亚洲季风区铁杉属现代分布区及其气候特征

铁杉属在亚洲主要分布于亚洲季风区东部,间断分布于中国大陆、台湾岛以及日本列岛（除北海道）。在中国大陆间断分布在西南山区（横断山区和部分东喜马拉雅）、中部山区（秦岭-大巴山区）和东南山区（华东山地）。亚洲季风区铁杉属的现代分布环境要求年降水量范围为720～2103mm,生长季降水量为635～1489mm;年均温范围为5．8～18．2℃,冬季月均温为-2．7～11．5℃,年最冷月均温为-3．7-10．9℃,年最暖月均温为13．0—28．2℃,气温年较差为9．7～25．4℃。亚洲季风区铁杉属的现代地理分布与气候要素值相互关系的分析表明,无铁杉分布的朝鲜半岛及其邻近的我国东北山地和山东山地降水量均大于700mm,基本上能够满足铁杉属生长的水分条件;而朝鲜半岛及其邻近的我国东北山地和山东山地的气温年较差明显大于铁杉属分布区的值,同时两地的冬季月均温和年最冷月均温明显低于铁杉属分布区的值,这也是这些地区没有铁杉属分布的主要气候原因。总之,这些亚洲季风区水分条件能够满足铁杉属生长需要,但冬季温度（包括冬季月平均温度和年最冷月均温）和气温年较差无法满足铁杉属生长需要,导致这些地区无铁杉属现代分布记录。     

北美东部8种温带树种向北分布的限制气候因子

利用分布于北美东部的 8种温带落叶树种为研究材料 ,应用反映热量和降水状况的 11个变量为气候指标 ,用标准差分析方法分析了这 8个树种的分布北界与气候因子之间的关系。结果表明 ,在所使用的气候指标中 ,同一树种不同地点的温暖指数或年生物温度的标准差最小 ,说明积温是限制这些树种向北分布的主要气候因子 ;温暖指数和寒冷指数随着年降水量的增大而减小 ,寒冷指数随着年较差的增加而增加 ,温暖指数随着水热综合指数的增加表现出减小的趋势。表明 ,除了生长季节的积温之外 ,降水和大陆性等气候条件对温带树种向北分布也起着重要作用     

The Relation between Distribution of Edificator and Companion in Zonal Vegetation and Water-Temperature Condition in Northeast China

The relation between distribution of edificator and companion in zonal vegetation and water-temperature condition in the Northeast China was studied based on the method of Kira’s warmth index (WI), coldness index (CI) and the humidity index (HI) proposed by the author. The results are as follows: 1. The whole range, optimum range of warmth index and the value of humidity indexes of 40 species in the Northeast China have been determined. 2. The 40 species are divided into the six groups according to their warmth index. The species of each group have the same range of warmth index. 3. The upper limit of the forest in Changbai Mountain is in 2100 m above sea level, the WI value of which is 15 ℃·month. 4. The border line between the coniferous forest in the cold-temperate zone and the mixed coniferous-broad leaves forest in the temperate zone has the WI value of 45 ℃·month. The border line between the mixed coniferous-broad leaves forest in the temperate zone and the deciduous broad leaves forest in the warm-temperate zone has the WI value of 85 ℃·month. The border line between the forest and the grassland has the HI value of 7.5 mm/℃·month. The border line between the forest, grassland and the steppe has the HI value of 5.2 mm/℃·month.     

中国云杉林的地理分布与气候因子间的关系

为了揭示中国云杉林的地理分布与气候因子间的关系, 在中国云杉林15个群系的地理分布范围内选取613个地理坐标点, 其中包括云杉各个种分布的海拔上限和下限坐标点各235和228个。通过数字地球系统确定每个点的海拔高程, 从中国气象插值数据库获取每个点的气候数据。数据分析分别采用线性回归、变异系数比较和主成分分析(PCA)法。结果显示, 中国云杉林分布范围内, 年平均气温、最冷月平均气温、最热月平均气温、≥5 ℃积温、≥0 ℃积温、年降水量、土壤水分含量和干燥指数α的平均值分别是3.38 ℃、-9.75 ℃、14.78 ℃、1 227.83 ℃·d、2 271.19 ℃·d、712.23 mm、80.02%和0.50; 各气候因子与中国云杉林垂直分布的上下限间均具有显著的回归关系; 除了年平均气温和最冷月平均气温变异系数较大外, 其他6个气候因子的变异系数均较小, 且彼此间无显著差异; 无论是云杉分布的上限还是下限, ≥5 ℃积温和≥0 ℃积温在PCA第一主分量具有较高的载荷, 而年降水量和土壤水分含量在第二、三主分量具有较高的载荷。影响中国云杉林分布的主要气候因子是生长季节积温, 其次是年降水量和土壤水分含量。     

东喜马拉雅和横断山地区常绿硬叶栎类的气候封套与现代分布

Evergreen sclerophyllous oaks （the E.S. oaks, Quercus section Heterobalanus） are the dominant species of the local ecosystem in the eastern Himalaya and the Hengduan Mountains, southwest China. In this study, we document the climatic envelope of the seven E.S. oak species and examine the relationships between climate and their distribution. This was done using a principal components analysis （PCA） and multiple regression analysis （MRA） of nine climatic indices. The main climatic envelope of the E.S. oaks were： mean temperature of the warmest month （MTW）= 12.0-19.5℃, warmth index （WI） = 33.2-88.9℃ month, annual biotemperature （BT）=-6.9- -0.3 ℃, coldness index （CI）=-30.4- -10.1 ℃ month, mean temperature of the coldest month （MTC）=-3.7-3.0℃ and annual precipitation （AP）=701-897 mm at the lower limits; and MTW=8.3-16.1℃, WI=15.7-59.1℃ month, BT=3.6-8.9℃, CI=-55.4-19.3℃ month, MTC=8.3-16.1 ℃ and AP=610-811 mm at the upper limits. The climatic range of the E.S. oaks is wide and includes two climatic zones, the cool-temperature zone and the subpolar zone. The PCA and MRA results suggest that the thermal climate plays a major role and precipitation plays a secondary role in controlling the large-scale distribution of the E.S. oaks, except Quercus monimotricha. In thermal regimes, BT and/or MTW are most important for both lower and upper limits of the E.S. oaks. Furthermore, our results indicate that the upper distribution limits of the E.S. oaks are less determined by low temperatures and their duration （CI） than by other factors.     

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.     

Long-term climate record inferred from early-middle Pleistocene amphibian and squamate reptile assemblages at the Gran Dolina Cave, Atapuerca, Spain

The Gran Dolina cave site is famous for having delivered some of the oldest hominin remains of Western Europe (Homo antecessor, ca. 960 ka). Moreover, the evidence of lithic industries throughout the long vertical section suggests occupation on the part of hominins from the latest early Pleistocene (levels TD3/4, TD5, and TD6) to the late middle Pleistocene (level TD10). The Gran Dolina Sondeo Sur (TDS) has furnished a great number of small-vertebrate remains; among them some 40,000 bones are attributed to amphibians and squamates. Although they do not differ specifically from the extant herpetofauna of the Iberian Peninsula, the overlap of their current distribution areas (= mutual climatic range method) in Spain can provide mean annual temperatures (MAT), the mean temperatures of the coldest (MTC) and warmest (MTW) months, and mean annual precipitation (MAP) estimations for each sub-level, and their change can be studied throughout the sequence. Results from the squamate and amphibian study indicate that during hominin occupation the MAT (10-13 °C) was always slightly warmer than at present in the vicinity of the Gran Dolina Cave, and the MAP (800-1000 mm) was greater than today in the Burgos area. Climatic differences between “glacial” and “interglacial” phases are poorly marked. Summer temperatures (MTW) show stronger oscillations than winter temperatures (MTC), but seasonality remains almost unchanged throughout the sequence. These results are compared with those for large mammals, small mammals, and pollen analysis, giving a scenario for the palaeoclimatic conditions that occurred during the early to middle Pleistocene in Atapuerca, and hence a scenario for the hominins that once lived in the Sierra de Atapuerca.     

Climate and vegetation in China V. Effect of climatic factors on the upper limit of distribution of evergreen broadleaf forest

In order to clarify correlations between the upper limit of distribution of evergreen broadleaf forest and climatic factors, 62 stands distributed at the upper limit of the forest were collected from various parts of China, and their thermal and precipitation factors were estimated. Among six thermal climatic indices, i.e., warmth index (WI), coldness index (CI) and annual mean (AMT), January mean (JMT), mean minimum (MMT) and minimum (MT) temperatures, the CI at the stands showed the smallest range of variance, and it was therefore considered to be the most significant for interpreting the upward distribution of the forest. However, the distribution of the forest in mountain areas in southwestern China could not be explained by lower temperatures in winter such as CI but by a cumulative temperature such as WI. The continentality and precipitation factors were also important for delimiting the distribution of the forest. In addition, the relation between the distribution of the forest and the MMT was noted, and it was concluded that the MMT was an effective thermal index for explaining the upper limit of distribution of evergreen broadleaf forest in China.     

Stand structure and carbon metabolism of coppice forests at a heavy snowfall climate region of central Japan. I. Dynamics of stand structure during 14 years

The coppice stands under the climatic conditions of Warmth Index (WI)≧85°C·month, and Coldness Index (CI)≦−10°C·month could be classified into aQuercus serrata type andCarpinus types at a heavy snowfall climate region of central Japan. The latter types tended to be more distributed on steeper slopes rather than the former. An analysis of tree forms and growth rates of species could reveal the characteristic of this habitat and vegetation type. The growth rates of stem diameter of dominant species on a gentle slope were higher than ones on a steep slope. However, the dominant species on the steeper slope had plasticity to basal bend with sustentacular roots, which is one form of tolerance of heavy snowfall on steep slopes. Average mortality during 14 years was 26%, irrespective of the gradient of a slope. More than one-third of dead shoots were induced by direct damage, while cracked or broken pieces of a stem were from heavy snowfall. The shoot density at smaller diameter classes tended to be higher at a stand having high horizontal variance of canopy surface caused by basal bend of big shoots. A frequency distribution function of individual tree weight (w), f(w)=pw^−3/2, could be obtained at each coppice stand by statistical analysis. The value of a parameter p, a packing factor, tended to be higher at stands on steeper slopes consisting of largely bent shoots and was constant irrespective of the growth.     

壳斗科栎属青冈亚属的花粉形态及其系统学意义

对栎属青冈亚属17个种及炭栎花粉形态进行扫描电镜观察。结果表明:青冈亚属和炭栎花粉均为三拟孔沟,球形至长球形,(15～28.0)μm×(15.0～25.6)μm。花粉外壁纹饰可划分为3种:(1)蠕虫状(包括皱波状)、(2)聚合颗粒状、(3)棒状或小刺状。各纹饰在组间镶嵌分布。棒状及小刺状纹饰为栎属花粉较为原始式样,并为青冈亚属与高山栎组及部分巴东组(sect.Cerris)共有,推测其间具有较其它栎类更为接近的亲缘关系,可能为广义栎属中较为基础的类群,而与栎亚属sect.Quercuss.s.,Lobatae和Protobalanus关系较远。花粉形态对广义栎属系统分类价值仅适用于较高分类阶元。     

The Geographic Distribution of a Tropical Montane Bird Is Limited by a Tree: Acorn Woodpeckers (Melanerpes formicivorus) and Colombian Oaks (Quercus humboldtii) in the Northern Andes

Species distributions are limited by a complex array of abiotic and biotic factors. In general, abiotic (climatic) factors are thought to explain species’ broad geographic distributions, while biotic factors regulate species’ abundance patterns at local scales. We used species distribution models to test the hypothesis that a biotic interaction with a tree, the Colombian oak (Quercus humboldtii), limits the broad-scale distribution of the Acorn Woodpecker (Melanerpes formicivorus) in the Northern Andes of South America. North American populations of Acorn Woodpeckers consume acorns from Quercus oaks and are limited by the presence of Quercus oaks. However, Acorn Woodpeckers in the Northern Andes seldom consume Colombian oak acorns (though may regularly drink sap from oak trees) and have been observed at sites without Colombian oaks, the sole species of Quercus found in South America. We found that climate-only models overpredicted Acorn Woodpecker distribution, suggesting that suitable abiotic conditions (e.g. in northern Ecuador) exist beyond the woodpecker’s southern range margin. In contrast, models that incorporate Colombian oak presence outperformed climate-only models and more accurately predicted the location of the Acorn Woodpecker’s southern range margin in southern Colombia. These findings support the hypothesis that a biotic interaction with Colombian oaks sets Acorn Woodpecker’s broad-scale geographic limit in South America, probably because Acorn Woodpeckers rely on Colombian oaks as a food resource (possibly for the oak’s sap rather than for acorns). Although empirical examples of particular plants limiting tropical birds’ distributions are scarce, we predict that similar biotic interactions may play an important role in structuring the geographic distributions of many species of tropical montane birds with specialized foraging behavior.     

Modelling Mediterranean oak palaeolandscapes using the MaxEnt model algorithm: The case of the NE Iberia under the Middle Holocene climatic scenario

The Mediterranean Basin is a global biodiversity hotspot, and oak tree species play an important role in it. Since the beginning of the Holocene (∼11.4 kyr BP), the distribution of forests has not occurred uniformly, resulting in diverse types of vegetation landscapes. In this study, we used a maximum entropy algorithm (MaxEnt) to obtain the ecological niche model (ENM) of two sub-Mediterranean oak species, Quercus pubescens Willd. (pubescent oak) and Quercus ilex subsp. ilex (holm oak), both in the present day in the Iberian Peninsula and within a Middle Holocene (8.2–4.2 kyr BP) climatic scenario in the NE Iberian Peninsula. Moreover, we used the locations of Neolithic archaeological sites containing anthracological data to analyze the relationship between human occupations and oak habitats. Our results suggest that the two oaks have responded differently to the climatic conditions that have occurred, and show changes in both potential distributions. The palaeolandscape vegetation map shows a denser vegetation cover in the lowlands and a more open landscape in the highlands, with a higher dominance of Quercus pubescens in the septentrional areas, while Quercus ilex was more restricted to certain coastal areas. Temperature and precipitation factors, mainly seasonal climatic conditions, have had a greater impact on the distribution of vegetation than other factors. We found a good overlap between the ENM of the two oaks and the locations of the Neolithic sites analysed, and determined that the distribution of Neolithic archaeological sites is not random. The Neolithic populations in the study area depended heavily on the resources of the deciduous and evergreen sub- Mediterranean forest, although they also exploited the resources of the mountain pine forest. Neolithic sites distribution suggests that Neolithic human groups were aware of the potential of forests and probably gathered woody resources in their surroundings.     

Evaluation of Endophytic Bacteria as Potential Biological-Control Agents for Oak Wilt

Endophytic bacteria were isolated from surviving live oaks (Quercus fusiformis) in Texas, where oak wilt is epidemic, and evaluated as potential biological control agents for the disease. Of the 889 bacterial isolates tested, 183 showed in vitro inhibition of the pathogen, Ceratocystis fagacearum. Six isolates were further evaluated for colonization of containerized Spanish oaks (Quercus texana) and live oaks. In general, when injected into the stems, Bacillus species colonized Spanish oaks more extensively than Pseudomonas species; whereas, the opposite was true for live oaks. In containerized live oaks inoculated with the oak wilt pathogen, preinoculation with Pseudomonas denitrificans 1-15 reduced the number of diseased trees by 50% and in the same trial decreased the percentage of crown loss by 17%. In a second trial, no reduction in numbers of diseased trees was observed; however, preinoculation with either strain 1-15 or Pseudomonas putida 5-48 significantly reduced crown loss. These studies indicate that a potential exists for the utilization of selected bacteria to control oak wilt.     

Phylogeography and evolution of two closely related oak species (<Emphasis Type="Italic">Quercus</Emphasis>) from north and northeast China

Orographic and climatic influences during the Pleistocene have had a crucial role on interspecific divergence and population demography during speciation. However, associations between demographic histories of closely related species and related climatic events, especially in north and northeast China, are still underexplored. Genetic analyses with four chloroplast DNA and two nuclear genes and species distribution modeling were used for two closely related oak species (Quercus liaotungensis and Quercus mongolica) to test if their interspecific divergence and phylogeographical histories were possibly related to the Pleistocene-era climatic events. Potential divergence of the two oak species was estimated at about 0.92–2.15 Ma. Species distribution models and genetic data showed varying phylogeographical histories and spatial population structures between the two oaks, leading to different patterns of interspecific gene flow between the chloroplast and nuclear genes. The results indicate that speciation event between the two species is recent and may have been triggered by geological and climatic fluctuations linked to the upheavals of the Qinghai-Tibetan Plateau at the Pliocene/Pleistocene boundary. The two closely related oaks possess varying population demography during the interglacial-glacial climatic oscillations of the Quaternary, probably due to the various niche adaptations among different distribution ranges across their species trajectories.     

中国栎属(壳斗科)基于叶表皮及叶结构特征的分支分析

以三棱栎为外类群,基于叶表皮及叶结构特征对中国栎属进行初步的分支分析,结果显示了栎属内稳定的“支”结构及其种间关系。栎亚属可划分为5个组,分别为巴东栎组,高山栎组,Jiang子栎组,麻栎组和槲栎组,Jiang子栎组是栎亚属常绿栎类与落叶栎类之间的过渡类群;高山栎组应为栎亚属常绿栎类中的特化类群,而非最原始类群,5个组中,仅巴东栎组不是单系类群,而是一个多系类群,青冈亚属较栎亚属早分支,可划分为3大类;无毛类,简单被毛类和复杂被毛类;无毛类最原始,复杂被毛类与栎亚属关系最近,因此,叶表皮及叶结构特征对解决栎亚属组间亲缘关系有重要的分类学意义。     

Spread and potential host range of the invasive oak lace bug [Corythucha arcuata (Say, 1832) – Heteroptera: Tingidae] in Eurasia

1. The North American oak lace bug feeds on leaves of ‘white oaks” in its native range. In Europe, it was first discovered in northern Italy in 2000. In recent years, it has subsequently spread rapidly and population outbreaks have been observed in several European countries. In the present study, we summarize the steps of its expansion.
2. To predict its potential host range, we checked 48 oak species in 20 sentinel gardens in seven countries between 2013 and 2018.
3. In total, 27 oak species were recorded as suitable hosts; 13 of them are globally new ones, 23 out of the 29 in section Quercus (～ white oaks, an intrageneric taxonomic unit within genus Quercus), including Asian oaks, native to Japan, Korea and China, and four out of five in section Cerris (another intrageneric unit of the same genus), were accepted as hosts. None of the species in section Lobatae (red oaks) or in the Ilex group was accepted.
4. Host records were also collected in forest stands of 10 countries. We found 11 oak species that were infested. Outbreak populations were most commonly found on Quercus robur, Quercus frainetto, Quercus petraea and Quercus cerris, comprising widespread and outstandingly important oaks species in Europe.
5. Based on our findings, we conclude that suitable hosts for oak lace bug are present in most of Europe and Asia. This means that a lack of hosts will likely not restrict further range expansion.

     

Function of defensive volatiles in pedunculate oak (Quercus robur) is tricked by the moth Tortrix viridana

The indirect defences of plants are comprised of herbivore‐induced plant volatiles (HIPVs) that among other things attract the natural enemies of insects. However, the actual extent of the benefits of HIPV emissions in complex co‐evolved plant‐herbivore systems is only poorly understood. The observation that a few Quercus robur L. trees constantly tolerated (T‐oaks) infestation by a major pest of oaks (Tortrix viridana L.), compared with heavily defoliated trees (susceptible: S‐oaks), lead us to a combined biochemical and behavioural study. We used these evidently different phenotypes to analyse whether the resistance of T‐oaks to the herbivore was dependent on the amount and scent of HIPVs and/or differences in non‐volatile polyphenolic leaf constituents (as quercetin‐, kaempferol‐ and flavonol glycosides). In addition to non‐volatile metabolic differences, typically defensive HIPV emissions differed between S‐oaks and T‐oaks. Female moths were attracted by the blend of HIPVs from S‐oaks, showing significantly higher amounts of (E)‐4,8‐dimethyl‐1,3,7‐nonatriene (DMNT) and (E)‐β‐ocimene and avoid T‐oaks with relative high fraction of the sesquiterpenes α‐farnesene and germacrene D. Hence, the strategy of T‐oaks exhibiting directly herbivore‐repellent HIPV emissions instead of high emissions of predator‐attracting HIPVs of the S‐oaks appears to be the better mechanism for avoiding defoliation.     

壳斗科的地质历史及其系统学和植物地理学意义

在收集整理现有壳斗科化石资料的基础上,讨论了壳斗科及其各属的起源时间、地史分布和地史 演替过程以及这些化石资料在系统学和植物地理学上的意义。白垩纪尚无壳斗科可靠的大化石记录, 微化石需要进一步研究才能确定亲缘关系以及古新世壳斗科已经分化出两个类群。从以上这些事实推 论壳斗科起源于白垩纪晚期,而壳斗科现代各属出现的时间应不晚于古新世。最早发现的壳斗科化石和现代栗亚科和水青冈亚科在形态结构上非常相似,这一事实表明,壳斗科分为两个亚科的观点更接近客观事实。在水青冈亚科中,三棱栎类的化石最早出现;在栎属中,青冈亚属更接近祖先类群;在地史中全缘栎类较具齿栎类出现早,粗齿的落叶栎类出现最晚。三棱栎属、栲属和石栎属的化石在老第三纪出现于北美和欧洲的事实说明,北美、欧洲和东亚在老第三纪时有一个相通的壳斗科植物区系。南美的三棱栎是通过北美进入南美的。中国横断山、欧洲地中海沿岸和北美西北部有一类形态特征相似、亲缘关系相近的硬叶栎类,它们之间有相同的地质演替历史,它们现代分布边界可能就是古地中海的边界。美洲的栎类有两个来源,常绿硬叶栎类是通过古地中海沿岸而经北美-欧洲陆桥到达的,落叶栎类则是在中新世以后通过白令海峡到达的。     

Fossils of the Fagaceae and their implications in systematics and biogeography

The fossil history of the Fagaceae from China and its systematic and biogeographic implications are discussed based on revisionary studies of the fossil records. No creditable macrofossil record of the Fagaceae exists in the Cretaceous deposits and all the Cretaceous microfossil reports remain equivocal and require further study. The Paleocene fossils show the appearance and diversification of the two groups corresponding to the subfamilies Fagoideae and Castaneoideae sensu Nixon. By the Eocene, all modern genera had been present. The oldest fagaceous fossils represent subfamily Fagoideae with affinities to the extant genus Trigonobalanus. The leaf fossil genus Berryophyllum, with affinities to Quercus subg. Cyclobalanopsis, has been documented by the early Eocene and might have occurred earlier than other fossils assignable to Quercus. The appearance of evergreen sclerophyllous Ouercus with entire leaves might have occurred earlier than those with toothed leaves. Deciduous, urticoid-leaved oak fossils (Quercus subg. Quercus sect. Quercus) had not appeared until the Miocene. Fossil equivalents of Trigonobalanus, Castanopsis and Lithocarpus had occurred in Europe and North America by the early Tertiary, suggesting that continuous distributions were achieved via the northern hemisphere land bridges. Three groups of evergreen sclerophyllous oaks of apparent close phylogenetic relationships occurred in the Hengduan mountains, the Mediterranean area and northwestern North America. Their fossil forms have become dominant elements of those vegetation zones since the Miocene. A shared fossil history indicates a possible biogeographic boundary formed by the ancient Mediterranean. The evidence suggests that the oaks might arrive in North America during two distinct geologic periods: evergreen sclerophyllous entire-leaved oaks appeared by the Early Tertiary, whereas thedeciduous oaks with urticoid leaves appeared in the Late Tertiary.     

An altitudinal transect study of the vegetation on Mount Kinabalu,Borneo

A quantitative transect analysis of altitudinal sequences of forest canopy species from 600 to 3400 m asl on Mt. Kinabalu (4101 m), Borneo, resulted in four discrete altitudinal vegetation zones. These were made up of mutually exclusive species groups for lowland (<1200 m asl), lower montane (1200 to 2000–2350 m asl), upper montane (2000–2350 to 2800 m asl), and subalpine (2800 to the forest line, 3400 m asl) zones. Zonal soil types were correlated with the vegetation zones. In upslope sequence, these were: lowland Oxisols, montane Histosol/Spodosol complex, and subalpine Inceptisols. The highest contents of organic carbon, extractable phosphorus, and exchangeable magnesium and potassium were recorded in the lower and upper montane zones. The upper boundaries of the lowland, upper montane and subalpine zones coincided with thermal thresholds of latitudinal bioclimatic zones: 18°C TMIN (Köppen's tropical), WI 85 (Kira's warm temperate), and WI 45 (Kira's cool temperate), respectively. The upper limit of the lower montane zone was correlated with an abrupt increase of water surplus estimated from the annual rainfall minus annual potential evaporation. These climatic characteristics appear to define ecological altitudinal turnover points, so called critical altitudes, where groups of associated species are displaced by other groups.Abbreviations asl = above sea level - DBH = diameter at breast height - PHQ = Park headquarters - TMAX = Mean daily maximum air temperature - TMIN = Mean daily minimum air temperature - TWINSPAN = Two-way indicator species analysis - WI = Warmth index