蕨类植物起源与系统发生关系研究进展

蕨类植物是一群非常古老的植物,在其漫长的演化历史中丢失了大量的系统发生信息,因此重建其系统演化关系是蕨类植物学家所面临的重要任务之一.近年来的分子系统学研究已经确立了一些类群的系统关系,但现存的蕨类分类系统和系统发生关系还有很多谜团有待揭开.化石记录表明蕨类植物最早出现于大约4亿多年前,但大部分现代真蕨类缺少足够的化石证据.它们的起源时间和方式及其内部各类群的系统发生关系等诸多问题的解决,还有待于大量形态学和不同分子水平上的实验数据的进一步积累,并结合其直接的化石证据进行综合研究.     

石松类和蕨类植物研究进展: 兼论国产类群的科级分类系统

综述了石松类和蕨类植物系统发育的最新研究成果。目前研究表明传统的蕨类植物概念(包括石松类和蕨类)需要修订, 一个关于蕨类植物的分类系统也已经发表。中国的植物多样性很丰富, 包括了世界上石松类和蕨类植物主要类群的代表。我们利用rbcL基因序列(包括国产蕨类63科中的62科179属184种)构建了系统发育树。基于rbcL序列分析所获得的石松类和蕨类各主要类群间的系统演化关系同以往对各个特定类群开展的较为密集的类群取样和多性状分析(形态学＋分子序列证据)的结果基本一致。在参考Smith等人系统的基础上, 我们尝试性地对中国石松类和蕨类植物进行了科级水平上的系统发育重排。     

蕨类植物的起源演化:对"古老"类群的重新审视

一般认为,蕨类植物是一群非常古老的植物,其演化历史可以追溯到4亿多年前。近十几年来,由于多种研究手段的应用,特别是分子系统学的异军突起,在蕨类植物起源演化方面取得了重要进展,使我们可以重新审视这一古老植物类群的演化历史。本文与传统观点相对比,简述了蕨类植物在分类范畴、起源演化及其重要类群间的系统演化关系方面的最新进展。     

中国现代石松类和蕨类植物分类系统概览

回顾了蕨类植物的系统分类简史,介绍了世界上石松类和蕨类植物分类研究的最新成果,即石松类和蕨类是维管植物中的两大类群,其系统发育关系逐渐明确,分类系统已经得到更新。基于新的分类系统,中国石松类有1亚纲、3目、3科;蕨类有4亚纲、11目、37科及12亚科。     

介形类系统分类与起源演化的形态、分子和化石证据

介形类(Ostracoda)因其丰富的化石记录和广布的海陆现生代表类群,而被认为是进化生物学中研究生物多样性产生机制和演变历程的颇具潜力的重要模式生物。介形类在甲壳亚门中的谱系发生位置、起源及其内部各类群间的系统关系还存在诸多争议。基于其体制构造的形态学特征,介形类被归入甲壳亚门下的颚足纲(Maxillopoda),但来自18S rDNA序列数据分析却显示Maxillopoda不是单系群。基于化石记录和壳体形态特征,高肌虫(Bradoriida)长期以来被认为是介形类的一个祖先类群,但保存有软躯体的早寒武世化石的研究表明,Bradoriida不是介形类甚至可能也不属于甲壳类。不同的研究者所强调的壳体和肢体形态特征各异,导致介形类最大的现生类群速足目(Podocopida)的四个超科之间的关系也存在诸多推测。壳体和肢体特征在系统演化意义上的不兼容,需要分子生物学等证据的介入。分子、形态和化石证据的积累及各种信息整合是系统演化研究的必然趋势。     

中国蕨类植物区系地理若干问题研究进展

在简述蕨类植物系统发生关系、传统分类系统与基于分子系统学的现代分类系统的基础上,概述了中国蕨类植物区系地理研究的现状及最新进展,并对今后的研究工作进行了展望。根据分子系统学研究,传统的蕨类植物门以及一些蕨类植物类群并不是自然类群。中国蕨类植物区系及分布的基本规律是东南多、西北少,华南、西南极为丰富,西南地区是许多温带属的分布中心。分析了中国蕨类植物区系研究的主要问题,指出关于蕨类植物在生态系统中发挥的作用,特殊生境研究、数值分析方法在蕨类区系地理中的应用,区系地理学与其他学科的交叉是今后可能的热点与亟待加强的几个领域。     

苔藓动物的起源与系统发生研究进展—形态学和分子生物学的证据

苔藓动物是后生动物中的一个重要类群。然而,和其它主要后生动物类群相比,长期以来对它的系统学研究却相对滞后。其起源,系统发生地位以及与其它后生物门类之间、其内部各高级分类群间的谱系发生关系一直存在争议。一般认为它是介于原口动物和后口动物之间的过渡类群。但是,近年来的分子系统学研究已经证实了它的原口归属。古生物学资料表明,虽然苔藓动物的大多数类群在奥陶纪已经分化出来,但它在寒武纪却缺乏任何化石记录。另外,苔藓动物起源的时间和方式、其内部各类群间的系统发生关系特别是现生类群和化石类群之间的关系等诸多问题的解决,还有待于大量的形态学和不同的分子数据的进一步积累,并结合其地层分布等各种相关资料进行综合研究。     

多足动物亚门主要类群的谱系发生与谱系年代分析——基于形态、化石和分子的综合数据

多足动物可能是最早入侵陆地环境的无脊椎动物之一。随着泛甲壳动物大类的确立,多足动物亚门的谱系发生问题已成为节肢动物研究中的焦点。基于表型性状的系统学研究中,多足动物亚门在节肢动物门中的系统分类和单系性等问题一直存在争议;化石记录的稀少又使多足动物的起源及演化历史变得迷雾重重。近年来分子系统学的进展为深入探讨多足动物谱系发生问题开辟了新的途径;分子定年的应用为探讨多足动物起源、登陆等早期演化问题提供了新的手段。谱系年代学综合了化石记录和分子定年两方面的优势,为更加精确地讨论多足动物起源、内部类群分歧时间及其地质背景奠定了基础。谱系年代分析显示,多足动物类起源于寒武纪晚期或更早;多足动物内部类群的分歧不晚于奥陶纪;而化石证据显示,多足动物的登陆事件也可能发生在这一时期,多足动物内部类群的分歧事件(及食性分化)可能与登陆过程有关。精确的多足动物亚门谱系发生关系以及谱系年代学细节还有待于进一步综合系统学、多基因和多重化石参照点的合理分析加以深入和完善,进而为早期陆地复杂生态系统的建立提供新的证据。     

壳斗演化的假说及分子系统学和化石证据

壳斗作为壳斗科Fagaceae的重要特征, 其起源、演化和多样性形成长期以来为系统学家们所关注, 有众多的观点和假说。本文在综合这些假说的基础上, 结合分子系统学和大化石证据, 讨论了壳斗演化的规律。研究结果支持壳斗具枝性性质, 来源于二歧聚伞花序最外侧小枝的假说。在比较3个壳斗演化模型的基础上, 根据4种分子系统关系的一致性重建了壳斗的演化历史: 壳斗是单系起源; 水青冈属的2果4裂瓣壳斗是最早分化的壳斗类型, 三棱栎型壳斗是较晚分化的类型; 并支持具有内裂瓣的金鳞果类型壳斗可能与其他横切面为圆形的壳斗有较近的共同祖先。这一结果也得到了多个大化石证据的支持。壳斗科多个分子系统关系的一致性和壳斗化石证据共同证实了壳斗演化的规律是从开裂向裂瓣融合、逐步简化、多向发展; 支持开裂壳斗、坚果三角形、较小、具棱或狭翅是祖征, 融合壳斗、坚果圆形、较大、不具棱是衍征。这些结果为进一步探索壳斗科的演化历史和系统发育提供了新的证据。     

分子系统学在微体古生物及相关领域中的应用

核酸（ＤＮＡ和ＲＮＡ）和蛋白等生物大分子（尤其是它们的序列资料）可作为重要的生物性状用于系统分类和演化等主题的研究。相对于形态学性状而言,分子性状不仅是前者的补充,而且具有许多前者无法比拟的优点;比如ＤＮＡ作为遗传信息的直接载体能较准确地反映生物类群之间的系统发生关系、信息量巨大、易于定量化和进行计算机分析等等。分子古生物研究包含两个方面：一、发掘化石生物分子,以提供历史生物界演化过程中的直接遗传学证据以及检验分子演化速率等方面的独特数据;二、利用现代分子生物学数据,探讨化石生物界的系统发生问题。上述两个研究方向均已成为当今演化生物学领域的热点。有孔虫等具有重要化石记录的微体生物的分子系统学研究已经开始。随着现生的和化石的生物分子资料的逐渐积累,预期在不久的将来,分子资料将成为微体古生物研究中不可缺少的重要数据之一。     

中国蕨类植物细胞分类学研究概况

中国蕨类植物类群十分丰富,由63科230属约2600种组成,其中约有10％的种类为中国特有。迄今为止,中国蕨类植物进行过染色体计数的种类约有395种,仅占中国蕨类植物总数的15％,其中48％为多倍体。本文对中国蕨类植物细胞分类学研究的历史和现状进行了综合评述。文中论述了蕨类植物染色体数目和染色体基数在系统学上的应用以及蕨类植物染色体组型分析、多倍化、无融合生殖、多倍体复合体及网状进化的研究概况。中国20多年的蕨类植物细胞分类学研究表明,蕨类植物的染色体数目、基数、组型分析、多倍化及其繁殖方式对蕨类植物的分类、起源和演化的研究都有重要作用,对种及种复合体的鉴定、科属的划分也有重要的参考价值。并对中国蕨类植物细胞分类学研究提出了一些建议。     

Pteridophytes as field indicators of natural biodiversity restoration in the Scottish flora

Summary

The pteridophyte flora of Scotland, as with that of the British Isles as a whole, is still recovering from the great reduction of its original diversity caused by the Pleistocene glaciations. However, the isolation of the British Isles, their range of suitable habitats and their climatic and edaphic opportunity has provided conditions in which processes of natural biodiversity restoration in pteridophytes can be demonstrated and monitored. The modern pteridophyte vegetation of Scotland is analyzed in this evolutionary perspective. Using the examples of both ferns and fern allies, it is suggested that it is at least as important to conserve suitable sites for the continued origin of novel taxa as it is to allow for the perpetuation of existing taxa and communities.     

Relative importance of water, energy, and heterogeneity in determining regional pteridophyte and seed plant richness in China

Environmental variables, such as ambient energy, water availability, and environmental heterogeneity have been frequently proposed to account for species diversity gradients. How taxon-specific functional traits define large-scale richness gradients is a fundamental issue in understanding spatial patterns of species diversity, but has not been well documented. Using a large dataset on the regional flora from China, we examine the contrast spatial patterns and environmental determinants between pteridophytes and seed plants which differ in dispersal capacity and environmental requirements. Pteridophyte richness shows more pronounced spatial variation and stronger environmental associations than seed plant richness. Water availability generally accounts for more spatial variance in species richness of pteridophytes and seed plants than energy and heterogeneity do, especially for pteridophytes which have high dependence on moist and shady environments. Thus, pteridophyte richness is disproportionally affected by water-related variables; this in turn results in a higher proportion of pteridophytes in regional vascular plant floras (pteridophyte proportion) in wet regions. Most of the variance in seed plant richness, pteridophyte richness, and pteridophyte proportion explained by energy is included in variation that water and heterogeneity account for, indicating the redundancy of energy in the study extent. However, heterogeneity is more important for determining seed plant distributions. Pteridophyte and seed plant richness is strongly correlated, even after the environmental effects have been removed, implying functional linkages between them. Our study highlights the importance of incorporating biological traits of different taxonomic groups into the studies of macroecology and global change biology.     

New Zealand's pteridophyte flora—Plants of ancient lineage but recent arrival?

A hypothesis is presented that most pteridophytes arrived in New Zealand relatively recently, by long-distance dispersal. The flora comprises 194 native species, of which 89 (46%) are endemic and 105 (54%) are widespread. Of the latter, 90% are shared with temperate Australasia, 53% with tropical regions, 14% with temperate southern Africa and 13% with the circum-Antarctic islands and South America. New Zealand has undergone such dramatic changes in location, land area, and topography since initial separation from Gondwana 85 Ma that it seems improbable that the 95 species shared with temperate Australasia could have remained conspecific throughout that time. Modern fossil and molecular evidence strongly suggest that many families of ferns had not even evolved prior to separation, and palynological evidence from New Zealand indicates that 78% of pteridophyte genera first appeared there only after separation from Gondwana. Present-day distributions in New Zealand suggest that ferns have greater dispersal potential than flowering plants, and that pteridophyte distributions are more heavily influenced by temperature, rainfall, and geothermal activity than by geological history. Most endemic pteridophyte species have a predominantly southern distribution pattern and are characteristic of cool, lowland to montane forest. Pteridophytes in the northern part of New Zealand show a lower level of endemism than elsewhere and tend to be widespread species that have arrived from temperate Australasian and tropical regions. There is also evidence that at least some pteridophytes have migrated from New Zealand to Australia. It is suggested that the hypothesis of long-distance dispersal of pteridophytes across the Tasman Sea could be tested by molecular techniques.     

The arbuscular mycorrhizas of pteridophytes in Yunnan, southwest China: evolutionary interpretations

 The percentage of arbuscular mycorrhizal pteridophytes among 256 pteridophyte species distributed in Yunnan (southwest China) was found to be lower than that in angiosperms. In the pteridophytes, the occurrence of arbuscular mycorrhizas was low in sporophytes of fern-allies and leptosporangiates, whereas in the eusporangiates it was relatively high. From the standpoint of mycotrophism, the evolutionary trend in the Filicineae may be from constantly mycorrhizal to facultative mycorrhizal and finally to nonmycorrhizal plants. Accepted: 21 September 2000     

Contrasting environmental and regional effects on global pteridophyte and seed plant diversity

Pteridophytes (ferns and fern‐allies) represent the second‐largest group of vascular plants, but their global biogeography remains poorly studied. Given their functional biology, pteridophytes are expected to show a more pronounced relation to water availability and a higher dispersal ability compared to seed plants. We test these assertions and document the global pattern of pteridophyte richness across 195 mainland and 106 island regions. Using non‐spatial and spatial simple and multiple regression models, we analyze geographic trends in pteridophyte and seed plant richness as well as pteridophyte proportions in relation to environmental and regional variables. We find that pteridophyte and seed plant richness are geographically strongly correlated (all floras: r=0.68, mainland: r=0.82, island floras: r=0.77), but that the proportions of pteridophytes in vascular plant floras vary considerably (0–70%). Islands (mean=15.3%) have significantly higher proportions of pteridophytes than mainland regions (mean=3.6%). While the relative proportions of pteridophytes on islands show a positive relationship with geographic isolation, proportions in mainland floras increase most strongly along gradients of water availability. Pteridophyte richness peaks in humid tropical mountainous regions and is lowest in deserts, arctic regions, and on remote oceanic islands. Regions with Mediterranean climate, outstanding extra‐tropical centres of seed plant richness, are comparatively poor in pteridophytes. Overall, water‐energy variables and topographical complexity are core predictors of both mainland pteridophyte and seed plant richness. Significant residual richness across biogeographic regions points to an important role of idiosyncratic regional effects. Although the same variables emerge as core predictors of pteridophyte and seed plant richness, water availability is clearly a much stronger constraint of pteridophyte richness. We discuss the different limitations of gametophytes and sporophytes that might have limited the ability of pteridophytes to extensively diversify under harsh environmental conditions. Our results point to an important role of taxon‐specific functional traits in defining global richness gradients.     

黄土高原蕨类植物区系特点的初步研究

对黄土高原的蕨类植物区系组成、分类及其特点进行了研究。结果表明,黄土高原有蕨类植物26科、53属、184种及3变种,种类较多的科有蹄盖蕨科（30种）、鳞毛蕨科（26种）、水龙骨科（20种）、铁角蕨科（12种）、卷柏科（12种）、中国蕨科（11种）及裸子蕨科（10种）等。这７科的种类共计１２１种,占本地区总种数的64.71%。种类较多的属有鳞毛蕨属（13种）、郑柏属（12种）、耳蕨属（11种）、铁角蕨属（11种）、蹄盖蕨属（10种）、瓦韦属（10种）、铁线蕨属（9种）及岩蕨属（9种）等。这8属的种类共计85种,占本地区总种数45.46%,黄土高原蕨类植物区系属的地理成分可划分为15个分布区类型。热带、亚热带分布类型的有18属,占总属数的41.86%,北温带及寒带分布类型的有22属,占总属数（世界分布属除外）的51.16%,其中,亚洲分布类型的最多,计有16属,占总属数的37.20%,表明黄封同原蕨类植物区系是温带类型,区系地理成分以华北区蕨类植物为主,同时也集中了华中、华东、东北、西北以及西南的蕨类植物。而中国特有的分布属只有1属。     

MYCORRHIZAE IN HAWAIIAN PTERIDOPHYTES: OCCURRENCE AND EVOLUTIONARY SIGNIFICANCE

Forty-five percent of the Hawaiian pteridophyte flora were examined for mycorrhizae. Vesicular-arbuscular mycorrhizae were present in 66 of the 89 species examined. Nonmycotrophy was significantly higher in Hawaiian pteridophytes than in non-Hawaiian pteridophytes. The highest mycorrhizal colonization occurred in the leptosporangiate families Dicksoniaceae, Dryopteridaceae, and Lindsaeaceae. Mycotrophy was lower in the eusporangiate families. Substrate strongly influenced the presence of mycotrophy. Mycorrhizae were present in 83% of terricolous species, in 86% of epilithic species, in 55% of epiphytic species, and absent from aquatic species. A phylogram of pteridophyte families is constructed incorporating the intensity of mycorrhizal colonization as one criterion for classification.     

Exploring the pteridophyte flora of the Eastern Afromontane biodiversity hotspot

In this review, we explore our current understanding of the fern and lycophyte diversity occurring in the Eastern Afromontane Biodiversity Hotspot (EABH). The review explores the species diversity of this region in the context of the Afromadagascan pteridophyte diversity based on an exhaustive species list assembled in the synopsis of Afromadagascan pteridophytes published by Roux in 2009. The list was updated by incorporating recent progress in our understanding of the taxonomy and phylogeny of these plants. Evidence for a distinct pteridophyte flora occurring in the East African mountain region was discovered using ordination and clustering analyses. This EABH floras shares links to other Afromadagascan pteridophyte floras such as the one in the tropical lowland forests of central and western Africa. These floras share the dominance of species that preferably occur in humid climates whereas other African pteridophyte floras tend to contain a higher proportion of xeric adapted ferns. The phylogenetic composition of the EABH pteridophyte flora was assessed by comparing global versus local proportion of orders, families, and genera. This analysis revealed distinct patterns that are partly caused by the radiation of Blotiella and Triplophyllum besides selective colonization of species pre-adapted to Afromadagascan climates. In situ speciation in the East African tropical mountains may have contributed to the global diversity of widespread genera such as Asplenium and Pteris. In summary, this is the first comprehensive attempt to assess the pteridophyte diversity of the East African mountains providing the framework for future studies on their conservation, ecology, and evolution.