Yong-Jiang Huang

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.

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.

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.

Late Miocene Palaeocarya (Engelhardieae: Juglandaceae) from Southwest China and its biogeographic implications

Fossil fruits of Palaeocarya (Juglandaceae) are described from late Miocene sediments of southeastern Yunnan, China. The fruits present a tri‐lobed wing consisting of an intact oblong‐ovate middle lobe and two lateral lobes. The lobes are apically obovate, and have pinnate venation. The middle lobe is thicker at the base and gradually tapers to the apex. The nutlet, located at the base of the winged fruit, is round and hispid, and is subdivided by a septum into two compartments. Based on extensive morphological comparisons to previously documented fossil fruits, we found that the fossil fruits align most closely with members of the genus Palaeocarya, but have a unique combination of characters. Thus, we describe the fossils as a new species, Palaeocarya hispida sp. nov. This species represents an important range expansion for low‐latitude occurrences of Palaeocarya in the late Miocene and therefore substantially improves our understanding of the biogeographic history of the genus. We propose that the wide distribution of Palaeocarya and relatively narrow distributions of close relatives, Engelhardia, Alfaropsis, and Oreomunnea, might be associated with a stepwise cooling and a major ice sheet expansion in the Antarctic and Arctic from the late middle Miocene to early Pliocene. In particular, the climatic oscillations during the Quaternary, such as the last glacial maximum, might have led to a decrease in the geographic distribution of Engelhardieae.

First occurrence of Cedrelospermum (Ulmaceae) in Asia and its biogeographic implications

Cedrelospermum (Ulmaceae) is an extinct genus with extensive fossil records in Europe and North America. However, no fossil of the genus has been reported from Asia. Here we describe Cedrelospermumasiaticum L.B. Jia, Y.J. Huang et Z.K. Zhou sp. nov. based on compressed fruits from the late Miocene of Yunnan, southwestern China. The fossil fruits are characterized by an ovate fruit body adjoined by double wings, with the veins on the primary wing converging toward a stigmatic area. According to the historical geographic distribution of the genus, we hypothesize that Cedrelospermum originated in North America where both single-winged and double-winged fruits were reported. The single-winged form subsequently spread into Europe via the North Atlantic land bridge and the double-winged form dispersed into Asia via the Bering land bridge. From the Eocene to Oligocene, a southward retreat of the genus distribution probably took place, which coincided with the global surface cooling initiated during the Eocene–Oligocene transition. The extinction of Cedrelospermum from Asia may be related to the intensification of the East Asian monsoon.

Late Pliocene diversity and distribution of Drynaria (Polypodiaceae) in western Yunnan explained by forest vegetation and humid climates

The palaeodiversity of flowering plants in Yunnan has been extensively interpreted from both a molecular and fossil perspective. However, for cryptogamic plants such as ferns, the palaeodiversity remains poorly known. In this study, we describe a new ferny fossil taxon, Drynaria lanpingensis sp. nov. Huang, Su et Zhou (Polypodiaceae), from the late Pliocene of northwestern Yunnan based on fragmentary frond and pinna with in situ spores. The frond is pinnatifid and the pinnae are entirely margined. The sori are arranged in one row on each side of the primary vein. The spores have a semicircular to bean-shaped equatorial view and a tuberculate surface. Taken together with previously described fossils, there are now representatives of three known fossil taxa of Drynaria from the late Pliocene of western Yunnan. These finds suggest that Drynaria diversity was considerable in the region at that time. As Drynaria is a shade-tolerant plant, growing preferably in wet conditions in the understory of forests, its extensive existence may indicate forest vegetation and humid climates in western Yunnan during the late Pliocene. This is in line with results from floristic investigations and palaeoclimatic reconstructions based on fossil floras.

New fossil seeds of Eurya (Theaceae) from East Asia and their paleobiogeographic implications

Eurya has an excellent fossil record in Europe, but it has only a few fossil occurrences in East Asia though this vast area houses the highest modern diversity of the genus. In this study, three-dimensionally preserved fossil seeds of Eurya stigmosa (Ludwig) Mai from the late Pliocene of northwestern Yunnan, southwestern China are described. The seeds are compressed and flattened, slightly campylotropous, and nearly circular to slightly angular in shape. The surface of the seeds is sculptured by a distinctive foveolate pattern, consisting of funnel-shaped and finely pitted cells. Each seed valve contains a reniform or horseshoe-shaped embryo cavity, a characteristic condyle structure and an internal raphe. These fossil seeds represent one of the few fossil records of Eurya in East Asia. This new finding therefore largely extends the distributional ranges of Eurya during Neogene. Fossil records summarized here show that Eurya persisted in Europe until the early Pleistocene, but disappeared thereafter. The genus might have first appeared in East Asia no later than the late Oligocene, and dispersed widely in regions such as Japan, Nepal, and southwestern China.

Peaches Preceded Humans: Fossil Evidence from SW China

Peach (Prunus persica, Rosaceae) is an extremely popular tree fruit worldwide, with an annual production near 20 million tons. Peach is widely thought to have origins in China, but its evolutionary history is largely unknown. The oldest evidence for the peach has been Chinese archaeological records dating to 8000–7000 BP. Here, we report eight fossil peach endocarps from late Pliocene strata of Kunming City, Yunnan, southwestern China. The fossils are identical to modern peach endocarps, including size comparable to smaller modern varieties, a single seed, a deep dorsal groove, and presence of deep pits and furrows. These fossils show that China has been a critical region for peach evolution since long before human presence, much less agriculture. Peaches evolved their modern morphology under natural selection, presumably involving large, frugivorous mammals such as primates. Much later, peach size and variety increased through domestication and breeding.