Contrasting patterns of genetic variation and structure in plant invasions of mountains

Aim To assess the population genetic consequences of the colonization of two species with contrasting mating systems, Solidago canadensis and Lactuca serriola, along altitudinal gradients in both their native and introduced ranges. Location Allegheny Mountains, West Virginia and Wallowa Mountains, Oregon, USA; Valais, southern Switzerland. Methods Leaf material was collected from populations along altitudinal gradients and genotyped at seven microsatellite loci for each species. Differences in variability between native and introduced areas and in relation to altitude were analysed using linear models. Differences in the genetic, geographical and altitudinal structure of populations between areas were analysed by AMOVA, cluster analysis and Mantel tests. Results Genetic variation within and across populations of S. canadensis was significantly reduced, while populations of L. serriola were significantly more variable, in the introduced area. Genetic diversity decreased significantly with altitude for S. canadensis but not L. serriola. Genetic structure of S. canadensis was similar in both areas, and populations were isolated by geographical but not altitudinal distance. By contrast, population structure of L. serriola was much weaker in the introduced area, and populations were not isolated by distance in either area. Main conclusions Solidago canadensis has experienced a strong genetic bottleneck on introduction to the Valais, but this has not prevented it from colonizing a wide altitudinal range. Variation in neutral markers is therefore not necessarily a good measure for judging the ecological behaviour of a species. By contrast, the greater variability of L. serriola in the introduced area, where it also occurs over a greater altitudinal range, can be explained by increased outcrossing among admixed populations. This suggests that the ecological amplitude of alien species might be enhanced after population admixture in the new range, especially for species with highly structured native populations. However, even genetically depauperate introduced populations can be expected to colonize the same environmental range that they occupy in the native area.     

Environment‐driven changes in diversity of riparian plant communities along a mountain river

The study of changes in species richness and composition along rivers has focused on large spatial scales. It has been ignored that in different sections of the river (high mountain area, middle zone, and mouth of the river) the specific environmental conditions can generate different longitudinal patterns of the species richness and composition. In this study, we determine whether species richness and composition of the riparian plant communities change along a mountain river and whether these changes are related to environmental variables. We expect an increase in species richness and turnover along the river, that the upstream communities would be a subset of the downstream communities, and that such would be related to edaphic and hydrologic conditions. To test this, we sampled three strata of the riparian vegetation (upper: individuals with <1 cm of ND, middle: individuals with >1 cm of ND, low: individuals with >1 m tall) in a set of 15 sites that we place along a mountain river. Additionally, we recorded topographic, hydrological, morphological, and soil variables. We performed correlation analyzes to determine whether changes in species richness and turnover were related to increased distance to the origin of the river. Also, we obtained the nestedness and evaluated the importance of environmental variables with GLM, LASSO regression, and CCA. With the increase in distance, the species richness decreases in the upper stratum, but not in the middle and the low stratum (although the highest values were observed near the origin of the river), the turnover increase in all strata and the upstream communities were not a subset of the downstream communities. The changes in species richness and composition were related to topographic (altitude), hydrological (flow), and edaphic (conductivity and pH) variables. Our results indicate that at small spatial scales the patterns of richness and composition differ from what has been found at larger spatial scales and that these patterns are associated with environmental changes in the strong altitude gradients of mountain rivers.     

The distribution of ground spiders (Araneae, Gnaphosidae) along the altitudinal gradient of Crete, Greece: species richness, activity and altitudinal range

Aim To study the altitudinal variation of ground spiders (Araneae, Gnaphosidae) of Crete, Greece, as far as species composition, species richness, activity and range of distribution are concerned. Location Altitudinal zones (0–2400 m) along the three main mountain massifs of the island of Crete. Methods Thirty‐three sampling sites were located from 0 to 2400 m a.s.l. on Crete, and sampled using pitfall traps. Material from the high‐activity period of Gnaphosidae (mid‐spring to mid‐autumn) was analysed. Sampling sites were divided into five altitudinal zones of 500 m each. Statistical analysis involved univariate statistics (anova ) and multivariate statistics, such as multidimensional scaling (MDS) and cluster analysis (UPGMA) using binomial data of species presence or absence. Results Species richness declines with altitude and follows a hump‐shaped pattern. The activity pattern of the family, as a whole, is not correlated with altitude and is highly species‐specific. In the highest zone, both species richness and activity decline dramatically. The altitudinal range of species distribution increases with altitude. On the Cretan summits live highly tolerant lowland species and isolated residents of the high mountains of Crete. Two different patterns of community structure are recorded. Main conclusions Communities of Gnaphosidae on Crete present two distinct structures following the altitudinal gradient, these being separated by a transitional zone between 1600 and 2000 m. This study supports previous results which show a hump‐shaped decline in species richness of Gnaphosidae along altitudinal gradients, leading to a peak at 400–700 m, where an optimum of environmental factors exists. This makes this zone the meeting point of the often opportunistic lowland species with the older and most permanent residents of the island. Rapoport's rule on the positive correlation of the altitudinal range of species distributions with altitude is also supported. The high activity recorded for the species that persist on the high mountains of Crete is indicative of a tolerant arachnofauna, and is considered to result from relaxation of competitive interactions with other species. This is related to a reduction in species numbers, shortening of the activity period on high mountains and the unique presence of high mountain species that thrive only there. As shown in our study, strategies to cope with altitude are species‐specific. Therefore, there cannot exist one single model to describe how animals react to the change in altitude, even under the same environmental conditions.     

祁连山北坡中段植物群落多样性的垂直分布格局

利用DCCA排序和海拔高程排序相结合的方法 ,对祁连山北坡中段植物群落物种多样性垂直分布格局进行了初步研究。结果表明 :1)植物群落草本层和灌木层物种丰富度和多样性在环境梯度上呈单峰曲线变化趋势 ,乔木层的物种丰富度和多样性在环境梯度上无变化。物种丰富度和多样性对环境梯度变化敏感程度的次序是草本层 >灌木层 >乔木层 ;2 )植物群落各层次均匀度在环境梯度上没有表现出一定的变化规律 ,均匀度可能更多地受制于群落自身动态的影响 ,而独立于生境的资源水平 ;3)草地群落物种多样性在DCCA环境梯度上曲线的拟合效果优于按海拔高程排序效果 ,灌木群落则相反 ;4)低海拔、中低海拔和中海拔地带的草本层物种丰富度和Shannon Wiener多样性指数 (H′)显著高于灌木层 (p <0 .0 1) ;高海拔地带草本层仅丰富度指数显著高于灌木层 (p <0 .0 5 )。在整个海拔范围内 ,草本层和灌木层的均匀度无显著差异。就资源的可利用性而言 ,研究区域植物群落物种多样性在垂直环境梯度上的变化规律表达了物种多样性与资源生产力的单调关系内涵。     

贺兰山东坡典型植物群落多样性垂直分布特征与土壤因子的关系

深入探究山地生态系统植物群落多样性与关键土壤因子的关系及土壤因子在植物群落构建与多样性维持中的重要作用是区域生物多样性保护和生态系统服务功能综合提升的基础。本文采用冗余分析（Redundancy analysis,RDA）和相关分析揭示了贺兰山东坡典型植物群落多样性垂直分布格局与土壤因子的关系。结果表明：（1）贺兰山东坡植物群落多样性具有明显的垂直分布格局,针阔叶混交林植物物种组成最为丰富,中域效应明显;不同层次植物多样性随海拔升高变化趋势不一致且各自在不同海拔间差异显著（P<0.05）,植物多样性大体表现为草本层 > 灌木层 > 乔木层。（2）研究区土壤因子沿海拔梯度变化趋势不同,均存在显著差异（P<0.05）,其中土壤含水量（SM）、有机碳（SOC）和全氮（TN）含量空间变异性较大;高山灌丛草甸土壤养分含量最高,高海拔区土壤P限制高。（3）9个土壤因子解释了研究区总体植物多样性89.68%的生态信息,与植物多样性具有较强的相关性,影响不同层次植物多样性垂直分异的关键土壤因子略有不同。综上,土壤有机碳、pH、碳磷比、含水量是影响贺兰山东坡植物多样性空间分布的主要驱动因子。     

丛枝菌根真菌群落沿高寒草原海拔梯度的变化特征

基于丛枝菌根(Arbuscular mycorrhizal,AM)真菌孢子形态学的鉴定,对沿不同海拔(4584、4628、4744、4880、4956 m)梯度采集的高寒草原建群植物根际土壤样品进行了分析。结果表明,高寒草原AM真菌属、种构成均较简单,Acaulospora、Claroideoglomus、Funneliformis、Glomus属见于各海拔梯度,海拔4744 m地带未见Pacispora属,海拔4744、4956 m地带无Scutellospora属分布,Rhizophagus属仅见于海拔4584 m地带。随海拔上升,AM真菌种数、物种丰度均呈显著下降;海拔4584—4880 m范围Shannon-Weiner指数(H)无显著差异,但在最高海拔时显著下降;优势种种数及所占比例与海拔梯度则呈显著正相关(Funneliformis geosporum、Claroideoglomus claroideum为不同海拔梯度优势种);沿海拔梯度,孢子密度基本呈单峰分布格局,峰值出现在海拔4744 m地带;海拔梯度对菌根侵染效应影响显著,菌根侵染率、侵染强度和丛枝丰度随海拔上升均呈显著下降趋势;不同海拔梯度高寒草原AM真菌群落相似度(Sorensen相似性系数0.821—0.969)较高,并在总体上表现出随海拔梯度增大而降低的趋势。土壤pH值、有效磷、有机碳、海拔对AM真菌的群落分布均产生显著影响,尤以海拔的影响最为显著。研究结果对预测高寒草原微生物的作用与影响,以及高寒草原环境对全球变化的响应等提供了理论依据。     

长白山溪流河岸带森林木本植物多样性沿海拔梯度分布规律

采用样带网格调查方法和α、β多样性指数分析方法,研究了长白山河岸带原始林和次生林群落木本植物多样性沿海拔梯度分布规律及其对采伐干扰的响应.结果表明：不同海拔区域河岸带原始林群落均由11～13个树种组成,其在群落中的地位随海拔升高而发生变化,阔叶树种优势地位逐步被针叶树种所取代,采伐干扰不仅改变了群落树种丰富度及其在群落中的地位,而且使针叶树种取代阔叶树种的趋势有所增强;河岸带原始林群落物种多样性沿海拔梯度呈现出中、低海拔区域相对较高且比较恒定(2.454～2.544),高海拔区域(2.250)下降的分布规律,采伐干扰改变了其沿海拔梯度分布格局(波动型),加大了不同海拔区域群落间的波动性(2.174～2.692);河岸带原始林群落树种沿海拔梯度的变动速率相对较低(1.5～3.5),且群落相似性较高(0.85～0.94),采伐干扰使次生林群落树种沿海拔梯度的变动幅度增大(0.5～6.0),群落相似性下降(0.68～0.91),但次生林群落沿海拔梯度分布仍具有较高连续性.     

Brain size variation along altitudinal gradients in the Asiatic Toad (Bufo gargarizans)

Size changes in brain and brain regions along altitudinal gradients provide insight into the trade‐off between energetic expenditure and cognitive capacity. We investigated the brain size variations of the Asiatic Toad (Bufo gargarizans) across altitudes from 700 m to 3,200 m. A total of 325 individuals from 11 sites and two transects were sampled. To reduce confounding factors, all sampling sites within each transect were within a maximum distance of 85 km and an altitudinal difference close to 2,000 m. Brains were dissected, and five regions were both measured directly and with 3D CT scan. There is a significant negative correlation between the relative whole‐brain volume (to snout‐vent length) and altitude. Furthermore, the relative volumes (to whole‐brain volume) of optic tectum and cerebellum also decrease along the altitudinal gradients, while the telencephalon increases its relative volume along the gradients. Therefore, our results are mostly consistent with the expensive brain hypothesis and the functional constraint hypothesis. We suggest that most current hypotheses are not mutually exclusive and data supporting one hypothesis are often partially consistent with others. More studies on mechanisms are needed to explain the brain size evolution in natural populations.     

Woodland mollusc communities along environmental gradients in the East Carpathians

Mollusc communities were sampled quantitatively at eleven sites representing different environmental conditions in the Bieszczady National Park (East Carpathians Mts, Poland). Overall 61 species were recorded. Alder forest in the valleys (AF; Alnetum incanae carpathicum, Caltho-Alnetum, secondary alder forest) hosted the richest fauna, with up to 41 species occurring sympatrically on 100 m² of forest floor and average density ca. 750 specimens m⁻². Three important ecological controllers of species composition and community structure were found. The main predictor of mollusc assemblage composition was calcium content; the first DCA axis of molluscs most significantly and highly correlated with calcium content in the leaf litter and organic matter in the upper layer. The second axis significantly correlated with altitude and negatively with annual temperature, and thus can be explained as an elevational gradient. We observed slope aspect to constitute the third significant gradient. On the basis of forward selection in CCA analysis organic matter in the upper layer of soil was the best predictor of species composition, which explained 26% of total variance. It comes to prove that in mountain forest on non-calcareous bedrock molluscs obtain calcium mainly from leaf litter.     

北京东灵山辽东栎(Quercus liaotungensis)林沿海拔梯度的物种多度分布

多度格局对理解群落结构具有重要意义。选用5个描述种-多度关系的生态位模型(断棍模型BSM、生态位优先占领模型NPM、生态位重叠模型ONM、随机分配模型RAM、优势优先模型DPM),分乔、灌、草3层分别对北京东灵山辽东栎林进行了研究。结果表明:在乔木层,沿海拔梯度从低海拔到高海拔,能很好地反映物种多度格局变异的模型有由RAM、NPM向DPM过渡的趋势;NPM和BSM均能较好地模拟灌木层的绝大多数海拔段,但NPM的效果更好;草本层以BSM的模拟效果最好。生态位模型可以反映出辽东栎林乔木层物种多度分布沿海拔存在的明显变异,而灌木层和草本层物种多度分布沿海拔梯度无明显变化或很难由生态位模型反映出来,是否引入新的模型方法,如中性模型、近中性模型,还有待于进一步研究。     

长白山北坡森林群落结构组成及其海拔变化

对长白山北坡海拔700～1900m的13个森林群落进行了群落组成、结构随海拔上升变化情况的研究。结果表明,随着海拔的升高,群落的建群种组成、层次结构等,都表现出由复杂多样逐渐向简单单一过渡的变化趋势,从700m至1700m,针叶树所占的比例随海拔的升高呈明显的增加趋势,阔叶树比例随之减少,到林线时针阔叶树种均急剧减少,沿海拔梯度存在明显的树种更替现象。从群落的平均胸径及胸高断面积和可知。最大值出现于900～1100m及1600m,这2个海拔段正是不同森林群落类型间的过渡区。除海拔1800m的岳桦林之外,其它群落径级结构都呈倒J形分布,即各径级树木中,更新幼苗、幼树在数量上占绝对优势,随着胸径的增大,立木株数逐渐减少,这种径级结构表明,长白山北坡各海拔群落更新良好,群落处于稳定的发展状态。     

Contrasting patterns of fig wasp communities along Mt. Wilhelm,Papua New Guinea

The fig (Moraceae) and pollinating fig wasp (Agaonidae) mutualism is best known as a model system for the study of coevolution in plant–pollinator interactions and its central role in shaping vertebrate communities in tropical forests. Figs also host myriad antagonistic parasitic fig wasps which impose costs on both partners threatening mutualism stability. Spatiotemporal variation in parasitic wasp abundance is a key factor in mitigating these effects. Because fig wasps are temperature sensitive and likely vary in their ability to traverse environmental gradients, we expect community assemblages and abundance of both pollinating and non-pollinating fig wasps to respond to changes along an elevational gradient. In the present study, we compare the fig wasp communities and abundance of three fig species growing along the slopes of the Mount Wilhelm altitudinal gradient in Papua New Guinea. We quantified wasps from over 100 male fig trees and calculated seed set for 55 female trees along each of the species’ distribution on the transect. Our results show that the abundance of both pollinating and non-pollinating fig wasps follow a mid-elevation peak, consistent with fig species richness found in the same transect. The patterns, however, are different according to the host's species distribution. Seed set remained relatively constant along the gradient for all species with some decrease along higher elevations, potentially affecting connectivity along the gradient. As suggested for insects in general, temperature and habitat diversity appear to play a fundamental role in the species richness and abundance of fig wasps.     

福建黄岗山东南坡和西北坡乔木物种多样性及群落特征的垂直变化

利用福建黄岗山东南坡30个样方和西北坡13个样方的资料,研究该地区乔木物种丰富度的垂直变化;东南坡共记录到乔木物种151种,隶属于42科73属;西北坡102种,隶属于32科54属。两坡面的乔木树种组成相差不大。物种丰富度随海拔的变化趋势是：随海拔升高,科、属、种的数量呈下降的趋势;东南坡科、属、种的数量在海拔800—1000m达到最大值,西北坡在海拔1500—1600m达到最大值。东南坡乔木物种Shannon—Wiener指数(H’)与海拔呈负相关;西北坡在海拔1200—1800m范围内H’高于东南坡;Screnson指数(IAc)在不同植被类型交替时出现上下波动,从常绿阔叶林向针阔混交林转化时,物种更替强烈,Screnson指数明显下降。乔木物种生长特征的分析表明,最大树高(Hmax)和最大胸径(DBHmax)出现在中海拔,在相同海拔范围内西北坡的Hmax和DBHmax高于东南坡;东南坡Hmax和DBHmax的峰值出现在海拔800—900m,西北坡出现在海拔1800m。东南坡立木密度在海拔1500m处达最高值,而西北坡立木密度变化不明显,仅在海拔1900m以上明显下降。将全部乔木种划分为常绿阔叶、针叶和落叶阔叶等三种生活型,分析不同生活型的生长特征发现,常绿阔叶种类的胸高断面积和(total basal area,TBA)在低海拔占比例大;针叶种类沿海拔出现两个峰值,与分布两种不同针叶林种类有关;落叶阔叶林的TBA在各海拔段均占一定比例,但在西北坡的比例要高于东南坡。两坡三种生活型的TBA沿海拔梯度变化特征基本上相似。     

Contrasting patterns of intraspecific trait variability in native and non-native plant species along an elevational gradient on Tenerife,Canary Islands

Background and AimsNon-native plant species are not restricted to lowlands, but increasingly are invading high elevations. While for both native and non-native species we expected variability of plant functional traits due to the changing environmental conditions along elevational gradients, we additionally assumed that non-native species are characterized by a more acquisitive growth strategy, as traits reflecting such a strategy have been found to correlate with invasion success. Furthermore, the typical lowland introduction of non-native species coming from multiple origins should lead to higher trait variability within populations of non-native species specifically at low elevations, and they might therefore occupy a larger total trait space.MethodsAlong an elevational gradient ranging from 55 to 1925 m a.s.l. on Tenerife, we collected leaves from eight replicate individuals in eight evenly distributed populations of five native and six non-native forb species. In each population, we measured ten eco-morphological and leaf biochemical traits and calculated trait variability within each population and the total trait space occupied by native and non-native species.Key ResultsWe found both positive (e.g. leaf dry matter content) and negative (e.g. leaf N) correlations with elevation for native species, but only few responses for non-native species. For non-native species, within-population variability of leaf dry matter content and specific leaf area decreased with elevation, but increased for native species. The total trait space occupied by all non-native species was smaller than and a subset of that of native species.ConclusionsWe found little evidence that intraspecific trait variability is associated with the success of non-native species to spread towards higher elevations. Instead, for non-native species, our results indicate that intermediate trait values that meet the requirements of various conditions are favourable across the changing environmental conditions along elevational gradients. As a consequence, this might prevent non-native species from overcoming abruptly changing environmental conditions, such as when crossing the treeline.     

Biomass and diversity of dry alpine plant communities along altitudinal gradients in the Himalayas

A non-linear relationship between phytodiversity and altitude has widely been reported, but the relationship between phytomass and altitude remains little understood. We examined the phytomass and diversity of vascular plants along altitudinal gradients on the dry alpine rangelands of Ladakh, western Himalaya. We used generalized linear and generalized additive models to assess the relationship between these vegetation parameters and altitude. We found a hump-shaped relationship between aboveground phytomass and altitude. We suspect that this is engendered by low rainfall and trampling/excessive grazing at lower slopes by domestic livestock, and low temperature and low nutrient levels at higher slopes. We also found a unimodal relationship between plant species-richness and altitude at a single mountain as well as at the scale of entire Ladakh. The species-richness at the single mountain peaked between 5,000 and 5,200 m, while it peaked between 3,500 and 4,000 m at entire Ladakh level. Perhaps biotic factors such as grazing and precipitation are, respectively, important in generating this pattern at the single mountain and entire Ladakh.     

Phylogenetic plant community structure along elevation is lineage specific

The trend of closely related taxa to retain similar environmental preferences mediated by inherited traits suggests that several patterns observed at the community scale originate from longer evolutionary processes. While the effects of phylogenetic relatedness have been previously studied within a single genus or family, lineage‐specific effects on the ecological processes governing community assembly have rarely been studied for entire communities or flora. Here, we measured how community phylogenetic structure varies across a wide elevation gradient for plant lineages represented by 35 families, using a co‐occurrence index and net relatedness index (NRI). We propose a framework that analyses each lineage separately and reveals the trend of ecological assembly at tree nodes. We found prevailing phylogenetic clustering for more ancient nodes and overdispersion in more recent tree nodes. Closely related species may thus rapidly evolve new environmental tolerances to radiate into distinct communities, while older lineages likely retain inherent environmental tolerances to occupy communities in similar environments, either through efficient dispersal mechanisms or the exclusion of older lineages with more divergent environmental tolerances. Our study illustrates the importance of disentangling the patterns of community assembly among lineages to better interpret the ecological role of traits. It also sheds light on studies reporting absence of phylogenetic signal, and opens new perspectives on the analysis of niche and trait conservatism across lineages.     

Changes in reproductive investment with altitude in an alpine plant

Aims In perennial species, the allocation of resources to reproduction results in a reduction of allocation to vegetative growth and, therefore, impacts future reproductive success. As a consequence, variation in this trade-off is among the most important driving forces in the life-history evolution of perennial plants and can lead to locally adapted genotypes. In addition to genetic variation, phenotypic plasticity might also contribute to local adaptation of plants to local conditions by mediating changes in reproductive allocation. Knowledge on the importance of genetic and environmental effects on the trade-off between reproduction and vegetative growth is therefore essential to understand how plants may respond to environmental changes.Methods We conducted a transplant experiment along an altitudinal gradient from 425 to 1?921 m in the front range of the Western Alps of Switzerland to assess the influence of both altitudinal origin of populations and altitude of growing site on growth, reproductive investment and local adaptation in Poa alpina .Important findings In our study, the investment in reproduction increased with plant size. Plant growth and the relative importance of reproductive investment decreased in populations originating from higher altitudes compared to populations originating from lower altitudes. The changes in reproductive investment were mainly explained by differences in plant size. In contrast to genetic effects, phenotypic plasticity of all traits measured was low and not related to altitude. As a result, the population from the lowest altitude of origin performed best at all sites. Our results indicate that in P. alpina genetic differences in growth and reproductive investment are related to local conditions affecting growth, i.e. interspecific competition and soil moisture content.