Introduction and Notes to R. Dahlgren's System of Classification of the Angiosperms

The present paper aims at introducting Dahlgren’s system of classification of the angiosperms. Phenetic and phylogenetic classifications are discussed. The basic principles and methods used by Dahlgren are explained. Dahlgren’s opinions on some important problems, such as the origin of angiosperms, the flowers of primitive angiosperms, the relation between the dicotyledons and monocotyledons, the origin of the monocotyledons, the treatment of the “Amentiferae” and of the orders of the “Sympetalae”, are all expressed. A brief comparison between Dahlgren’s system and three other current systems, viz. those of Takhtajan, Cronquist and Thorne is also given.     

General aspects of angiosperm evolution and macrosystematics

Taxonomy makes increasing use of significant results from many fields of research including the rapidly developing fields of micro– and macromolecular chemistry, ultrastructure and micromorphology in combination with macromorphology, anatomy, embryology, cytology, paleontology, biological interaction and distribution. Some of these results have contributed to make the current systems of classifications more concordant. Recent studies on Cretaceous fossils are related to present–day angiosperms and their floral types. It is concluded that pleiomerous flowers with helically arranged parts (corresponding to the Magnolia type, though probably less elaborate), on the basis of recent evidence can still be regarded as the probably earliest floral type in angiosperms. But the trimerous flowers must also have appeared very early, at least in the Albian. There is also evidence that the monocotyledons had differentiated as a separate group at that time. Similarities between certain extant monocotyledons and certain dicotyledons, in particular between some Dios–coreales and some Annonales–Aristolochiales, indicate that the monocotyledons had their roots in early Cretaceous pro–Magnoliiflorae. Fossil petaliferous flowers from Cenomanian layers, and later of a variety of flower types, such as the obdiplostemo–nous, petaliferous, epigynous Scandianthus (similar to extant saxifragaceous genera), or flowers with secondarily pleiomerous androecia of the theaceous type are discussed in relation to the distribution of corresponding floral types in extant dicotyledons. The main features of the author's classification of angiosperms are outlined with notes on important, though often neglected, aspects and critical problems. Finally, an updated table of classification down to family rank is presented.     

Reproductive development and nuclear DNA content in angiosperms

Correlates of nuclear DNA content in angiosperms have been noted previously for a range of features, cellular to geographic. A new hypothesis, the correlation between nuclear DNA content and reproductive developmental features (after Cavalier-Smith, Journal of Cell Science 34, 247–268, 1978) is posed and tested here. Of three features tested (megasporogenesis, microsporogenesis, and endosperm development), megasporogenesis alone was shown to be correlated with nuclear DNA amount. The hypothesis was examined in 107 families of angiosperms using nonparametric statistics, and in 53 families of monocotyledons and outgroups using a phylogenetic test of association. A correlation was found between large genomes and successive microsporogenesis for all angiosperms, but not for monocots and dicots analyzed separately, thus underlining the importance of taking into account phylogenetic relationships in such studies. A correlation between cellular endosperm and large genomes in dicotyledons needs to be confirmed in a phylogenetic context. A tendency for deviations from monosporic megasporogenesis to occur in taxa that have a nuclear DNA content of over 9.0 pg/C was demonstrated using both phylogenetic and nonphylogenetic tests. It is hypothesized that cytoskeleton dynamics are affected in reproductive cells, enabling decoupling between nuclear and cytoplasmic cell cycles and leading to variation in reproductive development.     

An updated angiosperm classification

DAHLGREN, C, 1989. An updated angiosperm classification. A new two-dimensional diagram, reflecting the system of classification of the angiosperms, is presented. It combines the dicotyledon diagram in G. Dahlgren (1989) and an adapted monocotyledon diagram after that in Dahlgren (1985) in a single diagram. An updated monocotyledon taxonomy is presented and the classification is appended.     

Angiosperm classification and phylogeny: a criticism

This is a criticism of Dahlgren's recent classification of angiosperms (1980) and his basic diagram of angiosperm phylogeny. The first is shown to include unnatural orders and super-orders; the second is accordingly as questionable. Basic factors for phylogenetic taxonomy in the angiosperms are considered. Leptocauly leads to parallel evolution and specific multiplication. Nomenclatural artifices for taxonomic equivalence are exposed; the name 'pseudon' is suggested for a conjectural taxon.     

Mature Phloem of Perennial Monocotyledons

Structural aspects of differentiating and mature sieve elements of perennial monocotyledons in general and of palms in particular are presented. As in other angiosperms, an immature sieve element undergoes a profound modification of the protoplast during differentiation. Intact, mature sieve elements lack nuclei, possess a parietal cytoplasm, empty lumen and sieve-plate pores that are free of obstructions. Such a structure is in general agreement with the physiological data obtained from exuding inflorescences of woody monocotyledons. Structural evidence and some tracer experiments indicate that sieve elements in perennial monocotyledons are long-lived and apparently function throughout the lifetime of the organ or the plant.     

Angiosperm origin and early stages of seed plant evolution deduced from rRNA sequence comparisons

Summary Complete or partial nucleotide sequences of five different rRNA species, coded by nuclear (18S, 5.8S, and 5S) or chloroplast genomes (5S, 4.5S) from a number of seed plants were determined. Based on the sequence data, the phylogenetic dendrograms were built by two methods, maximum parsimony and compatibility. The topologies of the trees for different rRNA species are not fully congruent, but they share some common features. It may be concluded that both gymnosperms and angiosperms are monophyletic groups. The data obtained suggest that the divergence of all the main groups of extant gymnosperms occurred after the branching off of the angiosperm lineage. As the time of divergence of at least some of these gymnosperm taxa is traceable back to the early Carboniferous, it may be concluded that the genealogical splitting of gymnosperm and angiosperm lineages occurred before this event, at least 360 million years ago, i.e., much earlier than the first angiosperm fossils were dated. Ancestral forms of angiosperms ought to be searched for among Progymnospermopsida. Genealogical relationships among gymnosperm taxa cannot be deduced unambiguously on the basis of rRNA data. The only inference may be that the taxon Gnetopsida is an artificial one, andGnetum andEphedra belong to quite different lineages of gymnosperms. As to the phylogenetic position of the two Angiospermae classes, extant monocotyledons seem to be a paraphyletic group located near the root of the angiosperm branch; it emerged at the earliest stages of angiosperm evolution. We may conclude that either monocotyledonous characters arose independently more than once in different groups of ancient Magnoliales or that monocotyledonous forms rather than dicotyledonous Magnoliales were the earliest angiosperms. Judging by the rRNA trees, Magnoliales are the most ancient group among dicotyledons. The most ancient lineage among monocotyledons leads to modern Liliaceae.     

Nuclear DNA content of monocotyledons and related taxa

Nuclear DNA content of 62 species of angiosperms including 52 monocotyledons and ten dicotyledons has been estimated by flow cytometry using Nicotiana tabacum var. Xanthi as the internal standard. These data, considered together with previous data on diploid species, suggest the following: 1) Most families and orders of monocotyledons have small genomes. Contrary to the general impression that monocotyledons are a group characterized by large genomes, genomes of over 20 pg/2C nucleus occur only in the Liliiflorae, Commelinales, Alismatales, and Araceae. 2) Variation within families ranges from two- to 56-fold, but is two- to fivefold in most families. Thus extraordinary variation in genome size appears to be limited to particular lineages, perhaps owing to some shared feature that facilitates such variation. 3) Endopolyploidy is not observed in the leaves of the species studied, although it has been reported to occur in the roots of several monocotyledons. This suggests that an examination of the basis for this difference between the roots and leaves of monocotyledons may provide clues to the mechanisms that regulate endopolyploidization in these organs.     

A preliminary review of the modern classification systems of the flowering plants

The present paper is divided into three parts: 1. The first part is devoted to the review of the historical origin of the modern classification systems of the flowering plants. Early systems of classification since the Aristotelian time provided a basis for the modern schemes of classification. This paper has reviewed briefly the history of plant systematics, which is divided into three periods: the period of mechanical systems of classification, of natural systems of classification, and of phylogenetic systems of classification. The historical development of the plant systems and the basic idea for three periods is discussed respectively. This paper also considers that the studies of the modern classification system have been entering a new period which primarily aims at overall evolutionary respects. 2. Comparison of the modern classification systems of the flowering plants. Four main modern systems (i. e. A. Cronquist 1979, A. Takhtajan 1980, J. Hutchinson 1973 and A. Engler^,s system as revised by H. Melchior in Engler^,s Syllabus der Pflanzenfamilien 1964) which have greatly influenced the systemstics of plants are compared as to their systematical principles, basic concepts and systematic positions of higher taxa (orders and families) of the flowering plants. The paper is of the opinion that there is still much important work to be done in every field. 3. A review for the modern classification systems of the flowering plants. The paper reviews the modern classification of angiosperms from six aspects: a). The flowers plants are originated from a common ancestral stock; b). The flowers of angiosperms are homogeneous, stamen and carpel are phylletic sporangiophores, all flowers of angiosperms are comparable; c). Monocotyledons are originated from primitive dicotyledons, and represent phylogenetically monophyletic branch; d). The idea advocating the subdivision of Dicotyledoneae into Lignosae and Herbaceae should be rejected as pointed out by many authors; e). The systematical position of Hamamelidae is still a crucial subject for further research; f). The evolutional trends and evaluation of taxonomic characters must be considered in connection with the cor-relation to other characters in question.     

被子植物的一个“多系-多期-多域”新分类系统总览

这篇文章是作者在1998年发表的两篇文章的续篇。在那两篇文章中作者论述了关于被子植物系统发育和演化问题的基本观点：本提出木兰植物门(被子植物)的一个新分类系统纲要,将木兰植物门分为8纲40亚纲202目和572科;其中命名了22新亚纲(Annonidae,Illiciidae,Ceratophyllidae,Lauridae,Calycanthidae, Chloranthidae, Aristolochiidae,Polygonidae,Plumbaginidae,Bromeliidae,Zingiberidae,Juncidae, Poaidae,Paeoniidae,Papaveridae,Trochodendridae,Betulidae,Malvidae,Ericidae,Myrtidae,Rutidae,Geraniidae)和6新目(Degeneriales,Aizoales,Platanales, Dipentodontales,Meliosmales,Balanitales)。并对每个科所包含的属、种数和地理分布作了说明。     

The origin of Darwin’s “abominable mystery”

The phrase “Darwin’s abominable mystery” is frequently used with reference to a range of outstanding questions about the evolution of the plant group today known as the angiosperms. Here, I seek to more fully understand what prompted Darwin to coin the phrase in 1879, and the meaning he attached to it, by surveying the systematics, paleobotanical records, and phylogenetic hypotheses of his time. In the light of this historical research, I argue that Darwin was referring to the origin only of a subset of what are today called angiosperms: a (now obsolete) group equivalent to the “dicotyledons” of the Hooker and Bentham system. To Darwin and his contemporaries, the dicotyledons’ fossil record began abruptly and with great diversity in the Cretaceous, whereas the gymnosperms and monocotyledons were thought to have fossil records dating back to the Carboniferous or beyond. Based on their morphology, the dicotyledons were widely seen by botanists in Darwin’s time (unlike today) as more similar to the gymnosperms than to the monocotyledons. Thus, morphology seemed to point to gymnosperm progenitors of dicotyledons, but this hypothesis made the monocotyledons, given their (at the time) apparently longer fossil record, difficult to place. Darwin had friendly disagreements about the mystery of the dicotyledons’ abrupt appearance in the fossil record with others who thought that their evolution must have been more rapid than his own gradualism would allow. But the mystery may have been made “abominable” to him because it was seen by some contemporary paleobotanists, most notably William Carruthers, the Keeper of Botany at the British Museum, as evidence for divine intervention in the history of life. Subsequent developments in plant systematics and paleobotany after 1879 meant that Darwin’s letter was widely understood to be referring to the abrupt appearance of all angiosperms when it was published in 1903, a meaning that has been attached to it ever since.     

Convergent evolution and adaptive radiation of beetle-pollinated angiosperms

A literature review of 34 families of flowering plants containing at least one species pollinated primarily by beetles is presented. While the majority of species are represented by magnoliids and basal monocotyledons specialized, beetle-pollinated systems have evolved independently in 14 families of eudicotyldons and six families of petaloid monocots. Four, overlapping modes of floral presentation in plants pollinated exclusively by beetles (Bilabiate, Brush, Chamber Blossom and Painted Bowl) are described. Chamber Blossoms and Painted Bowls are the two most common modes. Chamber Blossoms, found in magnoliids, primitive monocotyledons and in some families of woody eudicots, exploit the greatest diversity of beetle pollinators. Painted Bowls are restricted to petaloid monocots and a few families of eudicots dependent primarily on hairy species of Scarabaeidae as pollen vectors. In contrast, generalist flowers pollinated by a combination of beetles and other animals are recorded in 22 families. Generalist systems are more likely to secrete nectar and exploit four beetle families absent in specialist flowers. Centers of diversity for species with specialized, beetle-pollinated systems are distributed through the wet tropics (centers for Brush and Chamber Blossoms) to warm temperate-Mediterranean zones (centers for Painted Bowls and a few Bilabiate flowers). It is unlikely that beetles were the first pollinators of angiosperms but specialized, beetlepollinated flowers must have evolved by the midlate Cretaceous to join pre-existing guilds of beetlepollinated gymnosperms. The floras of Australia and western North America suggest that mutualistic interactions between beetles and flowers has been a continuous and labile trend in angiosperms with novel interactions evolving through the Tertiary.     

单子叶植物科级分类阶元形态性状分支分析

基于93个形态形状,采用13个被子植物基部类群做为外类群,对49个单子叶植物科级分类阶元进行了分支系统学分析。经过简约性分析,得到了1684棵同等最大简约分支树。严格一致树的分支结构图表明：1）古草本类植物和单子叶植物是姐妹群关系;2）具有网状脉的类群,薯蓣科,菝葜科,百部科是单子叶植物的最基部类群。由于性状状态间存在着较多的平行和逆转进化,这在一定程度上影响了系统发育重建的准确性;所选择的性状状态之间的演化很可能是平行的、多次的或者是特化的状态,因此这样复杂的演化关系的探索关键在于找到一些能确切反映其系统演化关系的形态性状。目前很难通过简约化的形态分支分析来解开整个单子叶植物的起源和演化之谜。为了避开对系统学分析造成干扰的误导性状,形态数据结合DNA序列分析很可能是必需的。     

The origin of angiosperms

The claim of monophyletic origin of angiosperms arose from the confusion of phylogenetic and taxonomic concepts. Unpreconceived studies of extant angiosperms point to more than one archetype. Several lines of angiosperms have simultaneously entered the fossil record; the monocotyledons, proto-Hamamelidales, proto-Laurales and “proteophylls” (possibly ancestral to the Rosidae) are recognized among them. Three groups of Mesozoic seed plants — the Caytoniales, Czekanowskiales and Dirhopalostachyaceae — are distinguished as major sources of angiosperm characters (proangiosperms). Other Mesozoic lineages probably also contributed to the angiosperm character pool. Angiospermization is related to Mammalization and other processes involved in development of the Cenozoic lithosphere and biosphere.     

Synopsis of a new “polyphyletic-polychronic-polytopic” system of the angiosperms

This work is continuation of two papers published by the present authors in 1998, in which our basic viewpoints on the phylogeny and evolution of angiosperms were presented. In this paper, a new outline of classification of the Magnoliophyta (angiosperms) is proposed. The Magnoliophyta are divided into 8 classes, 40 subclasses, 202 orders and 572 families. Among them, 22 new subclasses (Annonidae, Illiciidae, Ceratophyllidae, Lauridae, Calycanthidae, Chloranthidae, Aristolochiidae, Polygonidae, Plumbaginidae, Bromeliidae, Zingiberidae, Juncidae, Poaidae, Paeoniidae, Papaveridae, Trochodendridae, Betulidae, Malvidae, Ericidae, Myrtidae, Rutidae, Geraniidae) and 6 new orders (Degeneriales, Aizoales, Platanales, Dipentodontales, Meliosmales, Balanitales) are circumscribed. The number of genera and speciesin the families and each family’s geographical distribution are given.     

New evidence from the ultrastructural and micromorphological fields in angiosperm classification

Based on TEM investigations of some 1850 species and SEM examinations of about 6000 species of the Angiospermae, this is a survey of ultrastructural and micromorphological data (excluding pollen wall characters) which contribute valuable information for the classification of angiosperms. TEM characters predominately relate to phloem features, such as sieve–element plastids and crystalline P–protein, and to those equally present in other tissues, e.g. nuclear protein crystals and dilated ER–cisternae (DC). Of these, the sieve–element plastids with their types and subtypes (S, PI–PVI) and their over 20 forms represent the most thoroughly investigated TEM character. SEM characters mainly relate to epidermal surface features and can be grouped into four categories: (1) Cellular arrangement or cellular pattern; (2) Shape of cells (the “primary sculpture” of a surface); (3) Relief of outer cell walls (“the secondary sculpture” superimposed on the primary sculpture), caused mainly by cuticular striations and superficially visible wall inclusions and wall thickenings; (4) Epicuticular secretions (the “tertiary sculpture” superimposed on the secondary sculpture), i.e. mainly waxes and related substances. Ultrastructural evidence from sieve–element plastids for the classification of Mag–noliiflorae, Caryophylliflorae, Fabiflorae and the unity of the Monocotyledoneae is discussed, while further plastid data are listed for single families (e.g. Buxaceae, Cyrillaceae, Erythroxylaceae, Eucryphiaceae, Gunneraceae, Rhizophoraceae, Vitaceae). Crystalline P–protein dominates in Malviflorae, Violiflorae and Fabiflorae. Nuclear protein crystals are a specific feature of sieve elements of Boraginaceae. DC characterize Capparales s.lat. Micromorphological evidence derived from specific trichomes is presented as an aid to the characterization of Urticales and Loasales, while a detailed analysis of the micromorphology of the seed coat of Cactaceae and Orchidaceae provided new information for the classification of these families at the tribal and generic levels. As a completely new systematic feature for the classification of the Monocotyledoneae first results of micromorphological differences in wax crystalloids and their orientation patterns are presented: The Liliiflorae s. str. are clearly separated against the Zingiberiflorae–Commeliniflorae (incl. Velloziales, Bromeliales, Typhales) and Areciflorae, both characterized by two mutually exclusive and very specific wax types and delimited against taxa with unspecific waxes in the rest of the monocotyledons and all dicotyledons.     

花同源异型MADS-box 基因在被子植物中的功能保守性和多样性

MADS-box基因家族成员作为转录调控因子在被子植物花发育调控中发挥关键作用。本文以模式植物拟南芥(Arabidopsis thaliana) 和水稻 (Oryza sativa)为例, 综述了近10年来对被子植物(又称有花植物)两大主要类群——核心真 双子叶植物和单子叶植物花同源异型MADS-box基因的研究成果, 分析MADS-box基因在被子植物中的功能保守性和多样性,同时探讨双子叶植物花发育的ABCDE模型在多大程度上适用于单子叶植物。     

The distribution of ester-linked ferulic acid in the cell walls of angiosperms

Ester-linked ferulic acid occurs in the cell walls of two major groups of angiosperms, the commelinid monocotyledons and the ‘core’ Caryophyllales, at concentrations >3.5 mg g⁻¹ cell walls, and has been detected in primary cell walls by its autofluorescence using ultraviolet fluorescence microscopy. Both of these groups are resolved as monophyletic clades in phylogenetic trees constructed using gene sequences. In the primary cell walls of the commelinid monocotyledons, including the grasses (family Poaceae), the ferulic acid is ester-linked to the non-cellulosic polysaccharide glucuronoarabinoxylan. In contrast, in the ‘core’ Caryophyllales, the ferulic acid is ester-linked to the side chain arabinans and galactans of the pectic polysaccharide rhamnogalacturonan-1, at least in the family Amaranthaceae. In the walls of both angiosperm groups, a range of dehydrodiferulates have also been found. These are formed oxidatively via radical coupling and result in the cross linking of the polysaccharides to which they are attached. Much lower concentrations of ester-linked ferulic acid have been found in cell walls isolated from other angiosperms, although physiological stress conditions may cause increases in these concentrations. The polysaccharides to which the ferulic acid is attached to in the cell walls of these other angiosperms is unknown.     

Molecular phylogenies in angiosperm evolution

We have cloned and sequenced cDNAs for the glyceraldehyde-3-phosphate dehydrogenase of glycolysis, gapC, from a bryophyte, a gymnosperm, and three angiosperms. Phylogenetic analyses are presented for these data in the context of other gapC sequences and in parallel with published nucleotide sequences for the chloroplast encoded gene for the large subunit of ribulose-1,5-bisphosphate carboxylase/oxygenase (rbcL). Relative-rate tests were performed for these genes in order to assess variation in substitution rate for coding regions, along individual plant lineages studied. The results of both gene analyses suggest that the deepest dichotomy within the angiosperms separates not magnoliids from remaining angiosperms, but monocotyledons from dicotyledons, in sharp contrast to prediction from the Euanthial theory for angiosperm evolution. Furthermore, these chloroplast and nuclear sequence data taken together suggest that the separation of monocotyledonous and dicotyledonous lineages took place in late Carboniferous times [approximately 300 Myr before the present (Mybp)]. This date would exceed but be compatible with the late-Triassic (approximately 220 Mybp) occurrence of fossil reproductive structures of the primitive angiosperm Sanmiguelia lewisii.      

Occurrence of shikimic and quinic acids in angiosperms

Leaves of 83 angiosperms have been made surveyed for quinic and shikimic acids. The quinic acid content was higher in woody dicotyledons than in herbaceous dicotyledons or in the monocotyledons, substantiating the view that its presence may be correlated with the lignification in plants.