榕树-传粉者共生体系的协同进化与系统学研究进展及展望

 榕树-传粉者共生体系是目前植物与昆虫协同进化研究中的典型模式之一。国内外已经开展了大量的相关研究,从不同方面探讨了其特殊的一一对应的共生关系。榕树-传粉者的专一性互惠共生关系中蕴含了与系统发育有关的多因子协同进化的机理,因此,进行系统发育研究将有助于更好地揭示榕树-传粉者的协同进化历史和理解二者的专一性互惠共生关系。本文简单地介绍了目前榕树及其传粉者共生体系的研究状况之后,论述了榕树-传粉者协同进化的系统发育分子生物学研究成果。同时针对国际上在榕属植物的传统的系统与分类研究中存在的一些分歧及榕树传粉者亚科分类不匹配等问题,回顾了榕属的分类研究进展及其与传粉者的关系。最后,结合我国榕树与传粉者共生体系的研究状况对我国榕属的重新分类和系统发育研究作了展望。     

榕属与榕小蜂科的分类研究进展

简述了榕树与其传粉者的共生体系,讨论了榕属植物与榕小蜂科的分类历史和现状,指出了二者分 类研究中存在的分歧,并对榕属与榕小蜂科的分类不匹配问题进行了评述。结合我国榕属与榕小蜂科分类的 研究现状,展望今后研究的方向及前景。     

榕-传粉榕小蜂间的专一性与协同进化

共生体系的长期维持引发了一系列进化问题, 特别是共生双方的相互适应、协同进化成为进化理论的一大挑战。榕和传粉榕小蜂是目前所知的专性最强的共生体系之一, 两者异常丰富的物种组成以及宿主多样的生活型为以上问题的研究提供了理想的体系。早期认为榕与其传粉小蜂间均为狭义协同进化, 一对一原则在该体系中具有普适性。然而近年来发现越来越多的例外, 特别是宿主转移现象在某些地区、某些榕属类群中的普遍存在, 使榕及其传粉小蜂间严格的物种专一性及协同进化发生在物种水平的观点受到质疑, 最近提出了一个新的协同进化模式来解释榕与其传粉小蜂的对应关系。榕与传粉小蜂间的进化模式说明两者间既有狭义的协同进化, 也有发散协同进化关系, 从而导致它们之间物种专一性不同。目前, 两种协同进化模式在该系统中的相对重要性仍存在很大争议, 不同地区、不同榕属类群中两者的物种专一性程度和产生原因可能有很大差异, 榕与传粉榕小蜂系统的复杂性决定了不能将某些地区和某些类群的结论简单扩展, 有关该体系协同进化主导模式的正确评判有待于对不同地区和不同榕属类群对应传粉小蜂的物种组成、来源方式, 以及共生双方的系统发生关系进行更广泛、更深入的研究。     

榕树及其传粉榕小蜂的系统发育和协同进化研究现状及展望

榕属(Ficus)是有花植物中最大的木本属,全世界有750多种.榕树及其传粉榕小蜂形成的种类专一的互惠共生系统,长期以来被作为研究共生系统比较生物学和协同进化的模式材料.虽然从20世纪90年代才开始对榕树一榕小蜂体系开展分子系统发育研究,但由于这个体系的特殊性和分子技术的快速发展,越来越多的学者开始利用分子学手段来研究榕一蜂共生系统的一系列生物学问题.本文总结了近年来对榕树及其传粉榕小蜂开展的系统发育及协同进化方面的研究,并分析了中国此方面的研究现状,对未来的研究趋势和前景进行了展望.     

传粉细蛾与大戟科植物专性授粉的互惠共生体系

在已知的昆虫与植物所形成的专性授粉互惠共生体系中,榕树—榕小蜂、丝兰—丝兰蛾体系是经典实例,国内外学者已经从不同角度进行了大量的研究,为我们理解植物—传粉者互惠共生体系协同进化的机理和历史积累了宝贵的资料。近些年的研究发现鳞翅目细蛾科头细蛾属昆虫与大戟科植物之间也存在相似的协同进化关系。文章对国内外学者有关传粉细蛾与大戟科植物互惠共生协同进化的研究进行了整理。     

基于mtDNA COⅠ基因对榕属植物上粉蚧的分子鉴定和系统发育分析

【目的】榕属植物是我国华南地区普遍种植的园林植物,本研究旨在明确榕属植物上发生危害的粉蚧种类。【方法】测定榕属植物上发生为害的9种粉蚧的COⅠ基因序列,利用生物信息学软件分析其序列同源性、遗传结构及系统进化关系。【结果】测出38条粉蚧的COⅠ基因序列相似度达到99.25%～100.00%,9种粉蚧的种间遗传距离为0.070～0.173,其中堆蜡粉蚧与菠萝灰粉蚧遗传距离最大,而康氏粉蚧与橘小粉蚧遗传距离最小,说明康氏粉蚧与橘小粉蚧亲缘关系较近。构建的系统发育树中,每种粉蚧与GenB ank数据库下载的已知序列聚在一支,系统发育树结果与形态学鉴定结果一致。【结论】线粒体COⅠ基因序列能够快速准确鉴定榕属植物上的粉蚧,也揭示了榕属植物上粉蚧的遗传结构和系统进化关系,为农林业监测和精准防治粉蚧类害虫提供科学依据。     

大果榕亚组三种榕树传粉榕小蜂核型研究

研究了木瓜榕(Ficus auriculata Loureiro)、苹果榕(F.oligodon Miquel)及未知榕(Ficus sp.)3种榕树传粉榕小蜂的核型。结果表明:3种榕树对应的传粉榕小蜂具有非常相似的核型,染色体数目相同为2n=10,染色体全部为中着丝粒染色体;核型公式2n=2X=10=10M,核型均为Stebbins-1A型。以臂比不对称性系数Dc和相对长度不对称性系数Dt作二维散点图,Ceratosolen emarginatus Mayr(木瓜榕传粉榕小蜂)和C.emarginatus Mayr(苹果榕传粉榕小蜂)具有非常相近的核型不对称系数,核型不对称性大于Ceratosolen sp.(未知榕传粉榕小蜂),显示出两者更近的亲缘关系,而Ceratosolen sp.(未知榕)与前两者亲缘关系较远。此外,比较了Ceratosolen属和Blastophaga属在核型方面的差异,认为染色体数目包括染色体形态能够作为准确区分Ceratosolen属和Blastophaga属的依据,并对Ceratosolen属的核型进化进行了初步探讨。     

中国榕小蜂触角感受器形态特征及进化适应性分析

【目的】榕树与传粉榕小蜂体系是协同进化、专性传粉的经典系统,每种榕树上一般还生活有种类丰富、数目众多的非传粉榕小蜂。在选择压力下,榕小蜂为在榕果内生存产生了明显的适应性形态。触角感受器是昆虫通讯系统的单元,其形态是反映生态和进化适应性的最佳特征之一。本文旨在对中国部分传粉和非传粉榕小蜂的触角感受器的形态多样性和进化适应性进行系统研究。【方法】对来源于海南和云南15种榕树上24种榕小蜂54个型的触角感受器形态进行了扫描电镜观察,基于现有的分子系统发育树,对传粉榕小蜂触角感受器的形态特征进行了性状演化分析,并对榕小蜂的形态特征进行了进化适应性分析。【结果】榕小蜂触角感受器普遍存在雌雄二型现象。雌蜂触角感受器种类有毛状、锥状、板状、刺状、钟形、腔锥状和栓锥状等,数目丰富,并且进果产卵的传粉雌蜂和果外产卵的非传粉雌蜂之间、进果产卵的传粉雌蜂和进果产卵的非传粉雌蜂之间在形态上存在差异。传粉雄蜂和不具有雄性多型现象的非传粉雄蜂触角感受器极为退化,具有雄性多型的非传粉雄蜂触角感受器形态在种内不具有显著差异。性状演化分析表明进化路径相当复杂,可能存在多次独立进化过程。触角感受器的形态类型与其进化适应性相关。【结论】榕小蜂触角感受器类型多样,形态变化丰富,并为适应榕果内的生存而产生了进化适应性特点。雌蜂和雄蜂在榕果内受到了完全不同的选择压力,行使不同的生态功能,从而产生了不同的适应性形态。不同的适应性形态可能与雌蜂不同的产卵行为、雄蜂不同的交配策略具有一定联系。该文是首次对中国榕小蜂触角感受器形态进行系统研究的报道,有助于更好地理解榕小蜂的形态特征、进化路线、行为策略和生态关系。     

榕树(Ficus)在中国的分布及其在协同进化研究上的意义

中国有丰富的榕属资源(共99种),世界上6个亚属的榕树在中国均有不同程度的分布。因此,中国榕属植物的资料对从大地理尺度研究种和种间协同进化具有重要的参考价值。介绍了榕树在世界的分布规律,着重阐述了榕树在中国的分布及中国榕树分布种的研究现状与存在的问题,探讨了中国榕树的花序特点、开花物候及在协同进化中的意义,指出了中国榕树资料在世界研究中的地位和作用。     

论中国的榕树和相关动物的综合研究

榕属（Ficus）植物与其传粉昆虫榕小蜂之间有着一对一的专性共生关系,也与其他相关动物之间存在错综复杂的生态关系,已成为生态学领域的热点之一,榕属与榕小蜂亚科的系统分类、物种多样性、榕树雌雄异株的产生与维持、协同进化、个体生态学等方面的课题值得深入研究。中国生物区系历史悠久、气候带跨幅大、生境多样,有榕属植物98种（其中71种为雌雄异株榕树）,并具有丰富多样性的动物群落,能为进行共生生物学研究提供极好的素材和实验基地。迄今人们对其认识仍十分有限,可望利用中国的生物、地理资源优势,在这一领域里进行有规划的综合研究,促进榕树与相关动物的研究向纵深发展。     

Ficus (Moraceae) and Hymenoptera (Ghalcidoidea): Figs and their pollinators

Attempts are made to meet discrepancies between the botanical classification of Ficus and the zoological classification of its pollinators. They are discussed as Ficus elastica , subgen. Pharmacosycea , subsect. Palaeomorphe , dioecism in Ficus , the Ceratosolen complex, and sect. Conosycea. It seems from the botanical side that the evolution of Ficus into subgenera and sections preceded that of its insects into genera. On the whole there are no grounds for reclassification of Ficua , but leaf structure indicates that some rearrangement of the species of subgen. Urostigma sect. Conosycea may be needed to fit the entomology. Ficus asperiuscula poses a problem in sect. Sycidium.     

Phylogenetic relationships of functionally dioecious FICUS (Moraceae) based on ribosomal DNA sequences and morphology

Figs (Ficus, Moraceae) are either monoecious or gynodioecious depending on the arrangement of unisexual florets within the specialized inflorescence or syconium. The gynodioecious species are functionally dioecious due to the impact of pollinating fig wasps (Hymenoptera: Agaonidae) on the maturation of fig seeds. The evolutionary relationships of functionally dioecious figs (Ficus subg. Ficus) were examined through phylogenetic analyses based on the internal transcribed spacer (ITS) region of nuclear ribosomal DNA and morphology. Forty-six species representing each monoecious subgenus and each section of functionally dioecious subg. Ficus were included in parsimony analyses based on 180 molecular characters and 61 morphological characters that were potentially informative. Separate and combined analyses of molecular and morphological data sets suggested that functionally dioecious figs are not monophyletic and that monoecious subg. Sycomorus is derived within a dioecious clade. The combined analysis indicated one or two origins of functional dioecy in the genus and at least two reversals to monoecy within a functionally dioecious lineage. The exclusion of breeding system and related characters from the analysis also indicated two shifts from monoecy to functional dioecy and two reversals. The associations of pollinating fig wasps were congruent with host fig phylogeny and further supported a revised classification of Ficus.     

Phylogeny, biogeography, and ecology of Ficus section Malvanthera (Moraceae)

We conducted the first molecular phylogenetic study of Ficus section Malvanthera (Moraceae; subgenus Urostigma) based on 32 Malvanthera accessions and seven outgroups representing other sections of Ficus subgenus Urostigma. We used DNA sequences from the nuclear ribosomal internal and external transcribed spacers (ITS and ETS), and the glyceraldehyde-3-phosphate dehydrogenase (G3pdh) region. Phylogenetic analysis using maximum parsimony, maximum likelihood and Bayesian methods recovered a monophyletic section Malvanthera to the exclusion of the rubber fig, Ficus elastica. The results of the phylogenetic analyses do not conform to any previously proposed taxonomic subdivision of the section and characters used for previous classification are homoplasious. Geographic distribution, however, is highly conserved and Melanesian Malvanthera are monophyletic. A new subdivision of section Malvanthera reflecting phylogenetic relationships is presented. Section Malvanthera likely diversified during a period of isolation in Australia and subsequently colonized New Guinea. Two Australian series are consistent with a pattern of dispersal out of rainforest habitat into drier habitats accompanied by a reduction in plant height during the transition from hemi-epiphytic trees to lithophytic trees and shrubs. In contradiction with a previous study of Pleistodontes phylogeny suggesting multiple changes in pollination behaviour, reconstruction of changes in pollination behaviour on Malvanthera, suggests only one or a few gains of active pollination within the section.     

Pigeons as seed predators and dispersers of figs in a Malaysian lowland forest

Seven pigeon species fed on figs from at least 22 Ficus taxa at a lowland forest site in Peninsular Malaysia. Forest pigeons of the genus Treron are fig-eating specialists. Although these are important fig-seed predators, at least some of the population, and pigeons from other genera, potentially disperse Ficus seeds to distant sites.     

Patterns of diversification of Afrotropical Otiteselline fig wasps: phylogenetic study reveals a double radiation across host figs and conservatism of host association

We studied the phylogenetic relationships of Otiteselline fig waSPS associated with Ficus in the Afrotropical region using rDNA sequences. African fig species usually host two species of Otiteselline fig waSPS. Phylogenetic analyses reveal that this pattern of association results from the radiation of two clades of waSPS superimposed on the fig system. Within each clade, wasp species generally cluster according to their host classification. The phylogenies of the two clades are also generally more congruent than expected by chance. Together these results suggest that Otiteselline wasp speciation is largely constrained by the diversification of their hosts. Finally, we show a difference in ovipositor length between the two Otiteselline species coexisting in the same Ficus species, which probably corresponds to ecological differences. The diversification of ecological niches within the fig is probably, with cospeciation, one of the key factors explaining the diversification and maintenance of species of parasites of the fig/pollinator system.     

Phylogenetic relationships of fig wasps pollinating functionally dioecious Ficus based on mitochondrial DNA sequences and morphology

The obligate mutualism between pollinating fig wasps in the family Agaonidae (Hymenoptera: Chalcidoidea) and Ficus species (Moraceae) is often regarded as an example of co-evolution but little is known about the history of the interaction, and understanding the origin of functionally dioecious fig pollination has been especially difficult. The phylogenetic relationships of fig wasps pollinating functionally dioecious Ficus were inferred from mitochondrial cytochrome oxidase gene sequences (mtDNA) and morphology. Separate and combined analyses indicated that the pollinators of functionally dioecious figs are not monophyletic. However, pollinator relationships were generally congruent with host phylogeny and support a revised classification of Ficus. Ancestral changes in pollinator ovipositor length also correlated with changes in fig breeding systems. In particular, the relative elongation of the ovipositor was associated with the repeated loss of functionally dioecious pollination. The concerted evolution of interacting morphologies may bias estimates of phylogeny based on female head characters, but homoplasy is not so strong in other morphological traits. The lesser phylogenetic utility of morphology than of mtDNA is not due to rampant convergence in morphology but rather to the greater number of potentially informative characters in DNA sequence data; patterns of nucleotide substitution also limit the utility of mtDNA findings. Nonetheless, inferring the ancestral associations of fig pollinators from the best-supported phylogeny provided strong evidence of host conservatism in this highly specialized mutualism.     

半附生榕树和非附生榕树的细根性状

榕树作为热带雨林生态系统中的一个关键类群,在维持生物多样性方面发挥着重要作用。本研究以中国科学院西双版纳热带植物园内的榕树专类园区的10种榕属树种(5种半附生,5种非附生)为对象,采用根序法对其细根直径、根长、比根长、比表面积、分枝密度、组织密度、氮含量、碳含量和碳氮比等9个细根功能性状进行了研究,分析了半附生榕树和非附生榕树的细根性状差异;同时结合其原始分布生境,从植物的细根形态功能性状角度阐释其生存策略。结果表明:半附生榕的细根根长、细根直径显著高于非附生榕树(P!0.01),而其比根长和分枝密度显著低于非附生榕树,其余性状之间无显著差异(P>0.05); 10种榕树的细根形态特征与养分特征呈微弱或无相关关系,细根的直径与分枝密度、比根长和比表面积呈显著的负相关。研究认为,不同生活型榕树的根系结构和性状特征差异是对环境的适应表现,分布于山脊干旱环境的半附生榕树细根具有直径较粗、根系长、分枝密度和比根长低的特性,使其具有较好的水分吸收能力,并可能与菌根真菌有更紧密的互利共生关系,从而能适应相对干旱的山脊环境。