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1.
N. A. O'Connor 《Oecologia》1991,85(4):504-512
Summary Woody debris is a major structural component of south-eastern Australian lowland streams, and the decayed wood substrates provide a structurally complex habitat for macroinvertebrate colonization. I tested for the presence of a species richness-habitat complexity relationship for macroinvertebrate species inhabiting the surfaces of decayed submerged logs (snags) in a lowland stream in northern Victoria. The species-habitat complexity relationship is defined as the increase in species richness due to increased structural complexity of a habitat when area is held constant. The response of macroinvertebrates to seven treatments of artificial and natural substrates of differing levels and types of structural complexity were examined using cluster analyses and MANOVAs. These analyses revealed a significant species-habitat complexity relationship. In addition, a comparison of species evenness between simple and complex habitats supported the hypothesis that more complex habitats contained more species because they possessed more resources. Analysis of species richness, though informative, masked the complexity of species responses revealed by multivariate analyses of species abundances. These analyses showed that different species groups selected different microhabitats on snags, particularly in response to the level of sediment deposition, which was greater on more structurally complex snags. In comparison with the benthos, snags were significantly richer in species abundances, possibly related to low levels of dissolved oxygen in benthic habitats.  相似文献   

2.
Disturbance frequency, intensity, and areal extent may influence the effects of disturbance on biological communities. Furthermore, these three factors may have interacting effects on biological diversity. We manipulated the frequency, intensity, and area of disturbance in a full-factorial design on artificial substrates and measured responses of benthic macroinvertebrates in a northern Vermont stream. Macroinvertebrate abundance was lower in all disturbance treatments than in the undisturbed control. As in most other studies in streams, species density (number of species/sample) was lower in disturbed treatments than in undisturbed controls. However, species density is very sensitive to total abundance of a sample, which is usually reduced by disturbance. We used a rarefaction method to compare species richness based on an equivalent number of individuals. In rarefied samples, species richness was higher in all eight disturbed treatments than in the undisturbed control, with significant increases in species richness for larger areas and greater intensities of disturbance. Increases in species richness in response to disturbance were consistent within patches, among patches with similar disturbance histories, and among patches with differing disturbance histories. These results provide some support for Huston’s dynamic-equilibrium model but do not support the intermediate-disturbance hypothesis. Our analyses demonstrate that species richness and species density can generate opposite patterns of community response to disturbance. The interplay of abundance, species richness, and species density has been neglected in previous tests of disturbance models. Received: 20 July 1999 / Accepted: 26 January 2000  相似文献   

3.
1. Theory predicts that the stability of a community should increase with diversity. However, despite increasing interest in the topic, most studies have focused on aggregate community properties (e.g. biomass, productivity) in small‐scale experiments, while studies using observational field data on realistic scales to examine the relationship between diversity and compositional stability are surprisingly rare. 2. We examined the diversity–stability relationship of stream invertebrate communities based on a 4‐year data set from boreal headwater streams, using among‐year similarity in community composition (Bray–Curtis coefficient) as our measure of compositional stability. We related stability to species richness and key environmental factors that may affect the diversity–stability relationship (stream size, habitat complexity, productivity and flow variability) using simple and partial regressions. 3. In simple regressions, compositional stability was positively related to species richness, stream size, productivity and habitat complexity, but only species richness and habitat complexity were significantly related to stability in partial regressions. There was, however, a strong relationship between species richness and abundance. When abundance was controlled for through re‐sampling, stability was unrelated to species richness, indicating that sampling effects were the predominant mechanism producing the positive stability–diversity relationship. By contrast, the relationship between stability and habitat complexity (macrophyte cover) became even stronger when the influence of community abundance was controlled for. Habitat complexity is thus a key factor enhancing community stability in headwater streams.  相似文献   

4.
Dam removal is an approach for restoring rivers. However, there are increasing concerns about the impact of removal on downstream biota. We examined the short-term responses of benthic macroinvertebrates and their avian predator (Brown Dipper, Cinclus pallasii Temminck) in reaches downstream of a check dam after it was removed from a mountain stream in central Taiwan. The density and taxonomic richness of downstream macroinvertebrates decreased immediately after dam removal. The decreases were associated with scouring or burial by sediments from the upstream impoundment. Ten weeks post-removal, downstream macroinvertebrate densities, although marginally recovering, remained lower than both pre-removal and upstream densities. Substantial changes in community structure were not significantly associated with an increase in the proportion of taxa with short life spans. However, this small-scale disturbance had no strong effect on the abundance of their very mobile, avian predator. This study and other studies of dam removal have found that downstream sedimentation following dam removal can reduce macroinvertebrate densities and that they may recover over time. Thus, timescale must be considered when interpreting the consequences of dam removal, especially when the long-term goal is stream restoration.  相似文献   

5.
Benthic invertebrates from River Nyamweru, a tropical forest stream in western Uganda were sampled bimonthly between April and December 1997 using a modified Hess Sampler. A total of 3708 benthic fauna from thirteen taxonomic orders were collected. Benthic samples were dominated by Diptera (mainly Chironomidae) representing over 60% of all the organisms, followed by Ephemeroptera. Benthic invertebrate densities ranged from 63 ± 9.03 organisms/m2 to 300 ± 33.36 organisms/m2, with higher densities occurring during the dry season and lower densities during the wet season. The benthic community structure in River Nyamweru reflected mainly collectors and scrapers as the most important groups (83%), while predators were very rare (3.1%). River discharge influenced benthic abundance, with more invertebrates at lower discharge and fewer invertebrates at higher discharge. The applicability of the River Continuum Concept to tropical forest stream situation is discussed.  相似文献   

6.
1. The composition and spatiotemporal dynamics of biological communities are influenced by biotic processes, such as predation and competition, but also by physical disturbances, such as floods in running waters. However, the interplay of disturbance with predation is still poorly understood, especially in frequently disturbed streams. Further, different predator species can affect prey communities in different ways depending on their feeding mode and efficiency. 2. We investigated the individual and combined effects of flood‐induced bed disturbance and fish predation on the benthos for 4 weeks in 18 streamside channels fed by a flood‐prone New Zealand river. Bed movements caused by floods were simulated by tumbling the substratum in half the channels. Six channels each were stocked with introduced brown trout (Salmo trutta) or native upland bully (Gobiomorphus breviceps) or had fish excluded. We studied algal biomass and both invertebrate density and daytime activity on surface stones on several dates after the disturbance, invertebrate community composition in the substrata of the entire channels on day 28 and leaf decomposition rates over the 28‐day period. 3. Disturbance affected algal biomass and density, richness and activity of surface stone invertebrates, and overall density and richness of channel invertebrates. Presence or absence of fish, by contrast, did not influence overall invertebrate standing stocks when subsurface substrata were included but did affect invertebrate densities on surface stones in 45% of all analysed cases and invertebrate activity on surface stones in all cases. Leaf decomposition rates were not influenced at all by the experimental manipulations. 4. Native upland bullies featured more often than exotic brown trout in causing invertebrate density changes and equally often in causing changes to grazer behaviour. Overall, our results imply that fish predation can have strong effects on the benthic invertebrate community in frequently disturbed streams, especially via behavioural changes.  相似文献   

7.
Two experiments were performed in an upland stream to determine the effects of the frequency of physical disturbance on the relationship between an abundant glossosomatid trichopteran (Agapetus monticolus; Banks) and the epilithon upon which it feeds. Artificial cobbles with an established epilithic community were tumbled either every 1, 2 or 4 weeks. The first experiment failed to detect any significant effects of rock tumbling on the abundance of A. monticolus or the epilithon: a result due to several spates. The first experiment did reveal that disturbances may disrupt the ability of A. monticolus to locate patches of abundant food. The second experiment found that although the abundance of A. monticolus was not affected by the disturbances, periphyton abundance was significantly reduced. Increasing the frequency of disturbance did not magnify this effect. Comparisons of these results with other studies of disturbance in streams indicate that the effects of disturbance on herbivory may be highly variable. A variety of factors, such as the relative resistances of the herbivores and the epilithon, need to be examined before the effects of disturbances on lotic herbivorous interactions can be completely understood.  相似文献   

8.
In order to investigate any size-dependent differences between behavioural patterns, wild-caught Hart's rivulus Rivulus hartii of varying sizes were exposed to chemical alarm cues extracted from the skin of conspecifics or heterospecific Poecilia reticulata, or a tank water control, in a series of laboratory trials. In response to conspecific alarm cues, R. hartii subjects of the range of body sizes tested exhibited consistent, size-independent antipredator behaviours that were characterized by decreased locomotory activity and foraging levels and increased refuging behaviour. Conversely, focal R. hartii demonstrated significant size-dependent trends in response to heterospecific alarm cues, with smaller individuals exhibiting antipredator responses and larger individuals shifting their behaviour to increased levels of activity consistent with a foraging, or predatory, response. These results show that the behavioural responses of individual R. hartii to publicly available chemical alarm cues from heterospecifics are mediated by the size of the receiver.  相似文献   

9.
Forested headwater streams rely on their riparian areas for temperature regulation, woody debris inputs, and sediment retention. These products and services may be altered by disturbances such as timber harvest, windthrow, or development. This study investigated the effects of riparian forest disturbance by removing trees using 50 and 90% basal area harvests and by directly felling some trees into eight streams in eastern West Virginia. On summer afternoons, water temperature increased in the 50 and 90% BAH treatments at average rates of 0.18 and 0.79°C/100 m, respectively. The 90% BAH treatments had the potential to disrupt fish and invertebrate communities via increased water temperature. New roads and log landings associated with the riparian logging had no detectable effect on sedimentation or turbidity. Large woody debris (LWD) additions increased habitat complexity but no net increase in pool area was observed. Greater morphological instability was observed within the LWD addition sections as pools were both created and destroyed at significantly higher rates. Experimentally manipulating small riparian patches may be an analog for small-scale natural and anthropogenic disturbances. These common events are assumed to alter streams, but there are few experimental studies quantifying their effects.  相似文献   

10.
1. Temperature and oxygen are recognised as the main drivers of altitudinal limits of species distributions. However, the two factors are linked, and both decrease with altitude, why their effects are difficult to disentangle. 2. This was experimentally addressed using aquatic macroinvertebrates; larvae of Andesiops (Ephemeroptera), Claudioperla, (Plecoptera), Scirtes (Coleoptera) and Anomalocosmoecus (Trichoptera), and the amphipod Hyalella in an Ecuadorian glacier‐fed stream (4100–4500 m a.s.l.). The following were performed: (i) quantitative benthic sampling at three sites to determine altitudinal patterns in population densities, (ii) transplants of the five taxa upstream of their natural altitudinal limit to test the short‐term (14 days) effect on survival, and (iii) in situ experiments of locomotory activity as a proxy for animal response to relatively small differences in temperature (5 °C vs. 10 °C) and oxygen saturation (55% vs. 62%). 3. The transplant experiment reduced survival to a varying degree among taxa, but Claudioperla survived well at a site where it did not naturally occur. In the in situ experiment, Scirtes and Hyalella decreased their activity at lower oxygen saturation, whereas Andesiops and Anomalocosmoecus did so at a low temperature. The decrease in activity from a high to a low temperature and oxygen for the five taxa was significantly correlated with their mortality in the transplant experiment. 4. Together the present experiments indicate that even relatively small differences in temperature and oxygen may produce effects explaining ecological patterns, and depending on the taxon, either water temperature or oxygen saturation, without clear interacting effects, are important drivers of altitudinal limits.  相似文献   

11.
Climate change effects on some ecosystems are still poorly known, particularly where they interact with other climatic phenomena or stressors. We used data spanning 25 years (1981–2005) from temperate headwaters at Llyn Brianne (UK) to test three hypotheses: (1) stream macroinvertebrates vary with winter climate; (2) ecological effects attributable to directional climate change and the North Atlantic Oscillation (NAO) are distinguishable and (3) climatic effects on macroinvertebrates depend on whether streams are impacted by acidification. Positive (i.e. warmer, wetter) NAO phases were accompanied by reduced interannual stability (=similarity) in macroinvertebrate assemblage in all streams, but associated variations in composition occurred only in acid moorland. The NAO and directional climate change together explained 70% of interannual variation in temperature, but forest and moorland streams warmed respectively by 1.4 and 1.7°C (P<0.001) between 1981 and 2005 after accounting for NAO effects. Significant responses among macroinvertebrates were confined to circumneutral streams, where future thermal projections (+1, +2, +3°C) suggested considerable change. Spring macroinvertebrate abundance might decline by 21% for every 1°C rise. Although many core species could persist if temperature gain reached 3°C, 4–10 mostly scarce taxa (5–12% of the species pool) would risk local extinction. Temperature increase in Wales approaches this magnitude by the 2050s under the Hadley HadCM3 scenarios. These results support all three hypotheses and illustrate how headwater stream ecosystems are sensitive to climate change. Altered composition and abundance could affect conservation and ecological function, with the NAO compounding climate change effects during positive phases. We suggest that acidification, in impacted streams, overrides climatic effects on macroinvertebrates by simplifying assemblages and reducing richness. Climatic processes might, nevertheless, exacerbate acidification or offset biological recovery.  相似文献   

12.
Scaling the effects of predation and disturbance in a patchy environment   总被引:3,自引:0,他引:3  
Jill Lancaster 《Oecologia》1996,107(3):321-331
The effects of hydraulic disturbances on the impact of two predatory benthic invertebrates on their prey were examined in a stream at two distinct spatial scales. At the scale of small habitat patches (0.0625 m2), hydraulic patch type was an important determinant of the microdistribution of prey and predators. Prey abundances were similar across all patch types at baseflow, but local densities were higher in patches identified as low-flow refugia after periods of high and fluctuating flow. The microdistribution pattern of predatory larvae of a caddisfly, Plectrocnemia conspersa, was similar to that of its prey, whereas predatory larvae of an alderfly, Sialis fuliginosa, did not shift their microdistribution significantly with discharge and were always most abundant in lowflow refugia. There was little evidence of an aggregative response of predators with prey, even though both predators and prey are mobile. Both predator species showed similar patch-specific patterns of per capita consumption rates: uniform consumption rates across hydraulic patch types at low and moderate flows, but highest in flow refugia during high flows. Species-specific patterns, however, were apparent in the magnitude and direction of differences between consumption rates during disturbance events, and in comparable patches at base flow: At high flow, consumption rates for P. conspersa were exaggerated (3.9 times higher) in flow refugia but at par in other patches; for S. fuliginosa they were at par in flow refugia but reduced in other patches (up to 3.3. times lower). These differences may be related to species-specific foraging behaviours (search vs ambush predators) and the influence of prey movements on feeding success. Using the patch-scale results only, it is difficult to predict the effects of physical disturbance on predation intensity at the larger scales of whole habitats, populations or communities. At the large scale (>200 m2), net predator impacts were estimated over the stream reach, using a spatially explicit model that accounts, in an additive way, for habitat heterogeneity and patch-specific responses of predators and prey. The relationship between predator impact over the whole reach and hydraulic disturbance differed for the two predators. The predator impact of S. fuliginosa decreased with increasing hydraulic disturbance, as predicted by the harsh-benign hypothesis. There was no directional trend for P. conspersa, however, and maximum predator impact may occur at intermediate disturbance levels. For the prey community in this stream, predation pressure from S. fuliginosa appears to fluctuate directly with the discharge hydrograph, whereas predation from P. conspersa may be more persistent. Flow refugia may play a dual role in the sructure of stream communities by preventing catastrophic mortality of animals (predators and prey) from physical forces during disturbances, and by maintaining (or perhaps increasing) predation pressure. Summing the effects of species interactions in small habitat patches to the larger scale of a whole stream reach indicates that the scale of approach influences the observed patterns and their implied underlying process.  相似文献   

13.
Mutuku Mathooko  Jude 《Hydrobiologia》1997,362(1-3):211-218
Field experiments to examine the effect of continuous physical disturbance on the Ephemeroptera of the Naro Moru River, Kenya, wereundertaken from June 1993 to January 1994. Continuous disturbance wasadministered on a randomly selected subsite of the sediment surface.Artificial physical disturbance within the experimental subsite involvedcontinuous local displacement, shifting and stirring of the streambedsurface substrates (about 10 cm depth) by hand every one minute for 10 or 14min. Three control samples were also taken randomly from the sedimentsurface of an undisturbed stratified area of the study riffle at the startof each disturbance occasion. All samples were collected using a Hesssampler (surface sampling area of 3.142 dm2; meshsize 80μm). Seven mayfly species were particularly abundant and these included Afronurus sp., Afroptilum sudafricanum LESTAGE, Baetis s.l., Baetis(Nigrobaetis) sp. 1, Baetis (Nigrobaetis) sp. 2, Caenis sp. and Choroterpes(Euthraulus) sp. About 83,8% of the total mayfly density and88.1% of the biomass were removed from the streambed surface withinthe first three minutes of continuous physical disturbance. A mayfly biomassof 33.7391 mg dm2 and total density of 1357.6 inddm2 were collected from the disturbed subsite during the studyduration. Further, a biomass of about 42.8335 mg dm−2and total density of 2366 ind dm−2 were collected fromthe control sites. There was a near-complete depletion of mayflyindividuals from the topmost sediment layer within 14 min of continuousdisturbance. This revised version was published online in July 2006 with corrections to the Cover Date.  相似文献   

14.
1. Aquatic communities are structured by multiple forces, and identifying the driving factors over multispatial scales is an important research issue. The East Asian monsoon region is globally one of the richest environments in terms of biodiversity, and is undergoing rapid human development, yet the river ecosystems in this region have not been well studied. We applied a hierarchical framework to incorporate regional and local environmental effects on stream macroinvertebrate communities in this region. The knowledge gained is expected to improve the understanding of the importance of spatial scale on regional and local diversity in the East Asian monsoon region. 2. A national data set of benthic macroinvertebrates and environmental variables (geographical, land‐use, hydrological, substratum and physicochemical elements) in Korean rivers was used to determine the habitat preferences of macroinvertebrates. 3. Latitude, proportion of forest coverage, riffle habitat, silt substratum and temperature were the most important determinants for the ordinations of macroinvertebrate communities in each category evaluated by canonical correspondence analysis (CCA). The optimal habitats for stream macroinvertebrates are not the same for all species, and overall community metrics and abundance of sensitive species tended to be lower in open agricultural and urban streams than in forested streams. The sensitivity of mayflies and stoneflies to anthropogenic disturbances implicated them as good indicators to assess the effects of urban and agricultural activities. 4. A partial CCA was used to evaluate the relative importance of macrohabitat and microhabitat variables on community composition at three spatial scales (whole country, the large Han River basin and two small sub‐basins in the lowlands and highlands). The majority of community variation (17–22% for each environmental element) was explained by macrohabitat variables at the regional spatial scale. In contrast, large proportions (15–18%) were explained by microhabitat variables at the local spatial scale. 5. Our findings indicate that the relative importance of habitat scales should be determined by geographical size and that comprehensive understanding of multispatial scale patterns can be important for implementing sound biodiversity conservation programmes.  相似文献   

15.
1. Few studies have assessed the effects of macroconsumers, such as fishes and shrimps, on detritus and detritivores.
2. We used an underwater electric field to prevent macroconsumers from feeding in and on leaf packs in a lowland stream in Costa Rica and thus to determine their effects on the density of insect detritivores and decay rates of leaves.
3. Exclusion of macroconsumers resulted in significantly higher densities of small invertebrates inhabiting leaf packs. Most of these were collector–gatherers, none were shredders.
4. Despite the increase in invertebrate density, decay rates of leaves were not statistically different. These findings contrast with results from temperate streams showing that increases in the density of invertebrates in leaf packs typically result in an increased rate of decay.
5. Leaf decay rates and invertebrate densities were also compared between leaf packs placed in electric exclusion treatments and those placed in coarse (2 cm) plastic net bags (as used in many previous studies). Our results suggest that using such netting in tropical streams may deter macroconsumers, which can affect insect density and, potentially, decay rates of organic matter.  相似文献   

16.
Summary The role of predator density and plant detritus in determining patterns of insect abundance was tested in an Appalachian stream community (Reeds Creek, Pendleton Co, West Virginia). Insect colonization was followed over a 21-day period in field enclosures containing different densities of sculpins (Cottus bairdi and Cottus girardi). Sculpins caused no significant reductions in prey abundance; however, the combined effect of a guild of vertebrate predators caused a significant depression in the abundance of Chironomidae and the stonefly Leuctra. Since Chironomidae comprised approximately 85% of the total benthic fauna, vertebrate predation had an important role in the overall structure of the invertebrate community.Total insect abundance and diversity were correlated to the presence of plant detritus. However, this relationship was taxon specific, as abundance was highly correlated to detritus for only selected insect taxa. In general, abundance was best correlated with coarse particulate organic matter (CPOM), compared to fine particulate organic matter (FPOM) or total detritus (FPOM+CPOM+Whole leaf organic matter (LVOM)).This experiment illustrates that the influence of various potential structuring mechanisms is highly taxon dependent.  相似文献   

17.
SUMMARY. 1. Predation upon macroinvertebrates by the loach Oreonectes platycephalus Günther (Cobitidae) was studied using predator inclusion/exclusion cages in a series of pools along a Hong Kong stream. Treatments employed were predator exclusion, medium (approximately natural) predator densities (1 fish cage−1) and high predator densities (2 fish cage−1). Macroinvertebrate abundance in cages was monitored after 2 and 4-weeks exposure to predators.
2. The presence of fish was associated with significant declines in the total numbers of macroinvertebrates colonizing cages. However, taxa were influenced differently, with mayflies decreasing by a factor of two while the more mobile shrimps (Atyidae) were unaffected. Chironomid abundance (largely Chironominae) was unaffected by predator density and increased in week 4. Detritus acted as a confounding variable at this time because chironomid abundance was significantly correlated with the weight of accumulated detritus in cages.
3. While invertebrates were more abundant in cages lacking fish, there were no fewer invertebrates in cages with 2 fish than with 1 fish. This may indicate the presence of secure refuges among substrates in the cages, preventing the additional fish from depleting prey further, or a lack of precision of methods due to natural variations in prey densities and spatial patchiness.
4. No significant effects of predators on relative prey abundance or species richness were detected.
5. The impact of predation on prey abundance weakened on week 4, perhaps due to extra refuges among the accumulated detritus. However, drying of the stream increased fish densities in pools so that cages may have become zones of relative safety that were colonized readily by macroinvertebrates. This result highlights the need for year-round investigations to quantify predation effects in Hong Kong's seasonal tropical climate.  相似文献   

18.
1. The relationships between biological traits of macroinvertebrates and environmental characteristics were investigated in streams with contrasting physical, chemical or landscape level attributes. We used an ordination technique, RLQ analysis, which links an environmental table (R) with traits table (Q) through an abundance table (L) to investigate the relationship between habitat characteristics and biological traits.
2. A major environmental axis explaining the distribution of species and their distinctive biological features was obtained. This axis included variables of anthropogenic pressure (agricultural and urban uses) and natural variability (climatic and geologic) that are strongly intercorrelated in the study area, with a clear spatial component.
3. The attributes of species from frequently disturbed systems (small size, multivoltinism, diapause, ovoviviparity, etc.) were associated with semi-arid areas whereas traits common in more stable and favourable environments (large body size, semi-voltinism, isolated eggs, etc.) were found in upland forested areas.
4. The natural climatic variation was proposed as a disturbance axis of a theoretical habitat templet (driven by the intense hydrological disturbances typical of semi-arid streams), while anthropogenic pressure (mainly intensive agriculture) and high salinity, a natural consequence of geology, was proposed as an adversity axis. Different life-histories associated with contrasting environmental features were superimposed in this habitat templet.
5. The ecological–evolutionary scenario in which stream macroinvertebrates have evolved and by which their communities are organized, is closely linked to disturbance, environmental harshness and human pressure.  相似文献   

19.
20.
The effects of forest disturbance on diversity of tropical soil nematodes   总被引:8,自引:0,他引:8  
We provide the first account of the effects of forest disturbance on species richness of nematodes in tropical forest soils, from 24 sites along gradients of disturbance and regeneration in the Mbalmayo Forest Reserve, Cameroon. Species richness was very high. Samples of 200 nematodes from individual soil cores contained a maximum of 89 and an average of 61 species; in total we recorded 431 species and approximately 194 genera. The model of Siemann et al. (1996), predicting that species richness scales as the number of individuals I 0.5, underestimates nematode diversity 4–6 fold in these samples. Over 90% of specimens cannot be assigned to known species. Although nematode species richness declined with forest disturbance, statistically significant effects were detectable only under the most extreme conditions (active slash-and-burn agriculture and complete mechanical forest clearance) and even here remained at 40% of the richness of near primary sites. Impacts on trophic structure were also small, and there were no significant changes in the maturity index (MI) (Bongers 1990) with disturbance (mean MI across all treatments was very high, at 3.58). In the light of this study, the problems of completing reliable all-taxon inventories in tropical forests are briefly discussed. Received: 22 July 1996 / Accepted: 3 April 1997  相似文献   

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