Effects of monsoon on distribution patterns of tropical plants in Asia

Comparing with other regions, Asia is mostly dominated by the monsoon climate and tropical plants can be found at the furthest places away from the equator. Understanding the role of monsoon in the dispersal and evolution of tropical plants is helpful for exploring the distribution patterns of vegetation and mechanisms underlying the origin and maintenance of biodiversity in Asia. In summer, there are three types of monsoon in Asia, i.e. East Asia Monsoon, South Asia Monsoon, North-west Pacific Ocean Monsoon. The summer monsoon climate in Asia originated at about 40 Ma, when the early angiosperm evolved and started its diversification in Southeast Asia and South China. It suggested that the monsoon may facilitate the quick speciation and spread of early angiosperm. Monsoon climate facilitates the northward spread of Asia’s tropical plants and some tropical plants can be found even at Yarlung Zangbo River and the boundaries of Guizhou-Guangxi-Yunnan. Such effetcs largely change distribution patterns of zonal vegetation and even causes local vegetation types in some places with unusual topography such as tropical seasonal rainforests, monsoon rainforests, savanna and grassland along dry-hot valley in Southwest China, coastal savanna in West Hainan Island. The three summer monsoons interact at Southwest China and Indo-China Peninsula and these regions are dominated by limestone landscapes and high mountains with big rivers. Some Asia-endemic tropical taxa even formed a diversification and endemism center at this region, which may be a reason for the formation and maintenance of Indo-Burma biodiversity hotspots with global warming, the monsoon may further promote the northward spread of tropical plants and may have fundamental effects on biodiversity and flora evolution in South China.     

The diversification of the northern temperate woody flora – A case study of the Elm family (Ulmaceae) based on phylogenomic and paleobotanical evidence

Ulmaceae is a woody family widespread in northern temperate forests. Despite the ecological importance of this family, its phylogeny and biogeographic history are poorly understood. In this study, we reconstruct phylogenetic relationships within the family and infer spatio-temporal diversification patterns based on chloroplast genome (complete cpDNA) and nuclear ribosomal DNA sequences (nrDNA). The seven Ulmaceae genera are resolved in two main clades (temperate vs. tropical) by both cpDNA and nrDNA sequences. The temperate clade includes four genera, Hemiptelea, Zelkova, Planera, and Ulmus. The relationships among Planera and other genera are controversial because of inconsistent topologies between plastid and nuclear data. The tropical clade includes three genera ((Ampelocera, Phyllostylon), Holoptelea). Molecular dating and diversification analyses show that Ulmaceae originated in the Early Cretaceous (ca. 110–125 Ma) with the main lineages establishing from the Late Cretaceous to the early Eocene. The diversification rate slowed during the middle to the late Paleogene (ca. 23–45 Ma), followed by a rapid diversification of the East Asian temperate group in the Neogene, congruent with a global cooling event. The ancestral state optimization analysis suggests an East Asian origin of the temperate Ulmaceae clade during the Paleocene, which is consistent with the fossil record. Both phylogenomic and fossil evidence support East Asia as a center of origin and diversification for the temperate woody lineages.     

Both temperature fluctuations and East Asian monsoons have driven plant diversification in the karst ecosystems from southern China

Karst ecosystems in southern China are species‐rich and have high levels of endemism, yet little is known regarding the evolutionary processes responsible for the origin and diversification of karst biodiversity. The genus Primulina (Gesneriaceae) comprises ca. 170 species endemic to southern China with high levels of ecological (edaphic) specialization, providing an exceptional model to study the plant diversification in karsts. We used molecular data from nine chloroplast and 11 nuclear regions and macroevolutionary analyses to assess the origin and cause of species diversification due to palaeoenvironmental changes and edaphic specialization in Primulina. We found that speciation was positively associated with changes in past temperatures and East Asian monsoons through the evolutionary history of Primulina. Climatic change around the mid‐Miocene triggered an early burst followed by a slowdown of diversification rate towards the present with the climate cooling. We detected different speciation rates among edaphic types, and transitions among soil types were infrequently and did not impact the overall speciation rate. Our findings suggest that both global temperature changes and East Asian monsoons have played crucial roles in floristic diversification within the karst ecosystems in southern China, such that speciation was higher when climate was warmer and wetter. This is the first study to directly demonstrate that past monsoon activity is positively correlated with speciation rate in East Asia. This case study could motivate further investigations to assess the impacts of past environmental changes on the origin and diversification of biodiversity in global karst ecosystems, most of which are under threat.     

Neogene and Quaternary climate changes shaped the lineage differentiation and demographic history of Fokienia hodginsii (Cupressaceae s.l.), a Tertiary relict in East Asia

Historical climate oscillations and tectonic events have influenced the speciation and evolutionary history of many organisms. In this study, we chose Fokienia hodginsii (Dunn) A. Henry & H. H. Thomas (Cupressaceae s.l.), a Tertiary relict conifer, for the inference of the demographic history since the Neogene. Five chloroplast regions and two single-copy nuclear genes were amplified and sequenced in 497 individuals from 28 populations. The chloroplast data showed that F. hodginsii presented a high level of genetic diversity (H _T  = 0.860 ± 0.0279) and significant phylogeographic structure (N _ST  > G _ST , P < 0.05). According tobeast analysis, the divergence time of the two major lineages indicated by the phylogenetic construction produced from chloroplast and nuclear data could be dated to the early Miocene (ca. 19.34–19.95 Ma), which coincided with the onset and intensification of the Asian monsoon. During this time, environmental adaption under the different climatic conditions on either side of the Tanaka Line could have played important roles in maintaining and/or reinforcing the divergence of the two major lineages. The ecological niche modeling results showed that F. hodginsii experienced habitat fragmentation and strengthening of genetic barriers during the Last Glacial Maximum, followed by local expansion during postglacial periods. Our findings show that paleoclimate changes since the Neogene might have triggered the extinction of all but one Fokienia species and its intraspecific lineage differentiation. This study also suggests that Tertiary relicts in subtropical and tropical areas might have had a complex evolutionary history and their intraspecific differentiation time might have been earlier than expected.     

云南植物区系的起源与演化

云南是生物多样性高度富集的地区,其植物种类占中国植物种类的一半以上,几乎包含了欧亚大陆的各种主要植被类型,其错综复杂的生物区系是如何形成与演化的,这个基本问题目前尚未得到解决。本文结合云南的历史地质事件,从植物区系地理学研究上对云南植物区系的起源与演化研究进行了介绍。研究发现,云南植物区系具有一个远古的热带起源背景,在第三纪热带、亚热带性质的东亚植物区系基础上,随着喜马拉雅的隆升,世界性和北温带植物区系成分在北部地区渗透、迅速形成大量物种,使北部地区演化成为现今以世界性和北温带分布的科、属占优势的温带植物区系;在南部地区,因印度支那地质板块向东南亚逃逸,热带亚洲成分渗透、发展,演化成为以热带亚洲成分为主的热带植物区系;云南中部地区第三纪东亚植物区系成分有更多的保持与承袭。对云南南部、中部和北部植物区系系统发育关系(系统发育结构和β多样性格局)的研究支持了植物区系地理学上的推论。此外,思茅-兰坪(印度支那)地质板块自晚始新世以来发生顺时针旋转可能导致了云南西北部与云南东南部一些物种对应分布格局的形成;云南南部与东南部热带地区的植物区系可能因具有不同的地质背景和演化历程而发生了显著的生物地理分异。在晚第三纪各地质事件的影响下,云南植物区系在一个热带、亚热带性质的第三纪东亚植物区系基础上发生了歧化,演化成为现今的南-北、东-西生物地理分异明显的植物区系。     

NADH dehydrogenase subunit 1 and cytochrome c oxidase subunit I sequences compared for members of the genus Taenia (Cestoda)

Nine members of the genus Taenia (Taenia taeniaeformis, Taenia hydatigena, Taenia pisiformis, Taenia ovis, Taenia multiceps, Taenia serialis, Taenia saginata, Taenia solium and the Asian Taenia) were characterised by their mitochondrial NADH dehydrogenase subunit 1 gene sequences and their genetic relationships were compared with those derived from the cytochrome c oxidase subunit I sequence data. The extent of inter-taxon sequence difference in NADH dehydrogenase subunit 1 (5.9–30.8%) was usually greater than in cytochrome c oxidase subunit I (2.5–18%). Although topology of the phenograms derived from NADH dehydrogenase subunit 1 and cytochrome c oxidase subunit I sequence data differed, there was concordance in that T. multiceps, T. serialis (of canids), T. saginata and the Asian Taenia (of humans) were genetically most similar, and those four members were genetically more similar to T. ovis and T. solium than they were to T. hydatigena and T. pisiformis (of canids) or T. taeniaeformis (of cats). The NADH dehydrogenase subunit 1 sequence data may prove useful in studies of the systematics and population genetic structure of the Taeniidae.     

中国晚新近纪哺乳动物群落与东亚环境变化

根据已有的中国晚新近纪哺乳动物化石记录,综合新近研究成果,初步探讨中国晚新近纪哺乳动物群落演变进程及其对于气候与环境变化的响应。结果表明,自中中新世晚期至晚中新世(约13 Ma至7-8 Ma),中国北方哺乳动物群落组成上没有明显的分异;而在其后的晚中新世晚期发生了较为明显的区域性分化,分化为东部地区以适应潮湿的较为封闭的动物群落与西部地区适应干旱的开阔环境的群落,这种分异可能与东亚夏季风的加强密切相关; 青藏高原及其周边地区化石类群的扩散与交流资料也表明由青藏高原隆升而形成的地理阻隔也可能发生在该时期;上新世哺乳动物群落演化表现出对于持续的干旱化气候背景的适应,而晚上新世某些类群的分异与主导地位的确立与该时期气候环境背景的剧烈震荡相一致。由于化石资料的不完整,时段分布的不均一,以及地理分布区域的局限,全面认识中国晚新近纪哺乳动物群落组成以及与环境变化的关系尚有很长的一段历程。     

Biogeographical Divergence of the Flora of Yunnan,Southwestern China Initiated by the Uplift of Himalaya and Extrusion of Indochina Block

The floral composition of Yunnan is conspicuously linked to the biogeographical history of this extremely species-rich province in southwestern China. The floristic compositions of three representative regions in Yunnan were compared to reveal their variation with geography. From southern Yunnan, 4150 native species (including subspecies and varieties) from 1240 genera and 183 families of seed plants were recognized. From central Yunnan 3389 native species from 1095 genera and 167 families of seed plants were recognized. From northwestern Yunnan 6807 native species from 1296 genera and 166 families of seed plants were recognized. Although these three floras across Yunnan are similar in familial composition, similarities between the floras of southern and northwestern Yunnan are low at the generic and specific levels. The flora of northwestern Yunnan is dominated by families and genera with cosmopolitan and north temperate distributions, while the flora of southern Yunnan is dominated by tropical families and genera. Northwestern Yunnan is composed largely of temperate genera, of which the highest proportion has a north temperate distribution. In contrast, southern Yunnan has mainly tropical genera, of which most have a tropical Asian distribution. The flora of central Yunnan is a combination of southern and northwestern Yunnan. These three floras might be derived from a common Tertiary tropical or subtropical East Asian flora, but the geological history of each region has influenced its flora, and they have remained divergent since the late Tertiary. The flora of northwestern Yunnan has evolved with the uplift of the Himalayas and by gradual proliferation of mainly cosmopolitan and north temperate floristic elements, while the flora of southern Yunnan has evolved with extrusion of the Indochina block and the influence of mainly tropical Asian elements.     

Historical biogeography of the Southeast Asian and Malesian tribe Dissochaeteae (Melastomataceae)

The region of Tropical Southeast Asia and the Malay Archipelago is a very appealing area for research due to its outstanding biodiversity, being one of the most species-rich areas in the world with high levels of endemism, and due to its complex geological history. The high number of species in tribe Dissochaeteae (Melastomataceae) and their tendency to narrow endemism makethe tribe an ideal group for examining biogeographic patterns. We sampled 58 accessions spread over 42 accepted and two undescribed species of the Dissochaeteae. Two nuclear (ETS, ITS) and four chloroplast regions (ndhF, psbK-psbL, rbcL, rpl16) were used for divergence time estimation and ancestral area reconstruction. Results from the molecular dating analysis suggest that the diversity of Dissochaeteae in the Southeast Asian region resulted from a South American ancestor in the late Eocene. The ancestor of the Dissochaeteae might have migrated from South America to Southeast Asia via North America and then entered Eurasia over the North Atlantic land bridge during the Eocene. The origin and early diversification of the Dissochaeteae in Southeast Asia dates back to the middle Oligocene, and most of the genera originated during the Miocene. Indochina and Borneo are most likely the area of origin for the most recent common ancestor of the Dissochaeteae and for many of the early diverging clades of some genera within Southeast Asia.     

Animal‐mediated long‐distance dispersals and migrations shaping the intercontinental disjunctions of Celastrus (Celastraceae) among five continents

The mechanisms underlying the origin, evolution, and distributional patterns of organisms are a major focus of biogeography. Vicariance and long‐distance dispersal (LDD) are two important explanations for disjunctive distribution patterns among lineages. In‐depth biogeographic studies of taxa that exhibit wide‐ranging disjunctions can provide valuable information for addressing the relative importance of these biogeographic mechanisms. The genus Celastrus contains ca. 30 species that are disjunctly distributed in five continents of both the Northern and Southern Hemispheres, providing an excellent system for historical biogeographic analyses. Here, we used sequence data from five markers (nuclear external transcribed spacer and internal transcribed spacer, and plastid psbA‐trnH, rpl16, and trnL‐F) to reconstruct the phylogeny of Celastrus and investigate its phylogenetic relationships with Tripterygium, estimate clade divergence times using the fossil‐calibrated method, and infer its ancestral distribution range. Celastrus and Tripterygium were each supported as monophyletic. The morphology‐based classification systems were not supported by the phylogenetic results. The divergence time between Celastrus and Tripterygium was estimated to be 26.22 Ma (95% highest posterior density: 24.46–28.17 Ma), and the diversification of Celastrus were suggested to be linked to global warming events during the Miocene. Celastrus was suggested to have a tropical Asian origin, and dispersed to Central and South America, North America, Oceania, and Madagascar at different periods, most probably through LDD. Birds may have facilitated transoceanic migrations of Celastrus because of its bicolored fruits, which contain red and fleshy arils. Our results highlight the importance of key morphological innovations and animal‐mediated dispersals for the rapid diversification of plant lineages across vast distributional ranges.     

Out of the Andes: patterns of diversification in clearwing butterflies

Global biodiversity peaks in the tropical forests of the Andes, a striking geological feature that has likely been instrumental in generating biodiversity by providing opportunities for both vicariant and ecological speciation. However, the role of these mountains in the diversification of insects, which dominate biodiversity, has been poorly explored using phylogenetic methods. Here we study the role of the Andes in the evolution of a diverse Neotropical insect group, the clearwing butterflies. We used dated species-level phylogenies to investigate the time course of speciation and to infer ancestral elevation ranges for two diverse genera. We show that both genera likely originated at middle elevations in the Andes in the Middle Miocene, contrasting with most published results in vertebrates that point to a lowland origin. Although we detected a signature of vicariance caused by the uplift of the Andes at the Miocene–Pliocene boundary, most sister species were parapatric without any obvious vicariant barrier. Combined with an overall decelerating speciation rate, these results suggest an important role for ecological speciation and adaptive radiation, rather than simple vicariance.     

西双版纳自然保护区蚁生线虫草时空分布及寄主多样性

蚁生线虫草是一种重要的蚁科专化性虫生真菌。为明确双版纳自然保护区不同生境蚁生线虫草的分布规律,以期为蚁生线虫草的开发利用提供依据,于2017年7月-2018年9月对西双版纳自然保护区石灰岩山季雨林（高、中、低海拔带）、热带季雨林、季风常绿阔叶林、热带丛生竹林、暖热性针叶林5种不同生境中蚁生线虫草的分布及其寄主进行了系统调查。结果表明,在5种不同生境中,共采集蚁生线虫草1 449株,其主要分布于石灰岩山季雨林和热带丛生竹林,相对多度分别为51.1%和24.3%;其次为暖热性针叶林生境,相对多度为15.2%;热带季雨林和季风常绿阔叶林中分布相对较少,相对多度分别为5.7%和3.7%。从同一生境的石灰岩山季雨林不同海拔带的分布来看,蚁生线虫草主要分布于高海拔地带（33.7%）,而中海拔地带（9.7%）和低海拔（7.8%）相对较少。从季节性流行规律来看,西双版纳自然保护区蚁生线虫草的流行期为湿热季的6-10月及湿热季与雾凉季交替的11月。蚁生线虫草寄主多样性丰富,包括4亚科13属16种。该研究将为西双版纳自然保护区生物多样性及蚁生线虫草保护利用提供理论依据。     

Evolutionary Dynamics in the Southwest Indian Ocean Marine Biodiversity Hotspot: A Perspective from the Rocky Shore Gastropod Genus Nerita

The Southwest Indian Ocean (SWIO) is a striking marine biodiversity hotspot. Coral reefs in this region host a high proportion of endemics compared to total species richness and they are particularly threatened by human activities. The island archipelagos with their diverse marine habitats constitute a natural laboratory for studying diversification processes. Rocky shores in the SWIO region have remained understudied. This habitat presents a high diversity of molluscs, in particular gastropods. To explore the role of climatic and geological factors in lineage diversification within the genus Nerita, we constructed a new phylogeny with an associated chronogram from two mitochondrial genes [cytochrome oxidase sub-unit 1 and 16S rRNA], combining previously published and new data from eight species sampled throughout the region. All species from the SWIO originated less than 20 Ma ago, their closest extant relatives living in the Indo-Australian Archipelago (IAA). Furthermore, the SWIO clades within species with Indo-Pacific distribution ranges are quite recent, less than 5 Ma. These results suggest that the regional diversification of Nerita is closely linked to tectonic events in the SWIO region. The Reunion mantle plume head reached Earth’s surface 67 Ma and has been stable and active since then, generating island archipelagos, some of which are partly below sea level today. Since the Miocene, sea-level fluctuations have intermittently created new rocky shore habitats. These represent ephemeral stepping-stones, which have likely facilitated repeated colonization by intertidal gastropods, like Nerita populations from the IAA, leading to allopatric speciation. This highlights the importance of taking into account past climatic and geological factors when studying diversification of highly dispersive tropical marine species. It also underlines the unique history of the marine biodiversity of the SWIO region.     

Tectonics,climate and the diversification of the tropical African terrestrial flora and fauna

Tropical Africa is home to an astonishing biodiversity occurring in a variety of ecosystems. Past climatic change and geological events have impacted the evolution and diversification of this biodiversity. During the last two decades, around 90 dated molecular phylogenies of different clades across animals and plants have been published leading to an increased understanding of the diversification and speciation processes generating tropical African biodiversity. In parallel, extended geological and palaeoclimatic records together with detailed numerical simulations have refined our understanding of past geological and climatic changes in Africa. To date, these important advances have not been reviewed within a common framework. Here, we critically review and synthesize African climate, tectonics and terrestrial biodiversity evolution throughout the Cenozoic to the mid-Pleistocene, drawing on recent advances in Earth and life sciences. We first review six major geo-climatic periods defining tropical African biodiversity diversification by synthesizing 89 dated molecular phylogeny studies. Two major geo-climatic factors impacting the diversification of the sub-Saharan biota are highlighted. First, Africa underwent numerous climatic fluctuations at ancient and more recent timescales, with tectonic, greenhouse gas, and orbital forcing stimulating diversification. Second, increased aridification since the Late Eocene led to important extinction events, but also provided unique diversification opportunities shaping the current tropical African biodiversity landscape. We then review diversification studies of tropical terrestrial animal and plant clades and discuss three major models of speciation: (i) geographic speciation via vicariance (allopatry); (ii) ecological speciation impacted by climate and geological changes, and (iii) genomic speciation via genome duplication. Geographic speciation has been the most widely documented to date and is a common speciation model across tropical Africa. We conclude with four important challenges faced by tropical African biodiversity research: (i) to increase knowledge by gathering basic and fundamental biodiversity information; (ii) to improve modelling of African geophysical evolution throughout the Cenozoic via better constraints and downscaling approaches; (iii) to increase the precision of phylogenetic reconstruction and molecular dating of tropical African clades by using next generation sequencing approaches together with better fossil calibrations; (iv) finally, as done here, to integrate data better from Earth and life sciences by focusing on the interdisciplinary study of the evolution of tropical African biodiversity in a wider geodiversity context.     

中亚热带濒危植物毛红椿和南方红豆杉种内与种间竞争差异

濒危植物是全球生物多样性的重要组成部分,为了弄清中亚热带地区不同生长型濒危乔木树种的竞争模式及其机制,研究了江西九岭山国家级自然保护区毛红椿和南方红豆杉的种内与种间竞争差异。结果表明: 阳性落叶树种毛红椿种群的竞争压力主要来自于同种个体,种内竞争指数占总竞争指数的66.4%,耐阴常绿树种南方红豆杉种群主要受种间竞争的影响,种间竞争指数占总竞争指数的68.7%。2种濒危植物受到的种内、种间以及总竞争强度均随着林木径级的增大而逐渐减小,表明竞争压力由幼林向成熟林逐渐降低。在生长过程中,毛红椿以种内竞争引起的自疏作用为主导,南方红豆杉则由种间竞争引起的他疏作用为主导。毛红椿和南方红豆杉种群在更新过程中,小径级个体都要经历强度较大的种内或种间竞争选择才能发育为成树。由于主要竞争压力的来源差异,对于生活在中亚热带同一区域内不同生长型的濒危植物应该采取不同的生物多样性保育措施。     

Novel classification and biogeography of Leptolejeunea (Lejeuneaceae,Marchantiophyta) with implications for the origin and evolution of the Asian evergreen broad-leaved forests

Although recent molecular phylogenetic analyses have greatly improved our understanding of the classification of the large liverwort family Lejeuneaceae, the frequent incongruencies between morphology-based taxonomy and molecular phylogeny have hindered our understanding of evolutionary diversification within the group. Here we focus on Leptolejeunea (Spruce) Steph., a pantropical epiphyllous genus in Lejeuneaceae with 40 species. Phylogenetic studies on the genus have been hampered by insufficient taxon sampling, leaving the deep phylogenetic relationships within this group unresolved. We present the most comprehensive phylogenetic analysis of the genus to date with sampling of over 80% of species, including the enigmatic Leptolejeunea spinistipula (Mizut.) X.L.He endemic to Borneo. Based on data from three molecular markers with representatives of Leptolejeunea and its allies, Leptolejeunea appeared to be monophyletic following the exclusion of L. spinistipula and its transfer to Soella R.L.Zhu, L.Shu, Qiong He & Y.M.Wei. A total-evidence approach was taken to resolve the backbone phylogeny of Leptolejeunea and a first infrageneric classification of Leptolejeunea, including a new subgenus and three new sections, is proposed based on integrated molecular and morphological evidence. Reconstruction of the evolutionary history showed a wide ancestral area of Leptolejeunea during the Paleogene that arose in mainland Asia, followed by an accelerated speciation rate. Across the biogeographical history of Leptolejeunea, long-distance dispersal had profound effects on population expansion. Our findings suggest that Australasia is a source of biodiversity of Asian evergreen broad-leaved forests that have been established since the Oligocene and rose after the early Miocene.     

Plastome phylogenomic analysis of Torreya (Taxaceae)

Torreya Arn., a small genus of Taxaceae, consists of six species occurring in North America and eastern Asia. Several phylogenetic studies have previously been undertaken to reveal relationships within this genus, although only a few DNA segments or species were used. In the present study, we sequenced five Torreya plastomes and combined these with two existing plastomes from the genus to investigate plastome evolution and phylogenetic relationships within Torreya. All sequenced Torreya plastomes shared the same complement of 82 protein‐coding genes, 4 ribosomal RNA genes, and 31 transfer RNA genes. Phylogenetic inference using a maximum likelihood framework consisted of an 82‐gene, 17‐taxon dataset, including all species of Torreya, resolved Torreya as a monophyletic clade. Strongly supported relationships within the genus include the position of the early diverging T. jackii Chun, the two sister pairs T. fargesii Franch.–T. nucifera (L.) Siebold & Zucc. and T. grandis Fortune ex Lindl.–T. californica Torr., and the monophyly of the clade including T. fargesii var. yunnanensis, T. fargesii, and T. nucifera. In addition to the inference of species relationships, divergence time estimation and biogeographical analysis were carried out. The diversification of Torreya was estimated to be approximately 8.9 Ma. Ancestral state reconstruction of the geographical area suggested China/eastern North America as the most likely ancestral region for the six extant Torreya species.     

Evidence of extreme habitat stability in a Southeast Asian biodiversity hotspot based on the evolutionary analysis of neotenic net-winged beetles

The diversification of neotenic beetle lineages has not been studied, despite the potential for defining biodiversity hotspots and elucidating the history of regional faunas. Additionally, neotenics may provide insight into the process of speciation in small populations with extremely low dispersal ability and a limited range. Here, we used two rDNA and three mtDNA markers to investigate the phylogeny of Scarelus, a neotenic lineage endemic to Southeast Asian rainforests. Most genetic differentiation was associated with Palaeogene geographical divisions, which remain distinct despite temporary connections. Dispersal events were rare, with only two inferred for Scarelus: from Borneo to the Philippines 28.3 million years ago (Ma) and from Sumatra to Java 13.9 Ma. We suggest that speciation resulted from allopatric range fragmentation, and Scarelus diversified readily when conditions were favourable; in this case, at different times in the eastern (19.3-39.1 Ma) and western (3.5-13.9 Ma) parts of Sundaland. The observed strong phenotypic similarity was preserved under speciation through complete allopatry. Neotenic Lycidae have survived for a long time in very stable habitats, and extremely low dispersal activity has not limited their persistence; however, the long-term diversification rate of neotenics is low and diversification is nonexistent under stable conditions. The modern ranges of neotenic lineages are indicative of ancient rainforest refugia and may be used in biodiversity conservation management. Most neotenics are at risk of extinction because of their small ranges and a low dispersal potential.     

古地中海退却与喜马拉雅—横断山的隆起在中国喜马拉雅成分及高山植物区纱的形成与发展上的意义

横断山－喜马拉雅植物区系的开端是在晚白垩纪和早古近纪（早第三纪）。古植物资料在古近纪初期横断山－喜马拉雅植物区系是同古地中海沿岸一致的以照叶林为主的暖湿植物区系。古近纪后期和新近纪（新第三纪）以后古地中海气候逐步旱化。原来的暖湿植物区系在地中海地区逐步消失,而在横断山及喜马拉雅和东亚其他地区得以保存和发展,现代横断山及东喜马拉雅的亚热带森林即是其后裔。古近纪中期以后由于古地中海的逐步退却,气候变得干旱,原暖湿植物区系逐步被现代旱生的地中海植物区系所取代。新近纪以后,旱生的现代地中海植物区系由于喜马拉雅和横断山的隆起而转向适应高山环境,逐步分化形成了现代的中国－喜马拉雅成分。横断山－喜马拉雅地区硬中高山栎林的起源;铁筷子属,绿绒蒿属,芒苞草属,假百合属及马桑属的地中海,喜马拉雅－横断山间断分布的形成便是古地中海植物区系残遗的体现;黄花木属,独一味属等众多中国喜马拉雅成分就是古地中海祖先的后裔。这些代表类群的分析研究表明现代的喜马拉雅－横断山的高山植物区系以及中国－喜马拉雅成分中有相当的一部分是起源于新生代旱生的地中海植物区系。