Clarification of the relationship beteen Apiaceae and Araliaceae based on matK and rbcL sequence data

Apiaceae and Araliaceae (Apiales) represent a particularly troublesome example of the difficulty in understanding evolutionary relationships between tropical-temperate family pairs. Previous studies based on rbcL sequence data provided insights at higher levels, but were unable to resolve fully the family-pair relationship. In this study, sequence data from a more rapidly evolving gene, matK, was employed to provide greater resolution. In Apiales, matK sequences evolve an average of about two times faster than rbcL sequences. Results of phylogenetic analysis of matK sequences were first compared to those obtained previously from rbcL data; the two data sets were then combined and analyzed together. Molecular analyses confirm the polyphyly of apiaceous subfamily Hydrocotyloideae and suggest that some members of this subfamily are more closely related to Araliaceae than to other Apiaceae. The remainder of Apiaceae forms a monophyletic group with well-defined subclades corresponding to subfamilies Apioideae and Saniculoideae. Both the matK and the combined rbcL-matK analyses suggest that most Araliaceae form a monophyletic group, including all araliads sampled except Delarbrea and Mackinlaya. The unusual combination of morphological characters found in these two genera and the distribution of matK and rbcL indels suggest that these taxa may be the remnants of an ancient group of pro-araliads that gave rise to both Apiaceae and Araliaceae. Molecular data indicate that the evolutionary history of the two families is more complex than simple derivation of Apiaceae from within Araliaceae. Rather, the present study suggests that there are two well-defined "families," both of which may have been derived from a lineage (or lineages) or pro-araliads that may still have extant taxa.     

Evolution in Apiales: nuclear and chloroplast markers together in (almost) perfect harmony

Relationships within the angiosperm order Apiales have long been difficult to interpret. Traditionally, the order comprised two families, Apiaceae and Araliaceae. Recent studies, however, suggest three additional lineages should also be recognized in the order (Pittosporaceae plus two tribes segregated from Araliaceae, Mackinlayeae and Myodocarpeae), and that one taxon (Apiaceae subfamily Hydrocotyloideae) is polyphyletic. Nuclear data also support the placement of five enigmatic genera ( Aralidium , Griselinia , Melanophylla , Pennantia and Torricellia ) within an expanded Apiales. To date, detailed molecular studies of Apiales have relied largely on data derived from plastid sequences, especially mat K and rbc L. To test and complement the results of these studies, the 26S (large subunit) of nuclear ribosomal DNA was sequenced and analysed phylogenetically. Results from this study confirm that Apiales comprise five major lineages: core Apiaceae, core Araliaceae, Pittosporaceae, the Mackinlaya group and the Myodocarpus group. Moreover, using an expanded sampling of members of subfamily Hydrocotyloideae, the nature and extent of the polyphyly is confirmed, with members of this taxon found among four distinct clades within Apiales. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 123–147.     

Interspecific relationships in Fagopyrum (Polygonaceae) revealed by the nucleotide sequences of the rbcL and accD genes and their intergenic region

DNA sequences of the rbcL and accD 5' coding regions and their intergenic region (IGR) were determined in 20 taxa of 12 species of Fagopyrum. By comparing the sequence data and constructing phylogenetic trees, phylogenetic relationships among Fagopyrum species were investigated. The accD 5' coding region and the IGR have evolved about five times faster than the rcbL coding region; hence the phylogenetic trees based on the data of the accD 5' coding region, the IGR, and these data combined, had higher resolution than the trees based on rcbL alone. The classification of Fagopyrum based on the DNA sequences was almost perfectly consistent with the classification by Ohnishi and Matsuoka (1996, Genes & Genetic Systems 71: 383-390). New facts and minor differences are (1) polyploidization in F. cymosum might have occurred more than once, (2) F. lineare may not be a sister species of F. leptopodum, and (3) F. pleioramosum rather than F. capillatum may be a sister species of F. gracilipes.     

An ITS-based phylogenetic analysis of theEuryptera species group inLomatium (Apiaceae)

Lomatium, the largest genus of Apiaceae in western North America, includes many narrow endemics whose relationships are uncertain. Although no infrageneric classification exists forLomatium, several informal groups have been recognized. TheEuryptera group comprises seven narrowly endemic species distributed primarily in California. We conducted parsimony and maximum likelihood (ML) analyses using sequences of the internal transcribed spacers (ITS) of the nuclear ribosomal DNA from species of theEuryptera group and several other species ofLomatium. When considered with distribution, morphological, and cpDNA data, the ITS analyses are consistent with the monophyly of theEuryptera group and suggests that speciation in this group has occurred through geographical divergence. Inferences from ITS data also identify putative progenitors of the polyploidEuryptera species.     

Pollen Morphology of Hydrocotyloideae and Saniculoideae (Apiaceae) in China

Pollen grains of 30 species and 2 varieties from China, belonging to 5 genera (out of 36 species in 5 genera) of Hydrocotyloideae and Saniculoideae, were examined with light microscope and scanning electron microscope. 1.Six types of the pollen shape are recognized in Hydrocotyloideae and Saniculoideae from China, and their evolutionary trend is from rhomboidal→subspheroidal→ellipsoidal→subrectangular→superrectangular to equatorially-constricted. 2.Pollen grains of Hydrocotyloideae are mostly ellipsoidal with P / E ratio 1.07-1.6, the polar axis 22.5-46μm long, the equatorial axis 13.75-27.5μm long, and with the size index 21.1-31.8.They are angulapeturate in the great majority, and the exine sculpture is recticulate under SEM. Those of Saniculoideae are mostly superrectangular or rectangular, with P/E ratio 1.3-2.1, the polar axis 35-65μm long, the equatorial axis 17.5-42.5μm long, and with the size index 28.2-49.7. They are planaperturate in the great majority and the exine sculpture is striate-reticulate or cerebro-reticulate under SEM. Therefore, the latter might be more advanced than the former. 3.Pollen grains of Dickinsia Franch., endemic to China, are ellipsoidal,with P/E ratio 1.5,the polar axis 31.25-37.5μm long, the equatorial axis 20-25μm, and with the size index 27.81. They are angulaperturate, and the exine sculpture is cerebroid-reticulate under SEM. The genus is greatly similar to the members of Hydrocotyloideae in pollen morphology, which supports the treatment of the genus asa member of Hydrocotyloideae by Handel-Mazzetti (1933).     

鳞毛蕨科植物的系统发育: 叶绿体rbcL序列的证据

运用MEGA2和MrBayes 3.0b4软件包对105种鳞毛蕨类及近缘植物(其中新测定36种)的叶绿体DNA rbcL基因序列进行系统发育分析, 探讨了其主要分类群(属级水平)的系统演化关系。用最大简约法、邻接法和贝叶斯分析方法构建的系统树基本一致, 结果显示: (1)秦仁昌系统所定义的鳞毛蕨科Dryopteridaceae, 除了拟贯众属Cyclopeltis外, 均包含在两个单系群之中, 支持鳞毛蕨族Dryopterideae和耳蕨族Polysticheae的成立; 但是鳞毛蕨族还包含秦仁昌系统所定义的球盖蕨科Peranemaceae和三叉蕨科Tectariaceae肋毛蕨属Ctenitis的部分种类; 耳蕨族还包含产于美洲的Phanerophlebia属和Polystichopsis属; 确认石盖蕨属Lithostegia属鳞毛蕨族的成员, 且与复叶耳蕨属Arachniodes具有较近的亲缘关系。(2)拟贯众属与所分析的其他任何鳞毛蕨类植物的关系都比较疏远, 单独为一支。(3)秦仁昌系统所定义的球盖蕨科与肉刺蕨属Nothoperanema聚成一个分支, 属于鳞毛蕨族的成员。(4)鳞毛蕨属Dryopteris为多系类群, 耳蕨属Polystichum和贯众属Cyrtomium均为并系类群。(5)黔蕨属Phanerophlebiopsis、毛枝蕨属Leptorumohra和石盖蕨属与复叶耳蕨属构成一支; 柳叶蕨属Cyrtogonellum与Polystichum属和Cyrtomium属的部分种类聚成一支; 肉刺蕨属与球盖蕨科及鳞毛蕨属的部分种类聚成一支。对鳞毛蕨科的系统关系、球盖蕨科与鳞毛蕨科的系统关系、肋毛蕨属与鳞毛蕨科的系统关系以及中国或亚洲特有属(拟贯众属、肉刺蕨属、黔蕨属、毛枝蕨属、石盖蕨属和柳叶蕨属)的系统位置进行了讨论。     

Polyploid speciation inHedera (Araliaceae): Phylogenetic and biogeographic insights based on chromosome counts and ITS sequences

Variation in chromosome number and internal transcribed sequences (ITS) of nrDNA is used to infer phylogenetic relationships of a wide range ofHedera species. Polyploidy was found to be frequent inHedera, with diploid, tetraploid, hexaploid and octoploid populations being detected. Nucleotide additivity occurs in the ITS sequences of one tetraploid (H. hibernica) and two hexaploid species (H. maderensis, H. pastuchovii), suggesting that all three species originated by allopolyploidisation. ITS sequence polymorphism and nucleotide characters may indicate the presence of an ancient genome persistent only in some allopolyploid species. Phylogenetic analyses of ITS sequence data reveal two lineages ofHedera: one containing all sequences belonging to extant diploids plus the tetraploidH. algeriensis, and a second that includes this ancient ITS type and others exclusive to several polyploid species. The origin of the polyploids is evaluated on the basis of morphology, chromosome counts, ITS sequence polymorphism, and phylogenetic analyses. Reconstruction of reticulate evolution inHedera agrees with two allopolyploid areas on both sides of the Mediterranean basin. Morphological, molecular and cytological evidence also suggests an active dispersal ofHedera populations that may account for three independent introductions in Macaronesia.     

Phylogenetic relationships within Araucariaceae based on rbcL gene sequences

Phylogenetic relationships were determined in the Araucariaceae, which are now found mainly in the Southern Hemisphere. This conifer family was well diversified and widely distributed in both hemispheres during the Mesozoic era. The sequence of 1322 bases of the rbcL gene of cpDNA was determined from 29 species of Araucariaceae, representing almost all the species of the family. Phylogenetic trees determined by the parsimony method indicate that Araucariaceae are well defined by rbcL sequences and also that the monophyly of Agathis or Araucaria is well supported by high bootstrap values. The topology of these trees revealed that Wollemia had derived prior to Agathis and Araucaria. The rbcL phylogeny agrees well with the present recognition of four sections within Araucaria: Araucaria, Bunya, Eutacta, and Intermedia. Morphological characteristics of the number of cotyledons, position of male cone, and cuticular micromorphologies were evaluated as being phylogenetically informative. Section Bunya was found to be derived rather than to be the oldest taxon. Infrageneric relationships of Agathis could not be well elucidated because there are few informative site changes in the rbcL gene, suggesting the more recent differentiation of the species as their fossil records indicate. The New Caledonian Araucaria and Agathis species each formed a monophyletic group with very low differentiation in rbcL sequences among them, indicating rapid adaptive radiation to new edaphic conditions, i.e., ultramafic soils, in the post-Eocene era.     

基于叶绿体基因rbcL和psbA-trnH间区探讨石杉科植物系统关系

关于石杉科Huperziaceae植物的分类,一直存在一些争议。在旧的分类体系中石杉科植物被包含在一个混合的石松科Lycopodiaceae和多谱系的石松属Lycopodium中。本文利用叶绿体rbcL基因和psbA-trnH基因间区序列探讨石杉科植物的系统位置及石杉科内部的分类关系,用最大简约法和邻接法对自测序列结合由GenBank下载的rbcL及psbA-trnH基因间区序列进行系统发育分析。结果显示,石杉科与Phylloglossum属关系较近,与石松科关系较疏远。在石杉科中热带石杉属Huperzia植物和马尾杉属Phlegmariurus植物的关系要比它们与其他石杉属植物更近。所以,我们的rbcL基因数据不支持秦仁昌关于石杉科分为石杉属和马尾杉属的分类处理。但是,因为我们的psbA-trnH序列没有包括热带种类,对石杉属植物和马尾杉属植物的关系无验证。因此需要更多的样品和序列数据进一步探讨石杉科的演化关系。     

rbcL gene sequences reveal relationships among north-east Pacific species of Porphyra (Bangiales, Rhodophyta) and a new species, P. aestivalis

Phylogenetic analyses of the rbcL (chloroplast Rubisco large subunit) gene from 23 newly sequenced species of Porphyra, primarily from the north‐east Pacific, one Bangia and previously published sequences from both genera resolve relationships among most species of Porphyra and reveal five clades of species with Porphyra‐type morphologies among a number of Bangia lineages: (1) P. papenfussii V. Krishnam; (2) P. mumfordii S. C. Lindstrom et K. M. Cole and P. rediviva Stiller et Waaland together with a group of north Atlantic species, including the type of the genus, P. purpurea (Wahl‐enb.) C. Agardh; (3) P. cuneiformis (Setch. et Hus) V. Krishnam., P. occidentalis Setch. et Hus, P. schizo‐phylla Hollenb., and P. variegata (Kjellm.) Kjellm. and their Atlantic sibling species, all distromatic; (4) P. aestivalis sp. nov. and its north Atlantic sibling, P. birdiae C. D. Neefus et A. C. Mathieson; and (5) a speciose clade containing both Pacific and Atlantic representatives. Close relationships are confirmed between sibling species previously identified by iso‐zymes, morphology and chromosomal features. The morphologically similar dioecious P. pseudolanceolata V. Krishnam., P. conwayae (S. C. Lindstrom et K. M. Cole) stat. nov., and P. lanceolata (Setch. et Hus) G. M. Smith occur in a strongly supported subclade in clade 5 together with the monoecious P. fallax S. C. Lindstrom et K. M. Cole. Results presented here highlight the need for intensive taxon sampling and for examination of different parts of the genome to understand more fully relationships among species and higher level taxa in the Bangiales.     

线蕨属（水龙骨科）的系统发育研究：来自叶绿体rbcL和rps4-trnS序列的证据

线蕨属Colysis 植物主要分布于亚洲热带和亚热带地区,少数种类分布至非洲、澳大利亚(昆士兰)及新几内亚地区.自1849年成立以来,线蕨属的分类范畴和系统位置一直有待确定.本文利用叶绿体基因组的rbcL、rps4基因和rps4-trnS基凶间隔区序列,运用最大简约法和贝叶斯方法分析了线蕨属及其近缘类群的系统演化关系.研究结果显示:(1)线蕨属和薄唇蕨属Leptochilus(含似薄唇蕨属Paraleptochilus)组成一个支持率很高的单系分支(C-L Clade),但是薄唇蕨属的成员位于线蕨属的不同支系内,支持线蕨属和薄唇蕨属合并为一个属;(2)瘤蕨属Phymatosorus单独形成一个单系分支;(3)星蕨属Microsorum是一个多系类群,除Microsorium linguiforme、M.varians和M.pustulatum与马来群岛的Lecanopteris聚在一起外,其他的星蕨属成员均位于不同的支系上.本文的系统发育分析结果为线蕨属和薄唇蕨属的分类处理提供了分子系统学的证据.     

Phylogenetic relationships of Salix (Salicaceae) based on rbcL sequence data

Nucleotide sequences of the chloroplast-encoded rbcL gene were used to examine phylogenetic relationships of the genus Salix together with other allied genera of the family Salicaceae. Phylogenetic analyses of rbcL sequences strongly suggest the monophyly of three commonly recognized genera (Chosenia, Salix, and Toisusu). Two monophyletic groups are recognized within the larger monophyletic group. They do not correspond with any infrageneric taxa proposed so far. With regard to character evolution, it is thought that the reduction of stamen number from more than two stamens to two might occur in at least three lineages and that fused bud scales evolved several times and/or the reverse evolution occurred from fused to free. Some types of pollen surfaces are considered to have evolved independently.     

Choosing appropriate substitution models for the phylogenetic analysis of protein-coding sequences

Although phylogenetic inference of protein-coding sequences continues to dominate the literature, few analyses incorporate evolutionary models that consider the genetic code. This problem is exacerbated by the exclusion of codon-based models from commonly employed model selection techniques, presumably due to the computational cost associated with codon models. We investigated an efficient alternative to standard nucleotide substitution models, in which codon position (CP) is incorporated into the model. We determined the most appropriate model for alignments of 177 RNA virus genes and 106 yeast genes, using 11 substitution models including one codon model and four CP models. The majority of analyzed gene alignments are best described by CP substitution models, rather than by standard nucleotide models, and without the computational cost of full codon models. These results have significant implications for phylogenetic inference of coding sequences as they make it clear that substitution models incorporating CPs not only are a computationally realistic alternative to standard models but may also frequently be statistically superior.     

A phylogeny of the flowering plant family Apiaceae based on chloroplast DNA rpl16 and rpoC1 intron sequences: towards a suprageneric classification of subfamily Apioideae

The higher level relationships within Apiaceae (Umbelliferae) subfamily Apioideae are controversial, with no widely acceptable modern classification available. Comparative sequencing of the intron in chloroplast ribosomal protein gene rpl16 was carried out in order to examine evolutionary relationships among 119 species (99 genera) of subfamily Apioideae and 28 species from Apiaceae subfamilies Saniculoideae and Hydrocotyloideae, and putatively allied families Araliaceae and Pittosporaceae. Phylogenetic analyses of these intron sequences alone, or in conjunction with plastid rpoC1 intron sequences for a subset of the taxa, using maximum parsimony and neighbor-joining methods, reveal a pattern of relationships within Apioideae consistent with previously published chloroplast DNA and nuclear ribosomal DNA ITS based phylogenies. Based on consensus of relationship, seven major lineages within the subfamily are recognized at the tribal level. These are referred to as tribes Heteromorpheae M. F. Watson & S. R. Downie Trib. Nov., Bupleureae Spreng. (1820), Oenantheae Dumort. (1827), Pleurospermeae M. F. Watson & S. R. Downie Trib. Nov., Smyrnieae Spreng. (1820), Aciphylleae M. F. Watson & S. R. Downie Trib. Nov., and Scandiceae Spreng. (1820). Scandiceae comprises subtribes Daucinae Dumort. (1827), Scandicinae Tausch (1834), and Torilidinae Dumort. (1827). Rpl16 intron sequences provide valuable characters for inferring high-level relationships within Apiaceae but, like the rpoC1 intron, are insufficient to resolve relationships among closely related taxa.     

Reinstatement of Grateloupia subpectinata (Rhodophyta, Halymeniaceae) based on morphology and rbcL sequences

Morphological observations and molecular analyses of the north‐western Pacific species of the red algal genus Grateloupia (Halymeniaceae) indicate the presence of an entity, which is somewhat similar in gross morphology to G. asiatica Kawaguchi et Wang but is distinguished from the latter species by some morphological features. These include: (i) a somewhat fleshy texture; (ii) wider and much thicker (4.5–10 mm wide and up to 1300 μm thick) axes, of which an inner cortex consists of more (6–9) cells; (iii) generally longer (up to 17 cm), marginal and surface proliferations that are clearly constricted (terete) at bases; and (iv) much elongated, oblong auxiliary cells. Phylogenetic analysis using the ribulose‐l,5‐bisphosphate carboxylase/ oxygenase (rbcL) gene of G. asiatica and the alga in question shows them to be distantly related and strongly supports the differentiation of these two entities at the species level. Judging from the literature, this entity is actually Grateloupia subpectinata Holmes, which has been placed into synonymy under G. asiatica [as G. filicina (Lamouroux) C. Agardh] or G. prolongata J. Agardh in previous reports, and therefore the Holmes name is reinstated.