Current problems of environmental gradients and species response curves in relation to continuum theory

Abstract. Comparisons of the positions of species on Grimes'C-S-R triangular ordination model with their responses to individual environmental gradients indicates that the C-S-R model does not necessarily predict species ecological behaviour. The importance of the stress, productivity and disturbance gradients relative to other environmental gradients needs to be determined. In studies of species behaviour along a biomass/productivity gradient the collective vegetation property, biomass, has been confused with the environmental factor, fertility. Patterns of responses to biomass gradients e.g. Keddy's centrifugal model, should be examined in a two-dimensional environmental space to avoid such confounding effects. Assumptions regarding the shapes of species responses to environmental gradients remain untested. A recent model of species response functions to environmental gradients suggested that skewed responses curves show a pattern in the direction of the skew, always with the tail towards the presumed most mesic position on the gradient. Further evidence is presented to support this model for a temperature gradient in eucalypt forest in south-eastern Australia. 21 out of 24 species tested conform to the model.     

哀牢山中山湿性常绿阔叶林木径向生长季节动态及其对气候因子的响应

研究采用树木生长环在哀牢山中山湿性常绿阔叶林持续9年(2009—2017年)监测了2个常绿树种(厚皮香,Ternstroemia gymnanthera;南亚枇杷,Eriobotrya bengalensis)和2个落叶树种(西桦,Betula alnoides;珍珠花,Lyonia ovalifolia)的树干月生长量,采用逻辑斯蒂生长模型(Logistic model)模拟树木径向生长量和物候参数,并分析了年、季尺度上径向生长与主要气候因子的关系。结果表明:1)4个树种年平均生长量为6.3 mm,落叶树种年平均生长量(10.6 mm/a)显著高于常绿树种(3.0 mm/a);2)雨季(5—10月)是哀牢山中山湿性常绿阔叶林树木生长的主要时期,4个树种雨季平均生长量为5.9 mm,占全年总生长量的93%,其中落叶树种雨季生长量占全年的96%,而常绿树种雨季生长量占全年的86%;3)常绿树种生长季长度为169天,长于落叶树种(137天),而落叶树种最大生长速率(0.14 mm/d)显著高于常绿树种(0.03 mm/d),最大径向生长速率能很好地预测树种年生长量;4)低温、雾日和光合有效...     

Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.     

Diversity of forest vegetation across a strong gradient of climatic continentality: Western Sayan Mountains, southern Siberia

Southern Siberian mountain ranges encompass strong climatic contrasts from the relatively oceanic northern foothills to strongly continental intermountain basins in the south. Landscape-scale climatic differences create vegetation patterns, which are analogous to the broad-scale vegetation zonation over large areas of northern Eurasia. In their southern, continental areas, these mountains harbour forest types which potentially resemble the full-glacial forests recently reconstructed for Central Europe. To identify forest vegetation–environment relationships in the southern Siberian mountain ranges, forest vegetation of the Western Sayan Mountains was sampled on a 280 km transect running from the northern foothills with oceanic climatic features to the continental Central Tuvinian Basin in the south. Based on the species composition, vegetation was classified into hemiboreal forests, occurring at drier and summer-warm sites with high-pH soil, and taiga, occurring at wetter, summer-cool sites with acidic soil. Hemiboreal forests included Betula pendula-Pinus sylvestris mesic forest, Larix sibirica dry forest and Pinus sylvestris dry forest. Taiga included Abies sibirica-Betula pendula wet forest, Abies sibirica-Pinus sibirica mesic forest and Pinus sibirica-Picea obovata continental forest. Hemiboreal forests were richer in vascular plant species, while taiga was richer in ground-dwelling cryptogams. Vegetation–environment relationships were analysed by indirect and direct ordination. Winter and summer temperatures and precipitation exerted a dominant influence on species composition. Soil pH was also an important correlate of species composition, but this factor itself was probably controlled by precipitation. At a more local scale, the main source of variation in species composition was topography, producing landscape patterns of contrasting plant communities on slopes of different aspects and valley bottoms. The response of tree species to major environmental factors was expressed with Huisman–Olff–Fresco models. Larix sibirica appeared to be most resistant to drought and winter frosts, Pinus sibirica was adapted to low temperatures both in winter and summer, and Picea obovata had an intermediate response to climate. Betula pendula, Pinus sylvestris and Populus tremula were associated with the warmest sites with intermediate precipitation, while Abies sibirica was the most moisture-demanding species, sensitive to deep winter frosts.     

Climate and vegetation in China IV. Distribution of tree species along the thermal gradient

The aim of this study was to examine correlations between vegetational zones and thermal climate on the basis of analysis of the thermal distribution of main-component trees in each zone. The whole and main thermal distribution ranges for 73 tree species were obtained from their distributions along the warmth index (WI) gradicnt. Cluster analysis using the WI values at the upper and lower limits of the main distribution ranges as parameters gave seven species groups. The geographical distributions of the species in each group were described. Moreover, the frequency distribution curves of the seven species groups were determined along the WI grandient, and four borderlines (lines I, II, III and IV) were found among them. The WI values at the lines I, II and IV were 50, 90, 175°C month, respectively, and corresponded to boundaries between the subarctic and cool-temperate zones (I), the cool-and warm-temperate zones (II) and the warmtemperated and subtropical/tropical zones (IV), respectively. Line III corresponded to a WI of 135°C month. The WI values at the forest limit and at the lower limit of the subarctic forest were estimated, and their averages were 15.2 and 49.6°C month, respectively. In conclusion, thermal climatic zones in China could be divided by the following WI values: alpine (arctic) zone, WI<15; subarctic zone, 15–50; cool-temperate zone, 50–90; warm-temperate zone, 90–175; subtropical/tropical zone >175°C month.     

Radial growth response of four dominant boreal tree species to climate along a latitudinal gradient in the eastern Canadian boreal forest

To address the central question of how climate change influences tree growth within the context of global warming, we used dendroclimatological analysis to understand the reactions of four major boreal tree species –Populus tremuloides, Betula papyrifera, Picea mariana, and Pinus banksiana– to climatic variations along a broad latitudinal gradient from 46 to 54°N in the eastern Canadian boreal forest. Tree‐ring chronologies from 34 forested stands distributed at a 1° interval were built, transformed into principal components (PCs), and analyzed through bootstrapped correlation analysis over the period 1950–2003 to identify climate factors limiting the radial growth and the detailed radial growth–climate association along the gradient. All species taken together, previous summer temperature (negative influences), and current January and March–April temperatures (positive influences) showed the most consistent relationships with radial growth across the gradient. Combined with the identified species/site‐specific climate factors, our study suggested that moisture conditions during the year before radial growth played a dominant role in positively regulating P. tremuloides growth, whereas January temperature and growing season moisture conditions positively impacted growth of B. papyrifera. Both P. mariana and P. banksiana were positively affected by the current‐year winter and spring or whole growing season temperatures over the entire range of our corridor. Owing to the impacts of different climate factors on growth, these boreal species showed inconsistent responsiveness to recent warming at the transition zone, where B. papyrifera, P. mariana, and P. banksiana would be the most responsive species, whereas P. tremuloides might be the least. Under continued warming, B. papyrifera stands located north of 49°N, P. tremuloides at northern latitudes, and P. mariana and P. banksiana stands located north of 47°N might benefit from warming winter and spring temperatures to enhance their radial growth in the coming decades, whereas other southern stands might be decreasing in radial growth.     

Habitat associations with topography and canopy structure of tree species in a tropical montane forest on Mount Kinabalu,Borneo

Habitat associations with topography and canopy structure of 42 abundant tree species were studied in a 2.74-ha plot of tropical montane forest on Mount Kinabalu, Borneo. Many of these species belong to the same higher taxa including eight families and four genera. Analysis of intraspecific spatial distributions for stems ≥ 10 cm diameter revealed that 28 species (including all six species of Fagaceae) showed aggregated distributions at the 100-m² and/or 400-m² scales, and that 20 species showed habitat associations with topography by torus-translation tests; 17 species showed both characteristics. Species' associations with the local canopy structure were characterized by crown position index (CPI), which was defined relative to neighbour trees. The CPI differed greatly among individual stems at 10–40 cm diameter, and 19 species showed significantly different frequencies of crowns exposed vertically versus those shaded beneath the canopy. Mean growth rates at 10–40 cm diameter and size distributions of species were not related to topographic associations, but were explained by the associations with canopy structure; species with more exposed crowns grew faster and had less positively skewed distributions. Diversity in habitat associations was manifest between two genera (Syzygium and Tristaniopsis) in the family Myrtaceae and among species in these genera, but was less evident in other families and two genera (Garcinia and Lithocarpus). This revised version was published online in June 2006 with corrections to the Cover Date.     

Mollusc community patterns and species response curves along a mineral richness gradient: a case study in fens

Aim The goals of this study were to: (1) compare water conductivity and pH as proxy measures of mineral richness in relation to mollusc assemblages in fens, (2) examine the patterns of mollusc species richness along the gradient of mineral richness based on these factors, (3) model species–response curves and analyse calcicole–calcifuge behaviour of molluscs, and (4) compare the results with those from other studies concerning non‐marine mollusc ecology. Location Altogether, 135 treeless spring fen sites were sampled within the area of the Western Carpathians (east Czech Republic, north‐west Slovakia and south Poland; overall extent of study area was 12,000 km²). Methods Mollusc communities were recorded quantitatively from a homogeneous area of 16 m². Water conductivity and pH were measured in the field. The patterns of local species diversity along selected gradients, and species–response curves, were modelled using generalized linear models (GLM) and generalized additive models (GAM), both using the Poisson distribution. Results When the most acid sites (practically free of molluscs) were excluded, conductivity expressed the sites’ mineral richness and base saturation within the entire gradient, in contrast to pH. In the base‐rich sites, pH did not correlate with mineral richness. A unimodal response of local species diversity to mineral richness (expressed as conductivity) was found. In the extremely mineral‐rich, tufa‐forming sites (conductivity > 600 μS cm^?1) a decrease in species diversity was encountered. Response curves of the most common species showed clear differentiation of their niches. Significant models of either unimodal or monotonic form were fitted for 18 of the 30 species analysed. Species showed five types of calcicole–calcifuge behaviour: (1) a decreasing monotonic response curve and a preference for the really acid sites; (2) a skewed unimodal response curve with the optimum shifted towards the slightly acid sites; (3) a symmetrical unimodal model response curve with the optimum in the base‐rich sites, with no or slight tufa precipitation; (4) a skewed unimodal response curve but with the optimum shifted to the more mineral‐rich sites; and (5) an increasingly monotonic response curve, the optimum in the extremely base‐rich sites with strong tufa precipitation. Main conclusions Conductivity is the only reliable proxy measure of mineral richness across the entire gradient, within the confines of this study. This information is of great ecological significance in studies of fen mollusc communities. Species richness does not increase with increasing mineral richness along the entire gradient: only a few species are able to dwell in the extremely base‐rich sites. The five types of calcicole–calcifuge behaviour seen in species living in fens have a wider application: data published so far suggest they are also applicable to mollusc communities in other habitats.     

Effects of selective logging on tree diversity,composition and plant functional type patterns in a Bornean rain forest

Abstract. The effects of selective logging on tree diversity, changes in tree species composition and plant functional types were studied with the use of seven permanent plots in virgin and in logged forest. All plots were located in a lowland dipterocarp rain forest in East Kalimantan on the island of Borneo. Just after logging and during the following 20 yr tree diversity measured as Fishers’α was not significantly affected in logged forest plots. Temporal shifts in tree species composition were analysed with Principal Component Analysis (PCA). Logged forest plots had much larger changes over time than virgin forest plots. In the smallest diameter class, some logged forest plots showed a distinct trajectory in PCA space compared to virgin forest plots, while in larger diameter classes movement of logged plots in PCA space was random. This suggests that there is no predetermined community to which logged forest plots tend to shift when recovering from logging. We found a significant negative correlation between diameter increments and the species‐specific wood densities of tree species. Species‐specific wood density and potential tree height were used to assign species to five PFTs. As expected, logging increased the fraction of softwood stems in small diameter classes. In the largest diameter classes (≥ 50 cm DBH) a strong decrease of softwood emergent stems was found in logged forest plots. After more than 20 yr no recruitment was found of softwood emergent stems in selectively logged forest.     

千岛湖次生林优势种种群结构与分布格局

为了揭示亚热带地区常绿阔叶林次生演替机理,以砍伐后自然恢复40余年的浙江千岛湖姥山岛次生林为研究对象,分析了该群落优势种的种群结构、存活曲线和空间分布格局。结果表明:1)群落中重要值最大的是马尾松(Pinus massoniana)种群,达到50.47%,其后依次是石栎(Lithocarpusglabra)、苦槠(Castanopsis sclerophylla)和青冈(Cyclobalanopsis glauca)。2)马尾松的种群结构呈纺锤型,但1、2径级个体数极少,幼龄个体储备严重不足,已呈衰退趋势;石栎、苦槠、青冈的种群结构呈金字塔型。3)从静态生命表看,马尾松种群小径级的死亡率为负值,也说明马尾松种群幼苗严重缺乏,种群呈衰退趋势。4)马尾松种群的存活曲线为凸型,石栎、苦槠和青冈种群则近似直线型。5)4种优势种种群总体和各级个体的空间分布格局均呈聚集分布。总体来看,马尾松种群为稳定型趋向衰退型,而石栎、苦槠和青冈种群为稳定增长型,群落会继续向针阔混交林方向演替,进而演替成为常绿阔叶林。     

长白山阔叶红松林主要树种对干旱胁迫的生态反应及生物量分配的初步研究

以组成长白山温带阔叶红松林主要树种红松,水曲柳,胡桃楸,椴树和蒙古柞的幼苗,盆载于土壤含水量分别为田间持水量的85％－100％（CK）,65％－85％（MW）和45％－65％（LW）条件下4个月,研究不同土壤水分亏缺对不同树种生态反应和生物量再分配,结果表明,干旱处理显著降低供试树种的单叶面积,侧根数,根长等,降低叶比重,增加根比重,不同种对干旱响应应存在明显差异,椴树和水曲柳叶比重随水分亏缺的加剧而显著降低,蒙古柞和胡桃揪叶比重值在轻度水分亏缺时达到最高,但不同树种幼树茎比重并不随土壤水分含量的降低而发生明显变化,各树种生物量较对照组有显著差异,椴树,水曲柳,胡桃揪,红松幼树总生物量平均降低32．8,43．6,26．2和23．8％,蒙古柞幼 树生物量平均增加4．8％,耐旱性比其它4个供试树种强。     

Conifer seedling recruitment across a gradient from forest to alpine tundra: effects of species,provenance, and site

Background: Seedling germination and survival is a critical control on forest ecosystem boundaries, such as at the alpine–treeline ecotone. In addition, while it is known that species respond individualistically to the same suite of environmental drivers, the potential additional effect of local adaptation on seedling success has not been evaluated.

Aims: To determine whether local adaptation may influence the position and movement of forest ecosystem boundaries, we quantified conifer seedling recruitment in common gardens across a subalpine forest to alpine tundra gradient at Niwot Ridge, Colorado, USA.

Methods: We studied Pinus flexilis and Picea engelmannii grown from seed collected locally at High (3400 m a.s.l.) and Low (3060 m a.s.l.) elevations. We monitored emergence and survival of seeds sown directly into plots and survival of seedlings germinated indoors and transplanted after snowmelt.

Results: Emergence and survival through the first growing season was greater for P. flexilis than P. engelmannii and for Low compared with High provenances. Yet survival through the second growing season was similar for both species and provenances. Seedling emergence and survival tended to be greatest in the subalpine forest and lowest in the alpine tundra. Survival was greater for transplants than for field-germinated seedlings.

Conclusions: These results suggest that survival through the first few weeks is critical to the establishment of natural germinants. In addition, even small distances between seed sources can have a significant effect on early demographic performance – a factor that has rarely been considered in previous studies of tree recruitment and species range shifts.     

Species Abundance in a Forest Community in South China：A Case of Poisson Lognormal Distribution

Abstract: Case studies on Poisson lognormal distribution of species abundance have been rare, especially in forest communities. We propose a numerical method to fit the Poisson lognormal to the species abundance data at an evergreen mixed forest in the Dinghushan Biosphere Reserve, South China. Plants in the tree, shrub and herb layers in 25 quadrats of 20 m× 20 m, 5 m× 5 m, and 1 m× 1 m were surveyed. Results indicated that: (i) for each layer, the observed species abundance with a similarly small median, mode, and a variance larger than the mean was reverse J-shaped and followed well the zero-truncated Poisson lognormal; (ii) the coefficient of variation, skewness and kurtosis of abundance, and two Poisson lognormal parameters (& and μ) for shrub layer were closer to those for the herb layer than those for the tree layer; and (iii) from the tree to the shrub to the herb layer, the α and the coefficient of variation decreased, whereas diversity increased. We suggest that: (i) the species abundance distributions in the three layers reflects the overall community characteristics; (ii) the Poisson lognormal can describe the species abundance distribution in diverse communities with a few abundant species but many rare species; and (iii) 1/α should be an alternative measure of diversity.
(Managing editor: Ya-Qin HAN)     

Physiological synecology of tree species in relation to geographic distribution and ecophysiological parameters at the Atlantic forest periphery in Brazil: an overview

This paper provides a synthesis of the three papers published in the series Ecophysiology of selected tree species in different plant communities at the periphery of the Atlantic forest of SE – Brazil (Trees, this issue). We reviewed data obtained for seven tree species, belonging to the legume family and the genus Clusia of the Clusiaceae. These groups were chosen because legumes are the most abundant tree species in the Atlantic forest complex (except in the open restingas), and the genus Clusia is reportedly of utmost ecological importance as nurse plants in several habitats of the complex. These species were surveyed along one or more of several forest and open habitats at the margin of the Atlantic rain forest (open restingas, restinga dry forest, inselberg, semideciduous dry forest) and indicated the following general patterns: (1) at the intraspecific level, ecophysiological performance often (but not always) varied largely in both time and space; (2) at the interspecific level, ecophysiological performance was often (but not always) related positively to species dominance in the community; and (3) at both levels, ecophysiological performance was not related to the geographic distribution patterns, i.e., generalists and specialists did not form groups of similarly behaving plants in regard to ecophysiology. We discuss the implications of these findings, compare them with earlier studies, and suggest that ecophysiological research in the Atlantic rain forest complex could enhance its explanatory power by increasing spatial and temporal scales of observation.     

Gap-phase dynamics and coexistence of a long-lived pioneer and shade-tolerant tree species in the canopy of an old-growth coastal temperate rain forest of Chiloé Island, Chile

Aim A major question with regard to the ecology of temperate rain forests in south‐central Chile is how pioneer and shade‐tolerant tree species coexist in old‐growth forests. We explored the correspondence between tree regeneration dynamics and life‐history traits to explain the coexistence of these two functional types in stands apparently representing a non‐equilibrium mixture. Location This study was conducted in northern Chiloé Island, Chile (41.6° S, 73.9° W) in a temperate coastal rain forest with no evidence of stand disruption by human impact. Methods We assessed stand structure by sampling all stems within two 50 × 20 m and four 5 × 100 m plots. A 600‐m long transect, with 20 uniformly spaced sampling points, was used to quantify seedling and sapling densities, obtain increment cores, and randomly select 10 tree‐fall gaps. We used tree‐ring analysis to assess establishment periods and to relate the influences of disturbances to the regeneration dynamics of the main canopy species. Results Canopy emergent tree species were the long‐lived pioneer Eucryphia cordifolia and the shade‐tolerant Aextoxicon punctatum. Shade‐tolerant species such as Laureliopsis philippiana and several species of Myrtaceae occupied the main canopy. The stem diameter distribution for E. cordifolia was distinctly unimodal, while for A. punctatum it was multi‐modal, with all age classes represented. Myrtaceae accounted for most of the small trees. Most tree seedlings and saplings occurred beneath canopy gaps. Based on tree‐ring counts, the largest individuals of A. punctatum and E. cordifolia had minimum ages estimated to be > 350 years and > 286 years, respectively. Shade‐tolerant Myrtaceae species and L. philippiana had shorter life spans (< 200 years). Most growth releases, regardless of tree species, were moderate and have occurred continuously since 1750. Main conclusions We suggest that this coastal forest has remained largely free of stand‐disrupting disturbances for at least 450 years, without substantial changes in canopy composition. Release patterns are consistent with this hypothesis and suggest that the disturbance regime is dominated by individual tree‐fall gaps, with sporadic multiple tree falls. Long life spans, maximum height and differences in shade tolerance provide a basis for understanding the long‐term coexistence of pioneer and shade‐tolerant tree species in this coastal, old‐growth rain forest, despite the rarity of major disturbances.     

Responses of native and invasive plant species to selective logging in an Acacia koa‐Metrosideros polymorpha forest in Hawai'i

Questions: Is the introduced timber species Fraxinus uhdei invasive in Hawai'i? Has logging disturbance facilitated the spread of Fraxinus and other alien species? Location: Windward Mauna Kea, island of Hawai'i. Methods: We surveyed 29 plots which were established before selective logging of the native tree Acacia koa in 1971 to determine if Fraxinus spread beyond the borders of an existing plantation and if other alien species increased. We created gaps in the canopy of the Fraxinus plantation and measured seed rain and regeneration, and we sampled foliar and soil nutrients inside and around the plantation. Results: Basal area of Fraxinus increased from 0.7 m².ha^‐1in 1971 to 10.8 m².ha^‐1 in 2000. Fraxinus was not found in plots that were located more than 500 m from those where it occurred in 1971 except along a road. Basal area of Acacia koa decreased after logging but subsequently recovered. Occurrence of the alien vine Passiflora tarminiana and alien grass Ehrharta stipoides decreased. Seedling regeneration of Fraxinus was prolific in gaps but did not occur under the canopy. Basal area of Fraxinus did not correlate with soil nutrient concentrations. Conclusions: Fraxinus was able to regenerate following logging more rapidly than native tree species. Basal area growth of Fraxinus was great enough to offset a decline in native trees and cause an increase in forest productivity. If the Fraxinus plantation is harvested, managers should plan ways of favoring regeneration of the native Acacia which is more valuable both for timber and for conservation.     

Species Abundance in a Forest Community in South China: A Case of Poisson Lognormal Distribution

Case studies on Poisson lognormal distribution of species abundance have been rare, especially in forest communities. We propose a numerical method to fit the Poisson lognormal to the species abundance data at an evergreen mixed forest in the Dinghushan Biosphere Reserve, South China. Plants in the tree, shrub and herb layers in 25 quadrats of 20 m×20 m, 5 m×5 m, and 1 m×1 m were surveyed. Results indicated that: (i) for each layer, the observed species abundance with a similarly small median, mode, and a variance larger than the mean was reverse J-shaped and followed well the zero-truncated Poisson lognormal;(ii) the coefficient of variation, skewness and kurtosis of abundance, and two Poisson lognormal parameters (σ andμ) for shrub layer were closer to those for the herb layer than those for the tree layer; and (iii) from the tree to the shrub to the herb layer, the σ and the coefficient of variation decreased, whereas diversity increased. We suggest that: (i) the species abundance distributions in the three layers reflects the overall community characteristics; (ii) the Poisson lognormal can describe the species abundance distribution in diverse communities with a few abundant species but many rare species; and (iii) 1/σ should be an alternative measure of diversity.