First fossil record of an East Asian endemic genus Sladenia (Sladeniaceae) from its modern range: Implications for floristic evolution and conservation biology

Fossil records of endemic plants play an important role in recognizing the floristic history of East Asia and thereby facilitate the conservation of plant diversity in the region. However, the fossil record of many extant East Asian endemic genera remains poorly documented thus far. Here, we report an infructescence fossil of an East Asian endemic genus, Sladenia (Sladeniaceae), from the early Miocene of southeastern Yunnan, China. The fossil is characterized by: (i) dichasial cymes; and (ii) flask‐shaped ovary with dense subparallel ribs on the surface extending from the base to the distal end of the united style. It represents the first fossil record of Sladenia in Asia, showing that the genus was established in the region at least by the early Miocene. Given that a much older fossil record of Sladeniaceae has been reported from Africa and the sister group of Sladenia is distributed only in Africa, Sladenia is not likely of East Asian origin. The present endemic status of Sladenia was possibly achieved by regional extirpation in Africa and taking refuge in East Asia. This case thus supports the “Museum” rather than “Cradle” hypothesis for the genesis of high plant species in the flora of East Asia. A comparison of the present fossil with extant Sladenia infructescence shows morphological stasis from the early Miocene to present. Such evolutionary tardiness might have resulted in the reduced fitness of the genus, which further caused its current endangered situation.     

The fossil record of Eucommia (Eucommiaceae) in North America

Reproductive and vegetative remains of Eucommia from 25 localities in the United States, Canada, and Mexico document the wide distribution of this genus in North America during the Cenozoic. Autofluorescent elastic latex filaments bearing capitate termini are preserved in nearly all of the remains and provide conclusive evidence of their affinity to Eucommia. Four species of Eucommia are recognized on the basis of the characteristic samaras: E. eocenica from middle Eocene strata of the Mississippi Embayment in Missouri, Tennessee and Mississippi; E. montana from early Eocene to early Oligocene localities in British Columbia, Washington, Oregon, Utah, Colorado, and Montana; E. constans from Neogene rocks in central Mexico; and E. jeffersonensis n. sp. from the latest Eocene or earliest Oligocene John Day Formation of Oregon. Atypical specimens of E. montana and E. eocenica are the first records of two-seeded fruits for the genus. Eucommia leaves from Eocene localities in British Columbia and Mississippi are the first records of Eucommia foliage in North America whose identifications are confirmed by the presence of capitate latex strands. These leaves and a specimen from Oregon are referred to E. rolandii n. sp. Fruit evolution in Eucommia may have involved increases in samara size and symmetry, and reduction in seed number from two to one, perhaps as adaptations for wind dispersal. All fossil Eucommia samaras from North America are smaller and less symmetrical than those of the living species, E. ulmoides. Preliminary flight tests of E. ulmoides samaras and of models of the fossils suggest that E. ulmoides fruits are aerodynamically better suited for wind dispersal than the fossils.     

The role of the uplift of the Qinghai‐Tibetan Plateau for the evolution of Tibetan biotas

Biodiversity is unevenly distributed on Earth and hotspots of biodiversity are often associated with areas that have undergone orogenic activity during recent geological history (i.e. tens of millions of years). Understanding the underlying processes that have driven the accumulation of species in some areas and not in others may help guide prioritization in conservation and may facilitate forecasts on ecosystem services under future climate conditions. Consequently, the study of the origin and evolution of biodiversity in mountain systems has motivated growing scientific interest. Despite an increasing number of studies, the origin and evolution of diversity hotspots associated with the Qinghai‐Tibetan Plateau (QTP) remains poorly understood. We review literature related to the diversification of organisms linked to the uplift of the QTP. To promote hypothesis‐based research, we provide a geological and palaeoclimatic scenario for the region of the QTP and argue that further studies would benefit from providing a complete set of complementary analyses (molecular dating, biogeographic, and diversification rates analyses) to test for a link between organismic diversification and past geological and climatic changes in this region. In general, we found that the contribution of biological interchange between the QTP and other hotspots of biodiversity has not been sufficiently studied to date. Finally, we suggest that the biological consequences of the uplift of the QTP would be best understood using a meta‐analysis approach, encompassing studies on a variety of organisms (plants and animals) from diverse habitats (forests, meadows, rivers), and thermal belts (montane, subalpine, alpine, nival). Since the species diversity in the QTP region is better documented for some organismic groups than for others, we suggest that baseline taxonomic work should be promoted.     

The biogeography and phylogeny of schizothoracine fishes (Schizopygopsis) in the Qinghai‐Tibetan Plateau

Freshwater fish belonging to the genus Schizopygopsis are widespread in drainages throughout the Qinghai‐Tibetan Plateau and, thus, a model group with which to investigate how paleo‐drainage changes linked to historical uplifting within the Qinghai‐Tibetan Plateau influence speciation. To date, the phylogenetic and taxonomic relationships within Schizopygopsis remain controversial. In this study, we constructed a comprehensive molecular phylogeny of Schizopygopsis based on six mitochondrial gene sequences. We compared the taxonomic relationships revealed by this phylogeny with those obtained from morphological data. We also used this phylogeny to assess the extent to which the evolution of Schizopygopsis has been driven by paleo‐drainage changes linked to uplifting of the Qinghai‐Tibetan Plateau. Results indicated that all Schizopygopsis taxa formed a monophyletic group comprising five major clades, which were inconsistent with the taxonomic relationships based on morphology for this group. Our results also strongly supported the validity of S. anteroventris and S. microcephalus as distinct species within Schizopygopsis. Molecular calibrations showed that species within the middle Yangtze species diverged earlier (~4.5 Mya) than species within the Indus River (~3.0 Mya), the Mekong River (~2.8 Mya) and the Tsangpo + Salween rivers (~2.5 Mya). The most recent evolutionary splits occurred among species from the upper and lower Yangtze River, the Yellow River and the Qiadam Basin at about 1.8 to 0.3 Mya. Our molecular evidence and use of the molecular clock calibration have allowed us to associate speciation events within the genus Schizopygopsis to the formation and separation of paleo‐drainage connections caused by tectonic events during the uplifting of the Qinghai‐Tibetan Plateau (~4.5 Mya). This work underlines the dominant role of vicariance in shaping the evolutionary history of the genus Schizopygopsis. Further research using multiple loci and more extensive sampling will reveal a more complete picture of the phylogenetic relationships and biogeography of Schizopygopsis fishes.     

Phylogeography of Tibetan snowcock (Tetraogallus tibetanus) in Qinghai–Tibetan Plateau

Data from eight microsatellite loci were used to infer the evolutionary and past demographic processes in 97 Tibetan snowcocks sampled from eight different geographical locations on the Qinghai–Tibetan Plateau. Analysis of the microsatellite DNA markers indicated that Tibetan snowcock on the plateau were geographically structured, and that phylogenetic analyses identified three phylogroups, namely those from Xunhua, the Qilian Mountains and all others. The use of Bayesian Clustering and Population Assignment analyses of the microsatellite genotypes revealed clear differentiation among the eight sampled groups of Tibetan snowcock, indicating strong isolation of these sub-populations. Therefore, we suggested that the distribution pattern of Tibetan snowcock observed today resulted from adaptation to the climatic conditions and glacial cycles on the Qinghai–Tibetan Plateau. In addition, BOTTLENECK analysis indicated that Tibetan snowcock had recently passed through evolutionary bottlenecks. These results suggested that effective conservation measures should be undertaken to protect Tibetan snowcock from an increased probability of extinction.     

New Eocene fossil fruits and leaves of Menispermaceae from the central Tibetan Plateau and their biogeographic implications

Menispermaceae are a pantropical and temperate family with an extensive fossil record during the Paleogene period, especially in North America and Europe, but with much less evidence from Asia. The latest fossil evidence indicates a succession of tropical to subtropical flora on the central Tibetan Plateau during the Paleogene. However, the biogeographic histories of these floras are still unresolved. Here, we report on endocarps and leaves of Menispermaceae from the Middle Eocene of Jianglang village, Bangor County, central Tibetan Plateau. The endocarps belong to two genera: Stephania, which is characterized by a horseshoe-shaped endocarp and with one lateral crest ornamented by spiny to rectangular ribs, and a condyle area; and Cissampelos (s.l.), which has two characteristic lateral ridges and a conspicuous external condyle. Associated leaves belong to the genus Menispermites, and are characterized by actinodromous primary venation, brochidodromous secondary veins, entire margins, and the presence of marginal secondary veins. The biogeographic history of Menispermaceae is complex, but evidence from these new fossils indicates an early diversification of the group in Asia, probably in response to the warming climate during the Eocene. The Jianglang flora appears to be part of a boreotropical flora, connecting Asia with North American and European floras during the Middle Eocene. The modern distribution of menispermaceous taxa found in Jianglang, as well as other families represented in the Jianglang flora, show that a tropical to subtropical climate occurred during the Eocene in central Tibet.     

Phylogeny and biogeography of Zelkova (Ulmaceae sensu stricto) as inferred from leaf morphology, ITS sequence data and the fossil record

To address the evolution and geographical diversification of the genus Zelkova (Ulmaceae) a phylogenetic analysis of morphological data and the sequences of the internal transcribed spacers (ITS1 and ITS2) of nuclear ribosomal DNA were used. Cladistic analyses suggested that the Chinese species Z. schneideriana and Z. sinica are basal within Zelkova. The western Asian Z. carpinifolia either appears nested between the East Asian Z. schneideriana and Z. sinica and a clade formed by the Japanese Z. serrata and two Mediterranean species, Z. abelicea and Z. sicula (ITS), or forms a clade with Z. serrata that is sister to a clade Z. abelicea plus Z. sicula (morphology). Nucleotide data suggested that gene flow occurred between Z. schneideriana and Z. serrata, and Z. carpinifolia and a lineage ancestral to Z. abelicea/sicula. Character evolution in Zelkova appears to have gone from long leaves with numerous secondary veins, coarse to shallow teeth with blunt or slightly pointed apex and small stomata, to leaves that are either long or short with numerous or few secondary veins, coarse teeth with cuspidate or obtuse apex or conspicuously shallow teeth, and dimorphic stomata displaying ‘giant stomata’ surrounded by a ring of small stomata or uniform large stomata. These results are in agreement with fossil data. Early Cainozoic fossils attributed to Zelkova from North America and Central Asia closely resemble the modern Z. schneideriana and Z. carpinifolia. The genus could have originated in the northern Pacific area and migrated to Europe after the Turgai Strait was closed during the Late Oligocene. Geographical differentiation may have started with the isolation of Chinese populations (leading to modern Z. schneideriana and Z. sinica) from high‐latitude Eurasian (North American) populations. This widespread Early Cainozoic type may have diversified into the western Asian Z. carpinifolia and the more derived Japanese and Mediterranean species during the latest Cainozoic. The modern Japanese and European/western Asian species would have differentiated relatively late, while two locally endemic Mediterranean species are the result of the cooling and development of a Mediterranean climate belt in Europe during the Pleistocene. Fossils from the Miocene and Pliocene of Europe resemble modern Z. carpinifolia and Z. serrata. Differentiation of the two Mediterranean species Z. abelicea and Z. sicula in the Late Cainozoic cannot be traced by leaf morphology. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 147 , 129–157.     

Gradual genome size evolution and polyploidy in Allium from the Qinghai–Tibetan Plateau

Background and AimsGenome size is an important plant trait, with substantial interspecies variation. The mechanisms and selective pressures underlying genome size evolution are important topics in evolutionary biology. There is considerable diversity in Allium from the Qinghai–Tibetan Plateau, where genome size variation and related evolutionary mechanisms are poorly understood.MethodsWe reconstructed the Allium phylogeny using DNA sequences from 71 species. We also estimated genome sizes of 62 species, and determined chromosome numbers in 65 species. We examined the phylogenetic signal associated with genome size variation, and tested how well the data fit different evolutionary models. Correlations between genome size variations and seed mass, altitude and 19 bioclimatic factors were determined.Key Results Allium genome sizes differed substantially between species and within diploids, triploids, tetraploids, hexaploids and octaploids. Size per monoploid genome (1Cx) tended to decrease with increasing ploidy levels. Allium polyploids tended to grow at a higher altitude than diploids. The phylogenetic tree was divided into three evolutionary branches. The genomes in Clade I were mostly close to the ancestral genome (18.781 pg) while those in Clades II and III tended to expand and contract, respectively. A weak phylogenetic signal was detected for Allium genome size. Furthermore, significant positive correlations were detected between genome size and seed mass, as well as between genome size and altitude. However, genome size was not correlated with 19 bioclimatic variables.Conclusions Allium genome size shows gradual evolution, followed by subsequent adaptive radiation. The three well-supported Allium clades are consistent with previous studies. The evolutionary patterns in different Allium clades revealed genome contraction, expansion and relative stasis. The Allium species in Clade II may follow adaptive radiation. The genome contraction in Clade III may be due to DNA loss after polyploidization. Allium genome size might be influenced by selective pressure due to the conditions on the Qinghai–Tibetan Plateau (low temperature, high UV irradiation and abundant phosphate in the soil).     

Leaf and infructescence fossils of Alnus (Betulaceae) from the late Eocene of the southeastern Qinghai–Tibetan Plateau

Plant fossils from the Qinghai–Tibetan Plateau (QTP), China are critical to understand not only the diversification history of plants there, but also the paleoenvironmental conditions. Alnus are deciduous trees, mainly distributed in temperate and subtropical regions of Eurasia and North America, and they are well known in the fossil records throughout the Cenozoic in the Northern Hemisphere. We collected numerous well‐preserved Alnus leaf and infructescence fossils from the Lawula Formation (～34.6 Ma with ⁴⁰Ar/³⁹Ar dating) at the present elevation of 3910 m a.s.l. in the southeastern QTP. Based on detailed morphological comparisons with existing and fossil species, these fossils show closest affinity to Alnus ferdinandi‐coburgii C. K. Schneid., and we refer to these fossils as A. cf. ferdinandi‐coburgii. These specimens comprise the oldest megafossil record of Alnus in the QTP, and provide solid evidence for the distribution of Alnus there as early as the late Eocene. Extant A. ferdinandi‐coburgii is distributed in areas with mean annual temperature values between 9.7 °C and 16.9 °C, and mean annual precipitation values ranging from 896.2 mm to 1161.2 mm; therefore, fossils of A. cf. ferdinandi‐coburgii suggest a much warmer and wetter climate during the late Eocene than today in the southeastern QTP. This finding is consistent with other evidence for continued uplift of the southeastern QTP after the late Eocene that might be due to the eastward extension of the QTP.     

The fossil record and biogeography of the Ciehlidae (Actinopterygii: Labroidei)

The family Ciehlidae is a large group of tropical fishes in the order Perciformes, with an estimated number of living species exceeding 1400. The modern distribution of the family Ciehlidae is predominantly in fresh waters of Central and South America, Africa, Madagascar, India and the Middle East, with fossil members known from Africa, Saudi Arabia, the Levant, Europe, South America and Haiti. Many authors have referred to the distribution as being Gondwanan and have postulated that cichlids originated over 130 million years ago, in the Early Cretaceous. However, the suggested evidence for an Early Cretaceous origin of cichlids is equally or more compatible with a much younger age of origin. Based on the biology and distribution of modern and fossil cichlids, it is more probable that they arose less than 65 million years ago, in the Early Tertiary, and crossed marine waters to attain their current distribution.     

Nutlet micro‐morphology of the genus Microula (Boraginaceae) from the Qinghai–Tibetan Plateau,and its systematic implications

The micro‐morphology of nutlets in 30 taxa (28 species and 2 varieties) of Microula Benth. (Boraginaceae) was surveyed by light and scanning electron microscopy. A principal coordinate‐based ordination analysis (PCO) (8 morphological and 14 micro‐morphological characters) was applied to explore whether taxa clustered according to the current sectional taxonomy. The nutlet epidermis showed three major features: trichomes, papillae and lamellae. Two basic trichome types could be identified: branched and unbranched. Stellate trichomes (STT) and stipitate stellate trichomes (SST) can be considered as two subtypes of branched trichomes based on shape and branching location. Three kinds of unbranched trichomes were recognized by their shapes: slender trichomes (SLT), spear‐like trichomes (SPT) and conical trichomes (CT). Four types of papillae were revealed in this genus: central apiculate papillae (CAP), debris‐covered convex papillae (DCP), rugous papillae (RP) and tabular papillae (TP). The lamellae was classified in five types: glossy lamellae (GL), thread‐like lamellae (TL), rugous lamellae (RL), interrupted lamellae (IL) and debris‐covered lamellae (DCL). The variation in the nature of surface sculpturing, nutlet shape and size, dorsal aperture shape, and attachment scar positions proved useful diagnostic characters. Taxa belonging to the same section generally clustered together in the PCO analysis. A key based on the morphology and micro‐morphology of nutlets is provided to distinguish the sections of Microula. The presence of SLT, STT, DCP and DCL is consistent with the macro‐morphological classification and provides additional evidence for the subgeneric delimitations. The SST, SLT, CAP, DCP, RP, DCL, GL and RL are useful in discriminating species in Microula. Additionally, the distribution of some trichomes, papillae and lamellae types is correlated with the altitudinal distribution of the species.     

Polyploidy and new chromosome counts in Anaphalis (Asteraceae: Gnaphalieae) from the Qinghai‐Tibet Plateau of China

Abstract Anaphalis is one of the largest genera of the Asian Gnaphalieae (Asteraceae) and is most diversified on the Qinghai‐Tibet Plateau. The chromosome numbers and karyomorphology of six species from seven populations were investigated for the first time for this region. Chromosome numbers have been newly documented for four species: Anaphalis deserti (2n = 56 = 24 median [m]+ 24 submedian [sm]+ 8 acrocentric [st]); Anaphalis plicata (2n = 56 = 26m + 30sm); Anaphalis xylorhiza (2n = 28 = 2 median point [M]+ 14m + 12sm); and Anaphalis rhododactyla (2n = 56 = 32m + 24sm). Two further counts are in agreement with the previously reported numbers, that is, Anaphalis royleana (2n = 28 = 4M + 6m + 18sm) and Anaphalis margaritacea (2n = 28 = 12m + 16sm). A new polyploid cytotype (2n = 56 = 26m + 30sm) was found in the Zougong population of A. margaritacea. Polyploidy is for the first time reported for Anaphalis, with four out of seven counts being tetraploid. Our cytological results suggest that polyploids might have played an important role in the evolution of Anaphalis on the Qinghai‐Tibet Plateau.     

Patterns of species diversity and phylogenetic structure of vascular plants on the Qinghai‐Tibetan Plateau

Large-scale patterns of species richness and the underlying mechanisms regulating these patterns have long been the central issues in biogeography and macroecology. Phylogenetic community structure is a result of combined effects of contemporary ecological interactions, environmental filtering, and evolutionary history, and it links community ecology with biogeography and trait evolution. The Qinghai-Tibetan Plateau provides a good opportunity to test the influence of contemporary climate on shaping species richness because of its unique geological history, cold climate, and high biodiversity. In this study, based on high-resolution distributions of ˜9000 vascular plant species, we explored how species richness and phylogenetic structure of vascular plants correlate with climates on the highest (and species rich) plateau on the Earth. The results showed that most of the vascular plants were distributed on the eastern part of the plateau; there was a strong association between species richness and climate, even after the effects of habitat heterogeneity were controlled. However, the responses of richness to climate remarkably depended on life-forms. Richness of woody plants showed stronger climatic associations than that of herbaceous plants; energy and water availability together regulated richness pattern of woody plants; whereas water availability predominantly regulated richness pattern of herbaceous plants. The phylogenetic structure of vascular species clustered in most areas of the plateau, suggesting that rapid speciation and environment filtering dominated the assembly of communities on the plateau. We further propose that biodiversity conservation in this area should better take into account ecological features for different life-forms and phylogenetic lineages.     

The uplift of the Qinghai–Tibet Plateau and glacial oscillations triggered the diversification of Tetraogallus (Galliformes,Phasianidae)

The Qinghai–Tibet Plateau (QTP) plays an important role in avian diversification. To reveal the relationship between the QTP uplift and avian diversification since the Late Cenozoic, here, we analyzed the phylogenetic relationship and biogeographical pattern of the genus Tetraogallus (Galliformes, Phasianidae) and the probable factors of speciation in the period of the QTP uplift inferred from concatenated data of four nuclear and five mitochondrial genes using the method of the Bayesian inference. Phylogenetic analysis indicated that T. himalayensis had a close relationship with T. altaicus and conflicted with the previous taxonomy of dark‐bellied and white‐bellied groups. The molecular clock showed that the speciation of Tetraogallus was profoundly affected by the uplift of the QTP and glacial oscillations. Biogeographic analysis suggested that the extant snowcocks originated from the QTP, and the QTP uplift and glacial oscillations triggered the diversification of Tetraogallus ancestor. Specifically, the uplift of the mountain provided a prerequisite for the colonization of snowcocks Tetraogallus as a result of the collision between the Indian and the Arab plates and the Eurasian plate, in which ecological isolation (the glacial and interglacial periods alternate) and geographical barrier had accelerated the Tetraogallus diversification process. Interestingly, we discovered hybrids between T. tibetanus and T. himalayensis for the first time and suggested that T. tibetanus and T. himalayensis hybridized after a second contact during the glacial period. Here, we proposed that the hybrid offspring was the ancestor of the T. altaicus. In conclusion, the uplift of QTP and glacial oscillations triggered the snowcocks colonization, and then, isolation and introgression hybridization promoted diversification.     

Deyeuxia sorengii sp. nov. (Poaceae,Agrostidinae) from the Qinghai‐Tibetan Plateau

Deyeuxia sorengii B. Paszko & W. L. Chen, a new species of Deyeuxia from the Qinghai‐Tibetan Plateau, China, is described and illustrated. The new species is morphologically similar to D. zangxiensis P. C. Kuo & S. L. Lu, but differs from it by having a well‐developed and conspicuously hairy rachilla prolongation and a straight awn. Deyeuxia sorengii occurs in alpine meadows of central Xizang and central Qinghai. The first records of Deyeuxia zangxiensis are reported from western and eastern Xinjiang and northeastern Qinghai (China).