Correlations between leaf structural traits and the densities of herbivorous insect guilds

The functional composition of herbivorous insect assemblages was correlated with aspects of new and mature leaf surface features, anatomy and morphology across 18 co‐occurring plant species. Multivariate analyses of insects and leaf traits revealed that the functional composition of the herbivore assemblage was more strongly correlated with leaf structural traits than with leaf constituents. Leaf traits were more strongly correlated with the functional composition of the herbivore assemblage than with its taxonomic composition. Densities of sessile phloem feeders, ­rostrum chewers, and all herbivores were significantly negatively correlated with specific leaf weight, lamina and cuticle thickness, vascular tissue depth and stomate length, and were significantly positively correlated with stomate density. External chewer densities were significantly negatively correlated with percent lignified vein area, and significantly positively correlated with leaf surface area and the distance between lignified tissues. Spine‐like leaves were associated with significantly lower densities of sessile phloem feeders, external chewers and all herbivores compared to kite leaves (kite leaves are comprised of unfortified leaf tissue supported by a framework of vascular tissue). The presence of a thickened leaf hypodermis was associated with significantly lower densities of external chewers and rostrum chewers, while midrib protection was associated with significantly lower densities of external chewers. Leaf structural traits may not be the proximal factors influencing herbivorous insects, as leaf structural traits are correlated with many other plant traits such as photosynthetic rate, relative growth rate and leaf life‐span. Nonetheless, these results indicate that certain leaf structural traits may potentially be used to predict the functional structure of herbivorous insect assemblages. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77 , 43–65.     

Fossil Plants and Their Assemblages from the Early Carboniferous in Guangdong

This paper deals with the Lower Carboniferous fossil plants, 19 genera and 41 species including 10 new species, such as Sublepidodeneron wcwgtanense, Lepidodendron sophoroides, Adiantites lianpingensis, A. matouensis, Archaeopteridium shaoguanense, Rhacopteris angusta, Rhodeites lanceolata, Neuropteris shaoguanensis, Potoniea turbinata and P. racemicarpa. According to the distributive regularity of those fossils in stratigraphic section of Early Carboniferous from Guangdong area, the authors propose to divide the flora into three assemblages from lower to upper, namely, 1, Sublepidodendron mirabile assemblage; 2. Cardiopteridium spetsbergense-Adiantites gothanii assemblage; 3. Archaeopteridium-Sphenopteris obtusiloba assemblage, which separately represent the dispositions of Dahu Formation (the 1st assemblage), the lower part of Ceshui Formation (the 2nd assemblage) and upper part of Ceshui Formation or middle part of Zhongxin Formation (the 3rd assemblage).     

Aspects of carbonate sedimentation in the Ordovician of Baltoscandia

In the Baltoscandian epicontinental sea the normal type of Ordovician carbonate sediments was a mixture, in various proportions, of skeletal sand and carbonate mud. In some marginal areas sediments of Bahaman type were deposited during the latest part of the Viruan Epoch, and such sediments were widely distributed from the very latest Ordovician onwards. It is suggested that deposition mostly took place in a temperate climatic zone and that the bahamitic deposits indicate an increase of temperature to subtropical or tropical level.
The succession of the Ordovician lithofacies belts in Baltoscandia is mostly roughly symmetrical with regard to grain size, but asymmetrical as to the distribution of terrigenous mud. The position of the boundary between the terrigenous and carbonate mud belts may have been a result of competition in the material transport from the west and from the east. The direction of the lithofacies belts is not parallel to the axis of the Caledonian geosynclinal zone. The predominance of carbonate sediments in the Baltoscandian Ordovician epicontinental sea probably does not reflect particularly good conditions for production of carbonate material, but rather, a very low supply of terrigenous matter.
In the Central Baltic area changes of sea level were probably largely opposite to those in Västergötland. This indicates differential isostatic movements in the Baltoscandian cratonic shield during parts of the Ordovician Period.     

河北唐山西域山晚寒武世长山组的树形笔石

本文对东秦岭南部湖北郧西地区早石炭世四射珊瑚进行了研究,系统描述了11科中的24属和39种,其中包含2新属和21新种。本区早石炭世四射珊瑚拟划分为4个组合带:范家坪组上部Palaeosmilia-Lithostrotion组合带,下部Yuanophyllum组合带。袁家沟组上部Shennongia majus组合带,下部Zaphrentoides组合带。范家坪组是新建的组名。文中也讨论了Cystophrentis的分布时限,及其与Shennongia的亲缘关系。此外,Neoclisiophyllum在本区与岩关期的Shennongia共生,出现的时间早于华南各地。     

Stratigraphy and microfacies of the oligocene sequence at Gabal Bu Husah, Marada Oasis, South Sirte Basin, Libya

Summary Stratigraphic and microfacies investigations carried out on the Oligocene sequence exposed at Gabal Bu Husah, northwest of Marada Oasis, south of the Sirte Basin, Libya, showed that the Oligocene sequence conformably overlies the Late Eocene Thamat Formation and unconformably underlies the Early to Middle Miocene Marada Formation (Qarat Jahannam Member). The lithostratigraphic studies of the Oligocene sequence revealed the presence of two rock units, from base to top: 1- Umm Ad Dahiy Formation (Early Oligocene, Rupelian) and 2- Bu Hashish Formation (Late Oligocene, Chattian). The Oligocene sequence yields a rich foraminiferal assemblage with fifty-one benthonic and large formaminiferal species. The foraminiferal analysis allowed to subdivide the sequence into three local foraminiferal assemblage zones, arranged stratigraphically from base to top: 1)Elphidium minutum zone comprising the Umm Ad Dahiy Formation. 2)Miogypsinoides complanatus/Nonion granosus assemblage zone covering the lower two thirds of the Bu Hashish Formation. 3) Zone with abundantNummulites spp., including the upper part of the Bu Hashish Formation. The paleoenvironmental significance of the recorded species is described and discussed. The microfacies studies led to the recognition of eleven microfacies types. These microfacies indicate that the lower part of the Umm Ad Dahiy Formation was deposited in a shallow warm marine environment, but the conditions changed to fluviomarine in the uppermost part. The Bu Hashish Formation was deposited in a shallow-marine, inner-shelf environment (as indicated by the operculinid limestone) but a probable hiatus in its middle part is indicated by the presence of a nummulitic coquina and gypsum beds formed in a lagoonal environment. After returning to a shallow marine environment at the end of Oligocene the marine sedimentation ended.     

Eocene fossil rodent assemblages from the Erlian Basin (Inner Mongolia,China): Biochronological implications

Rodents from the Nuhetingboerhe-Huheboerhe area in the Erlian Basin, Inner Mongolia, differ stratigraphically: primitive ctenodactyloids such as Chenomys and Yuanomys are dominant in the upper part of the Nomogen Formation; Tamquammys dominates in the Arshanto Formation; Asiomys, Pappocricetodon, and Yuomys appear in the lower part of the Irdin Manha Formation and Tamquammys is rarer in this formation than in the Arshanto Formation. The assemblage of the upper part of the Nomogen Formation is similar to that of the Lingcha Formation of Hunan, the Wutu Formation of Shandong, the Yuhuangding Formation of Hubei, and the Bumban Member of the Nara-Bulak Formation of Mongolia. The assemblage in the upper part of the Arshanto Formation is correlated with that from the locality Andarak 2 in Kyrgyzstan. The assemblage from the lower part of the Irdin Manha Formation resembles that of the lower part of the Hetaoyuan Formation of Henan.On the basis of the comparison of the rodent assemblages, I consider that the age of upper part of the Nomogen Formation corresponds to the Bumbanian land mammal age. The Bumbanian, Arshantan, and Irdinmanhan land mammal ages are correlated respectively to the early Ypresian, the middle–late Ypresian, and the early Lutetian of the Geological Time Scale. The Bumbanian and Irdinmanhan land mammal ages are also correlated to the early Wasatchian and the early Uintan (or the later Bridgerian) of the North American Land Mammal Ages.     

西藏晚白垩世晚Campanian期至早Maastrichtian期菊石群——中国最年轻的菊石群

记述产自西藏南部仲巴县的一个中国目前所知最年轻的菊石群 ,含 2科 4属 6种 ,其中 4新种。据菊石群面貌和它们所产出的层位可以划分出上、下两个组合带 :上部Pachydiscuscf.hidakaensis Libycoceraszhongbaense组合带 ;下部Manambolitescujiangdingensis M .pivaeteaui组合带。产菊石的曲贝亚组的下段和上段的中、下部(82F5 0及其以下层位 )为晚Campanian期 ,上段上部 (即 82F5 2及其以上层位 )为早Maastrichtian期。     

The <Emphasis Type="Italic">Laluzia armini</Emphasis> (gen. et spec. nov.) ecosystem: understanding a deeper-water rudist lithosome from the Early Maastrichtian of Mexico

In the Lower Maastrichtian Cardenas Formation exposed at La Luz (State of San Luis Potosí, east-central Mexico), a shallowing-upwards mixed-clastic-carbonate sequence is exposed. The sequence passes from marls with thin siltstones, through lower and upper hippuritid-rudist-dominated intervals, and into a Durania-dominated interval. This succession shows an increase in grain size upwards and a progressive reworking of rudists upwards (preserved in life position, other than in distinct tempestites, in the lower part; invariably reworked/toppled in the upper part). Epibionts show a change from a serpulid–bryozoan assemblage in the lower sequence to a red algal–rhodolith assemblage in the upper part. Using these data, we argue that the sequence shows a change from a low-energy, relatively deep-water, nutrient-rich environment with low-light intensity in the lower part, to a high-energy, well-lit environment in the upper part. Two rudist species are present: a new multifold hippuritid rudist with cellular outer shell layer, Laluzia armini, that lacks pallial canals in its free valve and has a unique myocardinal pillar arrangement, in the lower part; and Durania in the upper part. Laluzia was adapted to low-energy, low-light, soft-bottom environments that were abundant within the lower part of the sequence exposed in the La Luz section—a very unusual environment for rudists.     

Ordovician radiolaria and conodonts from the peri-Gondwanan Karatau-Naryn microplate (Sarydzhaz, eastern Kyrgyzstan)

Cambrian-Ordovician sedimentary sequences crop out extensively in the area surrounding the Sarydzhaz River, a remote mountainous region situated in the eastern part of Kyrgyzstan. These sequences, composed essentially of fine siliciclastic lithologies, were deposited on a passive margin of the peri-Gondwanan Karatau-Naryn microplate. Palaeontological constraints for the age of these sequences are rare. Recent efforts of geological mapping in the area discovered reasonably well-preserved radiolarian and conodont faunas. The occurrence of conodonts was mentioned previously, but no taxonomic details were ever reported. Two conodont assemblages are identified from the Oldzhobai Formation; the first is dominated by Paracordylodus gracilis, which can be correlated with the upper Tremadocian to lower Floian; the second assemblage is more diverse and characterized by Prioniodus elegans, suggesting a mid Floian age. This is the first discovery of Ordovician Radiolaria from Kyrgyzstan. In spite of its rather moderately good to poor preservation, the fauna is interesting in many respects, especially due to the presence of a new radiolarian species (Inanigutta (?) kyrgyza nov. sp.). The latter is easily identifiable by the distinctive pore pattern of its cortical shell and has therefore the potential to become in the future a biostratigraphic marker species.     

澳大利亚Otway盆地白垩纪介形类化石的发现及其意义

一个白晚垩世的介形类化石组合首次被发现在Otway盆地的钻孔中。化石产于Eumeralla组的灰色砂泥岩中。Eumeralla组是依据孢粉材料而确定其时代为早白垩世,即相当于Barremain-Albian阶。介形类化石包括10属：Cytherelloidea sp.1,Bairdia cf.austracretacea bate,B?sp.juv.?,Eucythere?sp.,Cydocypris sp.,Darwinula cf．contracta Mandelstam,D.? sp.1,Cyprois sp.,Talicypridea sp.,Zigiphocypris sp.,Eacypris sp.及Candona sp.。它们代表一中、新生代混生的面貌;其时代应为晚白垩世。依据生态的差异,介形类化石又可分为两个不同的组合：海陆混合相和淡水－半咸水相。前者包括Cytherelloidea和Bairdia见于Crayfish 1A钻孔中,而后者Darwinula,Ziziphocypris和Cyclocypris产于Penola 1和Kkatnook 2钻孔中。海相属Cytherelloidea和Bairdia发现在540.4-543.1m(E umeralla组井深1957－477m在Crayfish 1A钻孔中）,代表着Eumeralla组上部发生的一个晚白垩世海侵。     

三峡水库蓄水后长江上游鱼类群聚沿河流-水库梯度的空间格局

研究对2005—2012年长江上游合江、木洞、万州和秭归江段的鱼类群聚结构进行了调查, 以分析三峡水库蓄水后长江上游鱼类群聚沿河流-水库梯度的空间格局。结果显示, 在三峡蓄水后, 在合江至秭归江段累计采集到土著鱼类368706尾, 合计132种, 隶属于17科, 其中鲤科鱼类为优势类群。沿河流-水库纵向梯度, 土著鱼类物种数下降而外来鱼类物种数增加。对应分析表明, 合江至秭归江段的鱼类群聚呈现出明显分化: 库区以上河段鱼类组成以流水性鱼类为主, 库区鱼类则以静水缓流型为主。指示物种分析进一步指出, 河流区鱼类以犁头鳅(Lepturichthys fimbriata)、红唇薄鳅(Leptobotia rubrilabris)、圆口铜鱼(Coreius guichenoti)、圆筒吻(Rhinogobio cylindricus)等22种鱼类为指示物种, 其功能群特征表现为偏好流水生境、生活史为均衡主义及食性为昆虫食性; 库区鱼类以宽口光唇鱼(Acrossocheilus monticolus)、宽鳍鱲(Zacco platypus)、鳊(Parabramis pekinensis)、鳙(Aristichthys nobilis)等16种鱼类为指示物种, 其功能群特征表现为偏好静水生境、食性为草食性或浮游食性。以上研究表明, 三峡水库蓄水导致的水环境变化是影响长江上游鱼类纵向格局的主要驱动力。建议相关管理部门根据不同河段鱼类群聚特征制定不同的渔业管理措施, 如保护土著鱼类资源、控制外来入侵鱼类。     

准噶尔盆地侏罗纪介形类新知

准噶尔盆地侏罗纪介形类已由众多中外地层古生物工作者进行过数十年的研究并有系统成果问世。本文仅介绍两个侏罗系化石点,作为前人工作之补充,充实已有化石记录,也提出了进一步划分的建议和依据。     

Soil factors influencing ferric hydroxide plaque formation on roots ofTypha latifolia L

The amount of ferric hydroxide plaque deposited onTypha latifolia roots varied between wetlands submerged throughout the growing season. Plaque formation was positively correlated with extractable iron in the substrate and pH, and negatively correlated with the percent organic matter and percent inorganic carbonates in the soil. All the above correlations were significant but weak, and in a stepwise regression analysis these four soil factors accounted for 72% of the variation in plaque deposition. Plaque formation was unrelated to soil texture.     

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.     

拉萨地区林周盆地设兴组上部(古近系)年代研究的新进展及意义

拉萨地区林周盆地典中那玛剖面设兴组上部新发现的孢粉组合以落叶、阔叶植物为主体,主要为桦科的Alnipollenites,Betulaepollenites,Carpiniptes,山毛榉科的Quercoidites,胡桃科的Juglanspollenites,榆科的Ulmipollenites,椴科的Tiliapollenites等,孢粉化石多为古近纪常见分子,未发现白垩纪的特征分子。古近纪早期大量出现的三孔沟,网面三孔沟等花粉少量出现;孢粉组合更接近于古近纪中晚期的面貌,其时代可能属于晚始新世。因此,设兴组上部的年代可能是始新世晚期。由此推论,设兴组和林子宗群之间的角度不整合不能代表白垩纪古近纪之间的构造运动,而是代表始新世晚期后的构造运动。同时,林周盆地可能沉积有古近纪的河湖相地层。     

Distribution of recent benthic foraminifera on the Sunda Shelf (South China Sea)

In this study we investigate the species composition and spatial distribution patterns of Rose Bengal stained and unstained benthic foraminifera from the central part of the Sunda Shelf in the south-western South China Sea in relation to environmental factors. The uppermost centimetre of the surface sediments (> 150 μm) from 45 sites from inner (60 m) to outer shelf (226 m) water depths revealed 584 species including 443 stained species.The univariate analyses of individual species abundances and community parameters and next canonical correspondence analysis were used to relate the faunal data to a set of measured environmental parameters. Four biofacies recognised on the Sunda Shelf are most strongly correlated to water depth, primary production and sediment type of the habitat. The inner shelf biofacies (CCA cluster A), defined by Ammomassilina alveoliniformis and Asterorotalia pulchella, occurs in fine grained sediments classified as modern terrigenous mud in the region with the highest primary production values. The high-energy inner shelf biofacies (CCA cluster B), defined by Heterolepa dutemplei and Textularia lythostrota, occurs in modern terrigenous sand and silt dominated sediments, northeast from the Natuna Island. The high-energy outer shelf biofacies (CCA cluster C), defined by Cibicidoides pachyderma and Textularia bocki, is sandwiched between assemblages of biofacies D. It occurs in the region characterised by neritic relict sand. In the shallow-waters on the Sunda Shelf the relationship of benthic foraminiferal faunal composition to grain size of sediments indirectly signals the prevailing bottom hydrodynamic conditions. The dominance of the epibenthic foraminifera attached to bigger particles (e.g. Cibicides lobatulus, Planulina arimiensis) and much higher abundances of empty tests suggest greater current velocities northeast of Natuna Island. The outer shelf biofacies (CCA cluster D) is defined by Facetocochlea pulchra and Bulimina marginata. It occurs in an area covered with modern terrigenous silt and mud and is characterised by lower annual primary production, but seasonally influenced by weak upwelling.     

Reefal and mud mound facies development in the Lower Devonian La Vid Group at the Colle outcrops (León province, Cantabrian Zone, NW Spain)

In the locality of Colle (Cantabrian Zone, NW Spain), the upper part of the Valporquero Shale Formation (Emsian, La Vid Group) contains an interval of shales and marlstones (barren, greenish-grey shales and fossiliferous, greenish-grey or reddish shales/marlstones) with beds and packages of homogeneous and cross-bedded skeletal limestones. Metre-scale mud mounds and coral biostromes occur encased in the fossiliferous reddish and greenish-grey shale/marlstones, respectively, with the coral biostromes overlying conspicuous skeletal limestone bodies. These rocks were deposited on a carbonate ramp, ranging from above storm wave base for the cross-bedded skeletal limestones to below the storm wave base for the remaining deposits, organic buildups included. The vertical stacking of these facies and the occurrence of the two types of buildups are interpreted to reflect the interplay among several (possibly 4th and 5th) orders of relative sea-level variations, during a 3rd-order highstand. Coral biostromes occur in early 5th-order transgressive system tracts developed within late 4th-order highstand, and are interpreted to have thrived on a stable granular substrate (skeletal limestones) in non-turbid waters, being later aborted by the onset of muddy sedimentation. Biostrome features suggest that they developed under environmental conditions essentially different from those related to the sedimentation of their granular substrate. Mud mounds occur in 5th-order transgressive and early highstand system tracts tied to early 4th-order sea-level rise. Field relationships suggest that mud mounds grew coevally with muddy sedimentation, with high-frequency variations in carbonate vs. terrigenous mud sedimentation influencing their development.An erratum to this article can be found at     

龙苍沟国家森林公园7种花楸属植物的叶解剖特征及其环境适应性

采用石蜡切片法对四川省龙苍沟国家森林公园内7种花楸属（Sorbus）植物的叶解剖特征进行研究,探究其结构特征与生境的相关性。结果显示：7种植物的叶片均为典型的背腹叶;叶片厚度介于108.16~208.21 μm,种间差异极显著（P<0.01）;上表皮厚度均大于下表皮厚度,且复叶物种的下表皮细胞均有乳突;栅栏组织由1~2层细胞构成,仅多对西康花楸（S. prattii var. aestivalis）的栅海比（栅栏组织与海绵组织的厚度比）为1.93,其余6种植物的栅海比均小于1;中脉维管束均呈心型,为典型的外韧型维管束,种间中脉突起度存在极显著差异（P<0.01）。各解剖结构中,上、下表皮可塑性最大,在生境中具有较强的潜在适应能力;中脉可塑性最小,整体结构较为稳定。栅栏组织、海绵组织和中脉组织是7种植物中种间差异最大的解剖结构。叶解剖结构与生境因子的相关性分析表明,栅栏组织厚度、栅海比和紧密度与年降水量、最暖季降水和海拔正相关（P<0.05）,与季节性温差负相关（P<0.05）;中脉直径和突起度与季节性温差呈正相关（P<0.01）,与年降水量、最暖季降水和海拔正相关负相关（P<0.05）。叶解剖结构性状的适应性变化,体现了7种花楸属植物在龙苍沟国家森林公园的生存策略。     

Experiments on epibenthic predation in the Wadden Sea

Field experiments were designed to evaluate the role of predators in the Wadden Sea, small predators like shore crabs, shrimps and gobies, and large ones like flatfish and birds. Exclosures, maintained in aCorophium volutator bed, an eelgrass bed, in a sandy and a muddy flat, protected the infauna from such epibenthic predation. The resulting changes in the macrofauna were recorded and compared with an unaffected control area. In sandy and muddy flats of the lower intertidal zone, cages (mesh size5 mm) altered abundance and composition of the infauna almost entirely. Nearly all species achieved higher population densities than in the control area, and in addition, the number of species increased as well. In the the scarcely populated mud flat a dense suspension-feeder assemblage emerged, associated with numerous tube-building polychaetes. The sand flat, normally dominated by deposit feeders, also became occupied by a dense suspension-feeder assemblage, mainly cockles. In contrast, narrowly meshed cages had only little effect in the beds of eelgrass and ofCorophium volutator. Both are positioned in the upper intertidal zone. Although a number of species still responded with significant increases in abundance, many remained indifferent or even tended to be less abundant within cages. Cages provided with a 20-mm mesh nylon net, excluding only birds, flatfish and the biggest crabs, increased significantly the survival of large-sized infaunal members. This was only apparent in the upper intertidal zone. It was concluded, that small sized epibenthic predators are the major determinants of the dynamic species abundance pattern of the lower intertidal flats. In the eelgrass bed, the meshwork of rootlets constitutes an important spatial refuge from these predators. The complex habitat structure causes a diversified faunal assemblage. TheCorophium bed in the uppermost intertidal zone is less accessable to predators like crabs, shrimp and gobies. The monotonous appearance of this faunal assemblage is assumed to be the outcome of competitive exclusion and of occasional harsh physical conditions.     

海南岛热带山地雨林演替后期幼树的叶片特征

植物的生理生态特征决定了植物在生态系统中的分布模式和物种的丰度。本文在开展样地调查的基础上,应用Li-6400光合测定系统研究了海南岛热带山地雨林原始林3个层次的6个优势种与9个伴生种幼树的光合作用参数,并测定了相应叶片的比叶面积（SLA）和氮、磷含量。15个种的SLA为38.2~143.7 cm²·g^-1、单位面积最大光合速率（A_-area）为2.77~7.61 μmol·m^-2·s^-1、单位干重最大光合速率（A_-mass）为21.2~83.4 μmol·kg^-1·s^-1,单位面积暗呼吸速率（R_d-area）为0.02~1.15 μmol·m^-2·s^-1、单位干重暗呼吸速率（R_d-mass）为0.19~12.61 μmol·kg^-1·s^-1,光补偿点（LCP）为2.5~32.2 μmol·m^-2·s^-1,表观量子效率（Ф）为0.006 6~0.042 8 μmol·μmol^-1 photon,叶片氮（LN）含量为7.98~23.63 mg·g^-1,叶片磷（LP）含量为3.98~13.40 mg·g^-1。中上层种的幼树具有较高的比叶面积、单位干重最大光合速率和表观量子效率;上层种的暗呼吸速率最低;幼树叶片的氮、磷含量随物种成树所在层次升高而减少,次序为下层种>中层种>上层种,但差异均不显著（P>0.05）。优势种和伴生种的光响应参数与SLA存在相关关系。伴生种的LN与SLA正相关（P<0.01）,LP与SLA负相关（P<0.05）;优势种的LN、LP与SLA不相关（P>0.05）。