横断山特有植物区系的物种多样性与生物地理学研究

前人对横断山区特有植物的宏生态学研究已有一定的资料积累，对横断山区在全球生物多样性保护中的关键地位和特有现象已有基本共识：（1）横断山区是全球生物多样性关键地区和生物多样性优先重点保护的热点地区之一；（2）横断山区不仅是中国的三大特有植物分布中心之一，而且是新特有中心和物种分化中心，特有现象的生态成因大于历史成因。横断山区种子植物种类丰富，且具有较高的特有现象，其中既有古特有成分也有新特有成分但更多的是新特有成分。但目前，关于横断山区特有现象与生态特征、特有种分布与物种丰富度、特有种丰富度与海拔梯度以及植被的关系等仍缺乏深入的研究。针对上述问题，本论文初步研究了横断山区（1）特有种的生活型、种子散布方式、传粉方式和繁育系统等生态特征，各生态特征之间的相关关系，以及它们的进化地位；（2）特有种的水平分布格局及环境成因；和（3）特有种和生态特征沿海拔梯度的变化规律。主要结论如下： 1．横断山区含特有植物2396个分类群（包括种、亚种和变种），隶属于372属90科。特有类群在372个属中的分布非常不均匀，约一半的特有类群（50.2%）集中分布在较少的属中（占总属数的6%），含特有类群的属大小差异很大，重要的大属是：马先蒿属、乌头属、杜鹃花属、翠雀属、紫堇属、虎耳草属、报春花属、小檗属、黄芪属、凤毛菊属、蚤缀属、橐吾属、龙胆属、景天属、柳属、苔草属、香茶菜属、凤仙花属、紫菀属、鼠尾草属和蝇子草属等，这些属多以横断山为分布和分化中心，为新特有成分。另外，地面芽植物和隐芽植物等占有绝对优势，而矮高位芽植物、高位芽植物、地面芽植物等占有较低的比例，反映了该特有区系的高山、亚高山植物区系特点，但又不乏与热带植物区系的渊源。与世界其它高山地区相比，该特有区系中的地面芽植物、隐芽植物和一年生植物有较高的比例。 2．草本、风力散布种子、昆虫传粉和两性花等为该特有植物区系的优势生态特征。除传粉方式和生活型间无明显的相互作用外，其它生态特征间都存在明显的正相关关系。定量化研究结果表明该特有区系特有种的传粉方式进化可塑性很小，处于保守（或原始）的进化地位。相反，种子散布方式和繁育系统的进化可塑性较大，处于衍生（或进化）的进化地位，是不同种系在历史发育过程中不同阶段的产物。生活型的进化地位在种间有较大的分异，具较小或较大可塑性的各占有一定的比例，因此生活型中既保留处于保守（或原始）的进化地位也有处于衍生（或进化）的进化地位，两种进化状态的生活型在该特有区系中并存，该特征不但是种系在进化早期决定的，同时也是不同种系在进化过程中获得的。 3． 特有种的分布不均匀，较高比例的特有种局限分布在较少的几个地区，北纬28-29°线是特有种丰富度重要的南北分界线。聚类分析结果表明其分布可以划分为3个物种聚集群：（1）北纬28-29°以北、北纬34°以南的藏东川西北物种聚集群；（2）北纬28-29°以南、北纬26°以北的藏东南—滇西北—川西南物种聚集群；和（3）北纬26°以南、北纬25°以北的滇西北物种聚集群。其中，藏东南—滇西北—川西南物种聚集群特有种丰富度最高，是横断山区特有种分布的核心区；滇西北物种聚集群特有种丰富度最低，藏东川西北物种聚集群特有种丰富度居中。藏东南—滇西北—川西南物种聚集群之所以特有种类丰富很有可能与该小区具有高的物种丰富度和落叶阔叶林、常绿/落叶针叶林的分布有密切的联系。在区域尺度上，横断山区总物种的分布中心与特有中心存在分异。尽管两者都处于北纬28-29°以南、北纬26°以北的藏东南—滇西北—川西南小区，但特有中心具有较小的分布范围，而总物种的丰富度中心分布范围较大。因此，我们推测特有分布中心不但与落叶阔叶林和常绿/落叶针叶林有密切的联系，而且也与物种丰富度有关。特有种与植被类型尤其是与常绿阔叶林和落叶阔叶林的关系研究对揭示特有现象的发生、性质和特点有一定的意义，有关研究尚需进一步开展。此外，海拔高差、区系物种丰富度、地理位置和单元面积为特有种丰富度地理分布的总体变异提供了74.7%的解释。 4．研究地区的南缘和东南缘有高比例的木本特有植物，而草本特有植物的比例相对最低，反映出该横断山区南缘的特有植物与热带植物区系的渊源关系，其成分多为一些古老的特有成分；而高比例和较高比例的草本特有植物在研究地区普遍存在，包括研究区域南、北的大部分地区，显然草本特有植物是横断山的优势特有成分。这种以草本特有植物为特征的高山、亚高山特有植物区系有别于我国其它两大特有植物分布中心。该特有中心的成因的最终解释还需要从物种形成（speciation）、灭绝（extinction）和扩散（biogeographic dispersal）——这三个直接影响一个地区物种数量变化的过程来考虑。 5．特有种丰富度沿海拔梯度呈“单峰” 变化曲线，在中海拔段达到峰值。与中国特有种、世界广布种相比，特有种的峰值最高，最适海拔分布范围最窄。特有率随海拔上升而逐渐增高，但在海拔5000-5500 m左右突然上升，在海拔6000 m左右达到最高值。说明（1）物种分布区和丰富度存在着正相关，即特有种具有小的分布区，而广布种的分布区较大；（2）特有率的线性变化规律反映了物种丰富度和特有现象之间的关系，即物种丰富度最大峰值以上海拔段物种数量减少，但特有率增加，说明了该特有区系的高海拔性质。 6．特有种沿海拔梯度表现出大小明显不同的物种聚集群，可能与横断山区的山地垂直气候及相应的植被类型有关。在2600-4600 m海拔段特有种数量最多，这可能与该海拔段分布着落叶阔叶林和常绿/落叶针叶林有密切的联系。特有植物沿海拔梯度可分为四个不同海拔段的物种聚集群：（1）200-1000 m；（2）1000-2600 m； （3）2600-4600 m；和（4）4600-6400 m。在对应海拔段依次分布着干热河谷稀树灌草丛、常绿阔叶林、落叶阔叶林和常绿/落叶针叶林、高山流石滩稀疏植丛。其中，分布在2600-4600 m海拔段的物种丰富度最高，1000-2600 m海拔段的物种丰富度次之，200-1000 m、4600-6400 m两个海拔段物种丰富度较低。 7．特有区系中占优势的生态特征包括草本、风力散布种子、虫媒传粉、两性花和克隆性等对海拔梯度的适应也呈“单峰” 变化曲线，但该主要特征与次要的生态特征（木本、动物散布种子、风媒传粉、单性花和非克隆性）相比，均具有较高的海拔峰值和较窄的最适分布范围，反映了主要生态特征对中、高海拔的适应特点。主要的生态特征与次要的生态特征并存构成了横断山区特有区系的基本生态特征。

中国蕨类植物保护生物地理学研究

保护生物地理学（Conservation biogeography）就是运用生物地理学的原理、理论和分析方法，特别是那些有关物种分布格局的信息，去解决生物多样性保护的相关问题，是当前生物多样性及其保护生物学研究领域的热点。通过构建中国蕨类植物物种及分布数据库得知，中国现有蕨类植物63科，221属，2456种，其中有1218种属于中国特有（中国蕨类总数约50%），3种蕨类植物为外来种。科的统计表明：最大的科为鳞毛蕨科、蹄盖蕨科、金星蕨科和水龙骨科4个科所包含的种类占全国总种数的51.5%。鳞毛蕨科、蹄盖蕨科、金星蕨科、水龙骨科、铁角蕨科、凤尾蕨科、碗蕨科、中国蕨科、叉蕨科、观音座莲科等十个大科所含种数占了中国特有种总数的80%。云南、四川、贵州、广西、台湾、湖南、西藏等省区是中国蕨类植物种类数量及中国特有种数量较多的地区；全国有22个省区拥有地方特有种，地方特有蕨类植物数量由西向东、由南向北逐渐递减，云南、台湾、西藏、海南等四省区的地方特有蕨类植物种数占各省区蕨类植物总数10%以上，反映了青藏高原隆起及岛屿现象对中国特有蕨类植物分化的影响。 中国蕨类植物与各地理、环境及气候因子的相关性分析结果表明：（1）随着经度、纬度的增加， 蕨类植物物种总数及中国特有蕨类植物总数具有减少的趋势；（2）蕨类植物在海拔1000-1600米之间、特有蕨类植物在海拔700-2100米之间具有最大的多样性，此区间外随着海拔的降低及升高，物种丰富度依次降低；（3）气候因子特别是一月均温、年均温、相对湿度、年降雨量等因素对蕨类植物及中国特有蕨类植物均具有较大的影响，与最热月均温成低度线性相关或相关不显著；（4）地形因子是影响蕨类植物多样性的一个独立因素，地形条件越复杂，蕨类植物种类及中国特有蕨类植物种类越丰富。 运用SPSS统计分析软件分析了中国蕨类植物在各省之间种的相似性，以省区为单元对中国各省区蕨类植物物种组成进行了聚类分析，最后结果划分成：①南方蕨类植物区（包括横断山—青藏高原亚区、华中亚区、华东亚区、华南亚区）和北方蕨类植物区（包括东北—华北—西北亚区、秦岭亚区、江淮亚区）。 中国蕨类植物区系地理成分分析结果表明：（1）中国—喜马拉雅分布（40.86%）、热带亚洲分布（30.95%）、东亚分布（12.77%）和中国—日本分布（8.24%）构成中国蕨类植物区系成分的主体；在洲际关系方面，中国蕨类植物与大洋洲的蕨类植物关系最为紧密，共有种最多，达94种；与南美洲共有的蕨类植物种类最少，仅32种。（2）基于地理成分的聚类分析结果表明，可将我国蕨类植物区系地理区划为代表温带亚洲成分的（特别是中亚、西亚地中海分布成分）西北温带亚洲植物区、代表古热带成分的华南古热带植物区和代表广义东亚成分的东亚植物区，东亚植物区可继续下分为中国东北—俄罗斯远东亚区（该区分布有大量的东亚－北美间断分布类型）、中国—日本植物亚区、中国—喜马拉雅植物亚区。 运用IUCN濒危物种等级对中国蕨类植物及其国家重点保护蕨类植物进行了现状评估，结果表明：可能灭绝（EX）的2种，极危（CR）的蕨类植物有33种，属于濒危（EN）的蕨类植物51种，属于易危（VU）的蕨类植物有109种，接近受危（NT）的蕨类植物158种，需予关注（LC）的845种，数据不足（DD）的1255种。对国家重点保护蕨类植物的保护级别提出部分变更的建议，对部分没有列入保护名录珍稀蕨类提出保护的建议，并就建议保护级别变更和增加保护的种类进行了必要的说明。

Cross-species chromosome painting in the golden mole and elephant-shrew: support for the mammalian clades Afrotheria and Afroinsectiphillia but not Afroinsectivora

Cross-species painting (fluorescence in situ hybridization) with 23 human (Homo sapiens (HSA)) chromosome-specific painting probes (HSA 1-22 and the X) was used to delimit regions of homology on the chromosomes of the golden mole (Ghrysochloris asiaticus) and elephant-shrew (Elephantulus rupestris). A cladistic interpretation of our data provides evidence of two unique associations, HSA 1/19p and 5/21/3, that support Afrotheria. The recognition of HSA 5/3/21 expands on the 3/21 synteny originally designated as an ancestral state for all eutherians. We have identified one adjacent segment combination (HSA2/8p/4) that is supportive of Afroinsectiphillia (aardvark, golden mole, elephant-shrew). Two segmental combinations (HSA 10q/17 and HSA 3/20) unite the aardvark and elephant-shrews as sister taxa. The finding that segmental syntenies in evolutionarily distant taxa can improve phylogenetic resolution suggests that they may be useful for testing sequence-based phylogenies of the early eutherian mammals. They may even suggest clades that sequence trees are not recovering with any consistency and thus encourage the search for additional rare genomic changes among afrotheres.

A multi-locus phylogeny of Nectogalini shrews and influences of the paleoclimate on speciation and evolution

Nectogaline shrews are a major component of the small mammalian fauna of Europe and Asia, and are notable for their diverse ecology, including utilization of aquatic habitats. So far, molecular phylogenetic analyses including nectogaline species have been unable to infer a well-resolved, well-supported phylogeny, thus limiting the power of comparative evolutionary and ecological analyses of the group. Here, we employ Bayesian phylogenetic analyses of eight mitochondrial and three nuclear genes to infer the phylogenetic relationships of nectogaline shrews. We subsequently use this phylogeny to assess the genetic diversity within the genus Episoriculus, and determine whether adaptation to aquatic habitats evolved independently multiple times. Moreover, we both analyze the fossil record and employ Bayesian relaxed clock divergence dating analyses of DNA to assess the impact of historical global climate change on the biogeography of Nectogalini. We infer strong support for the polyphyly of the genus Episoriculus. We also find strong evidence that the ability to heavily utilize aquatic habitats evolved independently in both Neomys and Chimarrogale + Nectogale lineages. Our Bayesian molecular divergence analysis suggests that the early history of Nectogalini is characterized by a rapid radiation at the Miocene/Pliocene boundary, thus potentially explaining the lack of resolution at the base of the tree. Finally, we find evidence that nectogalines once inhabited northern latitudes, but the global cooling and desiccating events at the Miocene/Pliocene and Pliocene/Pleistocene boundaries and Pleistocene glaciation resulted in the migration of most Nectogalini lineages to their present day southern distribution. (C) 2010 Elsevier Inc. All rights reserved.

A NEW SPECIES FROM SOUTHWESTERN CHINA IN THE AFRO-PALEARCTIC LINEAGE OF THE HORSESHOE BATS (RHINOLOPHUS)

A new species of horseshoe bat (Chiroptera: Rhinolophidae) is described from southwestern China. The presence of a wedge-shaped sella and pointed connecting process of the nose leaf aligns the new species to the landeri group in the Afro-Palearctic lineag

Phylogeny and zoogeography of the cyprinid hemicultrine group (Cyprinidae : Cultrinae)

Ying-Gui Dai and Jun-Xing Yang (2003) Phylogeny and zoogeography of the cyprinid hemicultrine group (Cyprinidae: Cultrinae). Zoological Studies 42(1): 73-92. The hemicultrine group consists of middle-sized cyprinids in Asia which taxonomically belong to the subfamily Cultrinae in the Cyprinidae (Cypriniformes), but there has been no convincing generic phylogenetic hypothesis proposal so far. On the basis of a morphological study of 65 specimens soaked in formalin and 14 skeletal specimens of 8 species in 6 genera within the hemicultrine group as an ingroup, a 70-character matrix was obtained. The generic phylogenetic relationships of the hemicultrine group are hypothesized with the matrix by the method of cladistic analysis. When the species Rasborinus lineatus or Cultrichthys erythropterus is used as the sole outgroup, the matrix gives the same single most-parsimonious tree of generic phylogenetic relationships within the hemicultrine group which shows that the hemicultrine group forms a monophyletic group. However, when Rasborinus lineatus is used as the sole outgroup and Cultrichthys erythropterus and Paralaubuca barroni are included in the ingroup, the hemicultrine group is validated to represent a paraphyletic group, and the hemicultrine group and the genus Paralaubuca form a monophyletic group. The tree of generic relationships and zoogeography of the monophyletic group comprising the hemicultrine group and the genus Paralaubuca suggest the following: (1) The monophyletic group comprises 2 smaller monophyletic groups: the genera Hemiculterella + Pseudohemiculter + Hainania and the genera Hemiculter + Paralaubuca + Pseudolaubuca + Toxabramis. (2) The sister groups of the monophyletic group show both overlapping and vicariant distribution patterns; therefore the generic distribution pattern of the monophyletic group maybe have resulted from both dispersal and vicariance events. (3) The monophyletic group probably originated on the Asian mainland from the Yangtze River to the Pearl River and on Hainan Island in China. (4) The monophyletic group probably originated after the Japanese Archipelago was separated from the Asian mainland at the beginning of the Quaternary Period in the Cenozoic but before Taiwan, Hainan Island, and Indonesia were completely isolated from the Asian mainland after the ice age in the Quaternary Period. (5) Speciation of the genus Hemiculter should have been the earliest, and those of the genera Paralaubuca, Pseudolaubuca, and Hainania ought to be the latest in the process of evolution of this monophyletic group. http://www.sinica.edu.tw/zool/zoolstud/42.1/73.pdf.

Relationships between Testate Amoebae Assemblages (Protozoa) and Geographic Factors in Yunnan Plateau Lakes, China

Knowledge of the effect of geographic factors on the assemblages of protozoan testate amoebae is still limited, despite there having been a number of studies on this fauna. We applied statistical analyses to data on the distribution of testate amoebae from nine major lakes in the Yunnan Plateau, southwest China. Cluster analysis, based on community structure, separated the lakes into two groups - the oligotrophic/mesotrophic lakes and the hypercutrophic lakes - confirming the idea that the testate amoebae assemblages in lakes are closely related to the trophic status. Additionally, within the oligotrophic/mesotrophic lakes, there was distinct geographic clustering. Linear regression analysis and the Mantel test both revealed that similarity of species composition decreased with increasing geographic distance among the oligotrophic/mesotrophic lakes.

Multiple invasions of europe by east asian cobitid loaches (Teleostei : Cobitidae)

While it has been widely suggested that freshwater fishes from East Asia invaded the western Palaearctic, details about this process are largely unknown. Here, using the cytochrome b gene, we evaluated the phylogenetic relationships of a small group of Eurasian primary freshwater fishes (Cobitidae), which are widely distributed and species rich in East Asia and Europe, with the purpose of inferring their invasion process of Europe from East Asia. Though phylogenetic relationships of cobitids were not well resolved, our analysis could identify three sister groups formed by the European and East Asian cobitids, which brought new insights into the biogeography of the genera Cobitis, Misgurnus, and Sabanejewia. The present results support the view that Asian cobitid fishes may have invaded Europe at least five times independently, and once reverse colonization of European cobitids to East Asia could also be found. Ancestral Sabanejewia might have been the first cobitids to cross Siberia and invade the EMZS (Euro-Mediterranean zoogeographic subregion) about 33.54 million years ago (MYA). One lineage of Cobitis and the ancestor of Misgurnus fossilis (Linnaeus) almost in the same time invaded the Europe, responding to 16.71 MYA and 16.59 MYA, respectively. Three different lineages of Cobitis were found to have invaded the EMZS from East Asia, and once reverse invasion to East Asia occurred to one subclade of European Cobitis. And our data also suggest that the diversity of East Asian cobitid fishes, especially of the genus Cobitis, is greatly underestimated.

Phylogeny of freshwater parasitic copepods in the Ergasilidae (Copepoda : Poecilostomatoida) based on 18S and 28S rDNA sequences

The phylogenetic relationships among the Ergasilidae genera are poorly understood. In this study, 14 species from four genera in the Ergasilidae including Sinergasilus, Ergasilus, Pseudergasilus, and Paraergasilus were collected in China, and their phylogenetic relationships were examined using neighbor-joining, maximum parsimony, maximum likelihood, and Bayesian inference methods based on partial sequences of 18S and 28S ribosomal deoxyribonucleic acid, respectively. All the analyses suggest that the Sinergasilus and Paraergasilus are both monophyletic, but the Ergasilus is polyphyletic rather than monophyletic. Considering the relationships among the four genera, the phylogenetic analyses and subsequent hypothesis tests all suggest that Pseudergasilus clustered with some Ergasilus species may have a closer relationship with Sinergasilus rather than with Paraergasilus. It is proposed that the Sinergasilus and the Pseudergasilus species might have evolved from Ergasilus species.

Phylogenetic relationships of the Chinese sisorid catfishes: a nuclear intron versus mitochondrial gene approach

Several recent molecular phylogenetic studies of the sisorid catfishes (Sisoridae) have challenged some aspects of their traditional taxonomy and cladistic hypotheses of their phylogeny. However, disagreement with respect to relationships within this family in these studies highlights the need for additional data and analyses. Here we subjected 15 taxa representing 12 sisorids genera to comprehensive phylogenetic analyses using the second intron of low-copy nuclear S7 ribosomal protein (rpS7) gene and the mitochondrial 16S rRNA gene segments both individually and in combination. The competing sisorid topologies were then tested by using the approximately unbiased (AU) test and the Shimodaira-Hasegawa (SH) test. Our results support previously suggested polyphyly of Pareuchiloglanis. The genus Pseudecheneis is likely to be nested in the glyptosternoids and Glaridoglanis might be basal to the tribe Glyptosternini. However, justified by AU and SH test, the sister-group relationship between Pseudecheneis and the monophyletic glyptosternoids cannot be rejected based on the second intron of rpS7 gene and combined data analyses. It follows that both gene segments are not suitable for resolving the phylogenetic relationships within the sisorid catfishes. Overall, the second intron of rpS7 gene yielded poor phylogenetic performance when compared to 16S rRNA gene, the evolutionary hypothesis of which virtually agreed with the combined data analyses tree. This phenomenon can be explained by the insufficient length and fast saturation of substitutions in the second intron of rpS7 gene, due to substitution patterns such as frequent indels (insertion/deletion events) of bases in the sequences during the evolution.

Molecular phylogeny of the specialized schizothoracine fishes (Teleostei : Cyprinidae), with their implications for the uplift of the Qinghai-Tibetan Plateau

Molecular phylogeny of three genera containing nine species and subspecies of the specialized schizothoracine fishes are investigated based on the complete nucleotide sequence of mitochondrial cytochrome b gene. Meantime relationships between the main cladogenetic events of the specialized schizothoracine fishes and the stepwise uplift of the Qinghai-Tibetan Plateau are also conducted using the molecular clock, which is calibrated by geological isolated events between the upper reaches of the Yellow River and the Qinghai Lake. Results indicated that the specialized schizothoracine fishes are not a monophyly. Five species and subspecies of Ptychobarbus form a monophyly. But three species of Gymnodiptychus do not form a monophyly. Gd. integrigymnatus is a sister taxon of the highly specialized schizothoracine fishes while Gd. pachycheilus has a close relation with Gd. dybowskii, and both of them are as a sister group of Diptychus maculatus. The specialized schizothoracines fishes might have originated during the Miocene (about 10 MaBP), and then the divergence of three genera happened during late Miocene (about 8 MaBP). Their main specialization occurred during the late Pliocene and Pleistocene (3.54-0.42 MaBP). The main cladogenetic events of the specialized schizothoracine fishes are mostly correlated with the geological tectonic events and intensive climate shift happened at 8, 3.6, 2.5 and 1.7 MaBP of the late Cenozoic. Molecular clock data do not support the hypothesis that the Qinghai-Tibetan Plateau uplifted to near present or even higher elevations during the Oligocene or Miocene, and neither in agreement with the view that the plateau uplifting reached only to an altitude of 2000 in during the late Pliocene (about 2.6 MaBP).

Comparative ontogenetic behavior and migration of kaluga, Huso dauricus, and Amur sturgeon, Acipenser schrenckii, from the Amur River

We conducted laboratory experiments with kaluga, Huso dauricus, and Amur sturgeon, Acipenser schrenckii, to develop a conceptual model of early behavior. We daily observed embryos (first life phase after hatching) and larvae (period initiating exogenous feeding) to day-30 (late larvae) for preference of bright habitat and cover, swimming distance above the bottom, up- and downstream movement, and diel activity. Day-0 embryos of both species strongly preferred bright, open habitat and initiated a strong, downstream migration that lasted 4 days (3 day peak) for kaluga and 3 days (2 day peak) for Amur sturgeon. Kaluga migrants swam far above the bottom (150 cm) on only 1 day and moved day and night; Amur sturgeon migrants swam far above the bottom (median 130 cm) during 3 days and were more nocturnal than kaluga. Post-migrant embryos of both species moved day and night, but Amur sturgeon used dark, cover habitat and swam closer to the bottom than kaluga. The larva period of both species began on day 7 (cumulative temperature degree-days, 192.0 for kaluga and 171.5 for Amur sturgeon). Larvae of both species preferred open habitat. Kaluga larvae strongly preferred bright habitat, initially swam far above the bottom (median 50-105 cm), and migrated downstream at night during days 10-16 (7-day migration). Amur sturgeon larvae strongly avoided illumination, had a mixed response to white substrate, swam 20-30 cm above the bottom during most days, and during days 12-34 (most of the larva period) moved downstream mostly at night (23-day migration). The embryo-larva migration style of the two species likely shows convergence of non-related species for a common style in response to environmental selection in the Amur River. The embryo-larva migration style of Amur sturgeon is unique among Acipenser yet studied.