Altitude effects on spatial components of vascular plant diversity in a subarctic mountain tundra

Environmental gradients are caused by gradual changes in abiotic factors, which affect species abundances and distributions, and are important for the spatial distribution of biodiversity. One prominent environmental gradient is the altitude gradient. Understanding ecological processes associated with altitude gradients may help us to understand the possible effects climate change could have on species communities. We quantified vegetation cover, species richness, species evenness, beta diversity, and spatial patterns of community structure of vascular plants along altitude gradients in a subarctic mountain tundra in northern Sweden. Vascular plant cover and plant species richness showed unimodal relationships with altitude. However, species evenness did not change with altitude, suggesting that no individual species became dominant when species richness declined. Beta diversity also showed a unimodal relationship with altitude, but only for an intermediate spatial scale of 1 km. A lack of relationships with altitude for either patch or landscape scales suggests that any altitude effects on plant spatial heterogeneity occurred on scales larger than individual patches but were not effective across the whole landscape. We observed both nested and modular patterns of community structures, but only the modular patterns corresponded with altitude. Our observations point to biotic regulations of plant communities at high altitudes, but we found both scale dependencies and inconsistent magnitude of the effects of altitude on different diversity components. We urge for further studies evaluating how different factors influence plant communities in high altitude and high latitude environments, as well as studies identifying scale and context dependencies in any such influences.     

北京东灵山区景观类型空间邻接与分布规律

景观由斑块实间镶嵌而成。如何从错综复杂的斑块镶嵌中发现景观类型的空间分布规律,是景观格局研究中亟待解决的问题。通过对北京东灵山地区景观类型空间邻接关系的研究,借鉴群落生态学方法对其空间分布规律进行了探讨。结果表明,各个景观类型的空间邻接特征明显差异,可依紧密邻接的类型数目和程度分为３类;决定景观类型邻接多样性的主要因素是邻接类型的数目;采用作者提出的以景观类型空间邻接长度替代群落学数据的群落分类方     

Beta diversity of aquatic invertebrates increases along an altitudinal gradient in a Neotropical mountain

Mountains harbor rich biodiversity and high levels of endemism, particularly due to changes in environmental conditions over short spatial distances, which affects species distribution and composition. Studies on mountain ecosystems are increasingly needed, as mountains are highly threatened despite providing ecosystem services, such as water supply for half of the human population. We aimed to understand the patterns and drivers of alpha and beta diversities of aquatic invertebrates in headwater streams along an altitudinal gradient in the second largest South American mountain range, the Espinhaço mountains. Headwater streams were selected at each 100 m of elevation along an altitudinal gradient ranging from 800 to 1400 m asl, where three substrate types per stream were sampled: leaf litter, gravel, and cobbles. Environmental variables were sampled to represent local riparian canopy cover, instream physical habitat, water quality, climatic data, and land use. Generalized linear models and mixed models were used to test relationships between altitude and the richness and abundance of invertebrates and to assess the influence of environmental variables on the same metrics. Patterns of spatial variation in aquatic invertebrate assemblages along the altitudinal gradient were assessed using multiplicative beta diversity partitioning. The richness and abundance of aquatic invertebrates decreased with increasing altitude, whereas beta diversity increased with increasing altitude. Significant differences in assemblage composition and in relative abundance of invertebrates were observed for both substrates and altitude. We thus show that the high regional beta diversity in aquatic ecosystems in the studied site is due to the high turnover among areas. Abstract in Portuguese is available with online material.     

Vegetation heterogeneity and diversity in flat and mountain landscapes of Patagonia (Argentina)

Abstract. We studied floristic and diversity patterns and their environmental controls in two landscapes of contrasting topography in the Patagonian steppe. The analyses were focused on the effects of water availability gradients and landscape configuration on plant species distribution and coexistence. Floristic variation was investigated using Correspondence Analysis. The relationship between floristic and environmental variation was analyzed using Canonical Correspondence Analysis and correlation tests. We explored diversity patterns by relating spatial distance to floristic dissimilarities. The floristic gradient was determined by shrub and grass species and was related to precipitation in the flat area, and to precipitation, elevation and potential radiation in the mountain area. Site species richness increased with water availability in both areas. Mean site species richness and species turnover in space was higher in the mountain than in the flat area. Landscape species richness and floristic gradients were more concentrated in the mountain than in the flat area. In contrast to shrubs and grasses, forb species distributions were uncoordinated and probably independent of any environmental gradient. Our results suggest (1) that landscape configuration affects species composition and diversity through its direct effect on abiotic environmental heterogeneity, and (2) that the environmental controls of the community composition vary depending on the plant functional type considered.     

新疆博格达山地面生苔藓植被的数量分类与排序研究

 应用数量分类(TWINSPAN)和排序(DCA)方法,对新疆博格达山苔藓植被进行了多元分析。TWINSPAN结果表明,该地区苔藓植被可分为6个主要群落类型,即牛角藓-真藓(Cratoneuron-Bryum)群落、扭口藓-墙藓(Barbula-Tortula)群落、提灯藓-细罗藓(Mnium-Leskeella)群落、对叶藓-提灯藓(Distichium_Mnium)群落、墙藓-连轴藓(Tortula_Schistidium)群落、扭口藓-紫萼藓(Barbula_Grimmia)群落,它们分别分布于山地荒     

中条山东段森林乔木树种分布的环境梯度分析与种组划分

利用CCA排序和TWINSPAN分类,探讨了中条山东段森林植被中乔木树种分布与环境梯度的关系,并根据树种在排序图上的相对位置,同时参考相关文献资料,进行了种组划分。结果表明,海拔梯度是影响乔木树种分布的最重要的环境因子,土壤肥力和土壤湿度指数也具有较大的作用。72个树种（含变种）归纳划分为6个种组,即低海拔暖湿型、中低海拔暖燥型、中低海拔温润型、中海拔湿沃型、中高海拔燥瘠型和高海拔冷湿型。每个种组内的树种在生态习性特别是在分布特征上显示出较大的相似性,介绍了各种组分布的生境特征。TWINSPAN分类结果与种组划分吻合较好。     

Quantitative analysis of endangered Acanthopanax senticosus communities in Dongling Mountain of Beijing

Based on the survey of community plots, a quantitative analysis of endangered Acanthopanax senti-cosus communities in Dongling Mountain was performed with two way indicator species analysis (TWINSPAN), detrended correspondence analysis (DCA) and canonical correspondence analysis (CCA). The communities of A. senticosus were classified into 9 types by TWINSPAN and the results were validated by DCA. On the DCA graph, the first axis reflected the gradient of altitude and the second axis reflected the aspect and slope. Most of A. senticosus were distributed in the thick forests at a high altitude with little light. With the exception of being a dominant species of shrub layers in a few communities, A. senticosus has a relatively scarce distribution. In accordance with DCA, the results of CCA also show the trend that the distribution of A. senticosus communities varied along with the gradient change of environmental factors. Altitude and light are the main factors affecting A. senticosus growth.     

东灵山濒危植物刺五加生存群落的数量分析

在样方调查的基础上,分别采用双向指示种分类法(TWINSPAN)、除趋势对应分析法(DCA)和典范对应分析(CCA)对东灵山地区濒危植物刺五加的生存群落进行数量分析。TWINSPAN分类将刺五加生存群落分为9类,分类结果在DCA二维排序图上得到了较好的验证,其中第一轴反映的是海拔梯度,第二轴反映了坡向、坡度的变化。刺五加多分布在位于海拔较高、光照较少的杂木林中,除在少数群落中成为灌木层的优势种外,相对分布较少。CCA排序结果与DCA的排序结果基本一致,反映出植物群落分布随环境因子梯度变化的趋势,揭示了海拔和光照是限制刺五加生长的主要环境因子。     

中条山混沟地区森林乔木种的数量分类与环境解释

通过对中条山混沟地区森林植被调查资料的多元分析——TWINSPAN分类、CCA排序与环境解释,划分了该地区的植被类型,给出了植被与环境因子的定量关系。结果如下:1)40个样地可划分为青檀林(Form.Pteroceltis tatarinowii)、栾树林(Form. Koelreuteria paniculata)、槲栎林(Form. Quercus aliena)、栓皮栎林(Form. Quercus variabilis)、鹅耳枥+葛萝槭林(Form. Carpinus turczaninowii+Acer grosseri)、元宝槭+千金榆林(Form. Acer truncatum+Carpinus cordata)和辽东栎林(Form. Quercus liaotungensis)7种群落类型,体现了中条山地带性植被类型为暖温带落叶阔叶林的特点。2)群落类型的划分主要反映了CCA排序第一、二轴的环境梯度,CCA排序第一轴突出反映了海拔与土壤养分梯度,第二轴与土壤pH值、湿度指数和坡度显著相关。总体来说,海拔和土壤因子是影响混沟地区乔木物种分布分异的最主要环境因子。3)环境因子和空间因子解释了物种格局变化的46.14%,其中环境因子占30.79%,空间因子占8.48%,空间因子和环境因子交互作用解释的部分占6.87%。良好的环境解释反映了调查取样和环境因子选取的合理性,同时也体现了混沟地区植被的原始性。     

Coupling effects of altitude and human disturbance on landscape and plant diversity in the vicinity of mountain villages of Beijing, China

Cone-cores discarded by Eurasian red squirrels were used to study the habitat selection of Korean pine-seeds hoarding, in forest patch Nos. 16 and 19 in Liangshui Nature Reserve, China. Ten transects with a total length of 15 km were uniformly set, and data from 343 valid samples were collected in a 369 hm² area. One hundred and eighty four were hoarding samples which were determined according to the cluster analysis based on the number of the cone-cores, while the other 159 were control samples. The principal component analysis, using 11 habitat factors, suggested that the distance from Korean pine forest, forest type, number of Korean pine seedlings, density and type of bush significantly influenced the habitat selection of hoarding by Eurasian red squirrels. The results of Bailey’s method indicated that the squirrels showed (1) preference for natural coniferous forest, natural fir and spruce forest and planted spruce forest; (2) avoidance of planted Korean pine forest and planted larch forest; and (3) random use of natural Korean pine forest. Moreover the distance from the Korean pines in the range of 150–600 m showed no effect on the habitat selection of hoarding by the Eurasian red squirrels. More than 50% of the cone-cores were discarded in either fringe or gap of the Korean pine forest with more cone-cores found at <300 m than at 300 m away (One-Way ANOVA; df = 3, 183, F = 5.76, p = 0.0009). This demonstrated that the Eurasian red squirrels could take the cone-cores out of the Korean pine forest. The density of bushes in samples of hoarding area was significantly lower than that in control samples (Kruskal–Wallis test; df = 1, χ² = 83.99, p < 0.0001). The number of the Korean pine seedlings in samples of hoarding area was significantly higher than that in the control samples (Kruskal–Wallis test; df = 1, χ² = 104.13, p < 0.0001). This illustrated that the hoarding habitat favoured the germination of the Korean pine seedlings. In conclusion the behavior of hoarding Korean pine seeds by the Eurasian red squirrels can promote the regeneration and dispersal of the Korean pines.     

Spatial distribution pattern of the riparian vegetation in a basin in the NW Spain

The riparian vegetation of a basin in the NW Spain was studied to establish its spatial variation pattern and to relate floristic and structural differences in the community to environmental factors. Eighty-seven sampling units in 43 sampling stations were used. Samples were classified in 5 groups using Two Way Indicator Species Analysis (TWINSPAN). Three groups represented reaches with riparian wood along their banks: Mediterranean alderwoods and shrubby willow woods. The remaining two corresponded to floodplains with vegetation colonizing moderately eutrophicated deposits of gravel. Structural characteristics of richness and diversity differentiated the alder woods. In these, the shaded environment created by the woody species limited herbaceous vegetation development. This was dominated by Carex acuta subsp. broteriana. Classification and CCA ordination results were compared. The TWINSPAN groups could be recognized in the CCA graph. The ordination was related to a pollution gradient associated with altitude. This pollution gradient involved bank ruderalization, incorporation of nitrophilous species and a decrease in the vegetation quality. The influence of the lithological features on vegetation was also evident.     

拉萨河流域亏组山植物物种丰富度和群落特征研究

为研究草地植物群落、物种丰富度及其分布格局与影响因子之间的关系,该文以拉萨河流域林周县卡孜乡亏组山为研究地点,对山体垂直样带(3900～5100 m)植物群落特征、植物物种丰富度与各影响因子之间的关系进行了研究.研究区域共设置了13个样带(每隔100 m设一个样带),每样带设置0.5 m×0.5 m的5个样方进行植被调...     

Coupling effects of altitude and human disturbance on landscape and plant diversity in the vicinity of mountain villages of Beijing, China

Villages represent the source of human disturbance in mountain regions, but how they alter surrounding landscapes and further affect plant diversity distribution along altitudinal gradient is still not well documented. Although the unimodal pattern along latitudinal gradient and the hypothesis of immediate disturbance (IDH) have been supported in many studies, their coupling effects on plant diversity distribution have been given less attention. In this paper, the coupling effects surrounding mountain villages were detected: (1) altitude determined the mountain landscape at regional scale, while human disturbance altered landscape fragmentation and diversity at local scale surrounding villages. (2) With the reducing human disturbance away from villages, plant diversity decreased, then increased, and finally decreased in different land uses. The plant diversity of shrubs reached the lowest. With the increasing altitude, plant diversity of the forest represented the unimodal trend, but other land use types had different properties. (3) The mechanism of the coupling effects is that the combination of topographic and soil factors determines plant diversity distribution at both landscape and plant community levels surrounding villages.     

Mountain summer farms in Røldal,western Norway –; vegetation classification and patterns in species turnover and richness

This paper discusses vegetation types and diversity patterns in relation to environment and land-use at summer farms, a characteristic cultural landscape in the Norwegian mountains. Floristic data (189 taxa) were collected in 130 4-m² sample plots within 12 summer farms in Røldal, western Norway. The study was designed to sample as fully as possible the range of floristic, environmental, and land-use conditions. Vegetation types delimited by two-way indicator species analysis were consistent with results from earlier phytosociological studies. Detrended correspondence analysis and canonical correspondence analysis show that rather than being distinct vegetation types, the major floristic variation is structured along a spatial gradient from summer farm to the surrounding heathland vegetation. Species richness (alpha diversity) was modelled against environmental variables by generalized linear modelling and compositional turnover (beta diversity) by canonical correspondence analysis. Most environmental factors made significant contributions, but the spatial distance-to-farm gradient was the best predictor of both species richness and turnover. While summer farms reduce mean species richness at the plot scale, the compositional heterogeneity of the upland landscapes is increased, thereby creating ‘ecological room’ for additional vegetation types and species. Within an overall similarity across scales, soil variables (pH, base saturation, LOI, phosphate and nitrogen) differed considerably in their explanatory power for richness and turnover. A difference between ‘productivity limiting’ factors and ‘environmental sieves’ is proposed as an explanation. Species turnover with altitude is relatively low in grasslands as compared to heaths.     

From forest to pasture: an evaluation of the influence of environment and biogeography on the structure of beetle (Scarabaeinae) assemblages along three altitudinal gradients in the Neotropical region

The objective of this study is to evaluate the effect of environmental (associated with the expansion of cattle ranching) and biogeographical factors on the diversity of dung beetle (Scarabaeinae) assemblages along three altitudinal gradients in the Neotropical region. One gradient is located in the Mexican Transition Zone, on the Cofre de Perote mountain, the other two are in the northern Andes (the Chiles Volcano and the Río Cusiana Basin). For the three gradients, the number of species and of individuals was similar in both forest and pasture, while species composition was different between habitats. On this mountain, species turnover in pastures was characterized by the addition of new species as altitude increased. In the northern Andes, species diversity was always greater in the forest than in the pasture, and species turnover between habitats was notably influenced by species loss with increasing altitude. As such the pasture fauna of the northern Andes was an impoverished derivative of the fauna present in the forests at the same altitude characterized by species of Neotropical affinity with a limited capacity for colonizing open, sunnier habitats. The opposite occurs in the areas used by cattle on the Cofre de Perote. This habitat has its own fauna, which is mainly comprised of Holarctic and Afrotropical species adapted to the prevailing environmental conditions of areas lacking arboreal vegetation. These results suggest that the impact on beetle communities caused by human activities can differ depending on the geographic position of each mountain and, particularly, the biogeographical history of the species assemblage that lives there.     

Weed vegetation ecology of arable land in Salalah,Southern Oman

This paper applies multivariate statistical methods to a data set of weed relevés from arable fields in two different habitat types of coastal and mountainous escarpments in Southern Oman. The objectives were to test the effect of environmental gradients, crop plants and time on weed species composition, to rank the importance of these particular factors, and to describe the patterns of species composition and diversity associated with these factors. Through the application of TWINSPAN, DCA and CCA programs on data relating to 102 species recorded in 28 plots and farms distributed in the study area, six plant communities were identified: I- Dichanthium micranthum, II- Cynodon dactylon–D. micranthum, III- Convolvulus arvensis, IV- C. dactylon–Sonchus oleraceus, V- Amaranthus viridis and VI- Suaeda aegyptiaca–Achyranthes aspera. The ordination process (CCA) provided a sequence of plant communities and species diversity that correlated with some anthropogenic factors, physiographic variables and crop types. Therefore, length of time since farm construction, disturbance levels and altitude are the most important factors related to the occurrence of the species. The perennial species correlated with the more degraded mountain areas of new farm stands, whereas most of the annuals correlated with old lowland and less disturbed farms.     

新疆东部天山蝶类多样性及其垂直分布

2006-2008年研究了新疆东部天山蝶类多样性和垂直分布.结果表明:研究区域内共记录蝴蝶7科43属63种,占新疆已记录蝶类种数的24.80％,区系组成主要是古北种,占73％;其次是广布种,占27％,没有发现东洋种.其中蛱蝶科的物种数最多,为11属19种,蚬蝶科的物种数最少,只有1属1种.按海拔将生境分为5个垂直自然带,包括低山灌木草原带、山地森林草原带、亚高山草甸带、高山草甸带、垫状植被带.蝶类物种数和个体数排序为亚高山草甸带＞山地森林草原带＞低山灌木草原带＞高山草甸带＞垫状植被带.采用Shannon-Wiener指数和G-F指数对蝶类物种和科、属的多样性进行了分析评价,结果显示亚高山草甸带的蝶类多样性最为丰富,其次是山地森林草原带和低山灌木草原带,而高山草甸带和垫状植被带的蝶类多样性相对较低,物种和科、属多样性分析结果均一致.蝶类垂直分布明显,物种数和个体数随海拔变化的趋势类似,均为先增加后下降.蝶类区系成分随着海拔升高发生改变,广布种的比例逐渐降低,高山草甸带和垫状植被带只有古北种分布.研究结果显示,生境改变对蝴蝶群落影响明显,保护生境是保护蝴蝶生存的最主要措施.     

Quantitative classification and ordination of forest communities in Pangquangou National Nature Reserve

Quantitative analysis of ecological relationships between vegetation and the environment has become an essential means in the field of research of modern vegetation ecology. In this article, based on data from 84 quadrates, forest communities in this reserve were investigated using TWINSPAN, DCA and DCCA. The results will be helpful in the construction and development of Pangquangou National Nature Reserve. Using TWINSPAN, the forest communities were classified into seven types. The distribution pattern of vegetation reflects the comprehensive influence of environments. The results of DCA and DCCA clearly reflect the relationship between the pattern of forest communities and environmental gradients. The ordination result of DCCA indicates that altitude is more important than other environmental factors because the change of altitude gradient will lead to changes in the temperature and humidity gradients. The first of the DCA ordination axes indicates the humidity gradient, and the second indicates the temperature gradient. All these results show that the main factors restricting the distribution of communities in this reserve are temperature and humidity. The ecological meaning of the ordination axis in DCCA is much clearer than that in DCA, and the species-environment correlation of DCCA is more obvious than DCA. The first DCCA axis indicates the altitude gradient among the communities, while the second is the gradient in aspect and slope among the communities. DCCA ordination can simultaneously express similarities of species and environment. Therefore, the quadrat location in the DCCA ordination figure is much closer than in the DCA.     

Altitudinal variation of dung beetle (Scarabaeidae: Scarabaeinae) assemblages in the Colombian Andes

Aim We describe the changes in species richness, rarity and composition with altitude, and explore whether the differences in Scarabaeinae dung beetle composition along five altitudinal transects of the same mountain range are related to altitude or if there are interregional differences in these altitudinal gradients. Location Field work was carried out on the eastern slope of the eastern Cordillera, Colombian Andes, between Tamá Peak to the north, in the Tamá National Park (07°23′ N, 72°23′ W) and the San Miguel River (00°28′ N, 77°17′ W) to the south. Methods Sampling was carried out between February 1997 and November 1999 in five regions spanning elevation gradients. In each gradient, six sites were chosen at 250 m intervals between 1000 and 2250 m a.s.l. Results We found a curvilinear relationship between altitude and mean species richness, with a peak in richness at middle elevations. However, the diversity of dung beetle assemblages does not seem to be related to the interregional differences in environmental conditions. The number of geographically restricted species is negatively and significantly related to altitude, with geographically restricted species more frequent at low altitude sites. Ordination delimited the two main groups according to altitude: one with all the highest sites (1750–2250 m a.s.l.) and a second group with the remaining sites (< 1750 m a.s.l.). Analysis of species co‐occurrence shows that these dung beetle assemblages seem to be spatially structured when all sites have the same probability of being chosen. In contrast, the spatial structure of species assemblages seems to be random when the probability of choosing any site is proportional to its altitude. Main conclusions The altitude of sites is the main factor that influences the diversity of these dung beetle assemblages. The peak in species richness at middle elevations, the higher number of geographically restricted species at lower altitudinal levels, and the compositional differences along these mountain gradients seem to result from the mixing at these altitudes of dung beetle assemblages that have different environmental adaptations and, probably, different origins. The relevance of altitude in these assemblages is related to the limited role of these Neotropical high altitude environments as centres of refuge and vicariance for a monophyletic group of warm‐adapted species, for which the vertical colonization of these high mountain environments by lineages distributed at lower altitudes would have been very difficult.     

Quantitative classification and ordination of forest communities in Pangquangou National Nature Reserve

Quantitative analysis of ecological relationships between vegetation and the environment has become an essential means in the field of research of modern vegetation ecology. In this article, based on data from 84 quadrates, forest communities in this reserve were investigated using TWINSPAN, DCA and DCCA. The results will be helpful in the construction and development of Pangquangou National Nature Reserve. Using TWINSPAN, the forest communities were classified into seven types. The distribution pattern of vegetation reflects the comprehensive influence of environments. The results of DCA and DCCA clearly reflect the relationship between the pattern of forest communities and environmental gradients. The ordination result of DCCA indicates that altitude is more important than other environmental factors because the change of altitude gradient will lead to changes in the temperature and humidity gradients. The first of the DCA ordination axes indicates the humidity gradient, and the second indicates the temperature gradient. All these results show that the main factors restricting the distribution of communities in this reserve are temperature and humidity. The ecological meaning of the ordination axis in DCCA is much clearer than that in DCA, and the species-environment correlation of DCCA is more obvious than DCA. The first DCCA axis indicates the altitude gradient among the communities, while the second is the gradient in aspect and slope among the communities. DCCA ordination can simultaneously express similarities of species and environment. Therefore, the quadrat location in the DCCA ordination figure is much closer than in the DCA.