Schmeissneria: An angiosperm from the Early Jurassic

The origin of angiosperms has been a focus of intensive research for a long time. The so-called preCretaceous angiosperms, including Schmeissneria, are usually clouded with doubt. To expel the cloud around the enigmatic Schmeissneria, the syntype and new materials of Schmeissneria collected previously in Germany and recently in China are studied. These materials include female inflorescences and infructescences. The latter are old materials but were under-studied previously. Light microscopy and scanning electron microscope observations indicate that the fruits in these infructescences have in situ seeds enclosed, and that the ovaries are closed before pollination. Thus the plants meet two strict criteria for angiosperms: angiospermy plus angio-ovuly. Placing Schmeissneria in angiosperms will extend the record of angiosperms up to the Early Jurassic, more compatible with many molecular dating conclusions on the age of angiosperms, and demanding a reassessment of the current doctrines on the origin of angiosperms. Although the phylogenetic relationship of Schmeissneria to other angiosperms apparently is still an open question, this study adds to research concerning the origin of angiosperms.     

侏罗纪的施迈斯内果(Schmeissneria)是不是被子植物?

中华施迈斯内果在辽西的发现为被子植物的历史添上了重要的一笔。施迈斯内果是一个具有近170年研究历史的植物化石属。新近发现的中华施迈斯内果中发现了以前没被认识到的新特征(未成熟的雌性器官的中央结构有一个纵向的隔壁和封闭的顶端)。新的特征促使人们重新审视该化石的系统位置。王鑫等人通过仔细分析被子植物的定义、结合新的化石特征认为施迈斯内果代表了一个具有被子植物特征的新类群。按照现有的被子植物定义,施迈斯内果可以归入被子植物。这样一来,被子植物的历史就会被前推到早侏罗世。当然也不排除未来被子植物的定义改变后为施迈斯内果建立一个与被子植物平行的种子植物新类群的可能性。文章简要地介绍了施迈斯内果的由来、特征以及王鑫等人如何判定其为被子植物的逻辑思维过程,以便大家论评。     

睡莲科系统位置评述

形态学研究显示睡莲科Nymphaeaceae具有许多原始性状。睡莲科又被称为“古草本”。最新的分子系统发育研究显示,睡莲科是现存被子植物系统树根部附近最早分异谱系的演化支之一,对研究被子植物(有花植物)的起源与早期进化具有重要价值,但有关睡莲科的范围和系统位置存在争议。被子植物的起源与辐射一直是植物学家关注的热点。本文对该科系统位置的研究历史与现状进行了评述。     

木兰藤科系统位置评述

木兰藤科(Austrobaileyaceae)含1属2种,是系统学上最孤立的科之一。其花粉类似于最古老的被子植物化石之一:晚白垩世的棒纹粉。最新的分子系统发育研究结果表明,木兰藤科是现存被子植物的基部类群之一,其对于被子植物的起源与早期进化的研究具有重要价值。被子植物(有花植物)的起源和辐射一直是植物学家关注的热点。有关木兰藤科的系统位置一直存在争议。本文对该科系统位置的研究历史与现状进行评述。     

Genome duplication and the origin of angiosperms

Despite intensive research, little is known about the origin of the angiosperms and their rise to ecological dominance during the Early Cretaceous. Based on whole-genome analyses of Arabidopsis thaliana, there is compelling evidence that angiosperms underwent two whole-genome duplication events early during their evolutionary history. Recent studies have shown that these events were crucial for the creation of many important developmental and regulatory genes found in extant angiosperm genomes. Here, we argue that these ancient polyploidy events might have also had an important role in the origin and diversification of the angiosperms.     

莲科系统位置评述

莲科含1属1种2亚种,以具有最古老的有活力的种子而著称。形态学研究显示,莲不仅具有双子叶植物特征,而且又具有单子叶植物的某些性状。因此,对研究被子植物(有花植物)的起源与演化以及单子叶植物的起源具有重要价值。被子植物(有花植物)的起源与辐射一直是植物学家关注的热点,有关莲科的系统位置存在争议。该文对该科系统位置的研究历史与现状进行评述。     

Summary statement

The contributions to the symposium on "New evidence for relationships and modern classifications in angiosperms" (held on 22 Aug 1981 during the XIII International Botanical Congress in Sydney) are surveyed, the present situation of this field of research is outlined. A ± monophyletic origin of the angiosperms is supported. Evidence for relationships among major taxa is still fragmentary and yet enormously expanding; synthetic efforts therefore become more and more imperative. A comparison of current classifications reveals similarities and convergencies, but also many areas of divergence and uncertainty. Ways towards a generally acceptable and more natural system of angiosperms are suggested.     

The ABC model and its applicability to basal angiosperms

BACKGROUND: Although the flower is the central feature of the angiosperms, little is known of its origin and subsequent diversification. The ABC model has long been the unifying paradigm for floral developmental genetics, but it is based on phylogenetically derived eudicot models. Synergistic research involving phylogenetics, classical developmental studies, genomics and developmental genetics has afforded valuable new insights into floral evolution in general, and the early flower in particular. SCOPE AND CONCLUSIONS: Genomic studies indicate that basal angiosperms, and by inference the earliest angiosperms, had a rich tool kit of floral genes. Homologues of the ABCE floral organ identity genes are also present in basal angiosperm lineages; however, C-, E- and particularly B-function genes are more broadly expressed in basal lineages. There is no single model of floral organ identity that applies to all angiosperms; there are multiple models that apply depending on the phylogenetic position and floral structure of the group in question. The classic ABC (or ABCE) model may work well for most eudicots. However, modifications are needed for basal eudicots and, the focus of this paper, basal angiosperms. We offer 'fading borders' as a testable hypothesis for the basal-most angiosperms and, by inference, perhaps some of the earliest (now extinct) angiosperms.     

A re-evaluation of Cercidiphyllum-like plant fossils from the British early Tertiary

The occurrence of Cercidiphyllum -like leaves, infructescences, fruits and seeds in the British early Tertiary is reviewed. Material previously described from the London Clay Formation, Lower Headon Beds (upper Barton Formation), Oldhaven Formation, and Woolwich and Reading Formation, are evaluated along with specimens from a new late Palaeocene locality in the Reading Beds. The leaves are assigned to Trochodendroides prestwichii; and infructescences, fruits and seeds, previously assigned to Carpolithus gardneri and Jenkinsella apocynoides , are united as a single species Nyssidium arcticum. The evidence that Trochodendroides and Nyssidium are parts of a single kind of fossil plant is supported, and the reconstructed ' Nyssidium plant' is compared with extant taxa, particularly extant Cercidiphyllum. The fossil plant shows a combination of characters not seen in any living genus, and the structure of the infructescence, as well as other features, precludes placing it within Cercidiphyllum. 'Nyssidium' and Cercidiphyllum are closely related. The morphological and sedimentological evidence indicates that 'Nyssidium' was an important colonizer of a variety of open flood-plain environments during the early Tertiary.     

The effect of unripe fruits on ripe fruit removal by birds in Pistacia terebinthus: flag or handicap?

The shrub Pistacia terebinthus produces crowded infructescences with up to several hundred fruits, which are bright red when unripe and turn green when ripe. Most fruits contain an empty seed and never reach maturity. More ripe fruits were removed by birds from experimental bicolored fruit displays (consisting of infructescences with ten ripe fruits and stripped of unripe fruits, paired with infructescences with only unripe fruits) than from monocolored ones (single infructescences with ten ripe fruits and stripped of unripe fruits). Thus, the presence of unripe fruits seems to increase the conspicuousness or attractiveness of fruit displays to fruit-eating birds. A second experiment compared ripe fruit removal from experimental infructescences having only ripe fruits, with that from control infructescences containing both ripe fruits and natural numbers of unripe fruits, all on P. terebinthus plants. Unlike the first experiment, each bicolored display in this case consisted of a single infructescence with both unripe and ripe fruits. A higher proportion of ripe fruits was removed by birds from infructescences free of unripe fruits. This result suggests that the presence of unripe fruits reduces the accessibility of ripe fruits for fruit-eating birds. This is further supported by field observations of bird foraging behavior.     

生花植物概念简介及其名称的商榷

在上个世纪最后的 2 0年里 ,系统学家应用形态性状对种子植物进行了大量的分支分析。其结果显示灭绝的五柱木属加上灭绝的本内苏铁目及尚存的买麻藤目是被子植物的姊妹群 ,形成一个强支 ,称之为生花植物支。生花植物假说对探讨被子植物起源有着重要影响 ,它激发人们讨论被子植物起源时间可能要提前到三叠纪甚至石炭纪 ,除了支持原有的真花学说外 ,还提出新恩格勒学说。但是 ,近年来对现存种子植物进行分子系统学研究的结果是 :(1)拒绝接受生花植物概念 ;(2 )买麻藤目并不是被子植物的姊妹群而是松柏目的姊妹群 ,甚至网结于松柏类而成为松科的姊妹群。这些结果并不使人惊讶 ,因为对探讨像包含许多灭绝类群的种子植物系统 ,决不可能是仅仅单独应用现代类群资料所能完成的。假如生花植物支是成立的 ,但其名称以AGPB支代替生花植物支可能较为合理。     

水盾草科系统位置评述

水盾草科(Cabombaceae)是双子叶植物。水盾草科包括2属: 水盾草属(Cabomba Aublet.)和莼菜属(Brasenia Schreb.)。形态学研究显示水盾草科具有许多原始性状, 而且在其适应水生环境的过程中经历了性状退化。水盾草科又被称为“古草本”。最新的分子系统发育研究显示, 水盾草科是现存被子植物系统树基部ANITA类群的成员之一。但有关水盾草科的系统位置存在争议。被子植物的起源与早期分化一直是植物学家关注的热点。本文对该科系统位置的研究历史与现状进行评述。     

水盾草科系统位置评述

水盾草科（Cabombaceae）是双子叶植物。水盾草科包括2属：水盾草属（Cabomba Aublet．）和莼菜属（Brasenia Schreb．）。形态学研究显示水盾草科具有许多原始性状,而且在其适应水生环境的过程中经历了性状退化。水盾草科又被称为“古草本”。最新的分子系统发育研究显示,水盾草科是现存被子植物系统树基部ANITA类群的成员之一。但有关水盾草科的系统位置存在争议。被子植物的起源与早期分化一直是植物学家关注的热点。本文对该科系统位置的研究历史与现状进行评述。     

Submarine pollination in the marine angiosperm Zostera marina (Zosteraceae). I. The influence of floral morphology on fluid flow

An understanding of the process of submarine pollination should provide insight into the evolutionary and reproductive ecology of the marine angiosperms (seagrasses). The flow around the reproductive organs of the seagrass Zostera marina L. (Potamogetonales) was, therefore, examined in a flow chamber. The phenological emergence of flowers during (1) pollen capture and (2) pollen release, and by fruit during (3) seed release, led to a reduction in flow rate toward the inflorescence. This change in flow due to floral emergence was associated with a 50% increase in the fluid shear stress [tau = (2.2 _ 0.3) x 10-3 Pa for an immature flower vs. tau = (3.1 _ 0.5) x 10-3 Pa for a receptive flower]. The Reynolds number (Re) and fluid shear stress around inflorescences and infructescences were comparable, indicating a dynamic similarity in the processes of pollen capture and fruit dehiscence [Re = 47 _ 5, tau = (1.6 _ 0.3) x 10-3 Pa for inflorescences; Re = 38 _ 5, tau = (1.3 _ 0.1) x 10-3 Pa for infructescences]. These results indicate that the emergence of reproductive organs leads to changes in fluid shear stress, which will affect the release, transport, and capture of particles including pollen. Theoretical considerations of these observations using aerosol-filtration theory suggest that pollen capture in Z. marina occurs through direct interception of pollen by stigmas.     

The floral genome: an evolutionary history of gene duplication and shifting patterns of gene expression

Through multifaceted genome-scale research involving phylogenomics, targeted gene surveys, and gene expression analyses in diverse basal lineages of angiosperms, our studies provide insights into the most recent common ancestor of all extant flowering plants. MADS-box gene duplications have played an important role in the origin and diversification of angiosperms. Furthermore, early angiosperms possessed a diverse tool kit of floral genes and exhibited developmental 'flexibility', with broader patterns of expression of key floral organ identity genes than are found in eudicots. In particular, homologs of B-function MADS-box genes are more broadly expressed across the floral meristem in basal lineages. These results prompted formulation of the 'fading borders' model, which states that the gradual transitions in floral organ morphology observed in some basal angiosperms (e.g. Amborella) result from a gradient in the level of expression of floral organ identity genes across the developing floral meristem.     

<Emphasis Type="Italic">Schmeissneria</Emphasis>: A missing link to angiosperms?

Background  

The origin of angiosperms has been under debate since the time of Darwin. While there has been much speculation in past decades about pre-Cretaceous angiosperms, including Archaefructus, these reports are controversial. The earliest reliable fossil record of angiosperms remains restricted to the Cretaceous, even though recent molecular phylogenetic studies suggest an origin for angiosperms much earlier than the current fossil record.     

Axial nature of the cupule-bearing organ in Caytoniales

Caytoniales are an important group of seed plants, and the nature of their female reproductive organ may influence interpretations of the seed plant phylogeny and the origin of angiosperms. Although not convincingly demonstrated by clear evidence, cupules on previously described specimens were interpreted as being distichously arranged, implying that the cupule-bearing organ in Caytoniales was a pinnate megasporophyll. Here a female reproductive organ of Paracaytonia hongtaoi gen. et sp. nov. (Caytoniales) is reported from Liaoning, China. The well preserved specimen clearly shows a spiral arrangement of cupules along the reproductive axis, suggesting that the cupule-bearing organ in Caytoniales is not a megasporophyll but a branch. This new information on the axial nature of the cupule-bearing organ in Caytoniales has significant implications on the placement of Caytoniales in the seed plant phylogeny and interpretation of the relationship between Caytoniales and angiosperms.     

Biodiversity and ecology of flower-associated actinomycetes in different flowering stages of <Emphasis Type="Italic">Protea repens</Emphasis>

Actinomycete bacteria have previously been reported from reproductive structures (infructescences) of Protea (sugarbush/suikerbos) species, a niche dominated by fungi in the genera Knoxdaviesia and Sporothrix. It is probable that these taxa have symbiotic interactions, but a lack of knowledge regarding their diversity and general ecology precludes their study. We determined the diversity of actinomycetes within Protea repens inflorescence buds, open inflorescences, young and mature infructescences, and leaf litter surrounding these trees. Since the P. repens habitat is fire-prone, we also considered the potential of these bacteria to recolonise infructescences after fire. Actinomycetes were largely absent from flower buds and inflorescences but were consistently present in young and mature infructescences. Two Streptomyces spp. were the most consistent taxa recovered, one of which was also routinely isolated from leaf litter. Lower colonisation rates were evident in samples from a recently burnt site. One of the most consistent taxa isolated from older trees in the unburnt site was absent from this site. Our findings show that P. repens has a distinct community of actinomycetes dominated by a few species. These communities change over time and infructescence developmental stage, season and the age of the host population. Mature infructescences appear to be important sources of inoculum for some of the actinomycetes, seemingly disrupted by fire. Increased fire frequency limiting maturation of P. repens infructescences could thus impact future actinomycete colonisation in the landscape. Streptomyces spp. are likely to share this niche with the ophiostomatoid fungi, which merits further study regarding their interactions and mode of transfer.     

Reconsiderations on two characters of early angiosperm Archaefructus

Archaefructus is a genus of considerable interest and importance in the study of early angiosperms. Three previously documented species have provided important information about early angiosperms, although some of their characters have been interpreted in various ways. Additional new materials presented in this paper along with the holotype of Archaefructus liaoningensis illustrate branching pattern, fruit arrangement, and seed attachment in Archaefructus. New observations indicate that Archaefructus has ovules/seeds attached to the midrib on the abaxial side of the fruits and a whorled/opposite arrangement for the fruits on the axis. New fossil material of Archaefructus sinensis demonstrates that fruit pairs are inserted on the infructescence axis oppositely. The diagnoses of Archaefructus and Archaefructaceae are emended, and their significance on early angiosperm evolution is discussed.     

Multigene analyses identify the three earliest lineages of extant flowering plants

Flowering plants (angiosperms) are by far the largest, most diverse, and most important group of land plants, with over 250,000 species and a dominating presence in most terrestrial ecosystems. Understanding the origin and early diversification of angiosperms has posed a long-standing botanical challenge [1]. Numerous morphological and molecular systematic studies have attempted to reconstruct the early history of this group, including identifying the root of the angiosperm tree. There is considerable disagreement among these studies, with various groups of putatively basal angiosperms from the subclass Magnoliidae having been placed at the root of the angiosperm tree (reviewed in [2-4]). We investigated the early evolution of angiosperms by conducting combined phylogenetic analyses of five genes that represent all three plant genomes from a broad sampling of angiosperms. Amborella, a monotypic, vessel-less dioecious shrub from New Caledonia, was clearly identified as the first branch of angiosperm evolution, followed by the Nymphaeales (water lillies), and then a clade of woody vines comprising Schisandraceae and Austrobaileyaceae. These findings are remarkably congruent with those from several concurrent molecular studies [5-7] and have important implications for whether or not the first angiosperms were woody and contained vessels, for interpreting the evolution of other key characteristics of basal angiosperms, and for understanding the timing and pattern of angiosperm origin and diversification.