应用GIS和最大熵模型分析秦岭冷杉潜在地理分布

秦岭冷杉(Abies chensiensis)为松科常绿针叶乔木,属国家濒危野生物种。本文利用获取的秦岭冷杉地理分布记录和14个气候指标、3个土壤指标和3个地形指标,通过最大熵模型(Max Ent)和Arc GIS空间分析构建秦岭冷杉潜在分布区。结果表明,影响秦岭冷杉潜在生境适宜性的主要环境指标为6个气候指标(年平均气温、年极端最高气温、1月平均气温、≥0℃积温、年干燥度指数和年日照时数)、1个土壤指标(p H)和1个地形指标(海拔)。研究区秦岭冷杉生长的最适宜区面积为19498.87 km2,适宜区面积为32219.61 km2,边缘适宜区面积为51874.76 km2,不适宜区面积为106307.97 km2,秦岭冷杉适宜生长区域狭窄,其中最适宜区和适宜区主要集中在甘肃东南部、陕西中南部、四川东北部,湖北西北部和河南西北部,适宜区内次生植被丰富、地形复杂。研究表明,基于最大熵模型与GIS空间分析构建的秦岭冷杉潜在地理分布,可以对我国秦岭冷杉适宜生长区做出科学的区划,为秦岭冷杉资源的保护和管理提供科学依据。     

濒危植物秦岭冷杉种群数量动态

为了对濒危植物秦岭冷杉种群数量动态评价和预测,通过样地调查和数据统计,研究了秦岭冷杉种群的年龄结构、静态生命表及其与环境因子关系,运用时间序列模型预测了种群数量动态.结果表明,多数秦岭冷杉种群幼龄级个体数较少,中老龄个体数量较大,呈衰退趋势.仅处于低海拔地区的秦岭冷杉-木蓝-苔草群丛中的种群(D种群)由于立地条件较好,幼龄级个体数量相对丰富,种群稳定.不同秦岭冷杉种群生命表和存活曲线的分析表明,尽管生境条件差异,但存活曲线基本接近DeeveyⅢ型;不同种群偏离典型存活曲线的程度与幼苗缺乏程度有关,一般Ⅲ～Ⅴ龄级死亡率较高.时间序列分析表明,在未来20、40和80年中,不同秦岭冷杉种群均会呈现老龄级株数先增后减的趋势,种群稳定性长期维持困难.对影响秦岭冷杉种群增长的10个环境因子通过主成分分析(PCA)发现,乔木层盖度、土壤有机质含量和空气湿度对种群发挥有利影响,而人为干扰和光照强度对秦岭冷杉种群增长发挥不利影响.应充分利用秦岭冷杉性喜荫、耐寒、种子活力较强的特点,加强现有林分就地保护,重点是具有结实能力的中老龄个体;在阴坡地带,对林下灌木比较密集的群丛,通过砍灌、清理林下活地被物等抚育措施,为幼苗发育创造良好的环境条件;就地采种育苗,扩大人工种群.     

青藏高原东缘林线杜鹃-岷江冷杉原始林的空间格局

为深入理解高山林线原始林中优势树种的空间结构特征和种间关系,以青藏高原东缘林线杜鹃-岷江冷杉原始林1 hm²样地调查数据为基础,采用成对相关函数g(r)函数,分析了优势种岷江冷杉(Abies faxoniana)和凝毛杜鹃(Rhododendron agglutinatum)各径级的空间分布格局和各径级间的空间关联性。结果表明:岷江冷杉和凝毛杜鹃径级分布连续,呈倒"J"型,均为进展种群。小尺度上,对于岷江冷杉活立木(小树+中树+大树)、小树、中树为显著聚集分布,大树为均匀分布,死树则为随机分布;凝毛杜鹃活立木(小树+中树+大树)、小树、中树、大树、死树均为显著聚集分布。小尺度上随径级增大岷江冷杉的分布格局从聚集分布趋于均匀分布。岷江冷杉大树与其小树的空间关联验证了Janzen-Connell假说中的距离制约效应。通过验证接受了随机死亡假说,发现种内的密度制约效应不明显。两个树种的枯立木对凝毛杜鹃小树均表现为排斥关系,而对岷江冷杉小树表现为相互独立的关系。两个优势种总体表现为相互独立,但种间不同径级间的空间关联性各异。     

秦岭地区15kaBP以来的植被与环境

通过对秦岭海拔2200m-山间盆地沼泽沉积剖面的花粉分析和^14C年代测定,结合对该区植被与表土花粉的研究,认为秦岭的暗针叶林带15ka BP以来植被和环境经历了4个发展阶段：(1)约15450～6760a BP,植被为冷杉(Abies)-云杉(Picea)林,气候寒冷潮湿;(2)约6760～2940a BP,沉积物中花粉稀少,以冷杉、云杉、桦(Betula)、栎(Quercus)为主,出现了喜暖型植物桤木(alnus),气候暖湿;(3)2940～40a BP,植被为冷杉林,气候冷湿;(4)0～40a BP,植被为桦、栎、榛(Corylus)、桤木、铁杉(Tsuga)和小片状分布的冷杉,松(Pinus)的数量极少,气候向暖干方向发展,这是近期人为活动干扰的结果。     

濒危植物秦岭冷杉结实特性的研究

秦岭冷杉 (Abieschensiensis) 是松科冷杉属常绿乔木, 为我国特有珍稀濒危植物。该文在秦岭冷杉分布最为集中的秦岭地区分高、低 2个海拔高度采集当年球果和种子, 通过野外观察、室内实验和对比分析, 研究秦岭冷杉的结实特征以及球果和种子的基本性状, 并探讨这些特征在不同海拔高度的差异。结果表明, 与冷杉属其它树种相比, 秦岭冷杉球果和种子较大、单果种子数较多, 但空粒多、饱满度低。自然状况下饱满度为 12.0 %, 人工目视分选后饱满度 2 1.6 %, 风选后可达到 38.1%, 其余种子大部分为空粒、涩粒, 有少量虫粒。随海拔高度增加, 秦岭冷杉天然群体成龄个体中结实母树比例降低, 结实量减少。秦岭冷杉总体结实母树比例为 5 5 %, 低海拔处结实母树比例较高, 为 76 %, 高海拔结实母树比例为 34%。秦岭冷杉单株母树的平均结实量为 113个, 结实量小于 10 0个的母树占 6 4.5 %, 其中 5 0 %的母树结实量在 5 0~ 10 0个, 结实量在 10 0~ 2 0 0个的占 16.1%, 2 0 0个以上的占 19.4 %。 2 0 0个以上球果的个体均生长在低海拔区间, 高海拔处结实量多在 10 0个以下。方差分析表明, 两个海拔高度间球果和种子特征差异是显著的, 绝大部分球果和种子性状指标的平均值在低海拔处大于高海拔处。因此, 从结实特性以及球果和种子的特征来看, 低海拔处秦岭冷杉生殖生长好于高海拔处。     

秦岭巴山冷杉(Abies fargesii)种群结构与动态

为了阐明秦岭巴山冷杉(Abies fargesii)种群的结构特征和动态规律,对位于秦岭中段南坡佛坪保护区内不同海拔分布的巴山冷杉种群的年龄结构、生命表以及存活曲线等进行了研究.结果表明: 在海拔2350、2500和2700 m,巴山冷杉种群都表现为衰退型,种群个体主要聚集在中龄级,幼龄级的个体都极度缺乏.在各海拔巴山冷杉种群生命表中,死亡量(dx)、死亡率(qx)和消失率(Kx)都为负值,反映了种群幼龄个体的缺乏程度.随着海拔的升高,幼龄级个体数量增加而大龄级个体数量减少.巴山冷杉种群生命表和存活曲线的分析表明,巴山冷杉种群个体存活量最多的年龄级和出现最大死亡率的年龄级都随着海拔的降低而依次后推.随着海拔的升高,巴山冷杉种群的分布格局由随机分布类型过渡为聚集分布类型.     

秦岭冷杉球果与种子的形态变异

在秦岭冷杉的天然分布区 ,分别选取核心区和边缘区进行取样 ,比较球果和种子的形态变异。核心区为秦岭地区 ,边缘区选在湖北神农架地区。根据秦岭冷杉分布特征 ,核心区沿海拔高度连续采样 ,边缘区选择两个不同海拔高度的种群 ,分单株采样 ,测量了球果、种子、种鳞、种翅等器官的 15个形态指标。相关分析表明 ,秦岭冷杉球果和种子形态特征 (变化 )与海拔高度显著相关 ,说明海拔因素影响着球果和种子形态变异。比较发现 ,核心区 (即秦岭地区 )的秦岭冷杉球果和种子的绝大部分指标均值显著大于边缘区 (即神农架地区 ) ,说明秦岭冷杉在核心区的生殖生长好于边缘区。相同海拔条件下 ,秦岭冷杉球果和种子形态在地区间差异显著 ,约 85 %的变异来自地区内个体间和个体内 ,说明秦岭冷杉的球果和种子形态特征变化受遗传控制更显著。在边缘区还比较了不同种群间、人工林和天然林在球果和种子形态特征的差异。球果和种子形态在同一地区两个种群间、在人工林与天然林间都有显著差异 ,大部分人工林球果和种子指标均值和变异幅度大于天然林 ,表明通过人为经营可以改善秦岭冷杉的一些生殖性状     

濒危植物秦岭冷杉生殖生态学特征

为了探索秦岭冷杉种群生殖生态学主要特征,通过样地调查、固定样地观测和室内实验分析,系统研究了秦岭地区5个秦岭冷杉（Abies chensiensis）种群生殖特性及其与环境因素的关系,结果表明,5个不同生境的种群都具有结实能力,海拔1500～1600m之间,秦岭冷杉球果的产量较高;各种群球果主要分布在树冠中上部和东、南、西方向;球果以中部产生饱满种子比率最高;秦岭冷杉种子千粒重为43．2g,其中具有生活力者占44．8％,种子的平均含水量为5．7％左右;在天然林散落条件下,种子库的萌发率仅为6．1％,霉坏、搬迁率93．9％;人工播种条件下,种子发芽率在苗圃内高于天然条件下,中低海拔高于高海拔地区。天然条件下,由种子转化成幼苗的过程是秦岭冷杉种群生活史的脆弱环节。秦岭冷杉林经营管理要以就地保护为主,促进种群生殖;要通过森林经营措施,为林下种子萌发和幼苗创造条件;种子脱落后,要及时扰动林下灌木、草本,使种子能够顺利落地,促其发芽成苗;种子大年要注意采种、育苗,扩大人工种群;速生丰产林培育应该以中低海拔的阴坡为主。     

秦岭细鳞鲑栖息地环境特征研究

为研究秦岭细鳞鲑（Brachymystax lenok tsinlingensis）的栖息地环境选择偏好,对陕西陇县秦岭细鳞鲑国家级自然保护区和甘肃秦州珍稀水生野生动物国家级自然保护区内25个样点进行了鱼类采样和生境测量。共采集到130尾秦岭细鳞鲑样本,样点的平均分布密度为（0.10±0.07）尾/m,两个保护区内秦岭细鳞鲑的密度存在显著性差异（P < 0.05）。与秦岭细鳞鲑密度分布呈正相关的因子依次为坡度、跌水区密度、海拔、粗糙度、遮蔽度、底栖动物生物量和流速,与密度呈负相关的环境因子依次为河宽、水深和溶氧。冗余性分析（RDA）筛选了坡度、粗糙度、遮蔽度、海拔和跌水区5个关键环境因子。基于关键环境因子的实测值建立了栖息地适合度曲线,秦岭细鳞鲑分布的最适坡度范围为5%-10%,海拔分布范围为1500-2000 m,粗糙度范围0.3-0.4;跌水区密度范围12-18个/100 m,遮蔽度在0.5以上。     

海拔梯度对长白山北坡岳桦水分利用效率的影响

以分布于长白山北坡海拔1800～2050 m的岳桦林为对象,以叶片碳稳定同位素(δ13C)值作为岳桦长期水分利用效率指示值,探讨海拔梯度对岳桦林水分利用效率的影响.结果表明:随海拔升高,岳桦林土壤体积含水量、比叶质量极显著增加,而叶片含水量和土壤温度显著降低;岳桦叶片δ13C值与海拔呈极显著正相关,增幅为1.013‰·(100 m)-1,与土壤体积含水量、比叶质量呈显著正相关,与生长季土壤平均温度、叶片含水量呈显著负相关.温度不是长白山林线的唯一限制性因子,海拔梯度上水热条件的差异及其交互作用可能造成岳桦生长期间的生理干旱.     

Study on reproduction ecology of endangered species Abies chensiensis

Qinling fir, Abies chensiensis, is endemic to China and has been listed in the China Plant Red Data Book as one of the second class conservation plants. It is found scattered only in the forested pockets of the Qinling, Bashan, and Shennongjia Mountains with altitude from 1 300 m to 2200 m in Shaanxi, Gansu, Henan, Sichuan, and Hubei Provinces. To conserve and effectively restore this species and explore its processes of reproduction, the production characteristics of cone and seed yields, seed bank dynamics, and seed germination rates of five Abies chensiensis populations (A, B, C, D, and E, respectively, in the Abies chensiensis-Pinus tabulaeformis-Sinarundinaria nitida-Carex lanceolata association, Abies chensiensis-Quercus aliena var. acutserrata-Litsea pungens-Carex lanceolata association, Abies chensiensis-Betula albo-sinensis-Sinarundinaria nitida-Duchesnea indica association, Abies chensiensis-Indigofera amblyantha-Carex lanceolata association and Abies chensiensis-Pinus tabulaeformis-Smilax stans-Carex lanceolata association) were studied in their natural habitats across 18 plots in the Qinling Mountain area. Experiments were carried out in laboratory and field conditions. The results showed that all the five populations had the potential to produce seeds, and higher yields occurred in the lower and middle altitude areas every three to five years. The majority of cones appeared on the upper and middle crown and the west, south, and east faces of the crown, and the majority of full seed appeared on the middle portion of the cone. The weight per 1000 seeds was 43.2 g, in which 44.8% had growing potential. The germination rate of seeds in the natural forest was 6.1% and the proportion of seeds lost or destroyed in different populations in the natural forest was 93.9%. The germination rate of seeds planted in the nursery was higher than that in the natural forest, and the germination rate in the lower or middle altitude populations was higher than that in the upper altitude populations. The course from seed to seedling was the critical period in the life cycle of A. chensiensis populations under natural conditions. Reproduction courses were influenced positively by an abundant coverage of tree layer, organic material in the soil, thick soil, and a dense population of parent trees, whereas they were influenced negatively by human disturbance and light. In situ conservation of A. chensiensis should be carried out in the future so as to promote expansion of the population. Thinning the shrub layer and grass layer will also help seeds to penetrate the soil, promoting seed germination. The artificial population should be expanded by collecting seeds in the harvest years. Fast-growing and high-yield plantations should be developed on cloudy slopes at lower or middle altitudes.     

Comparative analysis of genetic diversity and population genetic structure in Abies chensiensis and Abies fargesii inferred from microsatellite markers

Abies chensiensis Tieghem and Abies fargesii Franchet are two closely related tree species of Pinaceae endemic to China. A. chensiensis is usually found scattered in small forest fragments, whereas A. fargesii is a dominant member of coniferous forest. To evaluate the genetic effect of fragmentation on A. chensiensis, a total of 24 populations were sampled from the whole distribution of the two species. Seven nuclear microsatellite loci were employed to analyze comparatively the genetic diversity and population genetic differentiation. Both A. chensiensis and A. fargesii have high level within-population genetic diversity and low inter-population genetic differentiation. Low microsatellite differentiation (2.1%) between A. fargesii and A. chensiensis was observed. But microsatellite marker was able to discriminate most populations of these two species. Compared to A. fargesii, A. chensiensi has lower allelic diversity and higher genetic differentiation among populations. It suggested the existence of negative genetic impacts of habitat fragmentation on A. chensiensis.     

Species delimitation and phylogeography of the Abies chensiensis complex inferred from morphological and molecular data

Species delimitation in the Abies chensiensis complex, consisting of A. chensiensis, A. recurvata, A. ernestii and A. ernestii var. salouenensis, has been the subject of a long‐term dispute, as various taxonomic combinations have been proposed. We combined different lines of evidence, including morphological characters and mitochondrial and plastid DNA sequence data, to assess species delimitation. Plastid DNA sequence variation was highly consistent with the morphological result and separated A. recurvata from the other three taxa, but the mitochondrial DNA genealogy was more strongly related to geographical distribution rather than species delimitation, as proposed in previous studies. On the basis of morphological characters and plastid DNA sequence variation, the current species status of A. recurvata was well supported, and we also accepted A. ernestii and A. ernestii var. salouenensis as varieties of A. chensiensis. Furthermore, the phylogeographical history of A. chensiensis was surveyed by molecular data and ecological niche modelling. On the basis of molecular data, the current phylogeographical pattern of the A. chensiensis complex was probably shaped by recent rapid expansions from multiple refugia in the Hengduan Mountains and habitat contraction after the last glaciation. This hypothesis was supported by the results of ecological niche modelling and previous fossil records. Our findings suggest that these fir taxa might have continued to expand their range after the largest glaciation until the end of the last glaciation, but have contracted since then with the increasing temperature and habitat changes. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 175–188.     

Regeneration in subalpine coniferous forests

Death of canopy trees when gaps are formed was studied in a subalpine coniferous forest, central Japan, which was composed ofAbies, Tsuga, Picea, Betula, andSorbus. Typhoons were considered to be the most important cause of the death of canopy trees. The degree of disturbance in each of 16 plots (20 m×20 m) was represented by the percentage of the total basal area of dead trees to that of living and dead canopy trees (disturbance magnitude; MAG). The mortality of canopy trees increased as their dbh increase in the plots of lower MAG than 90%. The mortality varied among genera, andTsuga was characterized as having lower mortality than that of the other conifers. 418 dead trees were observed. The standing dead trees made up 10.7% of the trees, the stem broken trees 46.7%, and the uprooted trees 42.2%. The stem breaking was most frequent inAbies, and the uprooting was most frequent inTsuga, Picea, andBetula. Undeveloped forests, which have the L-shaped dbh distribution, were destroyed only in high degree (70%<MAG), while developed forests were destroyed in various degrees (30%<MAG<100%). The percentage of uprooted trees in basal area decreased with the development of the forest, from 60% to 10%.     

Structure of a pristine Picea abies forest in northeastern Europe

Abstract. The forest structure in a large, relatively homogeneous area of pristine Picea abies taiga in the southern boreal region west of the Ural mountains was studied along four 500-m long transects. P. abies dominated the forest in association with Abies sibirica and Betula spp. The mean volume of living trees was 216 m³/ha. This value varied among the four transects, from 138 - 252 m³/ha. Mean density of trees (DBH > 1 cm) (and variation over the transects) was 2 064/ha (1670 - 2710). Living trees classified as dying or seriously damaged made up 2.9 (2.5 - 3.5) % of the living tree volume. Betula was an important canopy component and made up 16% of the living tree volume. The number of dead standing trees varied from 195 - 325/ha, corresponding to a volume of 10.8 - 70.7 m³/ha. The density of trees with a broken stem was 90 - 170/ha and their estimated volume 7.6 - 41.3 m³/ha. Standing dead trees and trees with broken stems represented 10.4 and 8.9% of the total standing tree volume (living + dead), respectively. The mean volume of decaying logs on the forest floor was 117 (84.4–156.2) m³/ha, corresponding to 54 (35 - 113) % of the living tree volume. The canopy-forming trees were present in the understory as abundant saplings and suppressed individuals. The size distribution of the dominant tree species resembled the inverse J-shape. Generally, the forest was characterized by a high small-scale structural variation and a larger-scale relative homogeneity. This pattern is consistent with forest dynamics where the forest consists of a small-scale mosaic of patches in different stages of recovery following disturbance. Our results suggest that the ecology and dynamics of extensive areas of natural boreal forests can be driven by small-scale disturbance.     

Increased genetic divergence between two closely related fir species in areas of range overlap

Because of introgressive hybridization, closely related species can be more similar to each other in areas of range overlap (parapatry or sympatry) than in areas where they are geographically isolated from each other (allopatry). Here, we report the reverse situation based on nuclear genetic divergence between two fir species, Abies chensiensis and Abies fargesii, in China, at sites where they are parapatric relative to where they are allopatric. We examined genetic divergence across 126 amplified fragment length polymorphism (AFLP) markers in a set of 172 individuals sampled from both allopatric and parapatric populations of the two species. Our analyses demonstrated that AFLP divergence was much greater between the species when comparisons were made between parapatric populations than between allopatric populations. We suggest that selection in parapatry may have largely contributed to this increased divergence.     

The influence of savanna trees on nutrient,water and light availability and the understorey vegetation

In an East African savanna herbaceous layer productivity and species composition were studied around Acacia tortilis trees of three different age classes, as well as around dead trees and in open grassland patches. The effects of trees on nutrient, light and water availability were measured to obtain an insight into which resources determine changes in productivity and composition of the herbaceous layer. Soil nutrient availability increased with tree age and size and was lowest in open grassland and highest under dead trees. The lower N:P ratios of grasses from open grassland compared to grasses from under trees suggested that productivity in open grassland was limited by nitrogen, while under trees the limiting nutrient was probably P. N:P ratios of grasses growing under bushes and small trees were intermediate between large trees and open grassland indicating that the understorey of Acacia trees seemed to change gradually from a N-limited to a P-limited vegetation. Soil moisture contents were lower under than those outside of canopies of large Acacia trees suggesting that water competition between trees and grasses was important. Species composition of the herbaceous layer under Acacia trees was completely different from the vegetation in open grassland. Also the vegetation under bushes of Acacia tortilis was different from both open grassland and the understorey of large trees. The main factor causing differences in species composition was probably nutrient availability because species compositions were similar for stands of similar soil nutrient concentrations even when light and water availability was different. Changes in species composition did not result in differences in above-ground biomass, which was remarkably similar under different sized trees and in open grassland. The only exception was around dead trees where herbaceous plant production was 60% higher than under living trees. The results suggest that herbaceous layer productivity did not increase under trees by a higher soil nutrient availability, probably because grass production was limited by competition for water. This was consistent with the high plant production around dead trees because when trees die, water competition disappears but the high soil nutrient availability remains. Hence, in addition to tree soil nutrient enrichment, below-ground competition for water appears to be an important process regulating tree-grass interactions in semi-arid savanna.     

Pythium undulatum,cause of root rot of Abies procera Christmas trees and Pseudotsuga menziesii in Northern Germany

A devastating disease of Abies procera grown as Christmas trees in a forest situation in Northern Germany is reported. Disease symptoms began as a root rot with disintegration of the cortex of lateral and main roots, followed by rapid chlorosis and necrosis of the foliage which had a striking copper colour on dead and dying trees. All disease symptoms were reproducible in pathogenicity tests with potted A. procera and Pseudotsuga menziesii plants infected with the isolated pathogen. The symptoms were indistinguishable from those reported for Phytophthora spp., but the pathogen was identified by microscopy and ITS sequence analysis as Pythium undulatum, which has not previously been described as a cause of root rot of coniferous trees in outdoor situations. This fungus was isolated also from roots of A. amabilis, A. grandis and P. menziesii showing similar below-and above-ground symptoms. Trees of varying ages (6–22 years) were affected. Abies nordmanniana appeared not to be susceptible to P. undulatum even when in direct root contact with diseased A. procera or A. amabilis. The establishment and spread of the disease between 1999 and 2002 appeared to be correlated with poor soil drainage following a series of unusually wet summers.     

Early responses of bark and wood boring beetles to an outbreak of the hemlock looper Lambdina fiscellaria (Guenée) (Lepidoptera: Geometridae) in a boreal balsam fir forest of North America

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