A Phylogeny of Euphorbieae Subtribe Euphorbiinae (Euphorbiaceae) Based on Molecular Data

Phylogenetic relationships within Euphorbiinae were inferred from our analysis of the 3′; end of the chloroplast gene ndhF. A sampling of that subtribe covered 88 species; 3 closely related species from the subtribes Anthosteminae and Neoguillauminiinae and the tiribe Hippomaneae were included as outgroups. A phylogenetic assessment was carried out using the parsimony approach. The relationships revealed via these ndhF data supported the monophyly of subg.Esula, subg.Chamaesyce, subg.Euphorbia, and subg.Lacanthis. However, the polyphyly of subg.Agaloma, subg.Lyciopsis, and subg.Eremophyton also was strongly suggested. The African succulent Euphorbiinae can be divided into primarily two independent groups: 1) spiny succulents, which form a strongly supported clade with three subclades (subg.Euphorbia, subg.Lacanthis, andMonadenium+Synadenium); and 2) non-spiny succulents, which consist of sect.Meleuphorbia, sect.Medusae, sect.Anthacantha, sect.Trichadenia, sect.Pseudeuphorbium, sect.Treisia, and sect.Pseudacalypha. In the ndhF tree, the subg.Esula clade is placed as a sister to the rest of the Euphorbiinae. Thus, the origin of theEuphorbia s.I. should be sought within the herbaceous species of subg.Esula. The core North American endemicEuphorbia groups --Agaloma, Chamaesyce, andPoinsettia — are monophyletic and independent of the South American subg.Agaloma. Instead, they are derived from the AfricanEuphorbia subg.Lyciopsis andEremophyton. The Eurasian subg.Esula clade forms two subclades, which are concordant to sect.Esula and sect.Tithymalus.     

Giants and dwarfs: molecular phylogenies reveal multiple origins of annual spurges within Euphorbia subg. Esula

Euphorbia (Euphorbiaceae) comprises over 2150 species and is thus the second-largest genus of flowering plants. In Europe, it is represented by more than 100 species with highest diversity in the Mediterranean area; the majority of taxa belong to subgenus Esula Pers., including about 500 taxa. The few available phylogenetic studies yielded contrasting results regarding the monophyly of subg. Esula, and the phylogenetic relationships among its constituents remain poorly understood. We have sampled DNA sequences from the nuclear ribosomal internal transcribed spacer (ITS) and the plastid trnT-trnF region from about 100, predominantly European taxa of subg. Esula in order to infer its phylogenetic history. The plastid data support monophyly of subg. Esula whereas the ITS phylogeny, which is generally less resolved, is indecisive in this respect. Although some major clades have partly incongruent positions in the ITS and plastid phylogenies, the taxonomic content of the major terminal clades is congruent in both trees. As traditional sectional delimitations are largely not corroborated, an improved classification is proposed. Character state reconstruction illustrates that the annual life form developed independently several times in different clades of subgenus Esula from perennial ancestors, and that several morphological traits used in previous classifications of Euphorbia developed in parallel in different lineages.     

Seed morphology ofEuphorbia sectionTithymalopsis (Euphorbiaceae) and related species

Light and scanning microscopies were used to evaluate the sectional boundary of 17 species included by various authors inEuphorbia sectTithymalopsis. Based on seed shape, surface ornamentation, and mucilage formation, five seed morphological types were recognized in North American sectionTithymalopsis and related species. These results did not provide added support for recent treatment of the sectTithymalopsis s.s. However, a close relationship was suggested among species of theTithymalopsis subsectionCorollatae. In addition, including species from subsectionsInnocuae, Ipecacuanhae, andSphaerorhizae within sectTithymalopsis could not be justified, based on seed morphological data.     

Seed and gland morphology in Euphorbia (Euphorbiaceae) with focus on their systematic and phylogenetic importance, a case study in Iranian highlands

A comprehensive study based on gland and seed micromorphology in Euphorbia (Euphorbiaceae) for species distributed in Iranian highlands is presented. A total of 86 species were studied. The gland structure was examined by direct field observations. Taxonomically important characters of glands were observed and measured: size, texture, shape, color, and horns. For species out of Iran herbarium materials were studied. Seed characteristics were examined using scanning electron microscopy (SEM) as well as dissecting light microscopy. Significant features are: seed size, seed shape, presence of caruncle, shape of caruncle, and seed surface ornamentation. A phylogenetic study using Maximum Parsimony (MP) and Bayesian Inference (BI) was performed based on sequences of nuclear DNA internal transcribed spacers (ITS) for selected species representing the main clades known in Euphorbia and with special focus on the species distributed in Iranian highlands. ITS sequences for 20 accessions representing 19 species are provided for the first time, and 48 accessions of 47 species were used from GenBank. The topologies of both analyses were congruent. The results indicate: (1) four main clades with high supports in subgen. Esula which are appropriate to be recognized at sectional rank. (2) E. larica is nested within clade A including few members of subgen. Rhizanthium and is closely related to sect. Balsamis, which is suggested here to be transferred from subgen. Esula into subgen. Rhizanthium. (3) E. osyridea of the monotypic subsect. Osyrideae is closely related to E. buhsei and to the members of sect. Esula. Tracing morphological characters on the phylogenetic tree shows that several morphological characters, such as seed ornamentation applied in previous subgeneric classification of the subgen. Esula, are homoplasious, but the gland structure and capsule surface characters are more reliable for classification purposes.     

ITS sequences and speciation on far easternIndigofera (Leguminosae)

The internal transcribed spacer (ITS) regions of 18–26S nuclear ribosomal DNA was sequenced to address phylogenetic relationships and to measure the extent of differentiation among six species of the Far EasternIndigofera. ITS 1 had 230–240 base pairs (bp) long while ITS 2 had 211–213 bp long. The 5.8S rRNA coding region was 161 bp long. Sequence divergence calculated by Kimura's two parameter method between species ranged from 0.00 to 13.49%. A single most parsimonious tree was produced from 77 variable nucleotide sites, which had a consistency index of 0.97 and a retention index of 0.83. ITS sequence data suggested that the continental species ofI. kirilowii (2n=16) is diverged from the common ancestor of other species at first, and then the island species ofI. decora (2n=48) andI. venulosa and the Korean Peninsular species ofI. grandiflora (2n=16) andI. koreana (2n=32) are diverged from the ancestor. The molecular data supports that the speciation in the Far EasternIndigofera occurred with polyploidization from continental ancestor to peripheral peninsular and island species.     

Seed and capsule morphology of Iranian perennial species of Euphorbia (Euphorbiaceae) and its phylogenetic application

Of the 480 species of Euphorbia subgenus Esula, c. 290 occur in the Mediterranean and Irano‐Turanian regions. Turkey and Iran are the most species‐rich countries in Asia with 83 and 74 species, respectively. Following our previous paper on annual species of Iranian Euphorbia, we studied the quantitative and qualitative macro‐ and micromorphological traits of seeds and capsules of 47 perennial species, including E . ferdowsiana sp. nov. , E . sulphurea sp. nov. and E. glareosa, as a first report from Iran. A key for all Iranian perennial Euphorbia spp. based on seed and capsule morphology is provided. The phylogenetic relationships of Iranian species based on internal transcribed spacer (ITS) nuclear and ndhF plastid regions are updated and used for the characterization of the synapomorphies of each clade. Capsule shape, seed shape, seed surface and shape of the caruncle have been found to be homoplastic, whereas the presence or absence of granulate elements on seed surfaces represents a phylogenetically important trait for section delimitation. The capsule surface is synapomorphic for several sections, including Helioscopia (tuberculate‐verrucose), Myrsiniteae (vesiculate) and Esula (granulate), and seed shape is synapomorphic for sections Helioscopia (ellipsoidal), Myrsiniteae (ovoid‐quadrangular) and Herpetorrhizae (pseudo‐hexahedral). Reversals have also taken place in some features, including capsule surface (E. mazandaranica, E. altissima) and seed shape (E. densa, E. aleppica). It seems that ecarunculate seeds are plesiomorphic in sections Helioscopia (E. eriophora) and Herpetorrhizae (E. consanguinea and E. turczaninowii). © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 00 , 000–000.     

Taxonomic revision and phytogeographic studies in Euphorbia (Euphorbiaceae) in the Khorassan provinces of Iran

This study is focused on the genus Euphorbia L. in a part of northeast Iran, viz. the three Khorassan provinces. Since there are many taxa of Euphorbia in Iran which are used in different industries and have significant effects on human and non‐human life it is important to revise their taxonomy. With about 90 species, following Turkey with 91 species, Iran is the second richest country for Euphorbia in Asia. Of these, 30 species are distributed in the Khorassan provinces. This is the first comprehensive work on the genus in this region. According to ‘Flora Iranica’, there are 17 species of Euphorbia in northeast Iran, while according to our results, there are 30 species of Euphorbia in the Khorassan provinces alone. In addition to various new taxonomic and biogeographic results, a new species, viz. E. chamanbidensis, is described. Euphorbia chamanbidensis is closely related to E. aucheri, but seed micro‐morphological characters differentiate them. Two identification keys to the Euphorbia species of the studied area are provided, one based on macro‐morphological characters and another based on seed micro‐morphological characters. Phytogeographic analysis and distribution maps for all species are also presented.     

凤仙花属棒凤仙亚属(凤仙花科)植物花粉形态的分类学意义

棒凤仙亚属代表着凤仙花属最先分化出的一支,以多年生草本,多于5朵花的总状花序,侧生萼片4枚,3沟花粉,心皮四室且每室具1枚种子,果实棒锤状,种子椭圆形以及三沟花粉其极面观为三角形而区别于其它种类。该亚属主要分布于中国南部,少数种类辐射分布至华中和中南半岛地区。已有报道证实花粉的形态特征对凤仙花属植物的属下分类具有重要意义,但对棒凤仙亚属的种类少有涉及。为了进一步探讨该亚属花粉的形态特征及其分类学价值,该研究应用扫描电镜技术对国产棒凤仙亚属植物的14种以及凤仙花亚属的窄萼凤仙花的花粉进行了观察。结果表明：棒凤仙亚属的花粉以单粒存在,具3条萌发沟,辐射对称,极面观为三角形或三角状圆形,花粉粒外壁具均匀的网状纹饰,网脊平滑或具波状边缘,网眼中具不同密度的颗粒状突起。棒凤仙亚属的花粉形态与水角属的花粉形态相似,表现出了其原始性;其花粉形态特征与花部形态性状等相关性差,对该亚属的组级分类意义较小,但对于该亚属的种间界定具有重要意义;而其花粉的极面观形状、赤道面观形状以及网状纹饰等变异相对稳定,对于界定该亚属具有重要的分类学价值。     

陕西分布10种大戟属植物茎叶形态结构比较

利用石蜡切片法、叶表皮装片法对陕西分布10种大戟属植物茎叶的形态结构进行了比较观察。结果表明:(1)10种植物叶的上下表皮细胞形态、气孔分布及大小等指标存在明显差异,茎的结构在各种之间也存在一定差异。(2)用Statistica 6.0分析软件对所测的性状特征值进行了主成分分析和聚类分析,结果发现,在10种植物中,甘遂与其他种的亲缘关系最远,南大戟与华北大戟的亲缘关系较近,斑地锦与地锦草亲缘关系最近。该研究结果为大戟属植物的分类研究奠定了理论依据。     

SYSTEMATIC SIGNIFICANCE OF POLLEN NUCLEAR NUMBER IN EUPHORBIACEAE,TRIBE EUPHORBIEAE

Pollen nuclear number is determined in 139 species of 5 genera in the Euphorbieae, subtribe Euphorbiinae. The 111 new determinations are tabulated along with previous reports, and the results indicate that the distribution of binucleate (II) and trinucleate (III) pollen is strongly associated with the taxonomic groupings within the Euphorbieae. Although binucleate pollen is probably primitive within the tribe Euphorbieae, as suggested by the nuclear condition in Neoguillauminia, the situation in Euphorbia still requires further elucidation. Within Euphorbia, the morphologically most primitive species studied have III pollen despite the fact that II pollen is presumably the original condition for the subtribe Euphorbiinae. In Euphorbia, II pollen only is reported from nine sections and III pollen only from ten sections, while in four sections (Esula, Goniostema, Aphyllis, and Deuterocalli) both II and III pollen have been found. The New World species of Euphorbia nearly all have III pollen, whereas the vast majority of the African succulents have II pollen. The genera of New World origin, Chamaesyce and Pedilanthus, have III pollen, while the African genera Monadenium and Synadenium have II pollen. Independent derivations of III pollen from II pollen appear to have occurred in sections Goniostema, Aphyllis, and Deuterocalli (all of subg. Euphorbia). There is no evidence that reversals from III to II pollen have occurred.     

Phylogenetic systematics of New WorldIpomoea (Convolvulaceae) based on chloroplast DNA restriction site variation

Chloroplast DNA restriction site variation was studied in 31 New World species ofIpomoea, representing a majority of the New World sections and series within the genus. Using 14 endonucleases, a total of 124 phylogenetically informative restriction sites was detected. Dollo parsimony, Wagner parsimony, and bootstrap methods were employed to construct phylogenetic trees and evaluate confidence intervals of monophyletic groups. With a few exceptions, groups circumscribed on the basis of morphological variation are in agreement with groupings based on restriction site variation. Relationships between subgeneric groupings, however, disagree substantially with those proposed in the past. Although conflicting hypotheses for some intersectional relationships are not presently resolvable, cpDNA restriction site analyses propose the following refinements of existing classification schemes.Ipomoea ser.Setosae is divided into distantly related groups, as is sect.Pharbitis. SeriesTyrianthinae, a proposed segregate of sect.Pharbitis, is associated with sect.Calonyction and the Tricolor complex (subg.Quamoclit).Ipomoea sect.Batatas is segregated from other herbaceous groups of the heterogeneous subg.Quamoclit sensu lato and aligned as a derivative ofI. setosa, subg.Eriospermum. To test for homology of key characters weighted in traditional schemes, morphological features were studied with respect to their distribution on lineages defined by restriction site data. Characters such as setose sepals, foliose-pubescent sepals, and erect growth habit, among others, are interpreted as having multiple origins, while 3-locular ovaries, 4-locular ovaries, and long-haired seeds have evolved only once.     

ITS sequences from nuclear rDNA suggest unexpected phylogenetic relationships between Euro-Mediterranean, East Asiatic and North American taxa ofQuercus (Fagaceae)

Nucleotide sequences from the internal transcribed spacer (ITS) regions of the 18S–26S nuclear ribosomal DNA have been studied from ten species ofQuercus (representing four subgenera),Castanea sativa andFagus sylvatica, as a preliminary molecular contribution to the still poorly understood systematics and evolution ofFagaceae. The resulting matrix has been used to calculate pair-wise sequence divergence indices and to construct a maximum parsimony tree forQuercus coding indels as a fifth state. Divergence is greater forQuercus vs.Fagus than forQuercus vs.Castanea. The tree for theQuercus taxa studied reveals two clearly divergent clades. In clade I the evergreen W MediterraneanQ. suber appears in a basal position as sister to more distal deciduous taxa, i.e. the E MediterraneanQ. macrolepis and the E AsiaticQ. acutissima (all formerly united as different sections under the apparently polyphyletic subg.Cerris), andQ. rubra (a representative of the N American subg.Erythrobalanus), forming a pair withQ. acutissima. In clade II the evergreen southeastern N AmericanQ. virginiana is basal and sister to the remaining three branches, i.e. a pair of evergreen Mediterranean taxa withQ. ilex andQ. coccifera (subg.Sclerophyllodrys), the deciduous but otherwise plesiomorphic SE European/SW AsiaticQ. cerris (type species of subg.Cerris), and the related but more apomorphic European pairQ. petraea andQ. robur (subg.Quercus). These results partly conflict with current taxonomic classification but are supported by some anatomical and morphological characters. They document polyphyletic lines from evergreen to deciduous taxa and suggest Tertiary transcontinental connections within the genus.     

四种栎属青冈亚属植物叶绿体基因组特征及系统发育研究

栎属青冈亚属植物的系统发育地位长期存在着争议，部分种的种间关系不明确。为揭示宁冈青冈(Quercus ningangensis)、曼青冈(Q.oxyodon)、毛曼青冈(Q.gambleana)、竹叶青冈(Q.neglecta)的叶绿体基因组特征及系统发育关系，该研究选择以上4种栎属青冈亚属植物的成熟叶片进行二代测序，对其叶绿体基因组结构和特征进行分析，并结合相关类群进行系统发育研究。结果表明：(1)宁冈青冈、曼青冈、毛曼青冈、竹叶青冈的叶绿体基因组序列长度分别为160 906、160 883、160 832、160 784 bp,均编码133个基因，包括88个蛋白质编码基因、37个tRNA基因、8个rRNA基因。(2)4种栎属青冈亚属植物偏好以A/T结尾的密码子，质体基因组变异区域主要存在于非编码序列。(3)通过IR边界分析得出，4种栎属青冈亚属植物存在ycf1假基因且在IRb/SSC区域发生扩张。(4)系统发育分析显示，在壳斗科中，水青冈属(Fagus)和轮叶三棱栎属(Trigonobalanus)较早分化出来，栎亚属(subg.Quercus)未形成一个单系群，叶绿体基因组建树结...     

Phenetic investigation of non-nodulating African species ofAcacia (Leguminosae) using morphological and molecular markers

Morphological and RAPD markers were used to assess the relationships among nodulating and non-nodulating species of AfricanAcacia. Non-nodulating species of AfricanAcacia are only found within subg.Aculeiferum sect.Monacanthea. African species of sect.Monacanthea examined were found to form a group distinct from the other African species examined on a morphological and molecular basis. All lack the ability to nodulate, suggesting that non-nodulation may be used as a taxonomic tool. The species of sect.Aculeiferum were separated by RAPD and morphological analysis into two groups depending on whether they were armed with prickles in pairs and/or prickles in threes, or solitary. A third group of species was identified within sect.Acacia: further subdivision of this group was achieved into subsectt.Pluriseriae andUniseriae. The position ofA. albida relative to other AfricanAcacia species was found to be distinct but not totally independent of the genus. The partitioning and distribution of the genetic variability within the genus is further elucidated by the RAPD analysis of populations ofAcacia species. A population analysis ofA. polyacantha demonstrated geographical and site-specific variation.     

Corydalis hirtipes B.U. Oh et J.G. Kim: A new species ofCorydalis sectCorydalis (Fumariaceae) from Korea

Here we describe a new species ofCorydalis sectCorydalis (Fumariaceae), namely C.hirtipes B.U. Oh et J.G. Kim. This species, from the central part of Korea, is clearly distinguished from other species of sectCorydalis by its simple, unicellular hairs that are densely distributed on the stems, peduncles, and petioles. It is also distinctive from its close relative C.albipetala in having ternate leaves, lobate or parted leaflets, dentate or cleft bracts, and broad, fusiform fruits.     

Phylogenetic Relationships Within the Genus Potorous (Marsupialia: Potoroidae) Based on Allozyme Electrophoresis and Sequence Analysis of the Cytochrome b Gene

Controversy over the taxonomic designations within the genus Potorous has highlighted problems associated with using only morphological data. The recent rediscovery of Gilbert's Potoroo has allowed us to reexamine the relationship of this species to other extant potoroos. Phylogenetic relationships and estimated divergence times are presented, based on electrophoretic and cytochrome b sequence data. There was considerable concordance between the data sets. We conclude that Gilbert's Potoroo is a separate species and should be referred to by its original name, Potorous gilbertii (Gould 1841). Estimates for potoroid divergences are in line with those of Flannery (1989).     

The taxonomic relationships within the genus Excoecaria L. (Euphorbiaceae) based on leaf morphology and rDNA sequence data

The tropical Indo-Pacific genus Excoecaria L. (Euphorbiaceae) has several closely related species in Australia whose taxonomic relationships are unclear. The most widely reported species in Australia is the mangrove species Excoecaria agallocha L. (type species), whose taxonomic and geographic limits are difficult to define from its closely related species or sub-species. Two additional taxa have also been described but not clearly differentiated from the type species: Excoecaria dallachyana Baillon and Excoecaria ovalis Endl. This project aimed to determine the taxonomic relationships of the Australian Excoecaria species using both leaf morphological data and DNA sequence data from the internal transcribed spacer (ITS) region of ribosomal genes. The nucleotide differences in the examined ITS1 region show that E. agallocha from eastern Australia and E. ovalis from Western Australia respectively, are genetically uniform within species but differ from each other consistently, thus supporting species status. The leaf morphological data also support this view: single factor analysis of variance consistently separated E. ovalis from E. agallocha on the basis of leaf width, leaf length and length of petiole. In contrast, E. ovalis from the Gulf of Carpentaria differs only slightly from E. ovalis in Western Australia, but no evidence was found to suggest any leaf morphological differentiation within this species. The analysis also suggests that E. dallachyana is not closely related to either mangrove species E. agallocha or E. ovalis, despite superficial morphological similarities.     

Chloroplast DNA diversity in the genusRubus (Rosaceae) revealed by Southern hybridization

The variability in chloroplast DNA type of 20Rubus genotypes was examined by Southern hybridization. DNA extracted from theRubus accessions was digested with two restriction enzymes (EcoRI and EcoRV) and heterologous chloroplast DNA sequences from barley and pea were used as probes to detectRubus chloroplast DNA sequences on Southern blots ofRubus total DNA. Restriction fragment length polymorphism was detected and a total of 92 restriction fragments were generated by the probe/enzyme combinations examined. Cladistic principles based on the parsimony assumption were used to assemble a phylogenetic tree based on chloroplast restriction fragment length data. The phylogenetic tree grouped the taxonomically defined species and is in general agreement with information based on morphological criteria. However, the Japanese red raspberryR. illecebrosus was shown to have diverged considerably in terms of evolutionary time from other species in subg.Idaeobatus. Furthermore, the molecular approach provides a quantitative estimate of the relationship between species that is difficult to obtain from morphological data. In order to complement the chloroplast DNA information a ribosomal DNA probe was also included in the analysis and provided further information on the phylogenetic relationships withinRubus.