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Herbivore damage patterns on fossil leaves are essential to explore the evolution of plant-herbivore interactions under paleoenvironmental changes and to better understand the evolutionary history of terrestrial ecosystems. The Eocene–Oligocene transition (EOT) is a period of dramatic paleoclimate changes that significantly impacted global ecosyste...
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... Although the Markam and Lühe score sheets are already included in the online database (www.iprdatabase. eu), the ages of these two sites have been updated and more fossil species have been reported from assemblages in recent years (Su et al., 2019;Deng et al., 2020;Wu et al., 2022). Therefore, new analyses are conducted for these two sites. ...
The Eocene-Oligocene transition (EOT) marked a rapid global cooling event, often considered as the beginning of the modern icehouse world. Influenced by various factors, including tectonic activity and paleogeographic settings, the terrestrial records indicate a diverse response of fauna and vegetation to this global event. We examined nine macrofossil assemblages from seven fossil localities on the southeastern margin of the Tibetan Plateau and from the mid-latitudinal Europe ranging from the latest Bartonian and Priabonian (37.71–33.9 Ma) to the Rupelian (33.9–27.82 Ma). Our aims were to trace and compare the vegetation history of both regions in the late Eocene and early Oligocene. The results show that both regions experienced changes in vegetation composition in response to climate change, characterized by a decrease in the percentages of broad-leaved evergreen elements and distinctive changes in general vegetation types. A general change in the overall vegetation type from subtropical broad-leaved evergreen forests in the late Eocene to temperate broad-leaved mixed deciduous evergreen forests, or mixed mesophytic forests, in the early Oligocene is recognized in both regions. The results indicate a clear change in leaf architecture, leaf margin states, and secondary venation types in the mid-latitudinal Europe, while the results from the south-eastern margin of the Tibetan Plateau show a distinct reduction in leaf size. Our data suggest that both global and regional factors played key roles in shaping the vegetation in the two regions.
... The previously presumed Miocene strata (WGCPC, 1978; BGMRYP (Bureau of Geology and Mineral Resources of Yunnan Province), 1990, BGMRYP (Bureau of Geology and Mineral Resources of Yunnan Province), 1996; Xu et al., 2008;Li et al., 2015;Huang et al., 2016bHuang et al., , 2017Huang et al., , 2018 for many fossil floras in southwestern China has now been revised to be late Eocene/early Oligocene in age (Gourbet et al., 2017;Linnemann et al., 2018;Su et al., 2019;Tian et al., 2021). Evergreen representatives, including Fagaceae, Lauraceae and Fabaceae, are main components in these floras (Huang, 2017;Su et al., 2019;Deng et al., 2020;Tian et al., 2021;Wu et al., 2022b). The revised geological ages and components of fossil floras indicate that the temporal occurrence of EBLFs may have been advanced to the late Eoceneeearly Oligocene in southwestern China. ...
... Heterobalanus and conifers (Pinus, Abies, Picea, Tsuga) as the dominant elements (Tao and Kong, 1973;Huang et al., 2012Huang et al., , 2013Huang et al., , 2015Huang et al., , 2020Su et al., 2013;Zhu et al., 2015Zhu et al., , 2016. Some fossil assemblages (e.g., late Eocene Markam assemblage and early Oligocene Lühe assemblage in southwestern China, late Oligocene Noda assemblage in Japan and middle Miocene Shanwang assemblage in centraleastern China) represent evergreen-deciduous broadleaved mixed forests, dominated by the evergreen groups of Fagaceae, Magnoliaceae, Lauraceae and the deciduous groups of Betulaceae, Ulmaceae, Salicaceae (Hu and Chaney, 1940;Tanai and Uemura, 1991;Su et al., 2019;Deng et al., 2020;Wu et al., 2022b). Some fossil assemblages (e.g., early Oligocene Markam assemblage) represent alpine deciduous shrubs. ...
... Small-leaved shrubs such as Salix, Rosa and Quercus sect. Heterobalanus are main components (Su et al., 2019;Deng et al., 2020). Additionally, some fossil assemblages (e.g., middle Miocene Zhangpu assemblage and Guiping assemblage) represent tropical rainforests, dominated by tropical taxa including Annonaceae, Myrtaceae, and Burseraceae (Wang, 2018;Wang et al., 2021;Huang et al., 2021;Li et al., 2021b;Wu et al., 2021;Song et al., 2023). ...
Evergreen broad-leaved forests (EBLFs) are widely distributed in East Asia and play a vital role in ecosystem stability. The occurrence of these forests in East Asia has been a subject of debate across various disciplines. In this study, we explored the occurrence of East Asian EBLFs from a paleobotanical perspective. By collecting plant fossils from four regions in East Asia, we have established the evolutionary history of EBLFs. Through floral similarity analysis and paleoclimatic reconstruction, we have revealed a diverse spatio-temporal pattern for the occurrence of EBLFs in East Asia. The earliest occurrence of EBLFs in southern China can be traced back to the middle Eocene, followed by southwestern China during the late Eocene–early Oligocene. Subsequently, EBLFs emerged in Japan during the early Oligocene and eventually appeared in central-eastern China around the Miocene. Paleoclimate simulation results suggest that the precipitation of wettest quarter (PWetQ, mm) exceeding 600 mm is crucial for the occurrence of EBLFs. Furthermore, the heterogeneous occurrence of EBLFs in East Asia is closely associated with the evolution of the Asian Monsoon. This study provides new insights into the occurrence of EBLFs in East Asia.
... Prunus were estimated to be much younger in this study, about 39.27 Mya than the estimate by Chin et al., but much older than the estimate by Zhang et al. [14]. It has been suggested that the Eocene-Oligocene transition (EOT) around 30-40 Mya marked a period of global cooling that resulted in significant flora turnovers [40][41][42]. ...
Cherries (Prunus Subgenus Cerasus) have economic value and ecological significance, yet their phylogeny, geographic origin, timing, and dispersal patterns remain challenging to understand. To fill this gap, we conducted a comprehensive analysis of the complete chloroplast genomes of 54 subg. Cerasus individuals, along with 36 additional genomes from the NCBI database, resulting in a total of 90 genomes for comparative analysis. The chloroplast genomes of subg. Cerasus exhibited varying sizes and consisted of 129 genes, including protein-coding, transfer RNA, and ribosomIal RNA genes. Genomic variation was investigated through InDels and SNPs, showcasing distribution patterns and impact levels. A comparative analysis of chloroplast genome boundaries highlighted variations in inverted repeat (IR) regions among Cerasus and other Prunus species. Phylogeny based on whole-chloroplast genome sequences supported the division of Prunus into three subgenera, I subg. Padus, II subg. Prunus and III subg. Cerasus. The subg. Cerasus was subdivided into seven lineages (IIIa to IIIg), which matched roughly to taxonomic sections. The subg. Padus first diverged 51.42 Mya, followed by the separation of subg. Cerasus from subg. Prunus 39.27 Mya. The subg. Cerasus started diversification at 15.01 Mya, coinciding with geological and climatic changes, including the uplift of the Qinghai–Tibet Plateau and global cooling. The Himalayans were the refuge of cherries, from which a few species reached Europe through westward migration and another species reached North America through northeastward migration. The mainstage of cherry evolution was on the Qing–Tibet Plateau and later East China and Japan as well. These findings strengthen our understanding of the evolution of cherry and provide valuable insights into the conservation and sustainable utilization of cherry’s genetic resources.
... In addition, the Eocene-Oligocene transition (EOT, 30-40 Mya) in the Cenozoic witnessed a global cooling and led to a reorganization of organisms (Prothero, 1994;Sun et al., 2014;Deng et al., 2020). The diversification of clades 1 and 2 was herein estimated to be around 35.41 Mya (95% HPD: 26.51-43.39 ...
Prunus is an economically important genus widely distributed in the temperate Northern Hemisphere. Previous studies on the genus using a variety of loci yielded conflicting phylogenetic hypotheses. Here, we generated nuclear reduced representation sequencing data and plastid genomes for 36 Prunus individuals and two outgroups. Both nuclear and plastome data recovered a well-resolved phylogeny. The species were divided into three main clades corresponding to their inflorescence types, - the racemose group, the solitary-flower group and the corymbose group - with the latter two sister to one another. Prunus was inferred to have diversified initially in the Late Cretaceous around 67.32 million years ago. The diversification of the three major clades began between the Paleocene and Miocene, suggesting that paleoclimatic events were an important driving force for Prunus diversification. Ancestral state reconstructions revealed that the most recent common ancestor of Prunus had racemose inflorescences, and the solitary-flower and corymb inflorescence types were derived by reduction of flower number and suppression of the rachis, respectively. We also tested the hybrid origin hypothesis of the racemose group proposed in previous studies. Prunus has undergone extensive hybridization events, although it is difficult to identify conclusively specific instances of hybridization when using SNP data, especially deep in the phylogeny. Our study provides well-resolved nuclear and plastid phylogenies of Prunus, reveals substantial cytonuclear discord at shallow scales, and sheds new light on inflorescence evolution in this economically important lineage.
... The oldest fossil of the sub-genus Equisetum is E. bryanii Gould, which has well-preserved stems and leaf sheaths, from the Jurassic of south-eastern Queensland (Gould, 1968). Some fossilized tuberous rhizome parts of Equisetum have also been reported from Asia (Guo, 2000;Zhou et al., 2003;Sun et al., 2013;Yang et al., 2016;Deng et al., 2020), Europe (Denk et al., 2005), and North America (Skog and Dilcher, 1994). ...
... Four Miocene fossil species of Equisetum have been reported from the European continental region (Heer, 1868;Denk et al., 2005Denk et al., , 2011; Barr on and Postigo-Mijarra, 2011). A fossil species E. oppositum was recovered at different epochs (Palaeocene-Eocene, Oligocene, and Miocene) of the Cenozoic Era from China (Ma et al., 2012;Yang et al., 2016;Deng et al., 2020), which indicates that this Equisetum species used to grow in abundance during that era. Aung et al. (2020) al., 2021). ...
Equisetum L. (Equisetaceae) has long been a focus of attention for botanists and palaeontologists because, given its extensive and well-documented fossil record, it is considered the oldest extant vascular plant and a key element in understanding vascular plant evolution. However, to date, no authentic fossil evidence of Equisetum has been found from the Indian Cenozoic. Here, we describe a new fossil species, namely, E. siwalikum sp. nov., recovered from the middle Siwalik (Late Miocene) sediments of Himachal Pradesh, western Himalaya. We identified fossil specimens based on morphological and epidermal characters. In addition, X-Ray diffraction (XRD) analysis was used to determine the mineral composition of compressed stems of Equisetum. The close affinity of our recovered Siwalik fossils to Equisetum is supported by the presence of both macromorphological and epidermal characters. Because Equisetum generally grows in wet conditions around water reservoirs, our findings indicate that the fossil locality was humid and surrounded by swamp and lowland regions during deposition. Ample fossil evidence indicates that this sphenopsid once existed in the western Himalaya during the Siwalik period. However, at present Equisetum is confined to a particular area of our fossil locality, probably a consequence of severe environmental changes coupled with competition from opportunistic angiosperms. Our discovery of Equisetum fossils in appreciable numbers from the Siwalik sediments of the Himachal Himalayas is unique and constitutes the first reliable recognition of Equisetum from the Indian Cenozoic.
... Nonetheless, component communities in the fossil record have been widely discussed using damage types as proxies for herbivore taxa (Labandeira 1998(Labandeira , 2002D'Rozario et al. 2011;Slater et al. 2012Slater et al. , 2015Labandeira and Currano 2013;Labandeira et al. , 2016Labandeira et al. , 2018Ding et al. 2014Ding et al. , 2015Schachat et al. 2014Schachat et al. , 2015Feng et al. 2017;Kustatscher et al. 2018;Xu et al. 2018;Correia et al. 2020;). However, here too, there is reason for caution: even the fossil floras that have been most thoroughly sampled for insect herbivory contain various damage types that occur on only one specimen (Wilf et al. 2005(Wilf et al. , 2006Prevec et al. 2009;Wappler 2010;Knor et al. 2012;Wappler et al. 2012;Donovan et al. 2014;Adroit et al. 2018;Labandeira et al. 2018;Xu et al. 2018;Deng et al. 2020), indicating that many damage types remain unobserved due to incomplete preservation and sampling. Because we cannot find every damage type from a fossil assemblage, and because we cannot link damage types to the insect taxa in a one-to-one manner, the term "component community" as developed in the context of modern ecology may be somewhat inapplicable. ...
Studies of insect herbivory on fossilized leaves tend to focus on a few, relatively simple metrics that are agnostic to the distribution of insect damage types among host plants. More complex metrics that link particular damage types to particular host plants have the potential to address additional ecological questions, but such metrics can be biased by sampling incompleteness due to the difficulty of distinguishing the true absence of a particular interaction from the failure to detect it—a challenge that has been raised in the ecological literature. We evaluate a range of methods for characterizing the relationships between damage types and host plants by performing resampling and subsampling exercises on a variety of datasets. We found that the components of beta diversity provide a more valid, reliable, and interpretable method for comparing component communities than do bipartite network metrics and that the rarefaction of interactions represent a valid, reliable, and interpretable method for comparing compound communities. Both beta diversity and rarefaction of interactions avoid the potential pitfalls of multiple comparisons. Finally, we found that the host specificity of individual damage types is challenging to assess. Whereas bipartite network metrics are sufficiently biased by sampling incompleteness to be inappropriate for fossil herbivory data, alternatives exist that are perfectly suitable for fossil datasets with sufficient sample coverage.
... A number of fossil floras with well constrained geological ages have been discovered, such as the Bangor (~47 Ma; Su et al., 2020), Lunpola (50-20 Ma; He et al., 2012;Sun et al., 2014;Su et al., 2019b;Xiong et al., 2022), Markam (~34.6 Ma and~33.4 Ma; Su et al., 2019b;Deng W Y D et al., 2020), and Zanda (6.4-0.4 Ma; Wang X M et al., 2013;Huang et al., 2020) floras. ...
... The warm and humid airflow brought by the ISM can cross the southern Gangdese Mountains, so the tropical-subtropical flora and fauna would have been able to grow in the valley (Wu et al., 2017;Jia et al., 2018;Jiang et al., 2019;Liu et al., 2019;Su et al., 2019b;Deng et al., 2019a). Plant fossil assemblages and the latest stratigraphic chronology studies in the Markam Basin showed that the southeast margin of the QTP experienced uplift from the Eocene to Oligocene, resulting in the transformation of the vegetation from evergreen deciduous broadleaved mixed forest to alpine deciduous shrubs Deng W Y D et al., 2020). From the late Paleogene to Neogene, with the rapid uplift of the plateau, the low altitude basins would have gradually disappeared, with the tropicalsubtropical flora and fauna on the verge of extinction, and the glacial fauna adapted to a severe cold climate starting to appear (Wang et al., 2016;Deng et al., 2019b). ...
The growth of the Qinghai-Tibetan Plateau (QTP) during the Cenozoic drove dramatic climate and environmental change in this region. However, there has been limited comprehensive research into evolution of climate during this interval. Here we present a quantitative reconstruction using Bioclimatic Analysis (BA) and Joint Probability Density Functions (JPDFs) based on data available for 48 fossil floras, including macrofossils and palynological fossils collected in the QTP area from the Paleogene to Neogene (66–2.58 Ma). Both methods indicate that there was an overall decline in temperature and precipitation. Paleoclimatic simulations using Hadley Centre Coupled Model version3 (HadCM3) show that the most prominent climate change was very likely driven by QTP orographic evolution from the late Eocene, which was accompanied by a shift in temperature from a latitudinal distribution to a topographically controlled pattern. In addition, with the growth of the QTP, temperature and precipitation decreased gradually in the northeastern part of the plateau. Different sources of evidence, including plant fossil records, climate simulations and other proxies, indicate that the topographic evolution of the QTP and other geological events, in conjunction with global cooling, may have been the main factors driving climate change in this region. This research can provide insights into Cenozoic environmental change and ecosystem evolution.
... Because they experienced low winter temperatures the Markam and Lühe paleofloras contained a large number of deciduous components . In the early Oligocene the Markam area attained near its present elevation, and the vegetation changed from a subtropical forest to alpine shrubland Deng et al., 2020). In the late Miocene some winter temperatures in Yunnan approached those of present (Jacques et al., 2014), and the percentage of deciduous elements such as Betulaceae decreased, while the evergreen elements such as Fagaceae and Lauraceae became dominant in the middle Miocene Zhenyuan flora (Wang et al., 2019a and the late Miocene Xianfeng and Xiaolongtan floras (Zhou, 1985;Xia et al., 2009;Xing et al., 2012;Jacques et al., 2014;Li et al., 2015a;Zhang et al., 2019). ...
Quantifying the interactions between topography, climate and plant diversity within one of the world's biodiversity hotspots, the southeastern margin of the Tibetan Plateau, remains elusive due to few reliable quantitative paleoelevation reconstructions, precise geological age constraints and well-preserved plant fossils. The Lühe Basin, on the southeastern margin of Tibetan Plateau has yielded abundant plant fossils with a U-Pb age of 33–32 Ma, providing an opportunity to estimate the elevation of this region and plant diversity at that time. Fossil leaf physiognomy was used to reconstruct the paleoclimate and the paleoelevation of the basin was derived from moist enthalpy. The results show that the Lühe Basin, had attained it's present elevation (1.7 ± 0.9 km) by the early Oligocene and, compared to now, experienced a humid subtropical climate with a wetter dry season and lower precipitation seasonality in an overall wetter precipitation regime (1748.5 ± 606 mm). This was accompanied by a greater seasonal range in temperature, although the mean annual temperature (14.9 ± 2.3 °C) was similar to that of today (15.6 °C). Combined with previous studies, we conclude that the appearance of the modern flora across the southeastern margin of the Tibetan Plateau had started by the early Oligocene, corresponding with the establishment of modern topography at that time.
... For the coal mine section, biomarker analysis shows no marked cooling, which is similar to results from the town section (Lauretano et al., 2021, under review). Elsewhere in the nearby region, such as in Markam Deng et al., 2020) and Jianchuan (Wu et al., 2018), interpretations of climate and vegetation changes across the EOT are complicated because of possible contemporaneous regional uplift. While it seems that heterogenous EOT climate changes took place worldwide, regional environment in southwestern China appears to have been relatively stable. ...
Cupressaceae fossil tree stumps from the early Oligocene Lühe coal mine in southwestern China contain abundant quartz-petrified damage traces. The wood fossils were assigned to Taxodioxylon (very similar to extant Taxodium) based on wood anatomy analysis. Within the woods, three types of arthropods- and one fungus-mediated ichnofossils LHIF 1–4 (Lühe wood ichnofossils 1–4) were observed. The boring wood types for LHIF 1–3 are comparable to extant longicorn beetles (Cerambycidae), snout beetles (Curculionidae), and wood wasps (Siricidae). The polyporous structured traces of LHIF 4 were attributed to the invasion of stem canker fungus (Polyporaceae). This first-ever report of Taxodium-like fossil from the Oligocene of southwestern China points to Yunnan serving as a refugium for some lineage of gymnosperms at that time. Furthermore, the extensive traces of arthropods and fungus discovered from the fossil wood have filled a gap in fossil records for insect herbivory in this region. The wood stumps and decomposers suggest a swamp-like environment. The disappearance of Taxodium after the Oligocene supports incremental aridification and changes in winter season temperature conditions, which shifted the late Paleogene mixed deciduous broad-leaved and needle-leaved forest into the present evergreen broad-leaved forest.
... Our results of divergence time estimation indicated that the genus Parnassia began to diverge and spread to non-Pan-Himalayan areas during the Eocene. Combining with the Qinghai-Tibetan Plateau uplift, the global climate oscillations led to a significant change in plant diversity during the Eocene-Oligocene transition period (Abels et al., 2011;Deng et al., 2020). As a result, the differentiation Parnassia species began to appear at Oligocene in the Pan-Himalayan region, and then mountain building and Asian monsoon may have combined to promote species diversification and colonization in the early to middle Miocene (Xing and Ree, 2017;Ding et al., 2020). ...
Parnassia L., a perennial herbaceous genus in the family Celastraceae, consists of about 60 species and is mainly distributed in the Pan-Himalayan and surrounding mountainous regions. The taxonomic position and phylogenetic relationships of the genus are still controversial. Herein, we reassessed the taxonomic status of Parnassia and its intra- and inter-generic phylogeny within Celastraceae. To that end, we sequenced and assembled the whole plastid genomes and nuclear ribosomal DNA (nrDNA) of 48 species (74 individuals), including 25 species of Parnassia and 23 species from other genera of Celastraceae. We integrated high throughput sequence data with advanced statistical toolkits and performed the analyses. Our results supported the Angiosperm Phylogeny Group IV (APG IV) taxonomy which kept the genus to the family Celastraceae. Although there were topological conflicts between plastid and nrDNA phylogenetic trees, Parnassia was fully supported as a monophyletic group in all cases. We presented a first attempt to estimate the divergence of Parnassia, and molecular clock analysis indicated that the diversification occurred during the Eocene. The molecular phylogenetic results confirmed numerous taxonomic revisions, revealing that the morphological characters used in Parnassia taxonomy and systematics might have evolved multiple times. In addition, we speculated that hybridization/introgression might exist during genus evolution, which needs to be further studied. Similarly, more in-depth studies will clarify the diversification of characters and species evolution models of this genus.
