中国裸子植物的物种多样性格局及其影响因子

物种多样性的大尺度空间格局是宏观生态学和生物地理学研究的核心问题之一。本文利用中国裸子植物分布数据, 结合气候、地形等环境信息, 分析了中国裸子植物物种多样性的大尺度格局及其影响因素, 比较了不同类群之间物种多样性格局和主导因子的差异, 并探讨了裸子植物在植物区系中所占比重的地理格局。结果表明, 中国裸子植物的物种多样性总体上呈现南高北低的趋势, 物种多样性在横断山区最高。在裸子植物的三个主要类群中, 松柏亚纲的物种多样性格局与整体相似, 买麻藤亚纲的多样性高值区则出现在中国西北部的干旱地区, 苏铁亚纲的分布区较为狭窄, 主要集中在南方地区。线性回归分析结果表明, 空间异质性和降水因子对中国裸子植物多样性格局的解释率最高, 末次冰期以来的气温变化、海拔高差和能量因子次之。这表明中国裸子植物物种多样性的格局受到了多种因素的影响, 其中空间异质性和降水因子影响最大。进一步分析发现, 物种多样性格局的主导因子在不同类群之间具有显著差异, 这可能反映了这些类群的进化历史以及生理适应的差异。裸子植物与被子植物的比例具有明显的空间格局: 在东部、南部气候环境优越的地区, 裸子植物与被子植物的比例低于0.06; 而在西部、北部等气候环境比较恶劣的地区, 裸子植物的比例则显著上升。回归分析表明, 能量和水分因子显著影响了裸子植物与被子植物的比例。随着能量的降低和降水的减少, 裸子植物与被子植物的比例会显著升高, 这可能是由于被子植物在温暖湿润地区具有较强竞争优势, 但裸子植物对极端环境具有更好的适应。     

Tree height–diameter allometry across the United States

The relationship between tree height and diameter is fundamental in determining community and ecosystem structure as well as estimates of biomass and carbon storage. Yet our understanding of how tree allometry relates to climate and whole organismal function is limited. We used the Forest Inventory and Analysis National Program database to determine height–diameter allometries of 2,976,937 individuals of 293 tree species across the United States. The shape of the allometric relationship was determined by comparing linear and nonlinear functional forms. Mixed‐effects models were used to test for allometric differences due to climate and floristic (between angiosperms and gymnosperms) and functional groups (leaf habit and shade tolerance). Tree allometry significantly differed across the United States largely because of climate. Temperature, and to some extent precipitation, in part explained tree allometric variation. The magnitude of allometric variation due to climate, however, had a phylogenetic signal. Specifically, angiosperm allometry was more sensitive to differences in temperature compared to gymnosperms. Most notably, angiosperm height was more negatively influenced by increasing temperature variability, whereas gymnosperm height was negatively influenced by decreasing precipitation and increasing altitude. There was little evidence to suggest that shade tolerance influenced tree allometry except for very shade‐intolerant trees which were taller for any given diameter. Tree allometry is plastic rather than fixed and scaling parameters vary around predicted central tendencies. This allometric variation provides insight into life‐history strategies, phylogenetic history, and environmental limitations at biogeographical scales.     

Forest resilience to drought varies across biomes

Forecasted increase drought frequency and severity may drive worldwide declines in forest productivity. Species‐level responses to a drier world are likely to be influenced by their functional traits. Here, we analyse forest resilience to drought using an extensive network of tree‐ring width data and satellite imagery. We compiled proxies of forest growth and productivity (TRWi, absolutely dated ring‐width indices; NDVI, Normalized Difference Vegetation Index) for 11 tree species and 502 forests in Spain corresponding to Mediterranean, temperate, and continental biomes. Four different components of forest resilience to drought were calculated based on TRWi and NDVI data before, during, and after four major droughts (1986, 1994–1995, 1999, and 2005), and pointed out that TRWi data were more sensitive metrics of forest resilience to drought than NDVI data. Resilience was related to both drought severity and forest composition. Evergreen gymnosperms dominating semi‐arid Mediterranean forests showed the lowest resistance to drought, but higher recovery than deciduous angiosperms dominating humid temperate forests. Moreover, semi‐arid gymnosperm forests presented a negative temporal trend in the resistance to drought, but this pattern was absent in continental and temperate forests. Although gymnosperms in dry Mediterranean forests showed a faster recovery after drought, their recovery potential could be constrained if droughts become more frequent. Conversely, angiosperms and gymnosperms inhabiting temperate and continental sites might have problems to recover after more intense droughts since they resist drought but are less able to recover afterwards.     

Chilling rather than photoperiod controls budburst for gymnosperm species in subtropical China

低温而不是光周期调控中国亚热带裸子植物的出芽物候 摘要：被子植物春季物候的调控机制已经得到了广泛的研究。然而,裸子植物和被子植物在3亿年前就产生分化,裸子植物与被子植物的物候可能是受不同的因素所调控。亚热带植物物候的调节机制在很大程度上尚不明确,亚热带裸子植物物候是否由冷激需求和光照调控仍未得到验证。本研究在人工气候箱中设置了3个冷激处理和3 个光周期处理,并对切枝的出芽期进行了为期8周的研究。实验中我们测试了8种裸子植物：柳杉(Cryptomeria japonica)、杉木(Cunninghamia lanceolata)、柏树(Cupressus funebris)、银杏(Ginkgo biloba)、水杉(Metasequoia glyptostroboides)、马尾松(Pinus massoniana)、金钱 松(Pseudolarix amabilis)和罗汉松(Podocarpus macrophyllus),检验其出芽物候是否对光周期敏感或者是否具有较强的冷激需求,以及这两种因素哪个对促进出芽更为重要。研究结果表明,对于裸子植物,冷 激促进了出芽并提高了出芽率,而且裸子植物需要适度的低温天数来实现出芽。有趣的是,在同一森 林中裸子植物比被子植物对积温的需求更高。与德国温带裸子植物(194–600 d · °C)相比,亚热带裸子植 物(814–1150 d · °C)对积温的需求更高。光周期对裸子植物出芽的影响较小,说明冷激对裸子植物出芽的 促进作用大于光周期。这些结果表明,随着全球气候持续变暖,冬季气温的升高不仅会影响亚热带被子植物也会影响裸子植物的物候,从而可能导致春季出芽期的延迟。     

Cenozoic extinctions account for the low diversity of extant gymnosperms compared with angiosperms

We test the widely held notion that living gymnosperms are 'ancient' and 'living fossils' by comparing them with their sister group, the angiosperms. This perception derives partly from the lack of gross morphological differences between some Mesozoic gymnosperm fossils and their living relatives (e.g. Ginkgo, cycads and dawn redwood), suggesting that the rate of evolution of gymnosperms has been slow. We estimated the ages and diversification rates of gymnosperm lineages using Bayesian relaxed molecular clock dating calibrated with 21 fossils, based on the phylogenetic analysis of alignments of matK chloroplast DNA (cpDNA) and 26S nuclear ribosomal DNA (nrDNA) sequences, and compared these with published estimates for angiosperms. Gymnosperm crown groups of Cenozoic age are significantly younger than their angiosperm counterparts (median age: 32 Ma vs 50 Ma) and have long unbranched stems, indicating major extinctions in the Cenozoic, in contrast with angiosperms. Surviving gymnosperm genera have diversified more slowly than angiosperms during the Neogene as a result of their higher extinction rate. Compared with angiosperms, living gymnosperm groups are not ancient. The fossil record also indicates that gymnosperms suffered major extinctions when climate changed in the Oligocene and Miocene. Extant gymnosperm groups occupy diverse habitats and some probably survived after making adaptive shifts.     

Climate‐ and successional‐related changes in functional composition of European forests are strongly driven by tree mortality

Intense droughts combined with increased temperatures are one of the major threats to forest persistence in the 21st century. Despite the direct impact of climate change on forest growth and shifts in species abundance, the effect of altered demography on changes in the composition of functional traits is not well known. We sought to (1) quantify the recent changes in functional composition of European forests; (2) identify the relative importance of climate change, mean climate and forest development for changes in functional composition; and (3) analyse the roles of tree mortality and growth underlying any functional changes in different forest types. We quantified changes in functional composition from the 1980s to the 2000s across Europe by two dimensions of functional trait variation: the first dimension was mainly related to changes in leaf mass per area and wood density (partially related to the trait differences between angiosperms and gymnosperms), and the second dimension was related to changes in maximum tree height. Our results indicate that climate change and mean climatic effects strongly interacted with forest development and it was not possible to completely disentangle their effects. Where recent climate change was not too extreme, the patterns of functional change generally followed the expected patterns under secondary succession (e.g. towards late‐successional short‐statured hardwoods in Mediterranean forests and taller gymnosperms in boreal forests) and latitudinal gradients (e.g. larger proportion of gymnosperm‐like strategies at low water availability in forests formerly dominated by broad‐leaved deciduous species). Recent climate change generally favoured the dominance of angiosperm‐like related traits under increased temperature and intense droughts. Our results show functional composition changes over relatively short time scales in European forests. These changes are largely determined by tree mortality, which should be further investigated and modelled to adequately predict the impacts of climate change on forest function.     

Angiosperm origin and early stages of seed plant evolution deduced from rRNA sequence comparisons

Summary Complete or partial nucleotide sequences of five different rRNA species, coded by nuclear (18S, 5.8S, and 5S) or chloroplast genomes (5S, 4.5S) from a number of seed plants were determined. Based on the sequence data, the phylogenetic dendrograms were built by two methods, maximum parsimony and compatibility. The topologies of the trees for different rRNA species are not fully congruent, but they share some common features. It may be concluded that both gymnosperms and angiosperms are monophyletic groups. The data obtained suggest that the divergence of all the main groups of extant gymnosperms occurred after the branching off of the angiosperm lineage. As the time of divergence of at least some of these gymnosperm taxa is traceable back to the early Carboniferous, it may be concluded that the genealogical splitting of gymnosperm and angiosperm lineages occurred before this event, at least 360 million years ago, i.e., much earlier than the first angiosperm fossils were dated. Ancestral forms of angiosperms ought to be searched for among Progymnospermopsida. Genealogical relationships among gymnosperm taxa cannot be deduced unambiguously on the basis of rRNA data. The only inference may be that the taxon Gnetopsida is an artificial one, andGnetum andEphedra belong to quite different lineages of gymnosperms. As to the phylogenetic position of the two Angiospermae classes, extant monocotyledons seem to be a paraphyletic group located near the root of the angiosperm branch; it emerged at the earliest stages of angiosperm evolution. We may conclude that either monocotyledonous characters arose independently more than once in different groups of ancient Magnoliales or that monocotyledonous forms rather than dicotyledonous Magnoliales were the earliest angiosperms. Judging by the rRNA trees, Magnoliales are the most ancient group among dicotyledons. The most ancient lineage among monocotyledons leads to modern Liliaceae.     

Early angiosperm ecology: evidence from the Albian-Cenomanian of Europe

BACKGROUND AND AIMS: The mid-Cretaceous is a period of sudden turnover from gymnosperm to angiosperm-dominated floras. The aim was to investigate the fossil plant ecology in order to follow the spread of angiosperm taxa. METHODS: Floristic lists and localities from the latest Albian-Cenomanian of Europe are analysed with Wagner's Parsimony Method, a clustering method currently used in phylogeny (cladistics). KEY RESULTS: Wagner's Parsimony Method points out that (a) gymnosperms dominated brackish water-related environments while angiosperms dominated freshwater-related environments (e.g. swamps, floodplains, levees, channels), (b) angiosperms showed the highest diversity in stable, freshwater-related environments, (c) a single angiosperm, 'Diospyros' cretacea, is restricted to brackish water-related environments and (d) the families Lauraceae and Platanaceae were exclusive to disturbed, braided river environments, implying a opportunist strategy for early tree angiosperms. CONCLUSIONS: During the Mid-Cretaceous, European floras were characterized by (a) coastal gymnosperms, (b) highly diversified fluvial angiosperms and (c) the first European brackish water-related angiosperm.     

The tortoise and the hare: ecology of angiosperm dominance and gymnosperm persistence

Gymnosperms, and conifers in particular, are sometimes very productive trees yet angiosperms dominate most temperate and tropical vegetation. Current explanations for angiosperm success emphasize the advantages of insect pollination and seed dispersal by animals for the colonization of isolated habitats. Differences between gymnosperm and angiosperm reproductive and vegetative growth rates have been largely ignored. Gymnosperms are all woody, perennial and usually have long reproductive cycles. Their leaves are not as fully vascularized as those of angiosperms and are more stereotyped in shape and size. Gymnosperm tracheids are generally more resistant to solute flow than angiosperm vessels. A consequence of the less efficient transport system is that maximum growth rates of gymnosperms are lower than maximum growth rates of angiosperms in well lit, well watered habitats. Gymnosperm seedlings may be particularly uncompetitive since their growth depends on a single cohort of relatively inefficient leaves. Later, some gymnosperms attain a higher productivity than co-occurring angiosperm trees by accumulating several cohorts of leaves with a higher total leaf area. These functional constraints on gymnosperm growth rates suggest that gymnosperms will be restricted to areas where growth of angiosperm competitors is reduced, for example, by cold or nutrient shortages. Biogeographic evidence supports this prediction since conifers are largely confined to high latitudes and elevations or nutrient-poor soils. Experimental studies show that competition in the regeneration niche (between conifer seedlings and angiosperm herbs and shrubs) is common and significantly affects conifer growth and survival, Fast-growing angiosperms, especially herbs and shrubs, may also change the frequency of disturbance regimes thereby excluding slower-growing gymnosperms. Shade-tolerant and early successional conifers share similar characteristics of slow initial growth and low plasticity to a change in resources. Shade-tolerant gymnosperms would be expected to occur only where forest openings are small or otherwise unsuitable for rapid filling by fast-growing angiosperm trees, lianas or shrubs. The limited evidence available suggests that shade-tolerant conifers are confined to forests with small gap sizes where large disturbances are very rare. The regeneration hypothesis for gymnosperm exclusion by angiosperms is consistent with several aspects of the fossil record such as the early disappearance of gymnosperms from early successional environments where competition with angiosperms would have been most severe. However there are unresolved difficulties in interpreting process from paleoecological pattern which prevent the testing of alternative hypotheses.     

Are functional traits and litter decomposability coordinated across leaves,twigs and wood? A test using temperate rainforest tree species

Synthesis This study compared the decomposability of leaf, twig and wood litter from 27 co‐occurring temperate rainforest tree species in New Zealand. We found that interspecific variation in decomposition was not coordinated across the three litter types. Analysis of the relationships between functional traits and decomposition revealed that traits predictive of wood decomposition varied among the species independently from traits predictive of the decomposition of leaf and twig litter. We conclude that efforts to understand how tree species influence C, N and P dynamics in forested ecosystems through the decomposition pathway need to consider the functional traits of multiple plant structures. Plant functional traits are increasingly used to evaluate changes in ecological and ecosystem processes. However our understanding of how functional traits coordinate across different plant structures, and the implications for trait‐driven processes such as litter decomposition, remains limited. We compared the functional traits of green leaves and leaf, twig and wood litter among 27 co‐occurring tree species from New Zealand, and quantified the loss of mass, N and P from the three litter types during decomposition. We hypothesised that: a) the functional traits of green leaves, and leaf, twig and wood litter are co‐ordinated so that species which produce high quality leaves and leaf litter will also produce high quality twig and wood litter, and b) the decomposability of leaf, twig and wood litter is coordinated because breakdown of all three litter types is driven by similar combinations of traits. Trait variation across species was co‐ordinated between leaves, twigs and wood when angiosperm and gymnosperm species were considered in combination, or when angiosperms were considered separately, but trait coordination was poor for gymnosperms. There was little coordination among the three litter types in their decomposability, especially when angiosperms and gymnosperms were considered separately; this was caused by the decomposability of each of the three litter types, at least partially, being driven by different functional traits or trait combinations. Our findings indicate that although interspecific variation in the functional traits of trees can be coordinated among leaves, twigs and wood, different or unrelated traits predict the decomposition of these different structures. Furthermore, leaf‐level analyses of functional traits are not satisfactory proxies for function of whole trees and related ecological processes. As such, efforts to understand how tree species influence C, N and P dynamics in forested ecosystems through the decomposition pathway need to consider functional traits of other plant structures.     

Was the ANITA rooting of the angiosperm phylogeny affected by long-branch attraction? Amborella, Nymphaeales, Illiciales, Trimeniaceae, and Austrobaileya

Five groups of basal angiosperms, Amborella, Nymphaeales, Illiciales, Trimeniaceae, and Austrobaileya (ANITA), were identified in several recent studies as representing a series of the earliest-diverging lineages of the angiosperm phylogeny. All of these studies except one employed a multigene analysis approach and used gymnosperms as the outgroup to determine the ingroup topology. The high level of divergence between gymnosperms and angiosperms, however, has long been implicated in the difficulty of reconstructing relationships at the base of angiosperm phylogeny using DNA sequences, for fear of long-branch attraction (LBA). In this study, we replaced the gymnosperm sequences from the five-gene matrix (mitochondrial atp1 and matR, plastid atpB and rbcL, and nuclear 18S rDNA) used in our earlier study with four categories of divergent sequences--random sequences with equal base frequencies or equally AT- and GC-rich contents, homopolymers and heteropolymers, misaligned gymnosperm sequences, and aligned lycopod and bryophyte sequences--to evaluate whether the gymnosperms were an appropriate outgroup to angiosperms in our earlier study that identified the ANITA rooting. All 24 analyses performed rooted the angiosperm phylogeny at either Acorus or Alisma (or Alisma-Triglochin-Potamogeton in one case due to use of a slightly different alignment) and placed the monocots as a basal grade, producing genuine LBA results. These analyses demonstrate that the identification of ANITA as the basalmost extant angiosperms was based on historical signals preserved in the gymnosperm sequences and that the gymnosperms were an appropriate outgroup with which to root the angiosperm phylogeny in the multigene sequence analysis. This strategy of evaluating the appropriateness of an outgroup using artificial sequences and a series of outgroups with increments of divergence levels can be applied to investigations of phylogenetic patterns at the bases of other major clades, such as land plants, animals, and eukaryotes.     

Estimates of gene flow in forest trees

Qualitative and quantitative estimates of gene flow were obtained for fourteen gymnosperm and seven angiosperm forest tree species. High levels of gene flow were prevalent among gymnosperms while these levels varied from high to low among angiosperms. In both groups, species with greater pollen dispersal abilities appear to maintain high levels of gene flow. A detailed analysis of population structure in relation to gene flow was carried out on a gymnosperm species (Pinus rigida) and two angiosperm subspecies (Eucalyptus caesia ssp. caesia and ssp. magna). The results suggested that populations of many species may be concatenated systems bound by gene flow, and the overall levels of gene flow may be influenced by either single or clusters of populations. Different levels of gene flow was found between two closely related species of E. caesia growing under similar ecological conditions, suggesting a plausible link between pollinator behaviour and pollen flow.     

Global meta‐analysis of wood decomposition rates: a role for trait variation among tree species?

The carbon flux from woody debris, a crucial uncertainty within global carbon-climate models, is simultaneously affected by climate, site environment and species-based variation in wood quality. In the first global analysis attempting to explicitly tease out the wood quality contribution to decomposition, we found support for our hypothesis that, under a common climate, interspecific differences in wood traits affect woody debris decomposition patterns. A meta-analysis of 36 studies from all forested continents revealed that nitrogen, phosphorus, and C : N ratio correlate with decomposition rates of angiosperms. In addition, gymnosperm wood consistently decomposes slower than angiosperm wood within common sites, a pattern that correlates with clear divergence in wood traits between the two groups. New empirical studies are needed to test whether this difference is due to a direct effect of wood trait variation on decomposer activity or an indirect effect of wood traits on decomposition microsite environment. The wood trait–decomposition results point to an important role for changes in the wood traits of dominant tree species as a driver of carbon cycling, with likely feedback to atmospheric CO₂ particularly where angiosperm species replace gymnosperms regionally. Truly worldwide upscaling of our results will require further site-based multi-species wood trait and decomposition data, particularly from low-latitude ecosystems.     

东北温带森林20种乔木树种叶片干旱容忍性特征

全球范围内干旱频率和强度的增加严重影响树木生长,甚至导致森林大面积死亡。压力-容积(PV)曲线能够反映树木对干旱的容忍能力,但在局域尺度上尚未确定哪个PV曲线参数具有最优指示性。通过测定东北温带森林20种主要树种(包括16种被子植物和4种裸子植物)的PV曲线性状,包括质壁分离时的相对含水量(RWC_tlp)、失膨点叶水势(TLP)、饱和含水时的叶渗透势(π₀)、细胞弹性模量(ε)、叶水容(C_leaf)及叶结构性状(比叶面积和叶密度),研究局域尺度上叶片耐旱性的最佳指示性状,并分析叶片PV性状与结构性状间的相关性。结果表明: 被子植物的RWC_tlp 显著大于裸子植物,但其C_leaf 显著小于裸子植物,这表明用RWC_tlp和C_leaf可以指示东北温带森林不同功能型树种间耐旱性的大小。在被子植物中,TLP和π₀与叶密度呈显著负相关,且均与比叶面积呈显著正相关;而ε与比叶面积呈显著负相关。然而,裸子植物PV曲线性状与叶结构性状之间呈现与被子植物完全相反的趋势。裸子植物与被子植物树种之间PV曲线性状与叶结构性状关系的差异,可能归因于二者采取不同的干旱响应和适应策略。     

The evolutionary diversification of seed size: using the past to understand the present

The Devonian origin of seed plants and subsequent morphological diversification of seeds during the late Paleozoic represents an adaptive radiation into unoccupied ecological niche space. A plant's seed size is correlated with its life-history strategy, growth form, and seed dispersal syndrome. The fossil record indicates that the oldest seed plants had relatively small seeds, but the Mississippian seed size envelope increased significantly with the diversification of larger seeded lineages. Fossil seeds equivalent to the largest extant gymnosperm seeds appeared by the Pennsylvanian, concurrent with morphological diversification of growth forms and dispersal syndromes as well as the clade's radiation into new environments. Wang's Analysis of Skewness indicates that the evolutionary trend of increasing seed size resulted from primarily passive processes in Pennsylvanian seed plants. The distributions of modern angiosperms indicate a more diverse system of active and some passive processes, unbounded by Paleozoic limits; multiple angiosperm lineages independently evolved though the upper and lower bounds. Quantitative measures of preservation suggest that, although our knowledge of Paleozoic seeds is far from complete, the evolutionary trend in seed size is unlikely to be an artifact of taphonomy.     

Ontogenetic shifts in plant–plant interactions in a rare cycad within angiosperm communities

Gymnosperms and angiosperms can co-occur within the same habitats but key plant traits are thought to give angiosperms an evolutionary competitive advantage in many ecological settings. We studied ontogenetic changes in competitive and facilitative interactions between a rare gymnosperm (Dioon sonorense, our target species) and different plant and abiotic neighbours (conspecific-cycads, heterospecific-angiosperms, or abiotic-rocks) from 2007 to 2010 in an arid environment of northwestern Mexico. We monitored survival and growth of seedlings, juveniles, and adults of the cycad Dioon sonorense to evaluate how cycad survival and relative height growth rate (RHGR) responded to intra- and interspecific competition, canopy openness, and nearest neighbour. We tested spatial associations among D. sonorense life stages and angiosperm species and measured ontogenetic shifts in cycad shade tolerance. Canopy openness decreased cycad survival while intraspecific competition decreased survival and RHGR during early ontogeny. Seedling survival was higher in association with rocks and heterospecific neighbours where intraspecific competition was lower. Shade tolerance decreased with cycad ontogeny reflecting the spatial association of advanced stages with more open canopies. Interspecific facilitation during early ontogeny of our target species may promote its persistence in spite of increasing interspecific competition in later stages. We provide empirical support to the long-standing assumption that marginal rocky habitats serve as refugia from angiosperm competition for slow-growing gymnosperms such as cycads. The lack of knowledge of plant–plant interactions in rare or endangered species may hinder developing efficient conservation strategies (e.g. managing for sustained canopy cover), especially under the ongoing land use and climatic changes.     

Evolution of Reproductive Organs in Land Plants

LEAFY gene is the positive regulator of the MADS-box genes in flower primordia. The number of MADS-box genes presumably increased by gene duplications before the divergence of ferns and seed plants. Most MADS-box genes in ferns are expressed similarly in both vegetative and reproductive organs, while in gymnosperms, some MADS-box genes are specifically expressed in reproductive organs. This suggests that (1) the increase in the number of MADS-box genes and (2) the subsequent recruitment of some MADS-box genes as homeotic selector genes were important for the evolution of complex reproductive organs. The phylogenetic tree including both angiosperm and gymnosperm MADS-box genes indicates the loss of the A-function genes in the gymnosperm lineage, which is presumably related to the absence of perianths in extant gymnosperms. Comparison of expression patterns of orthologous MADS-box genes in angiosperms, Gnetales, and conifers supports the sister relationship of Gnetales and conifers over that of Gnetales and angiosperms predicted by phylogenetic trees based on amino acid and nucleotide sequences. Received 30 July 1999/ Accepted in revised form 9 September 1999     

Leaf energy balance modelling as a tool to infer habitat preference in the early angiosperms

Despite more than a century of research, some key aspects of habitat preference and ecology of the earliest angiosperms remain poorly constrained. Proposed growth ecology has varied from opportunistic weedy species growing in full sun to slow-growing species limited to the shaded understorey of gymnosperm forests. Evidence suggests that the earliest angiosperms possessed low transpiration rates: gas exchange rates for extant basal angiosperms are low, as are the reconstructed gas exchange rates for the oldest known angiosperm leaf fossils. Leaves with low transpirational capacity are vulnerable to overheating in full sun, favouring the hypothesis that early angiosperms were limited to the shaded understorey. Here, modelled leaf temperatures are used to examine the thermal tolerance of some of the earliest angiosperms. Our results indicate that small leaf size could have mitigated the low transpirational cooling capacity of many early angiosperms, enabling many species to survive in full sun. We propose that during the earliest phases of the angiosperm leaf record, angiosperms may not have been limited to the understorey, and that some species were able to compete with ferns and gymnosperms in both shaded and sunny habitats, especially in the absence of competition from more rapidly growing and transpiring advanced lineages of angiosperms.