Longitudinal patterns of plant diversity in the North American boreal forest

Spatial patterns of plant diversity in the North American boreal forest were examined according to three plant life forms (woody plants, herbaceous plants, and bryophytes) and two taxonomic levels (species and genus), using sixty 9-ha plots sampled in white spruce (Picea glauca (Moench) Voss) and black spruce (Picea mariana (P. Mill.) B.S.P.) ecosystems along a transcontinental transect from the Pacific coast eastwards to the Atlantic coast. The patterns of inventory diversity (represented by alpha diversity), differentiation diversity (represented by the similarity index, habitat-heterogeneity index, similarity decay rate, and length of the first axis in detrended correspondence analysis), and pattern diversity (represented by the mosaic diversity index) were assessed along the transect in both ecosystem types. At the stand level, central North America had the highest alpha diversity in terms of the number of species or genera, and western North America had a higher alpha diversity than eastern North America. At the continental scale, herbaceous plants had the highest beta diversity in terms of floristic change from the eastern to western North America, bryophytes had the lowest beta diversity, and woody plants were in the middle, regardless of ecosystem type and taxonomic level. Central North America had the lowest mosaic diversity across the boreal transect of North America. The white spruce ecosystems had a higher alpha diversity than the black spruce ecosystems regardless of plant life form, taxonomic level and geographic location. The white spruce ecosystems tended to have more bryophytes, less woody plants, and higher species:genus ratio than the black spruce ecosystems. In general, the white spruce and black spruce ecosystems shared the same patterns in diversity changes at different spatial scales, plant life forms, and taxonomic levels across the transect studied. The existing patterns of plant diversity in the North American boreal forest area resulted from a combination of ecological processes and spatial configuration.     

Composition and diversity of the spider fauna in the canopy of a montane forest in Tanzania

Spiders were sampled using insecticide knockdown in an African montane forest in the Uzungwa Mountains of Tanzania. The results are used to discuss the faunal composition at the site and in comparison to other sites, and the implications of the results for estimating spider diversity in Africa are discussed. A total of 5233 adults comprising 149 species were collected from 11 samples covering a total of 906 m² of projected area. Three species contributed 45% of the sample. Previous insecticide knockdown studies of tropical lowland forest canopies have shown a dominance of Theridiidae, Salticidae and Araneidae. In the present study Linyphiidae dominated in abundance and were the second most diverse in terms of species richness. Other abundant families were Oonopidae and Pholcidae, while Theridiidae, Salticidae and Araneidae were rich in species. This supports a previous study, which indicated that the importance of linyphiids increases with altitude. Species richness was predicted using a number of estimators, which produced relatively similar results. Using the abundance-based estimator, Chao 1, the predicted richness for the total area sampled is 183 ± 15 species. This indicates that at least 20% of the area's spider community remains unsampled. A high ratio of undescribed species (approximately 80%) and a relatively high species turnover compared to a site 20 km away within the same forest complex suggests that the number of spiders in Africa could well be much higher than the current, published estimate of 20000 species.     

Understorey vegetation in boreal Picea mariana and Populus tremuloides stands in British Columbia

Abstract. We compared the species composition and species density of vascular plants in the understorey vegetation of boreal forest between Picea mariana (Black spruce) and Populus tremuloides (Trembling aspen) stands in British Columbia, Canada, and related differences in species composition and species density between the two forest types to dominant canopy tree species as well as a wide variety of environmental factors. We analysed 231 stands, distributed in three different climatic regions representing drier, wetter, and milder variations of montane boreal climate. Of these stands 118 were dominated by P. mariana and 113 by P. tremuloides. P. tremuloides stands had higher species density than P. mariana stands in all climatic regions, but species density of each dominance type varied among climatic regions. The floristic composition of the understorey vegetation was markedly different for P. mariana and P. tremuloides dominated stands. A detailed study on the effect of canopy dominance and local environmental factors on the understorey vegetation of the boreal forest was conducted using 88 stands from one of the three climatic regions. Using a combination of ordination and variation partitioning by constrained ordination we demonstrated a small but unique effect of canopy dominance type on the understorey vegetation, while a larger amount of compositional variation was shared with other factors. Our results accord with a scenario in which differences in primary environmental factors and humus form properties, the latter accentuated by the canopy dominants themselves, are the most important causes of higher species density in P. tremuloides stands than in P. mariana stands, as well as differences in species composition among the two canopy dominance types. Processes and time scales involved in the small but significant direct and indirect effects of the canopy dominant on understo‐ rey species composition are discussed.     

Diversity of the understory vascular vegetation in 40 year-old and old-growth forest stands on Vancouver Island,British Columbia

Abstract. We studied plant diversity of the understory vascular vegetation in 40 yr-old plantations (immature stands) and old-growth forest stands on southwestern Vancouver Island, British Columbia, Canada. Site-specific comparisons using several indices of species diversity were made between: (1) immature stands segregated according to the canopy cover and dominant canopy tree species; and (2) immature and old-growth stands. There were no significant differences (P < 0.05) among immature stands in species richness (S) and the Shannon-Wiener index (H′), in relation to the canopy cover or in S, H′ and evenness (E) in relation to the dominant canopy tree species. Using the same indices, the plant diversity varied with edaphic conditions (represented by five site associations) and time (represented by two developmental stages). At both stand- and site levels, plant diversity increased with increasing soil moisture, from slightly dry to moist sites, and with increasing plant-available soil nitrogen in both immature and old-growth stands; and the plant diversity of immature stands across the sites studied was considerably lower than in old-growth stands, regardless of site association. The indices of plant diversity, floristic similarity indices, and species turnover rates indicated that the immature stands had their plant diversity at a minimum, but a drastic loss of diversity expected in the stem exclusion stage had not materialized. We attributed decline in plant diversity to the absence of old-growth structural features in immature stands. Several measures to foster the stand-level diversity were proposed.     

闽西北不同类型毛竹林养分分布及生物循环特征

以福建省永安市集约经营的毛竹纯林(Ⅰ)、竹阔混交林(Ⅱ)和竹针混交林(Ⅲ)3种类型林分为研究对象,对其N、P、K、Ca、Mg元素分布格局及生物循环特征进行了研究。结果表明,毛竹林N、P、K、Ca、Mg5种养分元素的排列顺序为土壤层乔木层凋落物层灌木层草本层,其中土壤层(0～60cm)养分累积量占系统养分总量99%。3种林分养分元素累积量的排列顺序为ⅡⅠⅢ,分别为146196.59、144466.35和105002.59kg.hm-2,其中植被层养分累积量的排列顺序为ⅠⅡⅢ,凋落物层和土壤层养分累积量排列顺序为ⅡⅠⅢ,竹阔混交林土壤层和凋落物层养分累积量最高,立地生产潜力最大。林分Ⅰ和Ⅱ的养分利用系数显著高于林分Ⅲ,反映了林分维持高生产力需要消耗更多的养分元素;循环系数排列顺序为ⅢⅡⅠ,竹针混交林和竹阔混交林维持林木生长所需的养分元素少于毛竹纯林;毛竹纯林养分周转时间最长,竹阔混交林次之,竹针混交林最短,分别为31.89年、12.39年和12.23年,混交林周转时间显著低于毛竹纯林。竹阔混交林既有较高的生产力,又有较强的养分归还能力,是一种较好的经营方式。     

芦芽山寒温性针叶林冠层下植被beta多样性格局及其成因

群落构建机制是生态学的中心议题之一。对山地植被beta多样性格局及其成因的探究有助于加深对此问题的认识。以芦芽山寒温性针叶林群落冠层下植被为研究对象,结合野外调查与室内实验获取的详细数据,运用Mantel检验、普通最小二乘回归和典范对应分析(CCA)等统计方法,探讨了林下植被的beta多样性格局及其成因,结果显示:(1)沿海拔梯度相邻群落间草本层物种周转率呈现递减格局,而灌木层变化规律不明显;(2)灌、草层beta多样性与海拔差异、地理距离呈显著正相关关系,而与局地环境异质性关系不显著。控制海拔作用后发现,灌、草层beta多样性与地理距离关系依然显著,而当消除地理距离的线性影响后,beta多样性与海拔关系也变得不显著(3)CCA模型中,环境因子共解释了物种组成变异的74.4%,其中,海拔、坡度、凋落物厚度、乔木密度与总干面积对林下灌、草植被物种组成具有显著影响,但土壤因子的作用未见显著。综上,生境筛滤与扩散限制共同主导了芦芽山寒温性针叶林冠层下群落构建过程,但扩散限制的影响强于生境筛滤作用。     

A floristic-structural gradient related to land forms in the southern Chihuahuan Desert

Abstract. The purposes of this study were to elucidate the floristic and structural characteristics of the vegetationin the Mapimi subdivision of the Chihuahuan Desert, Mexico, and to relate them to environmental variation. The main question addressed was: How does floristic composition, total species richness and life-form species richness vary in relation to environmental change? 154 sites, randomly selected and stratified over seven landscape units, were analyzed. Results showed the existence of a land form gradient along which vegetation types were ordered. Species richness varied along the gradient, the richest land form was bajada, indicating that the maximum species richness did not occur at one extreme of the water availability gradient but in a moderately poor situation. The lowest species richness was found in the playa land form. Cover-based life form spectra varied significantly with land forms, while presence-absence based life form spectra did not. It is suggested that this may be a consequence of the relatively young age of this desert.     

Structure and dynamics of an ancient montane forest in coastal British Columbia

Old-growth forests are common in the snowy, montane environments of coastal western North America. To examine dynamics of a stand containing four canopy tree species (Abies amabilis, Chamaecyparis nootkatensis, Tsuga mertensiana and T. heterophylla), we used four stem-mapped, 50 m ×50 m plots. From measurements of annual rings, we obtained ages from basal discs of 1,336 live trees, developed master chronologies for each species, reconstructed early growth rates, and delineated periods of release. The stand was ancient; individuals of all four species exceeded 900 years in age, and the oldest tree exceeded 1,400 years. The four plots differed in the timing of events, and we found no evidence of major, stand-level disturbance. Instead the stand was structured by small-scale patch dynamics, resulting from events that affected one to several trees and initiated episodes of release and relatively rapid early growth. The species differed in age structure and dynamics. A. amabilis and T. heterophylla had a classical reverse-J age structure indicative of stable populations, whereas C. nootkatensis and T. mertensiana appeared to rely on local episodes of increased recruitment, which were often separated by centuries, and were probably related to multiple-tree gaps that occurred infrequently. However, such gaps could be considered normal in the long-term history of the stand, and thus these species with their long life spans can persist. Most individuals of all four species grew extremely slowly, with trees typically spending centuries in the understory before reaching the canopy, where they were able to persist for additional centuries. Thus, the key features of this forest are the very slow dynamics dominated by small-scale events, and the slow growth of stress-tolerant trees.     

Variability of plant species diversity in Swedish natural forest and its relation to atmospheric deposition

Within a subprogram of Integrated Monitoring (IM), understorey vegetation in Swedish natural forests was observed at fifteen reference sites over the country for twelve seasons, 1982–1993. The main task of the subprogram was to assess the impact of atmospheric deposition, mainly sulphur and nitrogen, on natural vegetation through time. The present study is focused on the variability of plant species diversity at community level and the possible impact of sulphur and nitrogen deposition. Species richness, evenness and diversity varied greatly among the sites, and between years within each site. Regarding only coniferous forests the species richness was higher in the north than in the south. But the effects of site condition and atmospheric deposition were not clarified. Changes in species diversity through time differed from site to site. No overall temporal trend was found. The atmospheric deposition of sulphur and nitrogen demonstrated a clear geographical pattern being low in the north-west and high in the south-west. Sulphur deposition declined significantly in Southern Sweden during the period. We concluded that the species diversity of understorey vegetation at the Swedish IM sites was not significantly influenced by atmospheric deposition. The changes observed are explained as natural processes.     

Tropical forest diversity,environmental change and species augmentation: After the intermediate disturbance hypothesis

Abstract. It is not simple to predict how environmental changes may impact tropical forest species diversity. Published hypotheses are almost invariably too incomplete, too poorly specified and too dependent upon unrealistic assumptions to be useful. Ecologists have sought theoretical simplicity, and while this has provided many elegant abstract concepts, it has hindered the attainment of more practical goals. The problem is not how to judge the individual hypotheses and arguments, but rather how to build upon and combine the many hard-won facts and principles into an integrated science. Controversy is inevitable when the assumptions, definitions and applications of a given hypothesis are unclear. Elegance, as an end in itself, has too often been used to justify abstract simplification and a lack of operational definition. Clarifying and combining hypotheses while avoiding assumptions provides a potentially more useful, if less elegant, standpoint. An appraisal of Connell's intermediate disturbance hypothesis, and its application to long-term observations from a Ugandan forest illustrates these concerns. Current emphases encourage ecologists to exclude consideration of environmental instability and non-pristine ecosystems. In reality, many environmental changes and ecological processes contribute to both the accumulation and erosion of diversity, at all spatial and temporal scales. Site histories, contexts, long-term processes, species-pool dynamics, and the role of people require greater emphasis. These considerations reveal that many environmental changes, even those associated with degradation, can lead to a transient rise in species densities. Drawing on related studies, such as forest yield prediction, suggests that the formulation and calibration of simulation models provides the most tractable means to address the complexity of real vegetation. Simulation-based approaches will become increasingly useful both in unifying the study of vegetation dynamics and in providing improved predictive capacity. Quantification of the processes, scales and sensitivities of the dynamics of tropical forest communities remains a major challenge.     

Phytoplankton diversity (alpha,beta, and gamma) from the Araguaia River tropical floodplain lakes (central Brazil)

Alpha, beta and gamma are three components of species diversity. Knowing these attributes in floodplain lake phytoplankton communities is vital when selecting conservation areas. Species diversity is commonly used with other taxonomic groups, but rarely with phytoplankton. We compared the number of phytoplankton species (alpha diversity) from 21 Middle Araguaia River floodplain lakes in the 2000 and 2001 rainy and dry seasons. From these samples we estimated complete survey species richness (gamma diversity), quantified differences in species composition between lakes (beta diversity) and assessed the influence of abiotic variables on beta diversity. We recorded a total of 577 taxa. The Sjack1 estimator indicated that 62.31% of taxa were sampled in the 2000 rainy and 67.65% dry seasons, and 68.36% in the 2001 rainy and 73.5% dry seasons. In almost all seasons, alpha diversity negatively correlated with latitude. Beta diversity (β-1) was higher in high water periods, especially in 2000. This may have been caused by isolated heavy rainfall, which would have increased environmental heterogeneity and raised beta diversity. DCA showed differences in phytoplankton composition between rainy and dry seasons in 2000 and 2001, reflecting the influence of flood pulses on phytoplankton composition. The Mantel test indicated spatial distribution patterns where geographically more distant lakes had less-similar phytoplankton communities. Handling editor: J. Padisak     

The diversity of 15 ha of lowland mixed dipterocarp forest, Central Kalimantan

Fifteen hectares of primary forest in Central Kalimantan were enumerated and all trees 10 cm dbh tagged and identified to species as far as possible. Tree density, relative abundance of families and species diversity were calculated, as was a mean additive species/area curve for 1 ha and a species/area curve for all 15 ha. The Jaccard index of similarity was also calculated for the 15 plots. The 15 ha enumerated contain 8771 trees with a dbh of 10 cm, belonging to 1298 morphospecies in 56 families. On average a single plot had 583 trees and 205 species. These figures are within the upper range of species diversity for the Malesian region as a whole. Dipterocarpaceae make up 14% of all trees in the plots, closely followed by Euphorbiaceae with 13%. A surprisingly low coefficient of similarity is found between the plots, the lowest being 3% and the highest 30%. Species/area curve construction showed that the asymptote is not reached in any of the 1 ha plots or when the 15 plots are added together.     

The effects of forest disturbance on diversity of tropical soil nematodes

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     

Changes in plant species diversity along a chronosequence of vegetation restoration in the humid evergreen broad-leaved forest in the Rainy Zone of West China

Plant species diversity has been recognized as one of the vital attributes for assessing vegetation restoration. Changes in the diversity may be related to different stages of succession. In this study, 54 sites of humid, evergreen, broad-leaved forest were selected in the Rainy Zone of West China. A chronosequence of the sites was used to study the successive patterns of the diversity in the forest that had undergone natural regeneration for 5 to 350 years and to test the hypothesis that the diversity is maximized in mid-succession. Data were collected simultaneously at different stages of succession, and four α-diversity indices (species richness, Margalef index, Shannon-Wiener index, Pielou Evenness index) and two β-diversity indices (Whittaker index, Sørensen’s index) were calculated for each stratum in each plot. A total of 394 vascular plant species were recorded. From the β-diversity indices, the forest succession may be divided into the early-successional stage (before 50 years), mid-successional stage (from 50 to 300 years), and late-successional stage (after 300 years). In this community, the species diversity and richness were found to be the greatest at the mid-successional stage, followed by the late- and early-successional stages. The results of regression analysis indicated that the richness and Margalef index peaked around the 175th and 165th year, respectively. Shannon-Wiener index values also appeared to follow an approximately humped pattern of succession and were maximal around the 100th year. However, the species evenness did not show any significant relationship with successional age. Our results demonstrate (1) forest restoration is a long-term process and the formation of climax forest requires at least 300 years and (2) the forest has a strong capacity for restoration. Our results also suggest Lindera limprichitii and Machilus pingii as ideal tree species for afforestation because of their wide niche.     

Patterns of floristic diversity in semi-natural coastal vegetation of Lebanon and implications for conservation

The current understanding of the status of the vegetation in Lebanon is largely derived from herbarium data and associated floristic studies produced by early 20th century field botanists. In common with other areas in the Mediterranean, the Lebanese coastline is highly threatened by unregulated development, yet current patterns of species richness along the Lebanese coastal zone are little studied. The objective of this study was to assess the floristic richness of the Lebanese coastal zone and to provide baseline information for conservation planning. For this purpose, permanent sample plots (6m×100m) were established in 26 selected vegetation communities in coastal habitats. Monthly field collections of plant specimens were undertaken between October 1999 and July 2000. A total of 441 species were collected and identified. None of the recorded species are currently considered globally threatened, but two are Lebanese endemics (Matthiola crassifolia Boiss. & Gaill., Origanum ehrenbergii Boiss.). Species richness varied between communities, ranging from six species in a littoral limestone pavement community to 113 in an abandoned terrace community. The similarity between communities, based on Sorensen indices, was low and a large number of species were recorded only once. Cluster analysis showed a grouping of different communities within locations in some instances and the clustering of similar community types regardless of location in others. Species richness in riparian and littoral communities consisted mostly of habitat non-specific species. The low community similarity, patchy species distribution, and predominance of habitat non-specific species all point to the need to complement in situ conservation measures with ex situ conservation.     

Tree species richness complementarity, disturbance and fragmentation in a Mexican tropical montane cloud forest

Tropical montane cloud forest exhibits great heterogeneity in speciescomposition and structure over short geographic distances. In central Veracruz,Mexico, the conservation priority of seven cloud forest fragments was assessedby considering differences in woody plant species richness and complementarityof species among sites, forest structure, tree mortality, and timber andfirewood extraction as indicators of anthropogenic disturbance. Densities oftrees 5 cm dbh (360–1700 trees/ha) weredifferent among the sites, but basal area (35.3–89.3m²/ha) did not differ among fragments. The number of dead trees rangedfrom 10–30 to 170–200 trees/ha. The fragments'species composition was different but complementary. The Morisita–Hornindex indicated low similarity between fragments. A non-parametric estimator ofspecies richness indicated that more sampling effort would be necessary tocomplete the inventory (15 additional trees and two understory shrub species).Unfortunately, most of the fragments are threatened with deforestation. The numberof cut trees was similar among sites (0–15 cut trees/0.1 ha).Sites with immediate need for conservation were close to settlements, with highnumbers of cut trees and no legal protection. The selected sites represent thevariety of situations that exist in the region. Given the high complementarityobserved between fragments, a regional conservation approach will be required topreserve the last repositories of part of the tremendous biodiversity of theonce continuous forest in this region.     

Species richness patterns in the understorey of Pyrenean Pinus sylvestris forest

Abstract. Species richness was studied in the understorey of natural Pinus sylvestris forest in the eastern Pyrenees. Understorey plant species were grouped in three structural groups as woody species, herbs and mosses. The response curves of total species richness and species richness of each structural group were fitted against environmental and stand-structural parameters, using Generalized Linear Models. The results suggested that, to predict species richness, environmental parameters were more important than tree-canopy structural parameters, in particular incoming radiation and soil nutrient concentration. The species richness response curve was often humped in relation to soil nutrient concentration. Different patterns of species richness were found for each structural group.     

Patterns in the structure of grassland butterfly communities along a gradient of human disturbance: further analysis based on the generalist/specialist concept

Kitahara and Fujii, in 1994, analyzed the butterfly communities along a gradient of human disturbance by applying the generalist/specialist concept. Butterfly species were classified into generalist or specialist species based on their voltinism (seasonal time dimension) and potential larval resource breadth (food dimension). The community structure and species composition showed the systematic changes along the gradient. To verify the generality of those trends, we monitored five grassland butterfly communities with varying degrees of human disturbance twice a month during 1985 by the line transect method at the foot of Mt. Fuji, central Japan, and analyzed their structure in a manner similar to that employed by Kitahara and Fujii. Most results were consistent with the patterns recognized by Kitahara and Fujii. The route (community) order based on increasing human disturbance was strongly and negatively correlated with butterfly species richness but with neither butterfly species diversity (H′) nor evenness (J′). Also, the degree of human disturbance was significantly and negatively correlated with the number of specialist species, but not with that of generalists, in a community. Butterfly species richness was more strongly correlated with the number of specialist species than with that of generalists. Our analyses also showed that the generalist species were distributed more widely over the communities than were the specialists. However, in contrast to the trend revealed by Kitahara and Fujii, there was no significant difference in the population densities and in the spatial population variability between the two species groups. As a whole, our analyses confirmed the consistency of most community patterns detected by Kitahara and Fujii. The causes of the inconsistencies in some patterns were thought to be mainly the present habitat conditions with a relatively short growing season at high altitudes. Received: October 19, 1999 / Accepted: June 5, 2000     

Patterns of β‐diversity in a Mexican tropical dry forest

Abstract. Patterns of β‐diversity in a highly diverse tropical dry forest tree community are described; the contribution of environmental heterogeneity and distance to β‐diversity was assessed. Significant differences in elevation, insolation, slope and soil water holding capacity (p < 0.01), variables related to water availability, were found among 830 m × 100 m transects laid along contrasting slopes of a system of three parallel microbasins. A gradient in elevation and insolation was found within north‐facing transects, among 10 m × 10 m sites; south‐facing transects showed an elevation gradient while crest transects showed a gradient in water holding capacity. In total 119 species were registered, with 27 to 64 species per transect, and 4 to 16 species per site. A large β‐diversity was found among and within transects; two indices of β‐diversity consistently showed a higher β‐diversity within transects than among them. Among transects, 64% of the variance in species composition could be attributed to the environmental variables; an additional 22% to the spatial distribution of sites. Within transects, 42% of the deviance in β‐diversity values was explained by insolation, and 19% by distance. β‐diversity increased with distance and with difference in insolation among sites; north‐facing transects, those with most contrasting insolation conditions, had the steepest increase in β‐diversity with distance. Such increase was clearly associated with changes in species composition, not with changes in species richness.