Tao Su

LEAF MARGIN ANALYSIS: A NEW EQUATION FROM HUMID TO MESIC FORESTS IN CHINA

Abstract Leaf margin analysis (LMA) is a widely used method that applies present-day linear correlation between the proportion of woody dicotyledonous species with untoothed leaves (P) and mean annual temperature (MAT) to estimate paleotemperatures from fossil leaf floras. Previous works demonstrate that LMA shows regional constraints and to date, no equation has been modeled directly from Chinese forests. Fifty humid to mesic Chinese forests were chosen to understand the relationship between percentage of untoothed leaf species and MAT in China. Consistent with previous studies, the Chinese data indicate that P shows a strong linear correlation with MAT, but the actual relationship is a little different from those recognized from other regions. Among the several currently used LMA equations, the one resulting from North and Central American and Japanese data, rather than the widely used East Asian LMA equation, yields the closest values to the actual MATs of the Chinese samples (mean absolute error = 1.9 °C). A new equation derived from the Chinese forests is therefore developed, where MAT = 1.038 + 27.6 × P. This study not only demonstrates the similarity of the relationship between P and MAT in the Northern Hemisphere, but also improves the reliability of LMA for paleoclimate reconstructions of Chinese paleofloras.

Phylogeographic analysis reveals significant spatial genetic structure of Incarvillea sinensis as a product of mountain building

BackgroundIncarvillea sinensis is widely distributed from Southwest China to Northeast China and in the Russian Far East. The distribution of this species was thought to be influenced by the uplift of the Qinghai-Tibet Plateau and Quaternary glaciation. To reveal the imprints of geological events on the spatial genetic structure of Incarvillea sinensis, we examined two cpDNA segments ( trnH- psbA and trnS- trnfM) in 705 individuals from 47 localities.ResultsA total of 16 haplotypes was identified, and significant genetic differentiation was revealed (GST =0.843, NST = 0.975, P < 0.05). The survey detected two highly divergent cpDNA lineages connected by a deep gap with allopatric distributions: the southern lineage with higher genetic diversity and differentiation in the eastern Qinghai-Tibet Plateau, and the northern lineage in the region outside the Qinghai-Tibet Plateau. The divergence between these two lineages was estimated at 4.4 MYA. A correlation between the genetic and the geographic distances indicates that genetic drift was more influential than gene flow in the northern clade with lower diversity and divergence. However, a scenario of regional equilibrium between gene flow and drift was shown for the southern clade. The feature of spatial distribution of the genetic diversity of the southern lineage possibly indicated that allopatric fragmentation was dominant in the collections from the eastern Qinghai-Tibet Plateau.ConclusionsThe results revealed that the uplift of the Qinghai-Tibet Plateau likely resulted in the significant divergence between the lineage in the eastern Qinghai-Tibet Plateau and the other one outside this area. The diverse niches in the eastern Qinghai-Tibet Plateau created a wide spectrum of habitats to accumulate and accommodate new mutations. The features of genetic diversity of populations outside the eastern Qinghai-Tibet Plateau seemed to reveal the imprints of extinction during the Glacial and the interglacial and postglacial recolonization. Our study is a typical case of the significance of the uplift of the Qinghai-Tibet Plateau and the Quaternary Glacial in spatial genetic structure of eastern Asian plants, and sheds new light on the evolution of biodiversity in the Qinghai-Tibet Plateau at the intraspecies level.

Sequoia maguanensis, a new Miocene relative of the coast redwood, Sequoia sempervirens, from China: Implications for paleogeography and paleoclimate

UNLABELLED • PREMISE OF THE STUDY The paleogeographical origin of the relict North American Sequoia sempervirens is controversial. Fossil records indicate a Neogene origin for its foliage characteristics. Although several fossils from the Miocene sediments in eastern Asia have been considered to have close affinities with the modern S. sempervirens, they lack the typical features of a leafy twig bearing linear as well as scale leaves, and the fertile shoots terminating by a cone. The taxonomic status of these fossils has remained unclear.• METHODS New better-preserved fossils from the upper Miocene of China indicate a new species of Sequoia. This finding not only confirms the former presence of this genus in eastern Asia, but it also confirms the affinity of this Asian form to the modern relict S. sempervirens.• KEY RESULTS The principal foliage characteristics of S. sempervirens had already originated by the late Miocene. The eastern Asian records probably imply a Beringian biogeographic track of the ancestor of S. sempervirens in the early Neogene, at a time when the land bridge was not too cool for this thermophilic conifer to spread between Asia and North America.• CONCLUSIONS The climatic context of the new fossil Sequoia in Southeast Yunnan, based on other floristic elements of the fossil assemblage in which it is found, is presumed to be warm and humid. Following the uplift of the Qinghai-Tibet Plateau, this warm, humid climate was replaced by the present monsoonal climate with dry winter and spring. This change may have led to the disappearance of this hygrophilous conifer from eastern Asia.

Cenozoic plant diversity of Yunnan: A review

Yunnan in southwestern China is renowned for its high plant diversity. To understand how this modern botanical richness formed, it is critical to investigate the past biodiversity throughout the geological time. In this review, we present a summary on plant diversity, floristics and climates in the Cenozoic of Yunnan and document their changes, by compiling published palaeobotanical sources. Our review demonstrates that thus far a total of 386 fossil species of ferns, gymnosperms and angiosperms belonging to 170 genera within 66 families have been reported from the Cenozoic, particularly the Neogene, of Yunnan. Angiosperms display the highest richness represented by 353 species grouped into 155 genera within 60 families, with Fagaceae, Fabaceae, Lauraceae and Juglandaceae being the most diversified. Most of the families and genera recorded as fossils still occur in Yunnan, but seven genera have disappeared, including Berryophyllum, Cedrelospermum, Cedrus, Palaeocarya, Podocarpium, Sequoia and Wataria. The regional extinction of these genera is commonly referred to an aridification of the dry season associated with Asian monsoon development. Floristic analyses indicate that in the late Miocene, Yunnan had three floristic regions: a northern subtropical floristic region in the northeast, a subtropical floristic region in the east, and a tropical floristic region in the southwest. In the late Pliocene, Yunnan saw two kinds of floristic regions: a subalpine floristic region in the northwest, and two subtropical floristic regions separately in the southwest and the eastern center. These floristic concepts are verified by results from our areal type analyses which suggest that in the Miocene southwestern Yunnan supported the most Pantropic elements, while in the Pliocene southwestern Yunnan had abundant Tropical Asia (Indo–Malaysia) type and East Asia and North America disjunct type that were absent from northwestern Yunnan. From the late Miocene to late Pliocene through to the present, floristic composition and vegetation types changed markedly, presumably in response to altitude changes and coeval global cooling. An integration of palaeoclimate data suggests that during the Neogene Yunnan was warmer and wetter than today. Moreover, northern Yunnan witnessed a pronounced temperature decline, while southern Yunnan experienced only moderate temperature changes. Summer precipitation was consistently higher than winter precipitation, suggesting a rainfall seasonality. This summary on palaeoclimates helps us to understand under what conditions plant diversity occurred and evolved in Yunnan throughout the Cenozoic.

Distribution of Cenozoic plant relicts in China explained by drought in dry season

Cenozoic plant relicts are those groups that were once widespread in the Northern Hemisphere but are now restricted to some small isolated areas as a result of drastic climatic changes. They are good proxies to study how plants respond to climatic changes since their modern climatic requirements are known. Herein we look at the modern distribution of 65 palaeoendemic genera in China and compare it with the Chinese climatic pattern, in order to find a link between the plant distribution and climate. Central China and Taiwan Island are shown to be diversity centres of Cenozoic relict genera, consistent with the fact that these two regions have a shorter dry season with comparatively humid autumn and spring in China. Species distribution models indicate that the precipitation parameters are the most important variables to explain the distribution of relict genera. The Cenozoic wide-scale distribution of relict plants in the Northern Hemisphere is therefore considered to be linked to the widespread humid climate at that time, and the subsequent contraction of their distributional ranges was probably caused by the drying trend along with global cooling.

Resilience of plant-insect interactions in an oak lineage through Quaternary climate change

Abstract. Plant-insect interactions are vital for structuring terrestrial ecosystems. It is still unclear how climate change in geological time might have shaped plant-insect interactions leading to modern ecosystems. We investigated the effect of Quaternary climate change on plant-insect interactions by observing insect herbivory on leaves of an evergreen sclerophyllous oak lineage (Quercus section Heterobalanus, HET) from a late Pliocene flora and eight living forests in southwestern China. Among the modern HET populations investigated, the damage diversity tends to be higher in warmer and wetter climates. Even though the climate of the fossil flora was warmer and wetter than modern sample sites, the damage diversity is lower in the fossil flora than in modern HET populations. Eleven out of 18 damage types in modern HET populations are observed in the fossil flora. All damage types in the fossil flora, except for one distinctive gall type, are found in modern HET populations. These results indicate that Quaternary climate change did not cause extensive extinction of insect herbivores in HET forests. The accumulation of a more diverse herbivore fauna over time supports the view of plant species as evolutionary “islands” for colonization and turnover of insect species.

Evidence for insect-mediated skeletonization on an extant fern family from the Upper Triassic of China

Leaf skeletonization represents a distinctive form of insect feeding behavior. It commonly occurs in angiosperm leaves after their initial appearance during the Early Cretaceous. This type of feeding behavior rarely has been documented in pre-Mesozoic fossils. We describe the earliest evidence of insect-skeletonized leaves of Dictyophyllum nathorstii Zeiller, affi li- ated with the extant fern family Dipteridaceae in the Late Triassic Yipinglang fl ora from southwestern China. The skeletonization generally is located adjacent to the pinna rachilla of the distal free portion of the leaf. In the skeletonized area, the interveinal tissue is com- pletely removed, exposing the pinna rachilla, pinnule midveins, and lateral veins. Most non- vascular tissue has been removed between the vascular bundles, the latter forming polygo- nal meshes of varying size. Our report of insect-mediated skeletonization of fern leaves from southwestern China fi lls a spatiotemporal gap in the published data on the paleogeographi- cal distribution and stratigraphic occurrence of plant-arthropod associations, and indicates an antagonistic relationship between a fern host and its insect herbivore.

The oldest Mahonia (Berberidaceae) fossil from East Asia and its biogeographic implications.

Interpretation of the biogeography of the genus Mahonia (Berberidaceae) is limited by the lack of fossil records in East Asia. Compressed fossil foliage, described here as Mahonia mioasiatica sp. nov., were collected from the Upper Miocene Xiaolongtan Formation in Wenshan, Yunnan, southwest China. These specimens represent the oldest reliable fossil record of Mahonia in East Asia. This new fossil species shows a general similarity to Group Orientales and is most similar to the extant eastern Asian Mahoniaconferta. Considering other fossil evidence of Mahonia, we propose a migration route of this genus to Asia over the North Atlantic Land Bridge rather than the Bering Land Bridge. Our results also suggest that North America, Europe and East Asia have been successive centers of diversity for the genus, as a consequence of diversification in Group Orientales potentially related to historical climate change.

A new positive relationship between pCO2 and stomatal frequency in Quercus guyavifolia (Fagaceae): a potential proxy for palaeo-CO2 levels.

BACKGROUND AND AIMS The inverse relationship between atmospheric CO2 partial pressure (pCO2) and stomatal frequency in many species of plants has been widely used to estimate palaeoatmospheric CO2 (palaeo-CO2) levels; however, the results obtained have been quite variable. This study attempts to find a potential new proxy for palaeo-CO2 levels by analysing stomatal frequency in Quercus guyavifolia (Q. guajavifolia, Fagaceae), an extant dominant species of sclerophyllous forests in the Himalayas with abundant fossil relatives. METHODS Stomatal frequency was analysed for extant samples of Q. guyavifolia collected from17 field sites at altitudes ranging between 2493 and 4497 m. Herbarium specimens collected between 1926 and 2011 were also examined. Correlations of pCO2-stomatal frequency were determined using samples from both sources, and these were then applied to Q. preguyavaefolia fossils in order to estimate palaeo-CO2 concentrations for two late-Pliocene floras in south-western China. KEY RESULTS In contrast to the negative correlations detected for most other species that have been studied, a positive correlation between pCO2 and stomatal frequency was determined in Q. guyavifolia sampled from both extant field collections and historical herbarium specimens. Palaeo-CO2 concentrations were estimated to be approx. 180-240 ppm in the late Pliocene, which is consistent with most other previous estimates. CONCLUSIONS A new positive relationship between pCO2 and stomatal frequency in Q. guyavifolia is presented, which can be applied to the fossils closely related to this species that are widely distributed in the late-Cenozoic strata in order to estimate palaeo-CO2 concentrations. The results show that it is valid to use a positive relationship to estimate palaeo-CO2 concentrations, and the study adds to the variety of stomatal density/index relationships that available for estimating pCO2. The physiological mechanisms underlying this positive response are unclear, however, and require further research.

New U-Pb dates show a Paleogene origin for the modern Asian biodiversity hot spots

Yunnan, in southwestern China, straddles two of the world’s most important biodiversity hot spots (i.e., a biogeographic region that is both a reservoir of biodiversity and threatened with destruction) and hosts more than 200 fossiliferous sedimentary basins documenting the evolutionary history of that biodiversity, monsoon development, and regional elevation changes. The fossil biotas appear modern and have been assumed to be mostly Miocene in age. Dating has been by cross-correlation using palynology, magnetostratigraphy, and lithostratigraphy because numerical radiometric ages are lacking. Here we report the first unequivocal early Oligocene age (33–32 Ma) of a section in the Lühe Basin (25.141627°N, 101.373840°E, 1890 m above mean sea level), central Yunnan, based on U-Pb zircon dates of unreworked volcanic ash layers in a predominantly lacustrine succession hosting abundant plant and animal fossils. This section, located in Lühe town, is correlated with an adjacent section in the Lühe coal mine previously assigned to the upper Miocene based on regional lithostratigraphic comparison. Our substantially older age for the Lühe town section calls into question previous estimates for the surface uplift and climate history of the area, and the age of all other correlative basins. The modernization of the biota ~20 m.y. earlier than previously thought overturns existing concepts of vegetation history in southwestern China, and points to Paleogene modernization of the biota in Yunnan and associated Asian biodiversity hot spots.