Ecological and biogeographical studies on the tropical rain forest of south Yunnan,SW China with a special reference to its relation with rain forests of tropical Asia

Abstract. Ecological and biogeographic analyses of the tropical rain forest in south Yunnan were made using data from seventeen sample plots and floristic inventories of about 1000 species of seed plants. The rain forest is shown to be a type of true tropical rain forest because it has almost the same profile, physiognomic characteristics, species richness per unit area, numbers of individuals in each tree species and diameter classes of trees as classic lowland tropical rain forests. As the area is at the northern margin of monsoonal tropics, the rain forest differs from equatorial lowland rain forests in having some deciduous trees in the canopy layer, fewer megaphanaerophytes and epiphytes but more species of lianas as well as more species of microphylls. In its floristic composition, about 80% of total families. 94% of total genera and more than 90% of total species are tropical, of which about 38% of genera and 74% of species are tropical Asian. Furthermore, the rain forest has not only almost the same families and genera, but also the same families rank in the top ten both in species richness and in dominance of stems, as lowland forests in southeast Asia. It is indisputable that the flora of the rain forest is part of the tropical Asian flora. However, most of the tropical families and genera have their northern limits in south Yunnan and most have their centre of species diversity in Malesia. More strictly tropical families and genera have relatively lower species richness and importance compared with lowland rain forests in tropical southeast Asia. Thus, the flora also shows characteristics of being at the margin of the tropics. Based mainly on physiognomy and floristic composition the tropical rain forest of Yunnan is classified into two types, i.e. seasonal rain forest and wet seasonal rain forest, the latter is further divided into two subtypes, i.e. mixed rain forest and dipterocarp rain forest. From analysis of geographic elements it is also shown that the tropical rain forest of Yunnan occurs at a geographical nexus with its flora coming mainly from four sources, i.e. Malesia, south Himalayas, Indochina and China.     

云南热带雨林:特征、生物地理起源与演化北大核心CSCD

云南热带雨林具有与东南亚低地热带雨林类似的群落结构、生态外貌特征和物种多样性,是亚洲热带雨林的一个类型。它的植物区系组成中有90%的属和多于80%的种为热带分布成分,其中约40%的属和70%的种为热带亚洲分布型,它含属种较多的优势科和在群落中重要值较大的科也与亚洲热带雨林相似,是亚洲热带雨林和植物区系的热带北缘类型。云南西南部、南部与东南部的热带雨林在群落结构和生态外貌上类似,但在南部与东南部之间有明显的植物区系分异,它们经历了不同的起源背景和演化历程。云南的热带雨林在很大程度上由西南季风维持。喜马拉雅隆升导致西南季风气候形成和加强,在云南热带局部地区产生了湿润气候,发育了热带雨林植被。现在的云南热带雨林里或其分布地区有落叶物种或热带落叶林存在,这不仅是季节性气候的影响,推测在晚第三纪或第四纪更新世云南热带地区曾经历了干旱气候。云南热带雨林的分布主要受制于局部生境,并非地区性气候条件。     

云南热带季雨林及其与热带雨林植被的比较

在中国植物学文献中, 对热带季雨林的解释和运用是不一致的, 特别是易于把季雨林与热带雨林相混淆。季雨林是在具有明显干、湿季变化的热带季风气候下发育的一种热带落叶森林植被, 是介于热带雨林与热带稀树草原(savanna)之间的一个植被类型。云南的热带季雨林在分布生境、生态外貌特征、植物种类组成和地理成分构成上, 均与热带雨林有明显区别, 季雨林主要分布在海拔1 000 m以下的几大河流开阔河段两岸和河谷盆地, 其群落结构相对简单, 乔木一般仅有1至2层, 上层树种在干季落叶或上层及下层树种在干季都落叶; 在生活型组成上, 季雨林的木质藤本相对较少, 大高位芽植物及地上芽植物很少, 但地面芽植物很丰富, 地下芽植物和一年生植物也相对丰富; 在叶级和叶型特征上, 季雨林植物的小叶和复叶比例相对较高, 分别占到24%和44%; 在植物区系地理成分构成上, 季雨林的热带分布属合计也占绝对优势, 但以泛热带分布属的比例相对较高, 约占到总属数的30%, 热带亚洲至热带非洲分布属的比例也较高, 约占总属数的12%。季雨林的地理成分更为多样性, 起源与发展历史也更复杂和古老。     

滇东南马关古林箐热带雨林望天树群落的研究

云南东南部马关县古林箐的望天树群落以龙脑香科植物望天树为乔木层优势种,无患子科植物番龙眼为亚优势种,外貌以常绿大、中高位芽植物组成为特征,林内板根和茎花现象普遍,层间木质藤本和维管附生植物丰富,属于一种热带季节性雨林群落类型。在植物区系组成上,该群落以樟科、番荔枝科、楝科、大戟科、桑科、无患子科、橄榄科、柿树科等为主要组成科,在重要值统计上,以大戟科、无患子科、柿树科、龙脑香科、番荔枝科、樟科、楝科、橄榄科、桑科等为较优势的科,与东南亚热带雨林很接近,故为东南亚热带雨林的北缘类型。     

论滇南西双版纳的森林植被分类

本文基于多年研究成果的总结,对西双版纳森林植被的分类、主要植被类型及其特征进行了系统归纳,并讨论了它们与世界类似热带森林植被的关系。以群落的生态外貌与结构、种类组成和生境特征相结合作为植被分类的原则和依据,可以将西双版纳的热带森林植被分类为热带雨林、热带季节性湿润林、热带季雨林和热带山地常绿阔叶林四个主要的植被型,包括有至少二十个群系。热带雨林包括热带季节雨林和热带山地(低山)雨林二个植被亚型。热带季节雨林具有与赤道低地热带雨林几乎一样的群落结构和生态外貌特征,是亚洲热带雨林的一个类型,但由于发生在季风热带北缘纬度和海拔的极限条件下,受到季节性干旱和热量不足的影响,在其林冠层中有一定比例的落叶树种存在,大高位芽植物和附生植物较逊色而藤本植物和在叶级谱上的小叶型植物更丰富,这些特征又有别于赤道低地的热带雨林。热带山地雨林是热带雨林的山地亚型,是该地区热带山地较湿润生境的一种森林类型,它在植物区系组成和生态外貌特征上类似于热带亚洲的低山雨林,隶属于广义热带雨林植被型下的低山雨林亚型。热带季节性湿润林分布在石灰岩山坡中、上部,在群落外貌上类似热带山地常绿阔叶林但在植物区系组成上与后者不同,它是石灰岩山地垂直带上的一种植被类型。热带季雨林是分布在该地区开阔河谷盆地及河岸受季风影响强烈的生境的一种热带落叶森林,是介于热带雨林与萨王纳之间的植被类型。热带山地常绿阔叶林(季风常绿阔叶林)是西双版纳的主要山地植被类型,它分布在热带季节雨林带之上偏干的山地生境。它在植物区系组成上不同于该地区的热带季节雨林,在生态外貌特征上亦不同于热带山地雨林,是发育在受地区性季风气候强烈影响的热带山地的一种森林植被类型。     

云南中山湿性常绿阔叶林起源的探讨

云南中山湿性常绿阔叶林广泛分布于海拔1800~3400 m的山地,这类常绿阔叶林在生态外貌上具有亚热带常绿阔叶林特征,但其分布生境却是暖温带-温带气候,且植物区系组成与热带植物区系有古老的渊源。该文以研究得比较深入的哀牢山中山湿性常绿阔叶林为例,对其植物区系与其气候条件的不协调进行剖析。哀牢山的中山湿性常绿阔叶林记录了种子植物110科386属821种。在植物区系科的地理成分上,无论是含种数多的科,还是所有科的分布区类型,都是以热带分布科为主,体现了它的远古热带起源背景。在该种子植物区系中,热带分布属占总属数的47.75%,热带分布种占总种数的33.45%,在近代演化上仍体现出明显的热带亲缘特征。按照哀牢山中山湿性常绿阔叶林分布地区的暖温带-温带气候条件,它相当于中国东部的暖温带落叶阔叶林地区。在中国东部地区类似气候条件的地带性植被的植物区系中,热带分布属通常只占总属数的25%~27%,云南中山湿性常绿阔叶林植物区系的地理成分与所在地的温带气候明显不协调。对这种现象的解释,笔者认为结合哀牢山地区曾经是低海拔夷平面,在第四纪以来才迅速抬升形成的地质历史,可能这里曾经在第四纪以前的热带-亚热带性质的植物区系和常绿阔叶林随着地形的迅速抬升,逐渐适应了因海拔上升而形成的温带气候,成为在云南高海拔山地存在的所谓中山湿性常绿阔叶林,其植物区系和植被的热带-亚热带性质及常绿阔叶生态外貌并未因气候改变而发生改变。     

Extent and timing of floristic exchange between Australian and Asian rain forests

Aim We tested an entrenched concept – that the Australian rain forest flora is essentially a Gondwanan relict. We also assessed the role of regional‐level source–sink dynamics in the assembly of this flora. Location Eastern Australia. Methods To avoid potential biases inherent in selective studies undertaken to date, we used an analytical, whole‐of‐flora approach integrated with the fossil record. We identified disjunctions between woody Australian rain forest plant taxa and relatives on other land masses. To test the strength of the fossil evidence for the regional antiquity of this flora, we evaluated the proportion of these disjunct clades represented in the Australian fossil record, and to minimize the effects of biases in this record, we compared late Quaternary (i.e. late Pleistocene and Holocene, 126–0 ka), Pliocene and late Oligocene–early Miocene Australian pollen records interpreted as tropical rain forest. Using within‐species disjunctions as a proxy, we assessed the role of recent immigration from Asia into Australia. To assess the role of source–sink dynamics, we performed comparative analyses of disjunctions in major rain forest categories representing a north–south/climatic gradient. Results Southern Australian, cool temperate (microthermal) rain forests contain many floristic disjunctions with Gondwanan fragments and most of these clades have Gondwanan fossils. Disjunct clades in Australian mesothermal rain forest mostly occur in Asia/Malesia and a low proportion of these clades show pre‐Neogene records. Many clades in lowland tropical and ‘dry’ rain forest show disjunctions with Asia/Malesia and few have Australian fossil records. Rates of recent immigration from Asia/Malesia are high in these northern forests, and outweigh rates of recent emigration approximately nine‐fold. The late Quaternary fossil record has many more rain forest angiosperms than Oligocene–Miocene and Pliocene floras, consistent with extensive late Cenozoic immigration. Main conclusions The microthermal rain forests are largely Gondwanan relicts, but there is progressively greater, and more recent contribution from Asia/Malesia into more northern, and more lowland tropical rain forests. This variation reflects a strong gradient in geographic and ecological proximity between these forests and source floras in Asia/Malesia, and is consistent with a source–sink size model of immigration driven by late Cenozoic contractions and expansions of Australian rain forest.     

On the Classification of Forest Vegetation in Xishuangbanna,Southern Yunnnan

Xishuangbanna of southern Yunnan is a region of extremely interest to biologists and also a hotspot for biodiversity conservation . It is located in a transitional zone from tropical Southeast Asia to temperate East Asia biogeographically. The present paper reviewed vegetation types of Xishuangbanna and suggested a revised classification system based on theupdated study results over the last two decades . By combining physiognomic and floristic characteristics with ecological performances and habitats , the primary forest vegetation in Xishuangbanna can be organized into four main vegetation types: tropical rain forest, tropical seasonal moist forest, tropical montane evergreen broad-leaved forest and tropical monsoon forest. The tropical rain forest can be classified into two subtypes , i. e. tropical seasonal rain forest in the lowlands and tropical montane rain forest on higher elevations. The tropical seasonal rain forest in this region shows similar forest profile and physiognomic characteristics to those of equatorial lowland rain forests and is a type of world tropical rain forest. Because of conspicuous similarity on floristic composition , the tropical seasonal rain forest in Xishuangbanna is a type of tropical Asian rain forest . However , since the tropical seasonal rain forest occurs at the northern edge of tropical SE Asia, it differs from typical lowland rain forests in equatorial areas in maintaining some deciduous trees in the canopy layer , fewer megaphanerophytes and epiphytes but more abundant lianas and more plants with microphyll . It is a type of semi-evergreen rain forest at the northern edge of the tropical zone . The tropical montane rain forest occurs in wet montane habitats and is similar to the lower montane rain forests in equatorial Asia in floristic composition and physiognomy . It is a variety of lower montane rain forests at the northern tropical edges of tropical rain forests . The tropical seasonal moist forest occurs on middle and upper limestone mountains and is similar to the tropical montane evergreen broad-leaved forest of the region in physiognomy, but it differs from the latter in floristic composition. The monsoon forest in Xishuangbanna is a tropical deciduous forest under the influence of a strong monsoon climate and is considered to be a transitional vegetation type between tropical rain forest and savanna in physiognomy and distribution. The tropical montane evergreen broad- leaved forest is the main vegetation type in mountain areas . It is dominated by the tree species of Fagaceae , Euphorbiaceae , Theaceae and Lauraceae in majority. It differs from the tropical montane rain forests in lack of epiphytes and having more abundant lianas and plants with compound leaves . It is considered to be a distinct vegetation type in the northern margin of mainland southeastern Asia controlling by a strong monsoon climate, based on its floristic and physiognomic characteristics.     

滇南热带季雨林的一些问题讨论

通过分析云南南部的水、热条件及植被分布,讨论了季雨林植被类型及其特征,认为云南南部的季雨林是介于热带雨林与萨王纳之间的,在干季基本上是落叶的一种森林植被类型,符合Schimper (1903)定义的热带季雨林植被,并考虑它是一种生态学意义上的经向地带性植被,与该地区的纬向地带性植被热带季节雨林一起共同构成云南南部的水平地带性植被。在云南南部的石灰岩山坡分布的过去被认为是季雨林的森林植被,尽管也受到季节性干旱的影响而不同程度地具有落叶成分,但它在群落外貌上与典型的季雨林不相同,在植物区系组成上也明显不同于该地区非石灰岩山地的季风常绿阔叶林,在分布上亦是在石灰岩低山沟谷的热带季节性雨林水平地带性植被带之上,根据其生态外貌、植物区系组成和生境特点,我们建议用“热带季节性湿润林”来称呼这类石灰岩山地森林类型,在性质上属东南亚热带北缘石灰岩山地垂直带上的一种植被类型。     

Delving into the variations in tree species composition and richness across South American subtropical Atlantic and Pampean forests

Aims We analyse here the variations in species composition and richness and the geographic ranges of the tree species occurring in South American subtropical Atlantic and Pampean forests. Our goals were to assess (i) the floristic consistency of usual classifications based on vegetation physiognomy, climate and elevation; (ii) the leading role of temperature-related variables on the variations in species composition and richness; (iii) the predominance of species with tropical–subtropical ranges, possibly as a result of forest expansion over grasslands after the Last Glacial Maximum (LGM); (iv) the restriction of most subtropical endemics to stressful habitats as a possible result of past forest refuges during the LGM.Methods The region was defined by the Tropic of Capricorn to the north, the Rio de la Plata to the south, the Atlantic shoreline to the east and the catchment areas of the upper Paraná and Uruguay Rivers to the west. Multivariate analyses, multiple regression modelling and variance partition analyses were performed on a database containing 63 994 occurrence records of 1555 tree species in 491 forest sites and 48 environmental variables. All species were also classified according to their known geographic range.Important findings A main differentiation in species composition and richness was observed between the eastern windward coastlands (rain and cloud forests) and western leeward hinterlands (Araucaria and semi-deciduous forests). Pre-defined forest types on both sides were consistent with variations in tree species composition, which were significantly related to both environmental variables and spatial proximity, with extremes of low temperature playing a chief role. Tree species richness declined substantially towards the south and also from rain to seasonal forests and towards the highland summits and sandy shores. Species richness was significantly correlated with both minimum temperature and actual evapotranspiration. About 91% of the subtropical flora is shared with the much richer tropical flora, probably extracting species that can cope with frost outbreaks. The 145 subtropical endemics were not concentrated in harsher habitats.     

Beta‐Diversity of Termite Assemblages Among Primary French Guiana Rain Forests

Termites are an important component of tropical rain forests, and have been included in many studies focusing on the influence of human disturbance. Their distribution among primary rain forest has, however, rarely been investigated. Here we studied the termite fauna in seven mostly undisturbed forest sites, representing several rain forest types. Overall, approximately 70 percent of species were soil‐feeders and 25 percent were wood‐feeders, the remaining 5 percent being classified here as litter‐feeders. Termite species richness did not differ significantly among sites, but sites differed in termite abundance. The palm swamp and the low forest situated on the foothills of an inselberg, hosted different termite communities to the other sites. These two sites presented a singular physiognomy suggesting that forest type is an important factor influencing species composition. We found no correlation between termite species composition and distance between sites, highlighting that at the scale of our study (about 100 km), forest sites share a similar species pool.     

瑞丽莫里热带雨林种子植物区系的初步研究

初步分析了鲜为人知的滇西南瑞丽莫里的热带雨林植物区系组成与地理成分。该植物区系中热带和主产热带的科占总科数的80%以上,热带分布属占总属数的84.1%;典型热带分布种占总种数的82.1%,该区系在科、属和种水平上均以热带成分占优势,明显属于热带性质的植物区系。在其热带分布属中,又以热带亚洲分布属最多,占总属数的26.5%;典型热带分布种中也以热带亚洲分布及其变型的种占绝对优势,占总种数的72.9%,反映了该植物区系具有热带亚洲植物区系的性质特点。在其热带亚洲成分中,又具体以南亚—大陆东南亚成分比例最高,反映了滇西南的热带雨林植物区系由于地域邻接关系,受印度(喜马拉雅)—缅甸植物区系的强烈影响。     

砂仁种植对热带雨林植物多样性的影响探讨

通过对各种不同状况的林下种植砂仁的热带雨林片断与同样类型的原始热带雨林的比较研究 ,探讨了砂仁种植对热带雨林植物多样性各个方面的影响。砂仁种植必须大量清除小树、林下幼树、灌木层植物 ,致使种植了砂仁的热带雨林片断的乔木下层及几乎整个的幼树、灌木层遭严重破坏 ,物种和个体大量流失 ,群落结构变得非常不稳定。由于热带雨林的更新主要依赖于其林下层的幼苗库 ,林下层的幼树、苗库被清除或破坏 ,热带雨林就难以自我更新 ,故林下种植砂仁的更大的潜在危害是使得热带雨林不能自我更新。种植了砂仁的热带雨林片断 ,物种和个体数的减少或流失 ,主要是直接的人为清除的结果 ,清除强度大 ,种类和个体减少就多。热带雨林下砂仁种植后 ,那些在原乔木层中占多数的只有 1～ 2个个体的脆弱种类 (种群 )将首先消失     

海南岛霸王岭不同热带森林类型的种-个体关系

 比较分析了海南岛霸王岭自然保护区核心区热带低山雨林、山地雨林、云雾林、山地矮林等4种热带森林类型中不同大小径级树木的物种—个体关系。结果表明：各种植被类型中物种数与个体数对数成线性关系是一种普遍现象,且不受调查树木的径木级影响。在相同个体数的条件下,累积物种数随海拔增高而逐渐减少,也即物种数由热带低山雨林、山地雨林、云雾林到热带山地矮林逐渐减少。对于各种植被类型的种—多度关系,单个体、双个体的物种有相当高的比例,其后一般依个体数的增加,而逐渐降低物种的出现频率,呈典型的倒J型曲线,而且这一比例随树木径级的增加而增加。     

Tropical deciduous forest in Yunnan,southwestern China: Implications for geological and climatic histories from a little-known forest formation

In the southern mountain ranges of Yunnan province, China, deep valleys of several large rivers create rain shadows with hot dry summers, and are locally designated tropical; towards the north, notably in the Lancang (Upper Mekong) valley, these regions may experience frost during winter. The woody forest canopy of these valleys is predominantly deciduous, with evergreen elements in the north, where the canopy is open and the forest savanna-like. However, we here present tall forest with a closed deciduous canopy and semi-evergreen subcanopy observed in hot dry valleys of these rivers and their tributaries in the tropical south. The structure and physiognomy of these forests resemble the tall (moist) deciduous forest formation widespread in South Asia and Indo-Burma. Furthermore, these forests are largely composed of tropical elements at both the generic (80%) and the species level (>70%), indicating that these forests are indeed tropical. We originally hypothesized that these isolated forests represent refugia of a pre-Holocene extension of tall (moist) deciduous forest formation of South Asia and Indo-Burma. The sample plot we established to test this hypothesis confirmed that these forests share the structure and physiognomy of the tall (moist) deciduous forest formation; however, the plots also showed that these forests lack the characteristic and dominant species of the formation''s Indo-Burmese range. The tree flora, in particular, indicates that both deciduous and evergreen elements are instead mostly derived from the adjacent tropical semi-evergreen forests of tropical southern China; yet they also include an important endemic element, which implies that these forests have survived as refuges possibly since the Pliocene. The exceptional representation of evergreen elements in these forests indicates that they have rarely been subject to hot fires or domestic cattle browsing, adding to the unique nature of the forests and further justifying their strict conservation.     

Age and Long-term Growth of Trees in an Old-growth Tropical Rain Forest, Based on Analyses of Tree Rings and 14C1

In an old‐growth tropical wet forest at La Selva, Costa Rica, we combined radiocarbon (¹⁴C) dating and tree‐ring analysis to estimate the ages of large trees of canopy and emergent species spanning a broad range of wood densities and growth rates. We collected samples from the trunks of 29 fallen, dead individuals. We found that all eight sampled species formed visible growth rings, which varied considerably in distinctiveness. For five of the six species for which we combined wood anatomical studies with ¹⁴C‐dates (ring ages), the analyses demonstrated that growth rings were of annual formation. The oldest tree we found by direct ring counting was a Hymenolobium mesoamericanum Lima (Papilionaceae) specimen, with an age of ca. 530 years at the time of death. All other sampled individuals, including very large trees of slow‐growing species, had died at ages between 200 and 300 years. These results show that, even in an everwet tropical rain forest, tree growth of many species can be rhythmic, with an annual periodicity. This study thus raises the possibility of extending tree‐ring analyses throughout the tropical forest types lacking a strong dry season or annual flooding. Our findings and similar measurements from other tropical forests indicate that the maximum ages of tropical emergent trees are unlikely to be much greater than 600 years, and that these trees often die earlier from various natural causes.     

Geological History,Flora, and Vegetation of Xishuangbanna,Southern Yunnan,China1

Xishuangbanna of southern Yunnan is biogeographically located at a transitional zone from tropical Southeast (SE) Asia to subtropical East Asia, and is at the junction of the Indian and Burmese plates of Gondwana and the Eurasian plate of Laurasia. The flora of the region consists of a recorded 3336 native seed plant species, belonging to 1140 genera in 197 families, among which 83.5 percent are tropical genera and 32.8 percent are endemic to tropical Asia, suggesting a strong affinity to tropical Asian flora. The vegetation of Xishuangbanna is organized into four forest types: tropical rain forest, tropical seasonal moist forest, tropical montane evergreen broad‐leaved forest, and tropical monsoon forest. The tropical rain forest in Xishuangbanna has the same floristic composition of families and genera as some lowland equatorial rain forests in SE Asia, and is dominated (with a few exceptions) by the same families both in species richness and stem dominance. The exceptions include some deciduous trees in the canopy layer, fewer megaphanerophytes and epiphytes, and a higher abundance of lianas and microphyllic plants. We consider the tropical rain forest of Xishuangbanna as a type of tropical Asian rain forest, based on their conspicuous similarities in ecological and floristic characteristics.     

Studies in Neotropical paleobotany. XI. Late Tertiaryvegetation and environments of southeastern Guatemala: palynofloras fromthe Mio-Pliocene Padre Miguel Group and the Pliocene HerreriaFormation

Plant microfossils are reported from the late Tertiary Padre MiguelGroup and the Pliocene Herrería Formation of southeasternGuatemala. The most abundant palynomorphs are cf. Acrostichum(maximum 45%), Cyperaceae (29%), cf. Antrophyum(27%), monolete fern spores (16%), and Pinus(11%). Pollen grains of Picea, Juglans,Quercus, and Ulmus, primarily from the Padre Miguelflora, reveal the presence of a northern cool-temperate element. Theyounger Herrería flora is more lowland and warm temperate. Inthe absence of evidence for substantially higher elevations, thedifferences between the Neogene and modern vegetation are attributedmostly to climate. MAT (mean annual temperature) is estimated2°-3°C cooler than at present for the Padre Miguel flora,and ～3.5°C warmer for the Herrería flora. There islittle evidence for arid vegetation, and the tropical rain forest wasabsent or poorly represented. These data are consistent with those ofother fossil floras in the region and with trends suggested by isotopicpaleotemperature analysis and global sea-level changes. The Guatemalaassemblages further provide evidence that the current version of thetropical rain forest is recent in origin and has undergone considerablechange in its range and composition throughout the lateCenozoic.     

Wood Anatomy Reveals High Theoretical Hydraulic Conductivity and Low Resistance to Vessel Implosion in a Cretaceous Fossil Forest from Northern Mexico

The Olmos Formation (upper Campanian), with over 60 angiosperm leaf morphotypes, is Mexico''s richest Cretaceous flora. Paleoclimate leaf physiognomy estimates indicate that the Olmos paleoforest grew under wet and warm conditions, similar to those present in modern tropical rainforests. Leaf surface area, tree size and climate reconstructions suggest that this was a highly productive system. Efficient carbon fixation requires hydraulic efficiency to meet the evaporative demands of the photosynthetic surface, but it comes at the expense of increased risk of drought-induced cavitation. Here we tested the hypothesis that the Olmos paleoforest had high hydraulic efficiency, but was prone to cavitation. We characterized the hydraulic properties of the Olmos paleoforest using theoretical conductivity (K_s), vessel composition (S) and vessel fraction (F), and measured drought resistance using vessel implosion resistance and the water potential at which there is 50% loss of hydraulic conductivity (P₅₀). We found that the Olmos paleoforest had high hydraulic efficiency, similar to that present in several extant tropical-wet or semi-deciduous forest communities. Remarkably, the fossil flora had the lowest , which, together with low median P₅₀ (−1.9 MPa), indicate that the Olmos paleoforest species were extremely vulnerable to drought-induced cavitation. Our findings support paleoclimate inferences from leaf physiognomy and paleoclimatic models suggesting it represented a highly productive wet tropical rainforest. Our results also indicate that the Olmos Formation plants had a large range of water conduction strategies, but more restricted variation in cavitation resistance. These straightforward methods for measuring hydraulic properties, used herein for the first time, can provide useful information on the ecological strategies of paleofloras and on temporal shifts in ecological function of fossil forests chronosequences.