Characteristics of the Achnatherum splendens community along the Qinghai-Tibet Railway, China

The Achnatherum splendens community is an important type of warm steppe vegetation which plays a very important role in environmental protection in the area along the Qinghai-Tibet Railway. In order to under-stand its dynamic features, the characteristics of the A. splendens community, such as spatial distribution pattern, species richness and biomass, were examined near Haiyan County, Qinghai Province. Results showed that the A. splendens community consisted of two types of mosaic patches, the microaggregation dominated by A. splendens and the microaggregation dominated by Stipa sp. The number and size of the two patches showed an interesting distribution pattern. The A. splendens patch decreased with increasing distance away from the railway, while species diversity increased. The total coverage of A. splen-dens was significantly higher than that of Stipa sp. The number and size of patches may indicate the degradation of the A. splendens community, which could be acceler-ated by human activities. Based on the characteristics and status quo of the A. splendens community, we suggest that some measures be taken to control human activities and rodents to conserve the A. splendens community.     

青藏铁路沿线温性草原区芨芨草（Achnatherum splendens）群落特征

芨芨草群落是青藏铁路沿线一类重要的温性草原植被类型,对青藏铁路两侧及其四周的水土保持起着非常重要的作用。对青藏铁路西格段海晏县附近的芨芨草草原的空间分布特征、物种丰富度和生物量等特征进行了分析。结果显示,研究地区芨芨草群落由芨芨草斑块和针茅斑块镶嵌而成,两类斑块的数量、大小和斑块间大小变化幅度等特征在空间上表现出一定的规律;芨芨草斑块大小在样线上所占的比例比针茅斑块为小,且有随远离铁路而减小的趋势,而物种丰富度则出现相反的变化趋势;芨芨草斑块的盖度和单位面积上地上生物量显著高于针茅斑块。斑块的大小和数量在一定程度上反应了芨芨草群落的退化状况。人为活动往往会加剧芨芨草群落的斑块片段化。针对芨芨草群落的特征和现状,建议加强对这些区域内人为活动的限制力度以减少对芨芨草群落的干扰。     

The combined positive effects of two dominant species in an arid shrub-herbaceous community: implications from the performance of two associate species

To test the hypothesis that the species could be more beneficial in association with two-nurse species than with single-nurse species, we surveyed an arid shrub-herbaceous community primarily composed of two dominant species, Achnatherum splendens and Nitraria tangutorum and two associate species, Phragmites australis and Reaumuria soongorica in Northwest China. The performance (frequency, abundance, and size) of the two associate species in neighbored patches (patches formed by a dominant species which were closely adjacent to the patches formed by the other dominant species), isolated patches (patches formed by one dominant species with no neighbor), and open areas was compared to analyze the individual and combined effects of the two dominant species. A. splendens and N. tangutorum appeared to have reciprocal facilitation effects when growing adjacent to one another, as evidenced by the increased size of neighbored patches over isolated ones. The individual effects of A. splendens on the associate species should be generally neutral, since the frequency, abundance, size, and co-occurrence frequency of the two associate species in isolated A. splendens patches and open areas were the same (except for the significantly larger size of R. soongorica in isolated A. splendens patches). The individual effects of N. tangutorum were positive to the associate species R. soongorica, as evidenced by the fact that the frequency and abundance of R. soongorica was significantly higher (P < 0.05), and the size of two associate species was significantly larger (P < 0.05), in isolated N. tangutorum patches than in open areas. However, the frequency and abundance of P. australis, and the co-occurrence frequency of the two associate species in isolated N. tangutorum patches and in open areas were the same. In comparison to the isolated patches, there were significantly higher frequency and abundance (P < 0.05), larger sizes (P < 0.05), and higher co-occurrence frequency (P < 0.05) of the two associate species in neighbored N. tangutorum patches. Since the neighbored patches could be influenced by both dominant species, the combined effects of A. splendens and N. tangutorum were identified as positive and over-additive. We suggested that the indirect facilitation or synergistic effects could account for the combined effects of the two dominant species.     

内蒙古典型草原区芨芨草群落适生生境

芨芨草群落是干旱与半干旱地区重要的植被资源,内蒙古典型草原区常在地形稍低的部位镶嵌有芨芨草群落,且芨芨草群落常与低洼湿草甸群落相连。为了探究芨芨草群落的适生生境,在锡林郭勒盟典型草原区选取了4个代表性调查区,从芨芨草群落及其相邻群落分布的地形特征、调查区的地下水埋深、1 m深土层的土壤电导率、土壤pH值和土壤质地等指标上对芨芨草适生生境进行探究,结果表明,芨芨草群落在内蒙古典型草原区常分布在河漫滩和丘间蝶形洼地地区,呈现出由高程梯度导致的与地带性针茅群落、湿草甸群落带状分布的特点,导致这种成带分布的原因与地下水埋深及地表径流有关,芨芨草群落生境的地下水埋深一般在1-3 m且有地表径流的补充;芨芨草群落耐盐碱性强,其生境也与反映土壤盐分含量的土壤电导率和土壤pH值有关;但芨芨草群落生境的形成不是由土壤机械组成所致。     

Remnant shrubs in Mediterranean semi-arid steppes: effects of shrub size, abiotic factors and species identity on understorey richness and occurrence

Isolated shrub patches are a key element for community structure and dynamics in semi-arid ecosystems, and may act as “hot spots” of understorey species diversity. Despite its importance, the relationships between shrub patch characteristics and understorey species richness, and the relative importance of these characteristics against other factors driving understorey species richness, are still poorly understood. We studied perennial species richness under the canopy of late-successional shrubs in semi-arid Stipa tenacissima steppes of SE Spain. The main aim of this study was to evaluate the relationship between shrub size and understorey richness, and to test if this relationship is modified by species identity. We also aimed to know the relative importance of shrub patch characteristics and abiotic factors as predictors of the distribution of understorey species under shrub canopies. Altitude and geographical co-ordinates were able to significantly explain the patterns of shrub occurrence and abundance in the study area. Understorey species richness was significantly related to the size of individual shrubs according to a simple power relationship for all evaluated species. Slope values of linear regressions with log-transformed data, ranging between 0.22 and 0.37, did not differ between species. Site characteristics, patch characteristics and patch species identity significantly explained the occurrence of species on patches. However, the relative importance of site characteristics was higher than that of patch characteristics and species identity in explaining these patterns. Our results agree with the general expectations of the theory of island biogeography and complement previous studies that emphasise the importance of late-successional shrubs in semi-arid Stipa steppes. Environmental management activities within these steppes should promote the conservation of remnant shrubs, as well as its introduction with restoration activities.     

温带草原7种针茅植物根系特征及其对环境因子变化的适应

为比较针茅（Stipa）植物适应策略,以大尺度梯度下（>1600 km）温带草原主要针茅植物为对象,系统研究了8个样点7种针茅根系生物量、根冠比、解剖结构和生理调节物质的差异及其对环境因子的适应。在由东北至西南的区域上,随降水量下降针茅植物根系的抗旱特征增强或适应策略趋于复杂,不同针茅植物根系对水分变化（或旱季和雨季）有着不同的适应策略。综合分析表明贝加尔针茅（S. baicalensis）、大针茅（S.grandis）及克氏针茅（S.krylovii）（多伦样点）的生长受干旱制约,对降水高度敏感,雨季降水促进其生物量快速积累。沙生针茅（S.glareosa）、短花针茅（S.breviflora）、戈壁针茅（S.gobica）、本氏针茅（S.bungeana）等通过增大根冠比和渗透调节物质累积等途径提高根系吸水和保水能力,抵御干旱胁迫。偏相关分析显示实验区域针茅植物根系性状与降水量和海拔高度存在显著的相关性。     

Two-dimensional pattern analysis on dominant species and community in subalpine meadow of Luya Mountain, Shanxi Province, China

The spatial pattern analysis of population and community is important to understand community structure and has become one key topic in modern plant ecology. There are many techniques for analyzing one-dimensional pattern in ecological literature. Two-dimensional pattern analysis is better than one-dimensional analysis in the study on community spatial characteristics and structure. However, it is hard to analyze these two-dimensional patterns due to poor effective methodology. The two-dimensional sampling using two transects that meet at right angles was applied to get quadrat data in this work. And then the data from the two transects were analyzed separately by one-dimensional pattern analysis method, two-term local quadrat variance. The length, width, and area of patches at different scales of pattern for populations were obtained from the analysis. For community pattern, detrended correspondence analysis (DCA) was employed to summarize the species information firstly, and then the first DCA axis scores were analyzed to check its pattern. The application of this method to the pattern analysis on dominant populations and community for subalpine meadow (Comm. Polygonum viviparum + Carex rigescens + Kobresia bellardii) in the Luya mountains showed that it could release the characteristics of spatial pattern clearly and was a very effective technique. The method is easy to use and saves time with obvious advantages, compared with the two-dimensional pattern analysis methods presented in the literatures. In the study meadow, the patterns of the main dominant species, Polygonum viviparum, Carex rigescens, and Kobresia bellardii, were apparent and comparatively regular in shape with large areas of patches at the same scale compared with other species such as Festuca sp. and Thalictrum petaloideum. There were two or three scales of patterns for each plant population studied. This was related to population features, the interaction with environmental factors, and their dominant position in the community. The two scales of patterns for the meadow community were clear. The pattern of community was closely correlated with that of the main dominant species. The patches of dominant species were inter-distributed and overlapped, and formed the community pattern together. This is beneficial for utilization of resources, and keeping the community stable. __________ Translated from Acta Ecologica Sinica, 2005, 25(6): 1264–1268 [译自: 生态学报]     

青藏铁路沿线温性草原区芨芨草(Achnatherum splendens)群落特征

芨芨草群落是青藏铁路沿线一类重要的温性草原植被类型,对青藏铁路两侧及其四周的水土保持起着非常重要的作用。对青藏铁路西格段海晏县附近的芨芨草草原的空间分布特征、物种丰富度和生物量等特征进行了分析。结果显示,研究地区芨芨草群落由芨芨草斑块和针茅斑块镶嵌而成,两类斑块的数量、大小和斑块间大小变化幅度等特征在空间上表现出一定的规律;芨芨草斑块大小在样线上所占的比例比针茅斑块为小,且有随远离铁路而减小的趋势,而物种丰富度则出现相反的变化趋势;芨芨草斑块的盖度和单位面积上地上生物量显著高于针茅斑块。斑块的大小和数量在一定程度上反应了芨芨草群落的退化状况。人为活动往往会加剧芨芨草群落的斑块片段化。针对芨芨草群落的特征和现状,建议加强对这些区域内人为活动的限制力度以减少对芨芨草群落的干扰。     

Do seed and microsite limitation interact with seed size in determining invasion patterns in flooding Pampa grasslands?

Additive influences of the invasion ability of species (invasiveness) and the characteristics of the habitat which make it invasible (invasibility) cannot fully explain grassland invasion patterns. We tested the hypothesis that different species assemblages of grassland communities may partly result from interactive influences between the relative invasiveness of available species and community invasibility. During 10 months, we evaluated seed and microsite limitation of seedling emergence, survival, and recruitment of plants belonging to species with different seed size (large-seeded species vs. small-seeded species), with in a two-phase community mosaic typical of semi-natural grasslands in the southern flooding Pampa of Argentina. Seeds of large versus small-seeded species were sown either in species-poor patches dominated by a tall tussock grass (“pajonal”) or in species-rich patches dominated by short grasses (“matrix”), subjected to different levels of canopy disturbance (cut vs. uncut). Seed addition promoted seedling emergence for 7 out of the 10 species sown, and this effect was higher for large than for small-seeded species. After seed limitations were removed, interactive effects among seed size, community state and canopy disturbance reflected a strong positive influence of seed size on plant recruitment only in cut pajonal patches. Therefore, according to the stage of invasion process, relative species success may depend on non-interactive (seed-size effect on seed limitation to seedling emergence) or interactive influences among species invasibility and community invasiveness (from seedling emergence to plant recruitment). As a general conclusion, different assemblages of species are expected to successfully colonize spatially close grassland patches, according to both the available invasible species (seed size) and the community state and stage (species composition and canopy disturbance).     

Stability of the spatio-temporal distribution and niche overlap in neotropical earthworm assemblages

The spatial distribution of soil invertebrates is aggregated with high-density patches alternating with low-density zones. A high degree of spatio-temporal organization generally exists with identified patches of specific species assemblages, in which species coexist according to assembly rules related to competitive mechanisms for spatial and trophic resources occur. However, these issues have seldom been addressed. The spatio-temporal structure of a native earthworm community in a natural savanna and a grass–legume pasture in the Colombian “Llanos” was studied during a 2-year-period. A spatially explicit sampling design (regular grid) was used to discern the distribution pattern of species assemblages in both systems. Earthworms were collected from small soil pits at three different sampling dates. Data collected from 1 m² soil monoliths were also used in the present study. Data were analyzed with the partial triadic analysis (PTA) and correlograms, while niche overlap was computed with the Pianka index. The PTA and correlogram analysis revealed that earthworm communities displayed a similar stable spatial structure in both systems during the 2-year study period. An alternation of population patches where different species' assemblages dominated was common to all sampling dates. The medium-sized Andiodrilus sp. and Glossodrilus sp. exhibited a clear spatial opposition in natural savanna and the grass–legume pasture for the duration of the study. The Pianka index showed a high degree of niche overlapping in several dimensions (vertical distribution, seasonality of population density) between both species. The inclusion of space-time data analysis tools as the PTA and the use of classical ecological indices (Pianka) in soil ecology studies may improve our knowledge of earthworm assemblages' dynamics.     

<Emphasis Type="Italic">Fusarium matuoi</Emphasis> sp. nov. and its teleomorph <Emphasis Type="Italic">Cosmospora matuoi</Emphasis> sp. nov.

A group of Fusarium isolates from slime flux similar to F. aquaeductuum produced unique, strongly curved, aseptate, C-shaped conidia. They were found to be identical to F. splendens nom. nud. Dried specimens from which F. splendens was originally isolated were reexamined and characterized as a new species of Cosmospora. Cosmospora matuoi sp. nov. is proposed for the teleomorph, and Fusarium matuoi sp. nov. is proposed for its anamorph.     

Life-history, thallus ontogeny, and the effects of temperature, irradiance and salinity on growth of the edible green seaweed Gayralia spp. (Chlorophyta) from Southern Brazil

Gayralia K.L. Vinogr. is a monostromatic green alga of commercial importance in the southern Brazil, and its cultivation is being considered. This paper reports some basic aspects of the biology of this poorly known genus. Two populations of Gayralia spp., from outer and inner sectors of Paranaguá Bay, showed an asexual life history with a distinct pattern of thallus ontogeny. In one population (Gayralia sp. 1), zooids developed an expanded monostromatic blade directly, while in the other (Gayralia sp. 2) zooids produced an intermediate saccate stage, before giving rise to a monostromatic blade. Thalli of the two species differ in size and in cell diameter. The effects of temperature (16–30°C), irradiance (50–100 μmol photons m⁻² s⁻¹), and salinity (5–40 psu) on the growth of both populations were assessed. Plantlets of Gayralia sp. 1 from in vitro cultures showed a broader tolerance to all salinity and irradiance levels tested, with the highest growth rate (GR; mean 17% day⁻¹) at 21.5°C and 100 μmol photons m⁻² s⁻¹. Plantlets of Gayralia sp. 1 collected during the winter in the field showed higher GR, ranging from 5% day⁻¹ to 7.5% day⁻¹ in salinities from 20 to 40 psu, and 2.0% day⁻¹ and 4.3% day⁻¹ for plantlets collected during the summer. Gayralia sp. 2 from the field showed highest GR at salinity of 15 psu. These results suggest distinct physiological responses of the two species, in accordance with their distribution: Gayralia sp. 2 is limited to the inner areas of the estuary, while Gayralia sp. 1 grows in outer areas, where salinity values are higher than 20 psu. These data indicate that Gayralia sp. 1 has a higher potential for aquaculture than Gayralia sp. 2 due to its larger thalli, higher GR, and wider tolerance to environmental variations.     

Nitrogen fixation in mixed <Emphasis Type="Italic">Hylocomium splendens</Emphasis> moss communities

The pleurocarpus feather moss, Hylocomium splendens, is one of two co-dominant moss species in boreal forest ecosystems and one of the most common mosses on earth, yet little is known regarding its capacity to host cyanobacterial associates and thus contribute total ecosystem N. In these studies, we evaluated the N-fixation potential of the H. splendens–cyanobacteria association and contrasted the N-fixation activity with that of the putative N-fixing moss–cyanobacteria association of Pleurozium schreberi. Studies were conducted to: quantify N-fixation in H. splendens and P. schreberi in sites ranging from southern to northern Fennoscandia; assess N and P availability as drivers of N-fixation rates; contrast season-long N-fixation rates for both mosses; and characterize the cyanobacteria that colonize shoots of H. splendens. Nitrogen-fixation rates were generally low at southern latitudes and higher at northern latitudes (64–69°N) potentially related to anthropogenic N deposition across this gradient. Nitrogen fixation in H. splendens appeared to be less sensitive to N deposition than P. schreberi. The season-long assessment of N-fixation rates at a mixed feather moss site in northern Sweden showed that H. splendens fixed a substantial quantity of N, but about 50% less total N compared to the contribution from P. schreberi. In total, both species provided 1.6 kg fixed N ha⁻¹ year⁻¹. Interestingly, H. splendens demonstrated somewhat higher N-fixation rates at high fertility sites compared to P. schreberi. Nostoc spp. and Stigonema spp. were the primary cyanobacteria found to colonize H. splendens and P. schreberi. These results suggest that H. splendens with associated Nostoc or Stigonema communities contributes a significant quantity of N to boreal forest ecosystems, but the contribution is subordinate to that of P. schreberi at northern latitudes. Epiphytic cyanobacteria are likely a key factor determining the co-dominant presence of these two feather mosses across the boreal biome.     

Structure of vegetation patches in northwestern Patagonia, Argentina

Vegetation of arid and semiarid environments has in general a patchy distribution. Our objective was to (a) determine several qualitative and quantitative analytical characteristics of vegetation patches in an arid zone of Patagonia, Argentina, and (b) investigate relationships between them. Annual precipitation in this area was 200 mm during 1999–2005. Eight transects involving ten patches each were studied within a 15 × 15 km area. Mean (±1 SE) values (n = 80) in the vegetation patches were 315 ± 25 and 207 ± 16 cm for the greatest and lowest patch diameter, respectively; 23 ± 2 cm for mound height; 113 ± 12 cm for maximum vegetation height; and 170 ± 18 cm and 58 ± 2% for distance to the next vegetation patch and vegetation patch cover within a transect, respectively. Correlations between greatest and lowest diameters, mound height, maximum plant height and distance to the closest vegetation patch were all significant (P < 0.01; n = 80). In all vegetation patches, the greatest and lowest frequencies were found for the grass Stipa spp. (71.2%) and the shrub Grindelia chiloensis (Cornel.) Cabrera (12.5%). Stipa spp. and the shrub Atriplex lampa (Moq.) D. Dietr. showed the highest simultaneous frequency (50%). A reasonable association among species (>45%) was found for Stipa spp., Atriplex lampa and the shrubs Larrea divaricata Cav., Lycium chilense Miers ex Bertero and Junellia ligustrina (Lag.) Moldenke. Larrea divaricata and Atriplex lampa contributed more than 84% of the total patch standing crop (5,777 ± 435 g). Average patch size and specific diversity were 5.93 ± 0.33 m² and 1.31 ± 0.11, respectively. Aboveground standing crop of the two dominant shrubs decreased as plant species diversity increased (P < 0.05). Conservation of vegetation patches is crucial to prevent increased soil erosion and desertification in the study ecological system.     

Diversity and composition of macrobenthic community associated with sandy shoals of the Louisiana continental shelf

Along the Louisiana, USA continental shelf, sandy shoals are shallow, possibly oxygen-rich “islands” surrounded by deeper muddy deposits prone to hypoxia. Shoals also contain significant quantities of fine sand that may be mined in the future for coastal restoration. The ecological role of shoals remains poorly understood and we hypothesized that shoals provide critical habitat for benthic invertebrates. Using Ship Shoal as a model system, we assessed the diversity and structure of macrobenthic assemblages and how community structure varies with season and environmental parameters. High biomass (averaging 26.7 g m⁻²) and high diversity (161 species) of macrobenthos was found in 2006. Polychaetes (45%—72 species) and crustaceans comprised most of the species (28%—46 species); spionids and amphipods dominated the polychaete and crustacean groups respectively, both in terms of number of species and abundances. Sharp decreases in diversity, abundance and biomass occurred from spring to autumn. Species diversity and total abundance significantly increased with decreasing sediment grain size and increasing bottom water dissolved oxygen. Across seasons, mole crabs Albunea paretii and amphioxus Branchiostoma floridae typified the community and contributed most of the biomass. The polychaetes Nephtys simoni, Neanthes micromma, Dispio uncinata, Mediomastus californiensis and Magelona sp. A, the amphipod Acanthohautorius sp. A and the burrowing shrimp Ogyrides alphaerostris also contributed to variation in community composition. Cluster analyses quantified seasonal variation, mainly based on sharp decreases in abundance, as well as spatial differences in species composition oriented along both east–west and north–south gradients. Variation in benthic assemblages was correlated with water depth and sediment characteristics (mean grain size and percentage of gravel-sized shell debris). We conclude that Ship Shoal is an unrecognized biodiversity hotspot and a hypoxia refuge compared to the immediate surrounding area where the benthic community is affected by seasonal hypoxia events and we discuss how sand-mining may influence this community.     

Effect of two microalgae concentrations on body size and egg size of the rotifer <Emphasis Type="Italic">Brachionus calyciflorus</Emphasis>

The rotifer, Brachionus calyciflorus, was grown with two algae species (Chlorella sp. and Scenedesmus obliquus) at different concentrations (0.1, 1 and 10 × 10⁶ cells ml⁻¹). The body size (lorica biovolume) of individual rotifer and their egg size were measured when the populations were roughly in the exponential phase of population growth. The body size of the rotifers differed significantly (P < 0.05) among the two algae species used, however this effect was not observed for egg size. The body size of rotifers fed on higher densities of Chlorella sp. (10 × 10⁶ cells ml⁻¹) was significantly larger than for those fed on lower and medium densities (0.1 and 1 × 10⁶ cells ml⁻¹). Body size and egg size of rotifers fed with different amounts of Scenedesmus did not differ significantly. The egg size was significantly larger at higher food level of Chlorella. A significantly positive correlation was observed between the adult rotifer body size and their egg size.     

Water Relations Only Partly Explain the Distributions of Three Perennial Plant Species in a Semi-arid Environment

The water relations and stomatal conductances of three perennial plant species, Stipa tenacissima L., Anthyllis cytisoides L., and Retama sphaerocarpa (L.) Boiss., dominant on the upper slopes, mid-slopes and floor of a valley, respectively, in semi-arid south-east Spain, were investigated to test the hypothesis that differences in plant-soil water relations could account for the different distributions of each species in the catena. Diurnal measurements of water potential (Ψ_w), relative water content (RWC) and stomatal conductance (g_s) of leaves were made over one year. Leaf temperature, air humidity, wind-speed and incident quantum flux density were measured concurrently. Soil water content was determined gravimetrically at 0 – 5 cm and 15 – 20 cm depths. Measurements of Ψ_w, RWC and g_s were analysed according to meteorological conditions, based on the maxima for daily air temperature and atmospheric saturation water vapour deficit and on soil moisture content. The hypothesis that plant-soil water relations can explain the distribution of the three species along the catena from valley side to valley floor was rejected for Anthyllis and Stipa but confirmed for Retama. This revised version was published online in July 2006 with corrections to the Cover Date.     

Community structure and demography of small-mammal populations in insular montane forests in southern India

bstract Small mammals were studied in insular montane forest patches in the Upper Nilgiris in southern India from February 1994 to September 1996. Nine patches were selected at two sites, one with a single large 600-ha patch, the other with several small patches ranging in size from 0.2 to 60 ha. The population characteristics and community structure of small mammals were studied in relation to patch size and habitat structure within the patches. The two most abundant species were Rattus rattus (2–36 individuals/ha) and Suncus montanus (0–11 individuals/ha), while the abundance of seven other species recorded were very low. The population characteristics of R. rattus examined were density, biomass, proportions of juveniles, sub-adults and adults, mean weight of all animals, mean weight of adults, sex ratio and persistence. The proportion of adults, sub-adults, mean weight of animals, and mean weight of adult males were correlated with patch size. Persistence of sub-adult females in the large patch was especially high. The density and biomass of other small-mammal species were also studied. Two indices of diversity, species richness and proportion of R.␣rattus were compared as measures of community structure. Seven habitat characteristics were measured; of these, canopy cover, canopy height and tree density were correlated with the size of the patch. Density and biomass of species other than R. rattus and proportion of R. rattus were correlated with canopy height. Density and biomass of species other than R. rattus were highest in smaller patches. While the population characteristics of R. rattus may be affected by patch size, the density of rare species may be influenced by factors related to lower canopy height. Migration between patches may be an important factor in maintaining populations in these patches. Received: 5 January 1998 / Accepted: 23 March 1998