Interannual fluctuations in floral longevity, pollinator visitation and pollination limitation of an alpine plant (Gentiana straminea Maxim., Gentianaceae) at two altitudes in the Qinghai-Tibetan Plateau

Prolonged floral longevity and bumblebees as dominate pollinators in alpine ecosystem have been suggested to overcome pollination limitation of alpine plants arising from the decrease of pollinator activity with increasing altitude. However, this conclusion has never been examined in the Qinghai-Tibetan Plateau (QTP), the highest and largest plateau in the world. In this study, we intended to test year-to-year correlations between floral longevity, visiting frequency and pollen limitation of this species between two populations (at 3200 m and 4000 m) of Gentiana straminea in this plateau. Pollinator exclusion elongated both male and female phases greatly at both sites, and durations of both male and female phases in natural condition varied greatly over three years. The visiting frequency of bumblebees varied greatly at the higher altitude, but seemed to be stable at the lower altitude. Seed production was pollination limited in both populations in most studied years. The floral durations, pollinator frequency and pollination limitation showed no significant and consistent variations with the increase of altitude. The previous hypothesis that the prolonged floral longevity of alpine plants can compensate for low levels of pollinator visitation therefore could not be confirmed, and our results further suggested that in the QTP platform, the altitude shows no consistent effect on the reproductive performance of this species, despite that the fluctuation of visit frequency intensified at the higher altitude.

ISSR analysis of genetic diversity of the Qinghai-Tibet Plateau endemic, Rhodiola chrysanthemifolia (Crassulaceae)

Inter-simple sequence repeat markers (ISSR) were used to estimate genetic diversity within and among 10 populations of Rhodiola chrysanthemifolia along Nianqingtangula Mountains and Brahmaputra, a species endemic to the Qinghai-Tibet Plateau and an endangered medicinal plant. Of the 100 primers screened, 13 produced highly polymorphic DNA fragments. Using these primers, 116 discernible DNA fragments were generated of which 104 (89.7%) were polymorphic, indicating substantial genetic diversity at the species level. Genetic diversity measured by the percentage of polymorphic bands (PPB) at the population level ranged from 21.97% to 48.8%. Analysis of molecular variance (AMOVA) showed that the genetic variation was found mainly among populations (77.3%), but no regional differentiation was discernible. Variance within populations was only 22.7%. The main factor responsible for this high level of differentiation among populations is probably the historical geographical and genetic isolation of populations in a harsh mountainous environment. Concerning the management of R. chrysanthemifolia, the high genetic differentiation of populations indicates the necessity of conserving the maximum possible number of populations. (c) 2006 Elsevier Ltd. All rights reserved.

Mitochondrial DNA phylogeny of two morphologically enigmatic fishes in the subfamily Schizothoracinae (Teleostei : Cyprinidae) in the Qinghai-Tibetan Plateau

The complete mitochondrial DNA (mtDNA) cytochrome b gene (1140 bp) was sequenced in Herzenstein macrocephalus and Gymnocypris namensis and in 13 other species and sub-species (n = 22), representing four closely related genera in the subfamily Schizothoracinae. Conflicting taxonomies of H. macrocephalus and G. namensis have been proposed because of the character instability among individuals. Parsimony, maximum likelihood and Bayesian methods produced phylogenetic trees with the same topology and resolved several distinctive clades. Previous taxonomic treatments, which variously placed these two species of separate genera or as sub-species, are inconsistent with the mtDNA phylogeny. Both H. macrocephalus and G. namensis appear in a well-supported clade, which also includes nine species of Schizopygopsis, and hence should be transferred to the genus Schizopygopsis. Morphological changes are further illustrated, and their adaptive evolution in response to the local habitat shifts during the speciation process appears to be responsible for conflicting views on the systematics of these two species and hence the contrasting taxonomic treatments. These species are endemic to the Qinghai-Tibetan Plateau, a region with a history of geological activity and a rich diversity of habitats that may have result in the parallel and reversal evolution of some morphological characters used in their taxonomies. Our results further suggest that speciation and morphological evolution of fishes in this region may be more complex than those previously expected. (c) 2007 The Authors Journal compilation (c) 2007 The Fisheries Society of the British Isles.

Geographic variation in the leaf essential oils of Juniperus sabina L. and J-sablina var. arenaria (E.H. Wilson) Farjon

The composition of the leaf oils from seven populations of J. sabina L., one population of Juniperus sabina var. arenaria (E. H. Wilson) Farjon were examined for their geographic variation. In addition, the leaf oils of J. chinensis L. and J. davurica Pall. were compared to J. sabina. Juniperus sabina var. arenarla, the sand loving juniper, oil was found to be very similar to that of J. davurica, Mongolia, and J. sabina, on sand dunes in Mongolia. This suggests that J. sabina var. arenaria might be conspecific with J. davurica. Farjon's move (2001) of J. sabina var. arenaria out of J. chinensis is supported. Considerable differentiation was found in populations of J. sabina from the Iberian peninsula. Cedrol, citronellol, safrole, trans-sabinyl acetate, terpinen-4-ol and beta-thujone were found to be polymorphic in several populations.

Floral closure in response to temperature and pollination in Gentiana straminea Maxim. (Gentianaceae), an alpine perennial in the Qinghai-Tibetan Plateau

Floral closure may be induced by pollination and various other factors, but is rarely studied comprehensively. Different kinds of floral closure should have various effects on reproductive fitness of plants. Two contrasting types of floral closure were observed in the flowers of Gentiana straminea Maxim. in the eastern Qinghai-Tibetan Plateau. The first type occurred prior to pollination during both gender phases, in response mainly to decreasing air temperatures. Flowers closed when decreasing temperatures approached 20 degrees C and subsequently began to reopen the following day during mid-morning when air temperatures warmed to approximately 13-15 degrees C. This kind of floral closure can protect pollen grains on either stamens or stigmas, increasing fitness of both male and female. Following pollination, permanent floral closure occurred, although there was a delay between the dates of pollination and permanent closure, during which flowers continued to show temporary closure in response to low temperature episodes. The time required for permanent, pollination-induced closure varied according to the age of the gender phase, including a prolonged time before closure if pollination occurred early in the female phase. The retaining of permanent closed flowers increased both approaching (to inflorescences) and visiting (to unpollinated flowers) frequencies of individual plants when with fewer open flowers and the persisting corolla is further beneficial for seed sets of these pollinated flowers. Thus, two separate types of floral closure, one in response to environmental cues and the other in response to the age of each gender stage, appeared to have a strong influence on reproductive fitness in this species. These results revealed a different adaptive strategy of alpine plants in the sexual reproduction assurance in addition to the well-known elevated floral longevity, dominant role of more effective pollinators and increased reproduction allocation in the arid habitats.

Radiation and diversification within the Ligularia-Cremanthodium-Parasenecio complex (Asteraceae) triggered by uplift of the Qinghai-Tibetan Plateau

The Ligularia-Cremanthodium-Parasenecio (L-C-P) complex of the Tussilagininae (Asteraceae: Senecioneae) contains more than 200 species that are endemic to the Qinghai-Tibetan Plateau in eastern Asia. These species are morphologically distinct; however, their relationships appear complex. A phylogenetic analysis of members of the complex and selected taxa, of the tribe Senecioneae was conducted using chloroplast (ndhF and trnL-F) and nuclear (ITS) sequences. Phylogenetic trees were constructed from individual and combined datasets of the three different sequences. All analyses suggested that Doronicum, a genus that has been included in the Tussilagininae, should be excluded from this subtribe and placed at the base of the tribe Senecioneae. In addition, the Tussilagininae should be broadly circumscribed to include the Tephroseridinae. Within the expanded Tussilagininae containing all 13 genera occurring in eastern Asia, Tussilago and NSPetasites diverged early as a separate lineage, while the remaining I I genera comprise an expanded L-C-P complex clade. We suggest that the L-C-P clade, which is largely unresolved, most likely originated as a consequence of an explosive radiation. The few monophyletic subclades identified in the L-C-P clade with robust support further suggest that some genera of Tussilagininae from eastern Asia require generic re-circumscriptions given the occurrence of subclades containing species of the same genus in different parts of the phylogentic tree due to homoplasy of important morphological characters used to delimit them. Molecular-clock analyses suggest that the explosive radiation of the L-C-P complex occurred mostly within the last 20 million years, which falls well within the period of recent major uplifts of the Qinghai-Tibetan Plateau between the early Miocene to the Pleistocene. It is proposed that significant increases in geological and ecological diversity that accompanied such uplifting, most likely promoted rapid and continuous allopatric speciation in small and isolated populations, and allowed fixation or acquisition of similar morphological characters within unrelated lineages. This phenomenon, possibly combined with interspecific diploid hybridization because of secondary sympatry during relatively stable stages between different uplifts, could be a major cause of high species diversity in the Qinghai-Tibetan Plateau and adjacent areas of eastern Asia. (c) 2005 Elsevier Inc. All rights reserved.

Phylogenetic analyses of Saussurea sect. Pseudoeriocoryne (Asteraceae : Cardueae) based on chloroplast DNA trnL-F sequences

The genus Saussurea is distributed mainly in the temperate and subarctic regions of Eurasia and consists of about 300 species classified into six subgenera and 20 sections. Sect. Pseudoeriocoryne in the subgenus Eriocoryne comprises four species, and is delimited mainly by acaulescence and an inflorescence with congested capitula surrounded by a rosette of leaves. All of these species are endemic to the and Qinghai-Tibet Plateau. Sequences from the chloroplast DNA trnL-F region were obtained for the four species in this section and 26 other species from four subgenera of Saussurea to resolve phylogenetic relationships among these species and to determine whether the shared characters that define sect. Pseudoeriocoryne are synapomorphic or were acquired by convergent evolution. The resulting phylogenies indicated that Saussurea sect. Pseudoeriocoryne as traditionally defined does not constitute a monophyletic group and that each of its species belongs to separate clades. Furthermore, none of these species showed a close relationship with the other species of subgenus Eriocoryne. Our results further indicated that none of the investigated subgenera are monophyletic, and that species from different subgenera clustered together. All these conclusions are provisional and their confirmation would require stronger phylogenetic support. Two possible explanations are suggested for low sequence divergence, poor resolution of internal clades and clustering of species with the rather distinct morphology of Saussurea detected in the present study. The first is rapid radiation and diversification triggered by fast habitat fragmentation due to the recent lifting of the Qinghai-Tibet Plateau and the Quaternary climate oscillations. This could have led to rapid morphological divergence while sequences diverged very little, and also caused the convergent acquisition of similar characteristics in unrelated lineages due to similar selection pressures. The second possible explanation is that both introgressive hybridization and reticulate evolution might have caused the transferring of cpDNA sequences between morphologically dissimilar species, thus leading to homogenization of sequences between lineages. (C) 2004 Elsevier Ltd. All rights reserved.

Lack of altitudinal trends in phytochemical constituents of Swertia franchetiana (Gentianaceae)

Diverse biological characters commonly vary with altitude in species that have a wide altitudinal distribution, partly at least as a result of adaptation to differences in aridity, but whether such variation exists for phytochemical constituents remains unknown. Therefore, levels of seven important phytochemical constituents of Swertia franchetiana (swertiamarin, oleanolic acid, swertisin, mangiferin, 1,5,8-trihydroxy-3-methoxyxanthone, 1,8-dihydroxy-3,7-dimethoxyxanthone and 1,8-dihydroxy-3,5-dimethoxyxanthone) were studied and statistically compared, using materials collected from sites ranging from 2200 to 3960 m in altitude. Swertiamarin was the most abundant in all samples, then mangiferin, oleanolic acid and the other three xanthones. Throughout the distributional range of this species, no altitudinal trend was detected for other constituents except 1,8-dihydroxy-3,7-dimethoxyxanthone, which showed a negative correlation with altitude. However, the concentration of 1, 8-dihydroxy-3,7-dimethoxyxanthone and mangiferin showed a significantly latitudinal and longitudinal correlation. (C) 2004 Elsevier Ltd. All rights reserved.

Floral organogenesis and development of two taxa of the Solanaceae - Anisodus tanguticus and Atropa belladonna

Floral organogenesis and development of two Solanaceae species, Anisodus tanguticus and Atropa belladonna, were studied by using scanning electron microscopy (SEM) as part of a project on systematics and evolution in the tribe Hyoscyameae. These two species share the following common characters of floral organ initiation and development: (1) initiation of the floral organs in the two species follows Hofmeister's rule; (2) the mode of corolla tube development belongs to the "late sympetaly" type, namely, petals are initiated separately and later become joined by fusion of their basal meristem, then rise together and form a corolla tube; (3) primordia of the floral appendages are initiated in a pentamerous pattern and acropetal order: sepals are initiated first, followed by the petals and stamens, and finally the carpels. The whorl of five stamen primordia forms almost simultaneously and originates opposite the sepal primordia, but initiation of the sepal primordia shows different modes in the two species. The sepal primordia of Anisodus tanguticus have simultaneously whorled initiation, while those of Atropa belladonna have helical initiation. The systematic significance of the present results in the genera Anisodus and Atropa is discussed in this paper.

Karyomorphology of Biebersteinia Stephan (Geraniaceae) and its systematic and taxonomic significance

The systematic and taxonomic position of Biebersteinia Stephan has long been in dispute. The present paper describes for the first time the karyomorphology of two species in Biebersteinia Stephan. Both species commonly showed the interphase nuclei of the simple chromocenter type and the mitotic prophase chromosomes of the interstitial type. The karyotype formulae of both B. heterostemon and B. odora were 2n=10=2m(2sec)+8sm(2sec), belonging to the 3A type of Stebbins' classification. The karyotype of this genus is recorded for the first time. The basic chromosome numbers of four of the five known species of Biebersteinia have been recorded as x=5. The combination of resting nuclei of the simple chromocenter type, mitotic prophase chromosomes of the interstitial type, two pairs of chromosomes with four obvious secondary constrictions at the mitotic prophase and metaphase stages, and the peculiar 3A karyotype in Biebersteinia can be regarded as the karymorphological marker of this genus. The karyomorphological data presented here do not support the traditional grouping of this genus in Geraniaceae. The unique karyomorphology of Biebersteinia justifies its familiar or ordinal status, which is congruent with embryological, anatomical, chemical and molecular data. The systematic position of Biebersteinia needs further study.

湖南、广西泥盆纪硅岩地球化学与沉积构造背景

During the Devonian, a complicated carbonate platform-basin configuration was created through transtensional rifting in the context of opening of Devonian South China Sea; extensive bedded chert, commonly interbedded with tuffaceous beds, occurred in the narrow, elongate interplatform basins (or troughs) in South China, where they occurred earlier (Early Devonian) in southern Guangxi and later (early Late Devonian) in northern Guangxi-south central Hunan. In order to unravel the origin and distribution of the bedded chert successions, and their relationships to basement faulting activities during the opening of the Devonian South China Sea, studies of element (major, minor and REE) geochemistry and Rb-Sr, Sm-Nd isotopic systematics are carried out upon the chert deposits. These chert deposits commonly have high SiO2 contents and (average 94.01%) and low TFe2O3 (average 0.55%), together with other geochemical parameters, suggestive of both biogenic and hydrothermal origins. However, Fe/Ti ratio are high along the elongate interplatform basins(troughs) to the northwest along Wuxiangling-Zhaisha-Chengbu, and to the southeast along Xiaodong-Mugui-Xinpu, suggesting relatively intense hydrothermal activities there. They generally contain very low total REE contents (∑REE average 31.21ug/g) with mediate negative Ce anomalies (mean Ce/Ce*=0.83) and low Lan/Cen values (average 1.64), indicating an overall continental margin basin where they precipitated. The northward increases in Ce/Ce* values, particularly along the elongate troughs bounded both to the east and west of the Guangxi-Huanan rift basin, suggest a northward enhancement of terrigenous influences, thereby reflecting a gradual northward propagation of open marine setting. Generally low positive Eu anomalies in the chert, except for the apparently high Eu anomalies in the chert from Chengbu (Eu/Eu* up to 4.6), suggest mild hydrothermal venting activities in general, except for those at Chengbu. The initial 87Sr/86Sr (0) ratios of chert generally vary from 0.712000 to 0.73000 , suggesting influences both from terrigenous influx and seawater. The Nd isotopic model ages (tDM or t2DM) and initial εNd (0) values of chert vary mostly from 1.5 to 2.1 Ga, and from –16 to –21, respectively, implying that the silica sources were derived from the provenances of the Palaeoproterozoic crust relics at depth. The high εNd (0) values of chert (-0.22 to 14.7) in some localities, mostly along the elongate troughs, suggest that silica sources may have been derived from deeper-seated mantle, being channeled through the interplate boundary fault zones extending downwards to the mantle. At Wuxiangling, Nanning, chert occurs extensively from the Emsian through the Frasnian strata, both U/Th ratios and tDM ages of chert reached up to a maximum in the early Frasnian corresponding to the extensive development of chert in South China, pointing to a maximum extensional stage of Devonian South China basin, which is supported by the Ce/Ce* values as is opposed to the previous datasets as the coeval minimum values.

胶东地区金矿床成矿时代及其成矿地球动力学背景-兼论壳幔相互作用与成岩成矿

Jiaodong Peninsula is the largest repository of gold in China. Varieties of studies have been involved in the mechanism of metallogenesis. This thesis is a part of the project "Study of basic geology related to the prespecting of the supra-large deposits" which supported by National Climbing Program of China to Prof. Zhou. One of the key scientific problems is to study the age and metallogenic dynamics of ore deposit and to understand how interaction between mantle and crust constrains on metallogenesis and lithogenesis. As Jiaodong Peninsula to be study area, the Rb-Sr, Sm-Nd and Pb isotopic systematics of pyrite and altered rocks are measured to define the age and origin of gold. The elemental and Sr-Nd-Pb isotopic compositions of dikes and granites was studied to implicate the source and lithogenesis of the dike and granite and removal of lithosphere and the interaction between mantle and crust in the Jiaodong Peninsula. Considering the tectonic of Jiaodong Peninsula, basic on the time and space, this thesis gives a metallogenic dynamics of gold mineralization and discusses the constraints of the interaction between mantle and crust on the metallogenesis and lithogenesis. This thesis reports the first direct Rb-Sr dating of pyrites and ores using sub-sampling from lode gold deposit in Linglong, Jiaodong Peninsula and the results demonstrate this as a useful geochronological technique for gold mineralization with poor age constraint. The Rb-Sr data of pyrites yields an isochron age of (121.6-122.7) Ma, whereas, those of ore and ore-pyrite spread in two ranges from 120.0 to 121.8 Ma and 110.0-111.7 Ma. Studies of characteristic of gold deposit, microscopy of pyrite and quartz indicate that the apparent ages of ore and ore-pyrite are not isochron ages, it was only mixed by two end members, i.e., the primitive hydrothermal fluids and wall rocks. However, the isochron age of pyrite samples constrains the age of gold mineralization, i.e., early Cretaceous, which is in good consistence with the published U-Pb ages of zircon by using the SHRIMP technique. The whole rock Rb-Sr isochron age of altered rocks indicates that the age of gold mineralizing in the Xincheng gold deposit is 116.6 ± 5.3 Ma. The Sr, Nd and Pb isotopic compositions of pyrite and altered rocks indicate that the gold and relevant elements were derived from multi-sources, i.e. dikes derived from enriched lithospheric mantle and granites, granodiorites and metamorphic rocks outcropped on the crust. It also shows that the hydrothermal fluids derived from mantle magma degassing had play an important role in the gold mineralizing. The major and trace elements, Sr-Nd-Pb isotopic data of granites and granodiorites suggest that the Linglong Granite and Kunyushan Granite were derived from partial melting of basement rocks in the Jiaodong Peninsula at post-collision of North China Craton with South China Craton. Guojialing Granodiorite was considered to be derived from a mixture source, that is, mixed by magmas derived from an enriched lithospheric mantle and crust during the delamination of lithosphere induced by the subduction of Izanagi Plate and the movement of Tancheng-Lujiang Fault. There are kinds of dikes occurred in the Jiaodong Peninsula, which are accompanying with gold mineralization in time and space. The dikes include gabrro, diabase, pyroxene diorite, gabrrophyre, granite-porphyry, and aplite. The whole rock K-Ar ages give two age intervals: 120-124 Ma for the dikes that erupted at the gold mineralizing stage, and <120 Ma of the dikes that intruded after gold mineralizing. According to the age and the relationship between the dikes and gold mineralizing, the dikes could be divided into two groups: Group I (t = 120-124 Ma) and Group II (t < 120Ma). Group I dikes show the high Mg and K, low Ti contents, negative Nb anomalies and positive Eu anomalies, high ~(87)Sr/~(86)Sr and negative εNd(t) values and an enrichment in light rare earth elements, large ion lithosphile elements and a depletion in high field strength elements. Thus the elemental and isotopic characteristics of the Group I dikes indicate that they were derived from an enriched lithospheric mantle perhaps formed by metasomatism of the melt derived from the recycled crustal materials during the deep subduction of continent. In contrast, the Group II dikes have high Ti, Mg and K contents, no negative Nb anomalies, high ~(87)Sr/~(86)Sr and positive or little negative εNd(t) values, which indicate the derivation from a source like OIB-source. The geochemical features also give the tectonic constraints of dikes, which show that Group I dikes were formed at continental arc setting, whereas Group II dikes were formed within plate background. Considering the tectonic setting of Jiaodong Peninsula during the period of gold mineralizing, the metallogenic dynamics was related to the subduction of Izanagi Plate, movement of Tancheng-Lujiang Fault and removal of lithopheric mantle during Late Mesozoic Era.

山东西部中生代碳酸岩和火山岩的地球化学特征及其成因研究

On account of some very peculiar features, such as extremely high Sr and Nd contents which can buffer their primary isotopic signatures against crustal contamination, deep-seated origin within mantle, and quick ascent in lithosphere, carbonatites are very suitable for deciphering the nature of sub-continental lithospheric mantle(SCLM) and receiving widespread attentions all around the world. The Mesozoic carbonatites located in western Shandong was comprehensively investigated in this dissertation. The extremely high REE concentrations, similar spider diagrams to most other carbonatites around the world and high Sr. low Mn contents of apatite from carbonatites confirm their igneous origin. The K depletion of carbonatites from this studies reflect the co-existing of carbonatite melts with pargasite+phlogopite lherzolite rather than phlogopite lherzolite. Geological characteristics and their occumng without associated silicate rocks argue against their origin of fractionation of or liquid immisibility with carbonated silicate melts. In contrast to the low S7Sr/86Sr and high l43Nd/l44Nd of other carbonatites in the world, carbonatites of this studies show EMU features with high S7Sr/86Sr and low l4jNd/144Nd ratios, which imply that this enriched nature was formed through metasomatism of enriched mantle preexisted beneath the Sino-korean craton by partial melts of subducted middle-lower crust of Yangtze craton. In addition to carbonatites, the coeval Mesozoic volcanic rocks from western Shandong were also studied in this dissertation. Mengyin and Pingyi volcanic rocks, which located in the south parts of western Shandong are shoshonite geochemically. while volcanic rocks cropped out in other places are high-K calc-alkaline series. All these volcanic rocks enriched in LREE and LILE. depleted in HFSE, and show TNT(strong negative anomalies in Ta, Nb. Ti) patterns in spider diagrams which are common phenomena in arc-related volcanic rocks. The Sr-Nd-Pb isotopic systematics reveal that the volcanic rocks decrease gradually in 87Sr/86Sr, 206Pb/204Pb, 20SPb/204Pb and increase in TDM from south to north, suggesting the distinction of SCLM beneath Shandong in Mesozoic is more explicit in south-north trending than in east-west trending. The variable features of SCLM can be attributed to the subduction of Yangtze craton beneath Sino-Korean craton, and subsequent metasomatism of SCLM by partial melts of Yangtze lower crust in different extent.