A Middle Eocene lowland humid subtropical "Shangri-La" ecosystem in central Tibet.

Tibet's ancient topography and its role in climatic and biotic evolution remain speculative due to a paucity of quantitative surface-height measurements through time and space, and sparse fossil records. However, newly discovered fossils from a present elevation of ∼4,850 m in central Tibet improve substantially our knowledge of the ancient Tibetan environment. The 70 plant fossil taxa so far recovered include the first occurrences of several modern Asian lineages and represent a Middle Eocene (∼47 Mya) humid subtropical ecosystem. The fossils not only record the diverse composition of the ancient Tibetan biota, but also allow us to constrain the Middle Eocene land surface height in central Tibet to ∼1,500 ± 900 m, and quantify the prevailing thermal and hydrological regime. This "Shangri-La"-like ecosystem experienced monsoon seasonality with a mean annual temperature of ∼19 °C, and frosts were rare. It contained few Gondwanan taxa, yet was compositionally similar to contemporaneous floras in both North America and Europe. Our discovery quantifies a key part of Tibetan Paleogene topography and climate, and highlights the importance of Tibet in regard to the origin of modern Asian plant species and the evolution of global biodiversity.

Variations in morphological and epidermal features of shade and sun leaves of two species: Quercus bambusifolia and Q. myrsinifolia.

PREMISE: Microclimatic differences between the periphery and the interior of tree crowns result in a variety of adaptive leaf macromorphological and anatomical features. Our research was designed to reveal criteria for sun/shade leaf identification in two species of evergreen oaks, applicable to both modern and fossil leaves. We compared our results with those in other species similarly studied. METHODS: For both Quercus bambusifolia and Q. myrsinifolia (section Cyclobalanopsis), leaves from single mature trees with well-developed crowns were collected in the South China Botanical Garden, Guangzhou, China. We focus on leaf characters often preserved in fossil material. SVGm software was used for macromorphological measurement. Quantitative analyses were performed and box plots generated using R software with IDE Rstudio. Leaf cuticles were prepared using traditional botanical techniques. RESULTS: Principal characters for distinguishing shade and sun leaves in the studied oaks were identified as leaf lamina length to width ratio (L/W), and the degree of development of venation networks. For Q. myrsinifolia, shade and sun leaves differ in tooth morphology and the ratio of toothed lamina length to overall lamina length. The main epidermal characters are ordinary cell size and anticlinal wall outlines. For both species, plasticity within shade leaves exceeds that of sun leaves. CONCLUSIONS: Morphological responses to sun and shade in the examined oaks are similar to those in other plant genera, pointing to useful generalizations for recognizing common foliar polymorphisms that must be taken into account when determining the taxonomic position of both modern and fossil plants.

Asclepiadospermum gen. nov., the earliest fossil record of Asclepiadoideae (Apocynaceae) from the early Eocene of central Qinghai-Tibetan Plateau, and its biogeographic implications.

PREMISE: Apocynaceae is common in the fossil record, especially as seed remains from the Neogene of Europe and North America, but rare in Asia. Intrafamilial assignment is difficult due to the lack of diagnostic characters, and new fossil and modern data are needed to understand the paleobiogeography of this group. METHODS: We studied three Apocynaceae seed impressions from the Lower Eocene Niubao Formation, Jianglang village, Bangor County, central Qinghai-Tibetan Plateau. Morphological data from living and fossil species were phylogenetically mapped to enable systematic assignment. RESULTS: We describe a new genus, Asclepiadospermum gen. nov., and two new species, A. marginatum sp. nov. and A. ellipticum sp. nov. These species are characterized by an elliptical seed, a margin surrounding the central part of the seed, and polygonal, irregular, and small epidermal cells, and differ mainly in terms of the size of the margin and the shape of the apex. All these characters indicate that this new genus belongs to the subfamily Asclepiadoideae (Apocynaceae). CONCLUSIONS: These fossils represent the earliest fossil seed records of Asclepiadoideae. Asclepiadospermum indicates a humid tropical to subtropical flora during the early Eocene in central Tibet. Moreover, our discoveries indicate a close floristic connection between Eurasia and Africa during the early Eocene, which expands our knowledge of the floristic linkage between Tibet and other regions at that time.

The Eocene climate of China, the early elevation of the Tibetan Plateau and the onset of the Asian Monsoon.

Eocene palynological samples from 37 widely distributed sites across China were analysed using co-existence approach to determine trends in space and time for seven palaeoclimate variables: Mean annual temperature, mean annual precipitation, mean temperature of the warmest month, mean temperature of the coldest month, mean annual range of temperature, mean maximum monthly precipitation and mean minimum monthly precipitation. Present day distributions and observed climates within China of the nearest living relatives of the fossil forms were used to find the range of a given variable in which a maximum number of taxa can coexist. Isotherm and isohyet maps for the early, middle and late Eocene were constructed. These illustrate regional changing patterns in thermal and precipitational gradients that may be interpreted as the beginnings of the modern Asian Monsoon system, and suggest that the uplift of parts of the Tibetan Plateau appear to have taken place by the middle to late Eocene.

Coryphoid palms from the K-Pg boundary of central India and their biogeographical implications: Evidence from megafossil remains.

Ten palm leaf impressions are documented from the latest Maastrichtian (late Cretaceous) to early Danian (earliest Paleocene) sediments (K-Pg, c. 66-64 Ma) of the Mandla Lobe of the Deccan Inter-trappean Beds, Madhya Pradesh, central India. The palmate leaf shape along with a definite well-preserved costa support their placement in the subfamily Coryphoideae of the family Arecaceae. We place all recovered palm leaf specimens in the fossil genus Sabalites, report seven species of coryphoid palms and describe two new species namely, Sabalities umariaensis sp. nov. and Sabalites ghughuaensis sp. nov. The fossils indicate that coryphoid palms were highly diverse in central India by the latest Cretaceous. These and earlier reported coryphoid palm fossils from the same locality indicate that they experienced a warm and humid tropical environment during the time of deposition. These discoveries confirm the presence of a diversity of Coryphoideae in Gondwana prior to the India-Eurasia collision and provide information about coryphoid biogeographical history over geological time. Based on megafossil remains, we trace coryphoid palm migration pathways from India to mainland Southeast (SE) Asia and other parts of Asia after the docking of the Indian subcontinent with Eurasia early in the Paleogene.

Siwalik plant megafossil diversity in the Eastern Himalayas: A review.

The Eastern Himalayas are renowned for their high plant diversity. To understand how this modern botanical richness formed, it is critical to investigate past plant biodiversity preserved as fossils throughout the eastern Himalayan Siwalik succession (middle Miocene-early Pleistocene). Here, we present a summary of plant diversity records that document Neogene floristic and climate changes. We do this by compiling published records of megafossil plant remains, because these offer better spatial and temporal resolution than do palynological records. Analyses of the Siwalik floral assemblages based on the distribution of the nearest living relative taxa suggest that a tropical wet evergreen forest was growing in a warm humid monsoonal climate at the deposition time. This qualitative interpretation is also corroborated by published CLAMP (Climate Leaf Analysis Multivariate Program) analyses. Here, we also reconstruct the climate by applying a new common proxy WorldClim2 calibration. This allows the detection of subtle climate differences between floral assemblages free of artefacts introduced by using different methodologies and climate calibrations. An analysis of the Siwalik floras indicates that there was a gradual change in floral composition. The lower Siwalik assemblages provide evidence of a predominance of evergreen elements. An increase in deciduous elements in the floral composition is noticed towards the close of the middle Siwalik and the beginning of the upper Siwalik formation. This change reflects a climatic difference between Miocene and Plio-Pleistocene times. This review helps us to understand under what paleoenvironmental conditions plant diversity occurred and evolved in the eastern Himalayas throughout the Cenozoic.

Mercury isotopes show vascular plants had colonized land extensively by the early Silurian.

The colonization and expansion of plants on land is considered one of the most profound ecological revolutions, yet the precise timing remains controversial. Because land vegetation can enhance weathering intensity and affect terrigenous input to the ocean, changes in terrestrial plant biomass with distinct negative Δ199Hg and Δ200Hg signatures may overwrite the positive Hg isotope signatures commonly found in marine sediments. By investigating secular Hg isotopic variations in the Paleozoic marine sediments from South China and peripheral paleocontinents, we highlight distinct negative excursions in both Δ199Hg and Δ200Hg at Stage level starting in the early Silurian and again in the Carboniferous. These geochemical signatures were driven by increased terrestrial contribution of Hg due to the rapid expansion of vascular plants. These excursions broadly coincide with rising atmospheric oxygen concentrations and global cooling. Therefore, vascular plants were widely distributed on land during the Ordovician-Silurian transition (~444 million years), long before the earliest reported vascular plant fossil, Cooksonia (~430 million years).

A distinctive Eocene Asian monsoon and modern biodiversity resulted from the rise of eastern Tibet.

The uplift of eastern Tibet, Asian monsoon development and the evolution of globally significant Asian biodiversity are all linked, but in obscure ways. Sedimentology, geochronology, clumped isotope thermometry, and fossil leaf-derived numerical climate data from the Relu Basin, eastern Tibet, show at âˆ¼50-45 Ma the basin was a hot (mean annual air temperature, MAAT, ∼27 °C) dry desert at a low-elevation of 0.6 ± 0.6 km. Rapid basin rise to 2.0 ± 0.9 km at 45-42 Ma and to 2.9 ± 0.9 km at 42-40 Ma, with MAATs of âˆ¼20 and âˆ¼16 °C, respectively, accompanied seasonally varying increased annual precipitation to > 1500 mm. From âˆ¼39 to 34 Ma, the basin attained 3.5 ± 1.0 km, near its present-day elevation (∼3.7 km), and MAAT cooled to âˆ¼6 °C. Numerically-modelled Asian monsoon strength increased significantly when this Eocene uplift of eastern Tibet was incorporated. The simulation/proxy congruence points to a distinctive Eocene Asian monsoon, quite unlike that seen today, in that it featured bimodal precipitation and a winter-wet regime, and this enhanced biodiversity modernisation across eastern Asia. The Paleogene biodiversity of Asia evolved under a continually modifying monsoon influence, with the modern Asian monsoon system being unique to the present and a product of a long gradual development in the context of an ever-changing Earth system.

The rise and demise of the Paleogene Central Tibetan Valley.

Reconstructing the Paleogene topography and climate of central Tibet informs understanding of collisional tectonic mechanisms and their links to climate and biodiversity. Radiometric dates of volcanic/sedimentary rocks and paleotemperatures based on clumped isotopes within ancient soil carbonate nodules from the Lunpola Basin, part of an east-west trending band of basins in central Tibet and now at 4.7 km, suggest that the basin rose from <2.0 km at 50 to 38 million years (Ma) to >4.0 km by 29 Ma. The height change is quantified using the rates at which wet-bulb temperatures (Tw) decline at land surfaces as those surface rise. In this case, Tw fell from ~8°C at ~38 Ma to ~1°C at 29 Ma, suggesting at least ~2.0 km of surface uplift in ~10 Ma under warm Eocene to Oligocene conditions. These results confirm that a Paleogene Central Tibetan Valley transformed to a plateau before the Neogene.

Rapid Eocene diversification of spiny plants in subtropical woodlands of central Tibet.

Spinescence is an important functional trait possessed by many plant species for physical defence against mammalian herbivores. The development of spinescence must have been closely associated with both biotic and abiotic factors in the geological past, but knowledge of spinescence evolution suffers from a dearth of fossil records, with most studies focusing on spatial patterns and spinescence-herbivore interactions in modern ecosystems. Numerous well-preserved Eocene (~39 Ma) plant fossils exhibiting seven different spine morphologies discovered recently in the central Tibetan Plateau, combined with molecular phylogenetic character reconstruction, point not only to the presence of a diversity of spiny plants in Eocene central Tibet but a rapid diversification of spiny plants in Eurasia around that time. These spiny plants occupied an open woodland landscape, indicated by numerous megafossils and grass phytoliths found in the same deposits, as well as numerical climate and vegetation modelling. Our study shows that regional aridification and expansion of herbivorous mammals may have driven the diversification of functional spinescence in central Tibetan woodlands, ~24 million years earlier than similar transformations in Africa.

Fire-prone Rhamnaceae with South African affinities in Cretaceous Myanmar amber.

The rapid Cretaceous diversification of flowering plants remains Darwin's 'abominable mystery' despite numerous fossil flowers discovered in recent years. Wildfires were frequent in the Cretaceous and many such early flower fossils are represented by charcoalified fragments, lacking complete delicate structures and surface textures, making their similarity to living forms difficult to discern. Furthermore, scarcity of information about the ecology of early angiosperms makes it difficult to test hypotheses about the drivers of their diversification, including the role of fire in shaping flowering plant evolution. We report the discovery of two exquisitely preserved fossil flower species, one identical to the inflorescences of the extant crown-eudicot genus Phylica and the other recovered as a sister group to Phylica, both preserved as inclusions together with burned plant remains in Cretaceous amber from northern Myanmar (~99 million years ago). These specialized flower species, named Phylica piloburmensis sp. nov. and Eophylica priscastellata gen. et sp. nov., exhibit traits identical to those of modern taxa in fire-prone ecosystems such as the fynbos of South Africa, and provide evidence of fire adaptation in angiosperms.

Why 'the uplift of the Tibetan Plateau' is a myth.

The often-used phrase 'the uplift of the Tibetan Plateau' implies a flat-surfaced Tibet rose as a coherent entity, and that uplift was driven entirely by the collision and northward movement of India. Here, we argue that these are misconceptions derived in large part from simplistic geodynamic and climate modeling, as well as proxy misinterpretation. The growth of Tibet was a complex process involving mostly Mesozoic collisions of several Gondwanan terranes with Asia, thickening the crust and generating complex relief before the arrival of India. In this review, Earth system modeling, paleoaltimetry proxies and fossil finds contribute to a new synthetic view of the topographic evolution of Tibet. A notable feature overlooked in previous models of plateau formation was the persistence through much of the Cenozoic of a wide east-west orientated deep central valley, and the formation of a plateau occurred only in the late Neogene through compression and internal sedimentation.

Spatiotemporal maintenance of flora in the Himalaya biodiversity hotspot: Current knowledge and future perspectives.

Mountain ecosystems support a significant one-third of all terrestrial biodiversity, but our understanding of the spatiotemporal maintenance of this high biodiversity remains poor, or at best controversial. The Himalaya hosts a complex mountain ecosystem with high topographic and climatic heterogeneity and harbors one of the world's richest floras. The high species endemism, together with increasing anthropogenic threats, has qualified the Himalaya as one of the most significant global biodiversity hotspots. The topographic and climatic complexity of the Himalaya makes it an ideal natural laboratory for studying the mechanisms of floral exchange, diversification, and spatiotemporal distributions. Here, we review literature pertaining to the Himalaya in order to generate a concise synthesis of the origin, distribution, and climate change responses of the Himalayan flora. We found that the Himalaya supports a rich biodiversity and that the Hengduan Mountains supplied the majority of the Himalayan floral elements, which subsequently diversified from the late Miocene onward, to create today's relatively high endemicity in the Himalaya. Further, we uncover links between this Miocene diversification and the joint effect of geological and climatic upheavals in the Himalaya. There is marked variance regarding species dispersal, elevational gradients, and impact of climate change among plant species in the Himalaya, and our review highlights some of the general trends and recent advances on these aspects. Finally, we provide some recommendations for conservation planning and future research. Our work could be useful in guiding future research in this important ecosystem and will also provide new insights into the maintenance mechanisms underpinning other mountain systems.

Orographic evolution of northern Tibet shaped vegetation and plant diversity in eastern Asia.

The growth of the Tibetan Plateau throughout the past 66 million years has profoundly affected the Asian climate, but how this unparalleled orogenesis might have driven vegetation and plant diversity changes in eastern Asia is poorly understood. We approach this question by integrating modeling results and fossil data. We show that growth of north and northeastern Tibet affects vegetation and, crucially, plant diversity in eastern Asia by altering the monsoon system. This northern Tibetan orographic change induces a precipitation increase, especially in the dry (winter) season, resulting in a transition from deciduous broadleaf vegetation to evergreen broadleaf vegetation and plant diversity increases across southeastern Asia. Further quantifying the complexity of Tibetan orographic change is critical for understanding the finer details of Asian vegetation and plant diversity evolution.

The mid-Miocene Zhangpu biota reveals an outstandingly rich rainforest biome in East Asia.

During the Mid-Miocene Climatic Optimum [MMCO, ~14 to 17 million years (Ma) ago], global temperatures were similar to predicted temperatures for the coming century. Limited megathermal paleoclimatic and fossil data are known from this period, despite its potential as an analog for future climate conditions. Here, we report a rich middle Miocene rainforest biome, the Zhangpu biota (~14.7 Ma ago), based on material preserved in amber and associated sedimentary rocks from southeastern China. The record shows that the mid-Miocene rainforest reached at least 24.2°N and was more widespread than previously estimated. Our results not only highlight the role of tropical rainforests acting as evolutionary museums for biodiversity at the generic level but also suggest that the MMCO probably strongly shaped the East Asian biota via the northern expansion of the megathermal rainforest biome. The Zhangpu biota provides an ideal snapshot for biodiversity redistribution during global warming.

Cenozoic topography, monsoons and biodiversity conservation within the Tibetan Region: An evolving story.

The biodiversity of the Himalaya, Hengduan Mountains and Tibet, here collectively termed the Tibetan Region, is exceptional in a global context. To contextualize and understand the origins of this biotic richness, and its conservation value, we examine recent fossil finds and review progress in understanding the orogeny of the Tibetan Region. We examine the deep-time origins of monsoons affecting Asia, climate variation over different timescales, and the establishment of environmental niche heterogeneity linked to topographic development. The origins of the modern biodiversity were established in the Eocene, concurrent with the formation of pronounced topographic relief across the Tibetan Region. High (>4 km) mountains to the north and south of what is now the Tibetan Plateau bounded a Paleogene central lowland (<2.5 km) hosting moist subtropical vegetation influenced by an intensifying monsoon. In mid Miocene times, before the Himalaya reached their current elevation, sediment infilling and compressional tectonics raised the floor of the central valley to above 3000 m, but central Tibet was still moist enough, and low enough, to host a warm temperate angiosperm-dominated woodland. After 15 Ma, global cooling, the further rise of central Tibet, and the rain shadow cast by the growing Himalaya, progressively led to more open, herb-rich vegetation as the modern high plateau formed with its cool, dry climate. In the moist monsoonal Hengduan Mountains, high and spatially extensive since the Eocene but subsequently deeply dissected by river incision, Neogene cooling depressed the tree line, compressed altitudinal zonation, and created strong environmental heterogeneity. This served as a cradle for the then newly-evolving alpine biota and favoured diversity within more thermophilic vegetation at lower elevations. This diversity has survived through a combination of minimal Quaternary glaciation, and complex relief-related environmental niche heterogeneity. The great antiquity and diversity of the Tibetan Region biota argues for its conservation, and the importance of that biota is demonstrated through our insights into its long temporal gestation provided by fossil archives and information written in surviving genomes. These data sources are worthy of conservation in their own right, but for the living biotic inventory we need to ask what it is we want to conserve. Is it 1) individual taxa for their intrinsic properties, 2) their services in functioning ecosystems, or 3) their capacity to generate future new biodiversity? If 2 or 3 are our goal then landscape conservation at scale is required, and not just seed banks or botanical/zoological gardens.

Ancient orogenic and monsoon-driven assembly of the world's richest temperate alpine flora.

Understanding how alpine biotas formed in response to historical environmental change may improve our ability to predict and mitigate the threats to alpine species posed by global warming. In the world's richest temperate alpine flora, that of the Tibet-Himalaya-Hengduan region, phylogenetic reconstructions of biome and geographic range evolution show that extant lineages emerged by the early Oligocene and diversified first in the Hengduan Mountains. By the early to middle Miocene, accelerated diversification and colonization of adjacent regions were likely driven jointly by mountain building and intensification of the Asian monsoon. The alpine flora of the Hengduan Mountains has continuously existed far longer than any other alpine flora on Earth and illustrates how modern biotas have been shaped by past geological and climatic events.

Comment on "Revised paleoaltimetry data show low Tibetan Plateau elevation during the Eocene".

Botsyun et al (Research Articles, 1 March 2019, eaaq1436) have suggested that the Tibetan Plateau was low (substantially less than 3000 meters) during the Eocene, based on a comparison of oxygen isotope proxy data with isotope-enabled climate model simulations. However, we contend that their conclusions are flawed as the result of a number of failings of both the modeling and the data comparison.

Rising CO2 drives divergence in water use efficiency of evergreen and deciduous plants.

Intrinsic water use efficiency (iWUE), defined as the ratio of photosynthesis to stomatal conductance, is a key variable in plant physiology and ecology. Yet, how rising atmospheric CO2 concentration affects iWUE at broad species and ecosystem scales is poorly understood. In a field-based study of 244 woody angiosperm species across eight biomes over the past 25 years of increasing atmospheric CO2 (~45 ppm), we show that iWUE in evergreen species has increased more rapidly than in deciduous species. Specifically, the difference in iWUE gain between evergreen and deciduous taxa diverges along a mean annual temperature gradient from tropical to boreal forests and follows similar observed trends in leaf functional traits such as leaf mass per area. Synthesis of multiple lines of evidence supports our findings. This study provides timely insights into the impact of Anthropocene climate change on forest ecosystems and will aid the development of next-generation trait-based vegetation models.

Convergence in Maximum Stomatal Conductance of C3 Woody Angiosperms in Natural Ecosystems Across Bioclimatic Zones.

Stomatal conductance (g s) in terrestrial vegetation regulates the uptake of atmospheric carbon dioxide for photosynthesis and water loss through transpiration, closely linking the biosphere and atmosphere and influencing climate. Yet, the range and pattern of g s in plants from natural ecosystems across broad geographic, climatic, and taxonomic ranges remains poorly quantified. Furthermore, attempts to characterize g s on such scales have predominantly relied upon meta-analyses compiling data from many different studies. This approach may be inherently problematic as it combines data collected using unstandardized protocols, sometimes over decadal time spans, and from different habitat groups. Using a standardized protocol, we measured leaf-level g s using porometry in 218 C3 woody angiosperm species in natural ecosystems representing seven bioclimatic zones. The resulting dataset of 4273 g s measurements, which we call STraits (Stomatal Traits), was used to determine patterns in maximum g s (g smax) across bioclimatic zones and whether there was similarity in the mean g smax of C3 woody angiosperms across ecosystem types. We also tested for differential g smax in two broadly defined habitat groups - open-canopy and understory-subcanopy - within and across bioclimatic zones. We found strong convergence in mean g smax of C3 woody angiosperms in the understory-subcanopy habitats across six bioclimatic zones, but not in open-canopy habitats. Mean g smax in open-canopy habitats (266 ± 100 mmol m-2 s-1) was significantly higher than in understory-subcanopy habitats (233 ± 86 mmol m-2 s-1). There was also a central tendency in the overall dataset to operate toward a g smax of ∼250 mmol m-2 s-1. We suggest that the observed convergence in mean g smax of C3 woody angiosperms in the understory-subcanopy is due to a buffering of g smax against macroclimate effects which will lead to differential response of C3 woody angiosperm vegetation in these two habitats to future global change. Therefore, it will be important for future studies of g smax to categorize vegetation according to habitat group.