type. Soil samples were taken from all four sites under close to 100% homogeneous ground cover of each of two feathermoss species, i.e. Hylocomium splendens or Pleurozium schreberi, respectively. These mosses are ubiquitous in the boreal forest and constitute an abundant component of the forest floor vegetation over vast areas. Since there are no studies on how these mosses affect soil microbial community structure, our first aim was to investigate the effect of moss species on soil microbial communities. Our second aim was to investigate whether microbial communities differ among geographically separated forest sites with similar vegetation across vegetation zones. Soil microbial community structure differed between the study sites, although they appeared similar in terms of vegetation and abiotic soil conditions. Study site was the most important predictor of the variation in the PLFAs, more important than moss species, although there was a tendency for separation of microbial community structure between the two moss species.     

Invasive shrub distribution varies with distance to roads and stand age in eastern deciduous forests in Indiana, USA

We documented the relationship between densities of invasive exotic shrubs, distance to road, and successional age of the forest in 14 forested sites throughout central and southern Indiana. Roadways are increasingly abundant, human-made features that can be conduits for the spread of invasive exotic plants in a number of ecosystems. Little is known, however, about the role of roads in eastern deciduous forest ecosystems where road density is high. Further, it is not known whether the distribution of exotic plants along roads depends on the successional age of the forest. In this study, densities of four of seven exotic shrub species declined with increasing distance to the nearest road across all successional ages. Greater densities of exotic shrubs were found in young and mid-successional forests than mature forests. However, there was no interaction between distance to road and forest age, suggesting that the role of roads in the invasion process does not change across forest successional ages. We outline several potential mechanisms that may drive patterns of shrub distribution along roadside edges as a guide for future research.     

Significant and persistent impact of timber harvesting on soil microbial communities in Northern coniferous forests

Forest ecosystems have integral roles in climate stability, biodiversity and economic development. Soil stewardship is essential for sustainable forest management. Organic matter (OM) removal and soil compaction are key disturbances associated with forest harvesting, but their impacts on forest ecosystems are not well understood. Because microbiological processes regulate soil ecology and biogeochemistry, microbial community structure might serve as indicator of forest ecosystem status, revealing changes in nutrient and energy flow patterns before they have irreversible effects on long-term soil productivity. We applied massively parallel pyrosequencing of over 4.6 million ribosomal marker sequences to assess the impact of OM removal and soil compaction on bacterial and fungal communities in a field experiment replicated at six forest sites in British Columbia, Canada. More than a decade after harvesting, diversity and structure of soil bacterial and fungal communities remained significantly altered by harvesting disturbances, with individual taxonomic groups responding differentially to varied levels of the disturbances. Plant symbionts, like ectomycorrhizal fungi, and saprobic taxa, such as ascomycetes and actinomycetes, were among the most sensitive to harvesting disturbances. Given their significant ecological roles in forest development, the fate of these taxa might be critical for sustainability of forest ecosystems. Although abundant bacterial populations were ubiquitous, abundant fungal populations often revealed a patchy distribution, consistent with their higher sensitivity to the examined soil disturbances. These results establish a comprehensive inventory of bacterial and fungal community composition in northern coniferous forests and demonstrate the long-term response of their structure to key disturbances associated with forest harvesting.     

黄土高原羊圈沟小流域不同坡向、林龄刺槐人工林叶片尺度水碳交换及叶片养分差异

以黄土高原典型小流域(羊圈沟小流域)内刺槐人工林为对象,通过测定叶片尺度水碳交换过程及主要影响因素,探究了叶片尺度水碳交换随坡向和林龄的变化,并分析了叶片养分变化对水碳交换过程的潜在影响。结果表明:(1)11年生与27年生刺槐处于不同的生长阶段,前者水碳交换的强度高于后者。(2)半阳坡中光合固碳、水分利用效率及CO_2供应能力强的刺槐所占比例高于阳坡,说明半阳坡更适合刺槐生长。(3)叶片N、K含量在样方间的变化趋势为:半阳坡阳坡,27年生11年生,说明半阳坡刺槐的生长状况好于阳坡,高林龄刺槐的生长状况好于低林龄刺槐。同时,各样方间K含量具有显著差异,说明K对坡向及林龄的变化较为敏感。因此,与阳坡相比,该区域未来人工刺槐林建设应优先选择水热条件更为适宜的半阳坡,且在富含有效钾的土壤中进行植被恢复是提高植物水分利用效率的潜在途径,该研究结果对于未来干旱半干旱区退耕还林方案的改进及已有人工林的管理具有借鉴意义。     

Structure and function of anthropogenically altered microbial communities in coastal waters

Human-based (anthropogenic) nutrient and other pollutant enrichment of the world's coastal waters is causing unprecedented changes in microbial community structure and function. Symptoms of these changes include accelerating eutrophication, the proliferation of harmful microalgal blooms, excessive oxygen consumption (hypoxia, anoxia), increasing toxicity, altered routes and fluxes of organic and inorganic matter cycling, and disruption of food webs. Biogeochemical and trophic consequences are expanding on local, regional and global scales.     

Interactive effects of wildfire and permafrost on microbial communities and soil processes in an Alaskan black spruce forest

Boreal forests contain significant quantities of soil carbon that may be oxidized to CO₂ given future increases in climate warming and wildfire behavior. At the ecosystem scale, decomposition and heterotrophic respiration are strongly controlled by temperature and moisture, but we questioned whether changes in microbial biomass, activity, or community structure induced by fire might also affect these processes. We particularly wanted to understand whether postfire reductions in microbial biomass could affect rates of decomposition. Additionally, we compared the short‐term effects of wildfire to the long‐term effects of climate warming and permafrost decline. We compared soil microbial communities between control and recently burned soils that were located in areas with and without permafrost near Delta Junction, AK. In addition to soil physical variables, we quantified changes in microbial biomass, fungal biomass, fungal community composition, and C cycling processes (phenol oxidase enzyme activity, lignin decomposition, and microbial respiration). Five years following fire, organic surface horizons had lower microbial biomass, fungal biomass, and dissolved organic carbon (DOC) concentrations compared with control soils. Reductions in soil fungi were associated with reductions in phenol oxidase activity and lignin decomposition. Effects of wildfire on microbial biomass and activity in the mineral soil were minor. Microbial community composition was affected by wildfire, but the effect was greater in nonpermafrost soils. Although the presence of permafrost increased soil moisture contents, effects on microbial biomass and activity were limited to mineral soils that showed lower fungal biomass but higher activity compared with soils without permafrost. Fungal abundance and moisture were strong predictors of phenol oxidase enzyme activity in soil. Phenol oxidase enzyme activity, in turn, was linearly related to both ¹³C lignin decomposition and microbial respiration in incubation studies. Taken together, these results indicate that reductions in fungal biomass in postfire soils and lower soil moisture in nonpermafrost soils reduced the potential of soil heterotrophs to decompose soil carbon. Although in the field increased rates of microbial respiration can be observed in postfire soils due to warmer soil conditions, reductions in fungal biomass and activity may limit rates of decomposition.     

Soil aggregates in a tropical deciduous forest: effects on C and N dynamics, and microbial communities as determined by t-RFLPs

The aim of this study was to analyze C and N dynamics, as well as, soil bacterial community structure within soil micro- and macro-aggregates in a tropical deciduous forest in México. We measured, for three landscape positions and three seasons of the year: total, microbial and available forms of C and N; potential C and N mineralization; and soil bacterial communities by using t-RFLPs. The highest total C concentrations were found in the north-slopes and in the dry season (DS) samples. In general, micro-aggregates had higher concentrations than macro-aggregates of available C and N forms, and microbial C. Similarly, micro-aggregates had the highest potential C mineralization and net N mineralization. We detected 149 different OTUs (operational taxonomic units) from which 50% was shared by the two aggregate size fractions, 25% was exclusive to micro-aggregates and the 25% left was found only in macro-aggregates. Top-hills were richer in OTUs than north and south-slopes. The Unweighted Pair Group Method with Arithmetic mean (UPGMA) analysis indicated clear differences in community composition between the two aggregate size-fractions in relation to the presence of OTUs. These results suggest that the main difference between micro- and macro-aggregates is due to the community structure within each soil fraction and this difference could affect soil nutrients dynamics.     

Structure and function of microbial communities in soil from the southern Taiga

General regularities in the structure of the microbial communities of southern taiga soil ecosystems and taxonomic differences between the microbial communities of soils with different hydrothermal characteristics are discussed with reference to the main types of soils of the Central State Forest Biosphere Reserve.     

Soil fauna communities and microbial respiration in high Arctic tundra soils at Zackenberg, Northeast Greenland

The soil fauna communities were described for three dominant vegetation types in a high arctic site at Zackenberg, Northeast Greenland. Soil samples were extracted to quantify the densities of mites, collembolans, enchytraeids, diptera larvae, nematodes and protozoa. Rates of microbial respiration were also assessed. Collembolans were found in highest densities in dry heath soil, about 130,000 individuals m⁻², more than twice as high as in mesic heath soils. Enchytraeids, diptera larvae and nematodes were also more abundant in the dry heath soil than in mesic heath soils, whereas protozoan densities (naked amoeba and heterotrophic flagellates) were equal. Respiration rate of unamended soil was similar in soil from the three plots. However, a higher respiration rate increase in carbon + nutrient amended soil and the higher densities of soil fauna (with the exception of mites and protozoa) in dry heath compared to the mesic heath soils indicated a higher decomposition rate here.     

Changes in soil heterotrophic respiration,carbon availability,and microbial function in seven forests along a climate gradient

Soil microbial communities play an essential role in soil carbon (C) emission and C sequestration in forest ecosystems. However, little information is available regarding the relationship between soil C dynamics and microbial substrate utilization at large scales. Along the North–South Transect of Eastern China (NSTEC), seven forests representative of boreal, temperate and tropical biomes were examined. Soil heterotrophic respiration (R_h), soil dissolved organic C (DOC), microbial biomass C (MBC), and microbial community-level physiological profiles (CLPPs) were investigated using biochemical measurements, static chamber-gas chromatography analysis, and Biolog-Eco microplates, respectively. We found that soil R_h rates were significantly higher in subtropical and boreal forests than in temperate forests. Conversely, the concentrations of soil DOC and MBC, as well as microbial metabolic activity and functional diversity, were consistently higher in temperate forests than in subtropical forests. There were considerably different substrate utilization profiles among the boreal, temperate, and subtropical forests. Soil microorganisms from the temperate and boreal forests mainly metabolized high-energy substrates, while those from the subtropical forests used all substrates equally. In addition, soil R_h rates were significantly and negatively related to soil labile C concentrations, total metabolic activity, and the intensity of individual substrate utilization, indicating that soil microbes assimilated more soil substrates, thereby reducing CO₂ emissions. Overall, our study suggests that climate factors, as well as substrate availability, dominate the activities and functions of soil microbes at large scales.     

The microbial loop in a humic lake: seasonal and vertical variations in the structure of the different communities

Seasonal and vertical variations of the main microbial communities (heterotrophic bacteria, autotrophic picoplankton, auto- and heterotrophic nanoflagellates, ciliated protozoa and microalgae) and auto- and heterotrophic activities were estimated in a brown-colored humic and moderately acid lake in central France, the lake of Vassivière. The results demonstrated the dominant role of light in the vertical distribution of autotrophic and mixotrophic microorganisms which are confined to the 0–5 m layer during thermal stratification. The bacterial biomass was high throughout the water column probably because of the great availability of dissolved organic matter. Consequently, the predatory microzooplankton and particularly the various trophic groups of ciliated protozoa, were distributed in the water column according to the vertical distribution of the particular food resources (detritus, bacteria, algae). However, despite the great abundance of algae and bacteria, biomass of flagellated and ciliated protozoa was relatively weak. Most of the phytoplanktonic biomass was filamentous (Diatoms) or colonial (Cyanobacteria) and therefore almost probably difficult to ingest for algivorous microzooplankton. Regarding the low abundance of bacterivorous protozoa, the relation with the special physicochemical properties of this lake is discussed.

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Résumé Les variabilités saisonnière et verticale de l'abondance et de la biomasse des principles communautés de la boucle microbienne (bactéries hétérotrophes, picoplancton autotrophe, protozoaires flagellés auto- et hétérotrophes, protozoaires ciliés, microalgues et microcyanobactéries), et des activités auto- et hétérotrophes, ont été étudiées dans un lac à caractère humique et modérément acide du Massif Central Français, le lac de Vassivière.Le dénombrement des communautés de microorganismes a été réalisé en microscopie inversée et à épifluorescence après mise en oeuvre des fixations et des colorations adéquates. Les activités photosynthétique, photo- et chemohétérotrophes ont été mesurées à partir de l'assimilation de NaH¹⁴CO₃ et d'un mélange d'acides aminés tritiés grâce à une technique de double marquage. Les résultats obtenus mettent en évidence le rôle prépondérant du facteur lumineux dans la répartition verticale des microorganismes autotrophes et mixotrophes, dont l'essentiel de la biomasse est confiné dans la zone 0–5 m en période de stratification, alors que la biomasse bactérienne est élevée sur l'ensemble de la colonne d'eau en raison, sans doute, de la grande disponibilité en matiére organique dissoute. Consécutivement, le microzooplancton prédateur, et notamment les différents types trophiques de protozoaires ciliés, se répartit dans la colonne d'eau selon la distribution verticale des ressources nutritives particularies. Cependant, compte-tenu de l'abondance bactérienne et algae, la biomasse des protozoaires flagellés et ciliésest relativement peu importante. Concernant les espèces algivores, l'essentiel de la biomasse phytoplanctonique est de nature filamenteuse (Diatomées) ou coloniale (Cyanobactéries) est est donc sans doute difficilement ingérable pour le microzooplancton. Enfin, les relations entre les caractéristiques physico-chimiques spécifiques de ce lac et le faible développement des protozoaires bactérivores sont discutées.

     

Interactive influences of climate and parent material on soil microbial community structure in Bornean tropical forest ecosystems

Climate and parent material strongly control vegetation structure and function, yet their control over the belowground microbial community is poorly understood. We assessed variation in microbial lipid profiles in undisturbed forest soils (organic and surface mineral horizons) along an altitudinal gradient (700, 1,700, and 2,700 m a.s.l. mean annual temperature of 12–24°C) on two contrasting parent materials (acidic metasedimentary vs. ultrabasic igneous rock) in Mt. Kinabalu, Borneo. Soil organic carbon and nitrogen concentrations were generally higher at higher altitudes and, within a site, at upper soil horizons. Soil pH ranged from 3.9 to 5.3, with higher values for the ultrabasic soils especially at higher altitudes. The major shifts in microbial community structure observed were the decline in the ratio of fungal to bacterial lipid markers both with increasing soil depth and decreasing altitude. The positive correlation between this ratio with soil C and N concentrations suggested a strong substrate control in accord with the literature from mid to high-latitude ecosystems. Principal component analysis using seven groups of signature lipids suggested a significant altitude by parent material interaction—the significant difference in microbial community structure between the two rock types found at 2,700-m sites developed on weakly weathered soils diminished with decreasing altitude towards 700-m sites where soils were strongly weathered. These results are consistent with the hypothesis that parent material effect on soil microbial community (either directly via soil geochemistry or indirectly via floristic composition) is stronger at an earlier stage of ecosystem development.     

Biogenic volatile organic compounds as potential carbon sources for microbial communities in soil from the rhizosphere of Populus tremula

Catabolism of a (14)C-labelled volatile monoterpene compound (geraniol) to (14)CO(2) was investigated in soils taken from the rhizosphere at distances up to 200 cm from the trunks of three small Populus tremula trees growing at different sites in Slovenia. Emissions of limonene of up to 18 microg m(-2) h(-1) were detected from the soil surface at each site. Evolution of (14)C-labelled CO(2) was measured as a product of catabolism of (14)C-labelled geraniol introduced into the soil samples. Indigenous soil microorganisms degraded the geraniol rapidly. There was a significant difference in relative lag times and rates of catabolism along the gradient from the tree trunks, with relatively longer lag times and lower rates occurring in soil samples from the farthest point from the tree.     

Comparison of leaf litter ant communities in woodlands, lowland forests and montane forests of north-eastern Tanzania

Ants were extracted in Winkler bags from sifted leaf litter sampled in arange of forest and woodland types in and around Mkomazi Game Reserve innorth-eastern Tanzania, including the Eastern Arc Mountains of South Pare andWest Usambara. A total of 87 ant species were recorded, of which 32.2% were onlyrecorded from montane forests (1400–1850 m altitude), 6.9%only from lowland forest (540–810 m), 19.5% only fromwoodland (300–1080 m), and 16.1% in all three forest types.Of the 28 species recorded only from montane forests, 12 species were only foundin the Mkomazi forests, four only in the Pares and seven species only in theUsambaras. Sites of similar altitude grouped together in a cluster analysis, andspecies richness decreased with an increase in altitude. The lowland forest andclosed woodland sites did not form distinct communities. To ensure preservationof ant species, forests from a full range of altitudes need to be conserved.This study confirms the status of the West Usambara forests as having a highlyendemic biota, and the critical need to adequately conserve the remainingvestiges of montane forest within Mkomazi Game Reserve.