COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE VI. POMOIDEAE: ERIOBOTRYA,HETEROMELES, PHOTINIA,POURTHIAEA, RAPHIOLEPIS,STRANVAESIA

The pomoid genera, Eriobotrya, Photinia, Pourthiaea, Raphiolepis, Stranvaesia, and Heteromeles, have compound inflorescences and biovulate carpels which become papery at maturity. The carpels of all of these except Heteromeles are fused with one another. There are open sutures in the carpels of Heteromeles, Photinia, Pourthiaea, and Raphiolepis, and in these four genera the extent of fusion of the ovular bundle with the wing bundle is related directly to the state of tegumentary fusion and to the extent of fusion of the carpel with the floral cup. In those species of Eriobotrya and Stranvaesia with closed sutures the integuments tend to be fused, as do the ovular and wing bundles, and the carpels are adnate with the floral cup for a considerable distance; in species with open sutures the integuments tend to be free, the ovular and wing bundles tend to be separate, and the extent of fusion of carpel with floral cup tends to be shorter. In genera with connate carpels the wing bundles of adjoining carpels may also be fused. The greatest extent of fusion occurs in Eriobotrya and Raphiolepis, in which there may also be attenuation and disappearance of the wing bundles above the region of ovular insertion and even reduction and disappearance of the carpellary margin.     

Floral morphology of Maloideae (Rosaceae) and its systematic relevance

Flowers of 169 species of Rosaceae subfamily Maloideae, which were chosen to represent the taxonomic and geographic diversity of the group, were studied to ascertain their morphological variation and its systematic relevance. We describe and illustrate variation in size, indumentum, color, and macroscopic structural features. Most maloid species have syncarpous flowers with two to five carpels in which the ovary is at least three-quarters inferior, whereas species of other Rosaceae subfamilies have apocarpous or unicarpellate flowers with superior ovaries. However, maloid flowers show significant variation in the degree of carpel connation and of ovary adnation to the hypanthium. Cotoneaster, Heteromeles, and Pyracantha are completely apocarpous, and Dichotomanthes is perigynous with a completely superior ovary. Thus, no one floral character is sufficient to separate the Maloideae from other subfamilies of Rosaceae. Differences among their flowers support our recognition of Malus, Pyrus, and Sorbus as separate genera. Further, we argue for removal of Docyniopsis and Eriolobus from Malus, division of Sorbus into several genera, and union of Aronia, Photinia, and Stranvaesia. No floral characters support the traditional dichotomy of the subfamily into tribes Crataegeae and Sorbeae.     

猴樟叶绿体基因组特征分析

猴樟(Cinnamomum bodinieri)枝叶含有丰富的精油,是重要的园林绿化树种和经济树种,但目前有关猴樟基因组学的研究报道不多。为揭示猴樟叶绿体基因组特征及系统发育关系,该文基于高通量测序平台进行测序,从头组装了完整的猴樟叶绿体基因组,并对其基因组结构、基因构成及序列重复、密码子使用偏好性以及系统发育进行分析,结合樟亚科主要属物种叶绿体基因组数据构建系统发育树。结果表明:(1)猴樟叶绿体基因组全长152 727 bp,包括一对20 132 bp的反向重复(IRs)区、93 605 bp的大单拷贝(LSC)区和18 858 bp的小单拷贝(SSC)区,总GC含量为39.13%。(2)该基因组共编码127个基因,包括83个蛋白质编码基因(PCGs)、36个转运RNA基因(tRNAs)和8个核糖体RNA基因(rRNAs); 共鉴定出92个SSR位点,其中大部分是A/T组成的单核苷酸重复序列; 密码子适应指数(CAI)为0.166,有效密码子数(ENc)为54.68; 猴樟与近缘种的叶绿体基因组主要在IR区和2个SC区边界上存在一定的差异。(3)24种樟亚科植物的系统发育树显示,猴樟与樟树亲缘关系最近,同时支持了樟属-甜樟属分支(Cinnamomum-Ocotea Clade)、月桂属-新木姜子属分支(Laurus-Neolitsea Clade)、润楠属-鳄梨属分支(Machilus-Persea Clade)的建立。该研究丰富了猴樟遗传资源信息,进一步确定了樟亚科主要属的系统发育地位。     

Testing and using complete plastomes and ribosomal DNA sequences as the next generation DNA barcodes in Panax (Araliaceae)

Complete plastid genome (plastome) sequences and nuclear ribosomal DNA (nrDNA) regions have been proposed as candidates for the next generation of DNA barcodes for plant species discrimination. However, the efficacy of this approach still lacks comprehensive evaluation. We carried out a case study in the economically important but phylogenetically and taxonomically difficult genus Panax (Araliaceae). We generated a large data set of plastomes and nrDNA sequences from multiple accessions per species. Our data improved the phylogenetic resolution and levels of species discrimination in Panax, compared to any previous studies using standard DNA barcodes. This provides new insights into the speciation, lineage diversification and biogeography of the genus. However, both plastome and nrDNA failed to completely resolve the phylogenetic relationships in the Panax bipinnatifidus species complex, and only half of the species within it were recovered as monophyletic units. The results suggest that complete plastome and ribosomal DNA sequences can substantially increase species discriminatory power in plants, but they are not powerful enough to fully resolve phylogenetic relationships and discriminate all species, particularly in evolutionarily young and complex plant groups. To gain further resolving power for closely related species, the addition of substantial numbers of nuclear markers is likely to be required.     

Morphological and genomic evidence for a new species of Corallorhiza (Orchidaceae: Epidendroideae) from SW China

Corallorhiza sinensis, a new species of mycoheterotrophic orchid from western Sichuan, China, is described and illustrated based on molecular and morphological evidence. It is morphologically similar to Corallorhiza trifida, but can be distinguished by bigger flowers, both sepals and petals with 3 veins, and longer lateral lobes of lip. To distinguish the new Corallorhiza species and explore its phylogenetic position within subtribe Calypsoinae, this study employed sequences of the nuclear ribosomal DNA (nrDNA) and whole plastome assembled from the genome skimming approach. The plastome is 148,124 bp in length, including a pair of inverted repeats (IRs) of 26,165 bp, a large single-copy region (LSC) of 82,207 bp, and a small single-copy region (SSC) of 13,587 bp. Further, phylogenetic analyses were performed using nrDNA sequence and 79 coding sequences (CDSs) from 26 complete plastomes of subtribe Calypsoinae. The phylogenetic tree based on nrDNA sequence suggested that Corallorhiza is a monophyletic group, and strongly support C. sinensis as sister to the rest species of Corallorhiza. The plastid tree showed that 10 Corallorhiza species grouped into two clades and C. sinensis is most closely related to the North American C. striata and C. bentleyi instead of Oreorchis foliosa and O.angustata in the same clade. The plastid tree and nrDNA tree indicate Oreorchis is a paraphyletic. Although the topological conflicts are displayed between plastome and nrDNA phylogenies of C. sinensis, it is still the most closely related to Corallorhiza. Comparative analysis showed that C. sinensis populations are characteristic of the intermediate morphological traits between Corallorhiza and Oreorchis. The finding of this new species from China shed new light on the phylogeny of Oreorchis and Corallorhiza.     

Molecular phylogeny of Phoma and allied anamorph genera: Towards a reclassification of the Phoma complex

The present generic concept of Phoma is broadly defined, with nine sections being recognised based on morphological characters. Teleomorph states of Phoma have been described in the genera Didymella, Leptosphaeria, Pleospora and Mycosphaerella, indicating that Phoma anamorphs represent a polyphyletic group. In an attempt to delineate generic boundaries, representative strains of the various Phoma sections and allied coelomycetous genera were included for study. Sequence data of the 18S nrDNA (SSU) and the 28S nrDNA (LSU) regions of 18 Phoma strains included were compared with those of representative strains of 39 allied anamorph genera, including Ascochyta, Coniothyrium, Deuterophoma, Microsphaeropsis, Pleurophoma, Pyrenochaeta, and 11 teleomorph genera. The type species of the Phoma sections Phoma, Phyllostictoides, Sclerophomella, Macrospora and Peyronellaea grouped in a subclade in the Pleosporales with the type species of Ascochyta and Microsphaeropsis. The new family Didymellaceae is proposed to accommodate these Phoma sections and related anamorph genera. The present study demonstrated that Phoma radicina, the type species of Phoma sect. Paraphoma and Phoma heteromorphospora, the type species of Phoma sect. Heterospora can be assigned to the Phaeosphaeriaceae and Leptosphaeriaceae respectively.     

A set of plastid microsatellite loci for the genus Dyckia (Bromeliaceae) derived from 454 pyrosequencing

• Premise of the study: Phylogeographical analyses of Dyckia (Bromeliaceae) suffer from low levels of sequence variation. Plastid microsatellite markers were developed to achieve a better-resolved genus-wide plastid genealogy of Dyckia. • Methods and Results: Approximately 84% of the D. marnier-lapostollei plastome was sequenced using 454 technology. Flanking primer pairs were designed for 34 out of 36 chloroplast simple sequence repeats (cpSSRs) detected, and 12 loci were further characterized by genotyping Dyckia samples at the level of populations and species. Three, five, and six cpSSRs were polymorphic among four individuals of D. limae, 12 individuals of D. dissitiflora, and 12 of D. pernambucana, respectively, with two to three alleles per locus and species. All loci were polymorphic among 19 different Dyckia species, with three to 10 alleles per locus. Ten primer pairs cross-amplified with bromeliad genera from five subfamilies. • Conclusions: The set of 12 cpSSR markers provides a toolbox to analyze phylogeographical patterns of Dyckia species.     

Molecular systematics of the genus Astragalus L. (Fabaceae): Phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacers and chloroplast gene ndhF sequences

Comparative sequencing of internal transcribed spacers (ITS) and 5.8S gene of nuclear ribosomal DNA was carried out to examine phylogenetic relationships among subgenera and sections of Old World Astragalus as well as the recent segregate genera Barnebyella and Ophiocarpus. For a subset of these taxa (43 accessions), the nrDNA ITS data were supplemented by sequences from the chloroplast ndhF gene. Phylogenetic trees resulting from separate analyses of the nrDNA ITS and ndhF sequences were in conflict mainly on the position and relationships of Ophiocarpus aitchisonii, Astragalus hemsleyi, A. grammocalyx, A. coelicolor, A. capito, A. epiglottis and A. annularis. Excluding these taxa, phylogenetic analysis of a combined nrDNA ITS-ndhF data matrix was also conducted, so that in the resulting tree, most clades were more resolved and better statistically supported than those were in the separate analyses. Our results indicate that the monotypic segregate genera Barnebyella (= A. migpo), Ophiocarpus (= A. ophiocarpus) and morphologically isolated annual species A. dipelta (= Didymopelta turkestanica), A. schmalhausenii (= Sewerzowia turkestanica) and A. vicarius (= S. vicaria) are clearly nested within Astragalus. Our results confirm earlier studies that shows A. vogelii is allied with the genera Colutea and Oxytropis rather than with any Astragalus species. It is therefore excluded from Astragalus and elevated to the new generic rank and named as Podlechiella Maassoumi and Kazempour Osaloo. None of the eight traditionally recognized Astragalus subgenera Epiglottis, Trimeniaeus, Phaca, Hypoglottis, Calycophysa, Tragacantha, Cercidothrix and Calycocystis are monophyletic. Similarly, the monophyly of Podlechs new subgenera Trimeniaeus, Astragalus and Cercidothrix is not supported. Among the many species-rich sections analyzed here, only Cenanthrum, Chronopus, Laxiflori, Lotidium, Incani and Amodendron are monophyletic.     

Diversity and evolution of plastomes in Saharan mimosoids: potential use for phylogenetic and population genetic studies

Acacias (Mimosoideae) represent a major woody group in arid and subarid habitats of all tropical and subtropical regions. The genetic diversity and population dynamic of African species are still poorly investigated, in particular due to ploidy variation among and within species. Here, we aim to investigate the diversity of the plastid genome (or plastome) of Central Saharan mimosoids, in order to assess its potential utility for phylogenetic and population genetic analyses. We first used a genome skimming strategy to assemble the complete plastome plus the nuclear ribosomal DNA cluster of six species belonging to three genera (Vachellia, Senegalia, and Faidherbia). Phylogenetic relationships based on these data confirm the existence of three main evolutionary lineages in the Hoggar range (southern Algeria). An analysis of the plastome structure reveals an extension of the inverted repeat (IR) in Faidherbia albida as recently reported in two other genera of the same lineage (Inga and Acacia s.s.). Higher substitution rates are detected in this lineage, and our species sampling allows revealing genes (particularly accD, clpP, rps2, rps3, ycf1, ycf2, and ycf4) under positive selection following the IR extension. The reasons for this evolutionary transition need to be unraveled. We then develop 21 plastid microsatellites to be used on a large panel of mimosoid species. At a local scale, 18 of these loci reveal intra-specific polymorphism in at least one species. These markers may be useful to assess the genetic diversity of the plastome for comparative phylogeographies or population genetic studies.     

Phytochemical investigations in Photinia

Leaves of Photinia glabra, P. serrulata, P. glabra × serrulata, P. × fraseri, Heteromeles arbutifolia, Cotoneaster pannosa, and Eriobotrya japonica were investigated for oil and n-alkane content and for hydrogen cyanide production. Benzaldehyde proved to be the major volatile oil component in all taxa except Eriobotrya which yielded no oil upon steam distillation. A conjugated aldehyde was present in all taxa of Asian origin but essentially absent from the New World form Heteromeles. The presence of cyanogenic compounds was indicated in all species again excepting Eriobotrya. C ₂₄H ₅₀ to C ₃₃H ₆₈n-alkanes were presentin all taxa. The patterns of composition showed distinct differences along generally recognizable generic limits and again supported the separate placement of Heteromeles.     

The chicken or the egg? Plastome evolution and an independent loss of the inverted repeat in papilionoid legumes

The plastid genome (plastome), while surprisingly constant in gene order and content across most photosynthetic angiosperms, exhibits variability in several unrelated lineages. During the diversification history of the legume family Fabaceae, plastomes have undergone many rearrangements, including inversions, expansion, contraction and loss of the typical inverted repeat (IR), gene loss and repeat accumulation in both shared and independent events. While legume plastomes have been the subject of study for some time, most work has focused on agricultural species in the IR-lacking clade (IRLC) and the plant model Medicago truncatula. The subfamily Papilionoideae, which contains virtually all of the agricultural legume species, also comprises most of the plastome variation detected thus far in the family. In this study three non-papilioniods were included among 34 newly sequenced legume plastomes, along with 33 publicly available sequences, to assess plastome structural evolution in the subfamily. In an effort to examine plastome variation across the subfamily, approximately 20% of the sampling represents the IRLC with the remainder selected to represent the early-branching papilionoid clades. A number of IR-related and repeat-mediated changes were identified and examined in a phylogenetic context. Recombination between direct repeats associated with ycf2 resulted in intraindividual plastome heteroplasmy. Although loss of the IR has not been reported in legumes outside of the IRLC, one genistoid taxon was found to completely lack the typical plastome IR. The role of the IR and non-IR repeats in the progression of plastome change is discussed.     

Generic delimitation and biogeography of Maianthemum and Smilacina (Ruscaceae sensu lato): preliminary results based on partial 3′ matK gene and trnK 3′ intron sequences of cpDNA

The controversy over generic delimitation between Maianthemum and Smilacina has been unresolved for almost two centuries. Distributions of the two genera in the Northern Hemisphere also provide an excellent opportunity to further understand the disjunct distribution patterns of the North Temperate Flora. To test the generic delimitation and to investigate biogeographic patterns, we sequenced the partial 3′ matK gene and trnK 3′ intron of chloroplast DNA for 38 accessions, representing three species of Maianthemum, seven species of Smilacina, and four outgroup taxa. Maximum parsimony and neighbor-joining trees showed reciprocal monophyly of the two genera with very weak bootstrap support for each genus. Within each genus, relationships among species were poorly resolved. Despite its low resolution, this study shows that eastern Asian species of Smilacina and Maianthemum are generally more closely related to eastern North American taxa than to western ones. More detailed sampling of Smilacina from different geographic regions, especially from the two centers of diversity (southeastern Asia and Mexico/Central America), and additional sequences from cpDNA, as well as from nuclear DNA, are needed to test the reciprocal monophyly of the two genera and also to understand current distributions of disjunct taxa.     

Nuclear DNA from a 180‐year‐old study skin reveals the phylogenetic position of the Kinglet Calyptura Calyptura cristata (Passeriformes: Tyrannides)

The Kinglet Calyptura Calyptura cristata is one of the most enigmatic bird species in South America, known only from specimens collected in the 19th century and a few recent observations. Knowledge of its biology is scanty and its systematic position is obscure. Traditionally, Calyptura was placed in the Cotingidae, but associated with genera that are now known to fall outside the Cotingidae. In an attempt to clarify its phylogenetic position, sequence data from four nuclear markers were obtained from a 180‐year‐old museum study skin of Calyptura, and incorporated into a comprehensive dataset of tyrant flycatchers, cotingas, manakins and allies. Our analyses demonstrate that Calyptura is most closely related to Platyrinchus and Neopipo and that these three genera constitute a deep branch in the clade containing the Rhynchocyclidae (tody‐tyrants and flatbills) and Tyrannidae (typical tyrant flycatchers). The Calyptura specimen is one of the oldest avian museum specimens from which a substantial amount of nuclear DNA sequence data have been obtained, and highlights the immense value of museum collections for DNA‐based phylogenetic studies.     

Comparative spikelet morphology ofOryza schlechteri Pilger and related species ofLeersia andOryza (Poaceae)

Oryza schlechteri has only recently been rediscovered in Papua New Guinea. This species is quite distinct from otherOryza species but resembles closely related genusLeersia in its small panicle and profusely spreading habit. On the basis of spikelet morphology that has previously been used to distinguishLeersia andOryza species,O. schlechteri was compared withO. brachyantha. O. longiglumis, O. sativa, L. perrieri, L. tisseranti, andL. hexandra to clarify its generic position. Results corroborate that the presence of a sterile lemma and a striated spikelet epidermal (abaxial) surface lacking siliceous triads inO. schlechteri allies this species with otherOryza species rather than withLeersia.     

Taxonomic notes on Cimicifuga nanchuanensis (Ranunculaceae), a hitherto imperfectly known species from China

Cimicifuga nanchuanensis P. K. Hsiao (Ranunculaceae) has been regarded as an imperfectly known species. Based on critical examination of herbarium specimens and living plants we have determined that it is a morphologically distinctive species and that its geographical range is much wider than believed before, far from locally endemic to its type locality. Our cytological examination reveals the species to be a diploid (2n = 16). Our molecular phylogenetic analyses using nrDNA and cpDNA datasets confirm our morphological observations. As the generic concept and delimitation of Actaea L., in which Cimicifuga Wernisch. and Souliea Franch. are included, have also been strongly corroborated by our molecular phylogenetic analyses, here we adopt a broad Actaea and make a new combination for the species in question, i.e. Actaea nanchuanensis (P. K. Hsiao) J. P. Luo, Q. Yuan & Q. E. Yang.