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1.
Curr Biol ; 2024 May 10.
Article in English | MEDLINE | ID: mdl-38754424

ABSTRACT

A fundamental question in dinosaur evolution is how they adapted to long-term climatic shifts during the Mesozoic and when they developed environmentally independent, avian-style acclimatization, becoming endothermic.1,2 The ability of warm-blooded dinosaurs to flourish in harsher environments, including cold, high-latitude regions,3,4 raises intriguing questions about the origins of key innovations shared with modern birds,5,6 indicating that the development of homeothermy (keeping constant body temperature) and endothermy (generating body heat) played a crucial role in their ecological diversification.7 Despite substantial evidence across scientific disciplines (anatomy,8 reproduction,9 energetics,10 biomechanics,10 osteohistology,11 palaeobiogeography,12 geochemistry,13,14 and soft tissues15,16,17), a consensus on dinosaur thermophysiology remains elusive.1,12,15,17,18,19 Differential thermophysiological strategies among terrestrial tetrapods allow endotherms (birds and mammals) to expand their latitudinal range (from the tropics to polar regions), owing to their reduced reliance on environmental temperature.20 By contrast, most reptilian lineages (squamates, turtles, and crocodilians) and amphibians are predominantly constrained by temperature in regions closer to the tropics.21 Determining when this macroecological pattern emerged in the avian lineage relies heavily on identifying the origin of these key physiological traits. Combining fossils with macroevolutionary and palaeoclimatic models, we unveil distinct evolutionary pathways in the main dinosaur lineages: ornithischians and theropods diversified across broader climatic landscapes, trending toward cooler niches. An Early Jurassic shift to colder climates in Theropoda suggests an early adoption of endothermy. Conversely, sauropodomorphs exhibited prolonged climatic conservatism associated with higher thermal conditions, emphasizing temperature, rather than plant productivity, as the primary driver of this pattern, suggesting poikilothermy with a stronger dependence on higher temperatures in sauropods.

2.
Nat Commun ; 15(1): 3697, 2024 May 07.
Article in English | MEDLINE | ID: mdl-38714681

ABSTRACT

The transition from a humid green Sahara to today's hyperarid conditions in northern Africa ~5.5 thousand years ago shows the dramatic environmental change to which human societies were exposed and had to adapt to. In this work, we show that in the 620,000-year environmental record from the Chew Bahir basin in the southern Ethiopian Rift, with its decadal resolution, this one thousand year long transition is particularly well documented, along with 20-80 year long droughts, recurring every ~160 years, as possible early warnings. Together with events of extreme wetness at the end of the transition, these droughts form a pronounced climate "flickering", which can be simulated in climate models and is also present in earlier climate transitions in the Chew Bahir environmental record, indicating that transitions with flickering are characteristic of this region.

3.
Evol Hum Sci ; 6: e8, 2024.
Article in English | MEDLINE | ID: mdl-38516369

ABSTRACT

Previous work has proposed various mechanisms by which the environment may affect the emergence of linguistic features. For example, dry air may cause careful control of pitch to be more effortful, and so affect the emergence of linguistic distinctions that rely on pitch such as lexical tone or vowel inventories. Criticisms of these proposals point out that there are both historical and geographic confounds that need to be controlled for. We take a causal inference approach to this problem to design the most detailed test of the theory to date. We analyse languages from the Bantu language family, using a prior geographic-phylogenetic tree of relationships to establish where and when languages were spoken. This is combined with estimates of humidity for those times and places, taken from historical climate models. We then estimate the strength of causal relationships in a causal path model, controlling for various influences of inheritance and borrowing. We find no evidence to support the previous claims that humidity affects the emergence of lexical tone. This study shows how using causal inference approaches lets us test complex causal claims about the cultural evolution of language.

4.
Science ; 383(6687): 1130-1134, 2024 Mar 08.
Article in English | MEDLINE | ID: mdl-38452067

ABSTRACT

Anthropogenic climate change is increasing rapidly and already impacting biodiversity. Despite its importance in future projections, understanding of the underlying mechanisms by which climate mediates extinction remains limited. We present an integrated approach examining the role of intrinsic traits versus extrinsic climate change in mediating extinction risk for marine invertebrates over the past 485 million years. We found that a combination of physiological traits and the magnitude of climate change is necessary to explain marine invertebrate extinction patterns. Our results suggest that taxa previously identified as extinction resistant may still succumb to extinction if the magnitude of climate change is great enough.


Subject(s)
Climate Change , Extinction, Biological , Invertebrates , Animals , Aquatic Organisms , Biodiversity
5.
Sci Adv ; 9(45): eadf3326, 2023 11 10.
Article in English | MEDLINE | ID: mdl-37939193

ABSTRACT

The Arctic is among the most climatically sensitive environments on Earth, and the disappearance of multiyear sea ice in the Arctic Ocean is predicted within decades. As apex predators, polar bears are sentinel species for addressing the impact of environmental variability on Arctic marine ecosystems. By integrating genomics, isotopic analysis, morphometrics, and ecological modeling, we investigate how Holocene environmental changes affected polar bears around Greenland. We uncover reductions in effective population size coinciding with increases in annual mean sea surface temperature, reduction in sea ice cover, declines in suitable habitat, and shifts in suitable habitat northward. Furthermore, we show that west and east Greenlandic polar bears are morphologically, and ecologically distinct, putatively driven by regional biotic and genetic differences. Together, we provide insights into the vulnerability of polar bears to environmental change and how the Arctic marine ecosystem plays a vital role in shaping the evolutionary and ecological trajectories of its inhabitants.


Subject(s)
Ursidae , Animals , Ecosystem , Climate Change , Biological Evolution , Arctic Regions , Ice Cover
6.
Nat Commun ; 14(1): 5549, 2023 Sep 08.
Article in English | MEDLINE | ID: mdl-37684244

ABSTRACT

The Sahara region has experienced periodic wet periods over the Quaternary and beyond. These North African Humid Periods (NAHPs) are astronomically paced by precession which controls the intensity of the African monsoon system. However, most climate models cannot reconcile the magnitude of these events and so the driving mechanisms remain poorly constrained. Here, we utilise a recently developed version of the HadCM3B coupled climate model that simulates 20 NAHPs over the past 800 kyr which have good agreement with NAHPs identified in proxy data. Our results show that precession determines NAHP pacing, but we identify that their amplitude is strongly linked to eccentricity via its control over ice sheet extent. During glacial periods, enhanced ice-albedo driven cooling suppresses NAHP amplitude at precession minima, when humid conditions would otherwise be expected. This highlights the importance of both precession and eccentricity, and the role of high latitude processes in determining the timing and amplitude of the NAHPs. This may have implications for the out of Africa dispersal of plants and animals throughout the Quaternary.

7.
Nature ; 614(7949): 708-712, 2023 02.
Article in English | MEDLINE | ID: mdl-36792825

ABSTRACT

The latitudinal diversity gradient (LDG) is a prevalent feature of modern ecosystems across diverse clades1-4. Recognized for well over a century, the causal mechanisms for LDGs remain disputed, in part because numerous putative drivers simultaneously covary with latitude1,3,5. The past provides the opportunity to disentangle LDG mechanisms because the relationships among biodiversity, latitude and possible causal factors have varied over time6-9. Here we quantify the emergence of the LDG in planktonic foraminifera at high spatiotemporal resolution over the past 40 million years, finding that a modern-style gradient arose only 15 million years ago. Spatial and temporal models suggest that LDGs for planktonic foraminifera may be controlled by the physical structure of the water column. Steepening of the latitudinal temperature gradient over 15 million years ago, associated with an increased vertical temperature gradient at low latitudes, may have enhanced niche partitioning and provided more opportunities for speciation at low latitudes. Supporting this hypothesis, we find that higher rates of low-latitude speciation steepened the diversity gradient, consistent with spatiotemporal patterns of depth partitioning by planktonic foraminifera. Extirpation of species from high latitudes also strengthened the LDG, but this effect tended to be weaker than speciation. Our results provide a step change in understanding the evolution of marine LDGs over long timescales.


Subject(s)
Aquatic Organisms , Biodiversity , Foraminifera , Geographic Mapping , Plankton , Spatio-Temporal Analysis , Aquatic Organisms/classification , Aquatic Organisms/isolation & purification , Biological Evolution , Foraminifera/classification , Foraminifera/isolation & purification , Genetic Speciation , History, Ancient , Phylogeography , Plankton/classification , Plankton/isolation & purification , Temperature , Time Factors , Water/analysis , Hydrobiology
8.
Nature ; 613(7943): 292-297, 2023 01.
Article in English | MEDLINE | ID: mdl-36631651

ABSTRACT

The recovery of long-term climate proxy records with seasonal resolution is rare because of natural smoothing processes, discontinuities and limitations in measurement resolution. Yet insolation forcing, a primary driver of multimillennial-scale climate change, acts through seasonal variations with direct impacts on seasonal climate1. Whether the sensitivity of seasonal climate to insolation matches theoretical predictions has not been assessed over long timescales. Here, we analyse a continuous record of water-isotope ratios from the West Antarctic Ice Sheet Divide ice core to reveal summer and winter temperature changes through the last 11,000 years. Summer temperatures in West Antarctica increased through the early-to-mid-Holocene, reached a peak 4,100 years ago and then decreased to the present. Climate model simulations show that these variations primarily reflect changes in maximum summer insolation, confirming the general connection between seasonal insolation and warming and demonstrating the importance of insolation intensity rather than seasonally integrated insolation or season duration2,3. Winter temperatures varied less overall, consistent with predictions from insolation forcing, but also fluctuated in the early Holocene, probably owing to changes in meridional heat transport. The magnitudes of summer and winter temperature changes constrain the lowering of the West Antarctic Ice Sheet surface since the early Holocene to less than 162 m and probably less than 58 m, consistent with geological constraints elsewhere in West Antarctica4-7.

9.
Curr Biol ; 33(1): 109-121.e3, 2023 01 09.
Article in English | MEDLINE | ID: mdl-36549298

ABSTRACT

Past responses to environmental change provide vital baseline data for estimating the potential resilience of extant taxa to future change. Here, we investigate the latitudinal range contraction that terrestrial and freshwater turtles (Testudinata) experienced from the Late Cretaceous to the Paleogene (100.5-23.03 mya) in response to major climatic changes. We apply ecological niche modeling (ENM) to reconstruct turtle niches, using ancient and modern distribution data, paleogeographic reconstructions, and the HadCM3L climate model to quantify their range shifts in the Cretaceous and late Eocene. We then use the insights provided by these models to infer their probable ecological responses to future climate scenarios at different representative concentration pathways (RCPs 4.5 and 8.5 for 2100), which project globally increased temperatures and spreading arid biomes at lower to mid-latitudes. We show that turtle ranges are predicted to expand poleward in the Northern Hemisphere, with decreased habitat suitability at lower latitudes, inverting a trend of latitudinal range contraction that has been prevalent since the Eocene. Trionychids and freshwater turtles can more easily track their niches than Testudinidae and other terrestrial groups. However, habitat destruction and fragmentation at higher latitudes will probably reduce the capability of turtles and tortoises to cope with future climate changes.


Subject(s)
Turtles , Animals , Turtles/physiology , Climate Change , Ecosystem , Fresh Water , Probability
10.
Curr Biol ; 33(1): 206-214.e4, 2023 01 09.
Article in English | MEDLINE | ID: mdl-36528026

ABSTRACT

The ascendancy of dinosaurs to become dominant components of terrestrial ecosystems was a pivotal event in the history of life, yet the drivers of their early evolution and biodiversity are poorly understood.1,2,3 During their early diversification in the Late Triassic, dinosaurs were initially rare and geographically restricted, only attaining wider distributions and greater abundance following the end-Triassic mass extinction event.4,5,6 This pattern is consistent with an opportunistic expansion model, initiated by the extinction of co-occurring groups such as aetosaurs, rauisuchians, and therapsids.4,7,8 However, this pattern could instead be a response to changes in global climatic distributions through the Triassic to Jurassic transition, especially given the increasing evidence that climate played a key role in constraining Triassic dinosaur distributions.7,9,10,11,12,13,14,15,16 Here, we test this hypothesis and elucidate how climate influenced early dinosaur distribution by quantitatively examining changes in dinosaur and tetrapod "climatic niche space" across the Triassic-Jurassic boundary. Statistical analyses show that Late Triassic sauropodomorph dinosaurs occupied a more restricted climatic niche space than other tetrapods and dinosaurs, being excluded from the hottest, low-latitude climate zones. A subsequent, earliest Jurassic expansion of sauropodomorph geographic distribution is linked to the expansion of their preferred climatic conditions. Evolutionary model-fitting analyses provide evidence for an important evolutionary shift from cooler to warmer climatic niches during the origin of Sauropoda. These results are consistent with the hypothesis that global abundance of sauropodomorph dinosaurs was facilitated by climatic change and provide support for the key role of climate in the ascendancy of dinosaurs.


Subject(s)
Biological Evolution , Dinosaurs , Animals , Dinosaurs/anatomy & histology , Ecosystem , Fossils , Biodiversity , Phylogeny
11.
Sci Bull (Beijing) ; 67(21): 2245-2258, 2022 11 15.
Article in English | MEDLINE | ID: mdl-36546000

ABSTRACT

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.


Subject(s)
Biodiversity , Climate , Tibet , Asia, Eastern , Seasons
12.
Nature ; 610(7931): 313-318, 2022 10.
Article in English | MEDLINE | ID: mdl-36198797

ABSTRACT

Pterosaurs, the first vertebrates to evolve powered flight, were key components of Mesozoic terrestrial ecosystems from their sudden appearance in the Late Triassic until their demise at the end of the Cretaceous1-6. However, the origin and early evolution of pterosaurs are poorly understood owing to a substantial stratigraphic and morphological gap between these reptiles and their closest relatives6, Lagerpetidae7. Scleromochlus taylori, a tiny reptile from the early Late Triassic of Scotland discovered over a century ago, was hypothesized to be a key taxon closely related to pterosaurs8, but its poor preservation has limited previous studies and resulted in controversy over its phylogenetic position, with some even doubting its identification as an archosaur9. Here we use microcomputed tomographic scans to provide the first accurate whole-skeletal reconstruction and a revised diagnosis of Scleromochlus, revealing new anatomical details that conclusively identify it as a close pterosaur relative1 within Pterosauromorpha (the lagerpetid + pterosaur clade). Scleromochlus is anatomically more similar to lagerpetids than to pterosaurs and retains numerous features that were probably present in very early diverging members of Avemetatarsalia (bird-line archosaurs). These results support the hypothesis that the first flying reptiles evolved from tiny, probably facultatively bipedal, cursorial ancestors1.


Subject(s)
Dinosaurs , Fossils , Phylogeny , Animals , Dinosaurs/classification , Ecosystem , Models, Biological
13.
Nat Commun ; 13(1): 3787, 2022 07 01.
Article in English | MEDLINE | ID: mdl-35778378

ABSTRACT

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.


Subject(s)
Ecosystem , Plants , Animals , Forests , Mammals , Phylogeny , Tibet
14.
Sci Adv ; 8(6): eabj0944, 2022 Feb 11.
Article in English | MEDLINE | ID: mdl-35138908

ABSTRACT

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.

15.
Proc Natl Acad Sci U S A ; 118(45)2021 11 09.
Article in English | MEDLINE | ID: mdl-34725155

ABSTRACT

While paleoclimate records show that the Earth System is characterized by several different tipping points, their representation within Earth System models (ESMs) remains poorly constrained. This is because historical observations do not encompass variations large enough to provoke such regime changes, and paleoclimate conditions are rarely used to help develop and tune ESMs, which potentially ignores a rich source of information on abrupt climate change. A critical example is the early to mid-Holocene "greening" and subsequent rapid desertification of the Sahara, which most ESMs fail to reproduce, casting doubt on the representation of land-atmosphere coupling and monsoon dynamics. Here, we show that this greening and abrupt termination can be successfully simulated with one ESM after optimizing uncertain model components using both present-day observations and crucially mid-Holocene (6,000 y before present) reconstructions. The optimized model displays abrupt threshold behavior, which shows excellent agreement with long paleoclimate records that were not used in the original optimization. These results suggest that in order to realistically capture climate-system thresholds, ESMs first need to be conditioned with appropriate paleoclimate information.

16.
Natl Sci Rev ; 8(1): nwaa091, 2021 Jan.
Article in English | MEDLINE | ID: mdl-34691550

ABSTRACT

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.

17.
Sci Data ; 8(1): 228, 2021 08 27.
Article in English | MEDLINE | ID: mdl-34453060

ABSTRACT

Curated global climate data have been generated from climate model outputs for the last 120,000 years, whereas reconstructions going back even further have been lacking due to the high computational cost of climate simulations. Here, we present a statistically-derived global terrestrial climate dataset for every 1,000 years of the last 800,000 years. It is based on a set of linear regressions between 72 existing HadCM3 climate simulations of the last 120,000 years and external forcings consisting of CO2, orbital parameters, and land type. The estimated climatologies were interpolated to 0.5° resolution and bias-corrected using present-day climate. The data compare well with the original HadCM3 simulations and with long-term proxy records. Our dataset includes monthly temperature, precipitation, cloud cover, and 17 bioclimatic variables. In addition, we derived net primary productivity and global biome distributions using the BIOME4 vegetation model. The data are a relevant source for different research areas, such as archaeology or ecology, to study the long-term effect of glacial-interglacial climate cycles for periods beyond the last 120,000 years.

18.
Proc Biol Sci ; 288(1957): 20211342, 2021 08 25.
Article in English | MEDLINE | ID: mdl-34403638

ABSTRACT

Amidst long-term fluctuations of the abiotic environment, the degree to which life organizes into distinct biogeographic provinces (provinciality) can reveal the fundamental drivers of global biodiversity. Our understanding of present-day biogeography implies that changes in the distribution of continents across climatic zones have predictable effects on habitat distribution, dispersal barriers and the evolution of provinciality. To assess marine provinciality through the Phanerozoic, here we (a) simulate provinces based on palaeogeographic reconstructions and global climate models and (b) contrast them with empirically derived provinces that we define using network analysis of fossil occurrences. Simulated and empirical patterns match reasonably well and consistently suggest a greater than 15% increase in provinciality since the Mesozoic era. Although both factors played a role, the simulations imply that the effect of the latitudinal temperature gradient has been twice as important in determining marine provinciality as continental configuration.


Subject(s)
Biodiversity , Climate Change , Ecosystem , Fossils
19.
Proc Natl Acad Sci U S A ; 118(18)2021 05 04.
Article in English | MEDLINE | ID: mdl-33903233

ABSTRACT

Abiotic niche lability reduces extinction risk by allowing species to adapt to changing environmental conditions in situ. In contrast, species with static niches must keep pace with the velocity of climate change as they track suitable habitat. The rate and frequency of niche lability have been studied on human timescales (months to decades) and geological timescales (millions of years), but lability on intermediate timescales (millennia) remains largely uninvestigated. Here, we quantified abiotic niche lability at 8-ka resolution across the last 700 ka of glacial-interglacial climate fluctuations, using the exceptionally well-known fossil record of planktonic foraminifera coupled with Atmosphere-Ocean Global Climate Model reconstructions of paleoclimate. We tracked foraminiferal niches through time along the univariate axis of mean annual temperature, measured both at the sea surface and at species' depth habitats. Species' temperature preferences were uncoupled from the global temperature regime, undermining a hypothesis of local adaptation to changing environmental conditions. Furthermore, intraspecific niches were equally similar through time, regardless of climate change magnitude on short timescales (8 ka) and across contrasts of glacial and interglacial extremes. Evolutionary trait models fitted to time series of occupied temperature values supported widespread niche stasis above randomly wandering or directional change. Ecotype explained little variation in species-level differences in niche lability after accounting for evolutionary relatedness. Together, these results suggest that warming and ocean acidification over the next hundreds to thousands of years could redistribute and reduce populations of foraminifera and other calcifying plankton, which are primary components of marine food webs and biogeochemical cycles.


Subject(s)
Adaptation, Physiological/genetics , Biological Evolution , Climate Models , Foraminifera/physiology , Plankton/physiology , Climate Change , Ecosystem , Foraminifera/genetics , Fossils , Humans , Seawater/microbiology , Temperature
20.
Sci Adv ; 7(5)2021 01.
Article in English | MEDLINE | ID: mdl-33571113

ABSTRACT

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.

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