Karyomorphology of Taiwanese Begonia (Begoniaceae): taxonomic implications

 The karyomorphology of all 14 species of Taiwanese Begonia was investigated to elucidate their chromosome features and chromosomal evolution. Among all species investigated, differences in chromosome features are found in: (1) chromosome number 2n = 22, 26, 36, 38, 52, 60, 64, 82, and (2) frequencies of chromosomes with secondary, tertiary, and/or small constrictions of polyploids, ranging from 23% to 63%, which is higher than the expected value of about 9%. It is suggested that after polyploidization from the diploid species (i.e., 2n = 22 and frequencies of chromosomes with secondary, tertiary, and/or small constrictions of polyploids of about 9%), chromosome translocations occurred, followed by a decrease in chromosome number, and subsequently stabilized genomes were formed in various species in Taiwan. The karyomorphological evidence also suggested that the chromosome morphology has evolved in parallel in the begonias belonging to different sections in Taiwan. The variation in chromosomal features is more complex than the variation in floral and fruit morphologies. Karyomorphological data also supports the recognition of five new species in Taiwan: Begonia bouffordii, B. chuyunshanensis, B. pinglinensis, B. tengchiana, and B. wutaiana. Based on detailed karyomorphological analyses, the taxonomic implications, speciation, and chromosomal evolution in Taiwanese Begonia are discussed. Received: January 22, 2002 / Accepted: March 4, 2002     

Karyomorphology of Taiwanese Begonia (Begoniaceae): taxonomic implications

The karyomorphology of all 14 species of Taiwanese Begonia was investigated to elucidate their chromosome features and chromosomal evolution. Among all species investigated, differences in chromosome features are found in: (1) chromosome number 2 n = 22, 26, 36, 38, 52, 60, 64, 82, and (2) frequencies of chromosomes with secondary, tertiary, and/or small constrictions of polyploids, ranging from 23% to 63%, which is higher than the expected value of about 9%. It is suggested that after polyploidization from the diploid species (i.e., 2 n = 22 and frequencies of chromosomes with secondary, tertiary, and/or small constrictions of polyploids of about 9%), chromosome translocations occurred, followed by a decrease in chromosome number, and subsequently stabilized genomes were formed in various species in Taiwan. The karyomorphological evidence also suggested that the chromosome morphology has evolved in parallel in the begonias belonging to different sections in Taiwan. The variation in chromosomal features is more complex than the variation in floral and fruit morphologies. Karyomorphological data also supports the recognition of five new species in Taiwan: Begonia bouffordii, B. chuyunshanensis, B. pinglinensis, B. tengchiana, and B. wutaiana. Based on detailed karyomorphological analyses, the taxonomic implications, speciation, and chromosomal evolution in Taiwanese Begoniaare discussed.     

Molecular phylogeny of Coelogyne (Epidendroideae; Orchidaceae) based on plastid RFLPS, matK, and nuclear ribosomal ITS sequences: evidence for polyphyly

To evaluate the monophyly of Coelogyne (Epidendroideae; Orchidaceae) and reveal sectional relationships and relations to allied genera in subtribe Coelogyninae, we collected PCR (polymerase chain reaction) amplified restriction fragment length polymorphisms (RFLPs) from 11 plastid regions for 42 taxa (28 Coelogyne species and 14 representatives of other genera) and three outgroups from Bletiinae and Thuniinae. We also sequenced a large portion of the plastid trnK intron (mostly matK) and the nuclear ribosomal DNA internal transcribed spacers ITS1 and ITS2 (including the 5.8S gene). Separate phylogenetic analyses on each data set using maximum parsimony produced mainly congruent (except for the position of Panisea) but weakly supported clades. Parsimony analysis of the combined data clearly identified three main clades in Coelogyninae. Whereas Coelogyninae are monophyletic, Coelogyne is polyphyletic, with species falling into at least two well-supported clades. The utility of morphological characters used in previous classifications was explored by reconstructing character state evolution on one of the four molecular trees. Lip base and petal shape were homoplasious, whereas ovary indumentum and flower number were congruent with well-supported groups. The implications of our results for the classification of Coelogyne are discussed, and a reorganization of the genus by including Neogyna and Pholidota and removing several species is proposed.     

A phylogeny of the Lampropeltis mexicana complex (Serpentes: Colubridae) based on mitochondrial DNA sequences suggests evidence for species-level polyphyly within Lampropeltis

The systematic relationships of snakes in the Lampropeltis mexicana complex (L. mexicana, L. alterna, and L. ruthveni) are poorly known despite several taxonomic studies over the last 80 years. Mitochondrial DNA sequences were used to infer the phylogeny of the L. mexicana complex. At least one representative sample from the nine currently recognized species of Lampropeltis was sequenced. Our results suggest that a deep basal split resulted in the divergence of two groups of Lampropeltis, with one group occupying the upland areas of western United States and most of western and central Mexico, and the other northeastern Mexico and the lowland areas of the southern United States. Results also revealed that the L. mexicana complex and Lampropeltis triangulum are polyphyletic, with taxa from both groups nested together in deeply divergent northern and southern clades. These results are incongruent with previous hypotheses of phylogenetic relationships based on morphology, and suggest that morphological characters shared among the various tri-colored Lampropeltis (e.g., hemipenal structure and tri-colored pattern) may be difficult to interpret phylogenetically.     

世界秋海棠属侧膜组植物新资料

记述了秋海棠属侧膜组11新种,即星果草叶秋海棠(BegoniaasteropyrifoliaY.M.ShuietW.H.Chen)、耳托秋海棠(B.auritistipulaY.M.ShuietW.H.Chen)、桂南秋海棠(B.austroguangxiensisY.M.ShuietW.H.Chen)、水晶秋海棠(B.crystallinaY.M.ShuietW.H.Chen)、须苞秋海棠(B.fimbribracteataY.M.ShuietW.H.Chen)、巨叶秋海棠(B.gigaphyllaY.M.ShuietW.H.Chen)、黄氏秋海棠(B.huangiiY.M.ShuietW.H.Chen)、长柱秋海棠(B.longistylaY.M.ShuietW.H.Chen)、扁果秋海棠(B.platycarpaY.M.ShuietW.H.Chen)、喙果秋海棠(B.rhynchocarpaY.M.ShuietW.H.Chen)、多变秋海棠(B.variifoliaY.M.ShuietW.H.Chen);并报道了3新变种及1新名称,即疏毛越南秋海棠(B.boniiGagnep.var.remotisetulosaY.M.ShuietW.H.Chen)、密毛龙州秋海棠(B.morseiIrmsch.var.myriotrichaY.M.ShuietW.H.Chen)、簇毛伞叶秋海棠(B.umbr-aculifoliaY.WanetB.N.Changvar.flocculosaY.M.ShuietW.H.Chen)及彩纹秋海棠(B.variegataY.M.ShuietW.H.Chen)。     

中国秋海棠属等翅组植物订正

根据海南秋海棠 (BegoniahainanensisChunetF .Chun)、多花秋海棠 (B sinofloribundaL .J.Dorr)及一新种———长柄秋海棠 (B sublongipesY .M .Shui)的特征比较 ,订正了中国秋海棠属等翅组 [Sect Petermannia (Klotzsch)A .DC .],使该组在中国增至 3个种。     

Correlated evolution of leaf shape and trichomes in Begonia dregei (Begoniaceae)

Structural features of leaves, including size, shape, and surfaces, vary greatly throughout the plant kingdom. In both functional and phylogenetic analyses of leaves, the various morphological aspects are often considered independently of each other, although it is likely that many combinations of features do not occur at random due to either functional constraint or genetic correlation. The distribution of variation in leaf morphology in the highly variable Begonia dregei species complex was examined in natural populations and in F(2) offspring from a cross between plants from two populations. Leaf shape was quantified using several morphometric measures, and trichomes on leaves were counted and measured. Correlations between leaf shape and the numbers and size of trichomes were examined. There were significant correlations between the shapes of leaves and the presence, number, and size of trichomes among populations and in hybrid plants. Deeply incised leaves had larger numbers of longer trichomes at the sinuses. Higher numbers of trichomes on upper leaf surfaces occurred together with trichomes at the petiole and on the abaxial surface. The potential for independent evolution of leaf shape and trichomes in this group is limited. Hypotheses to explain the correlated development of leaf shape and trichomes are discussed.     

Lophotrochozoan phylogeny assessed with LSU and SSU data: evidence of lophophorate polyphyly

Of the three major bilaterian clades, Lophotrochozoa has the greatest diversity and disparity of body forms and is the least understood in terms of phylogenetic history. Within this clade, small nuclear ribosomal subunit (SSU or 18S) studies have failed to provide resolution and other molecular markers have insufficient taxon sampling. To examine relationships within Lophotrochozoa, we collected and complied complete SSU data and nearly complete (>90%) large nuclear ribosomal subunit (LSU or 28S) data totaling approximately 5kb per taxon, for 36 lophotrochozoans. Results of LSU and combined SSU+LSU likelihood analyses provide topologies more consistent with morphological data than analyses of SSU data alone. Namely, most phyla recognized on morphological grounds are recovered as monophyletic entities when the LSU data is considered (contra SSU data alone). These new data show with significant support that "Lophophorata" (traditionally recognized to include Brachiopoda, Phoronida, and Bryozoa) is not a monophyletic entity. Further, the data suggest that Platyzoa is real and may be derived within lophotrochozans rather than a basal or sister taxon. The recently discovered Cycliophora are allied to entoprocts, consistent with their initial placement based on morphology. Additional evidence for Syndermata (i.e., Rotifera+Acanthocephala) is also found. Although relationships among groups with trochophore-like larvae could not be resolved and nodal support values are generally low, the addition of LSU data is a considerable advance in our understanding of lophotrochozoan phylogeny from the molecular perspective.     

New and noteworthy species in Begonia section Gireoudia (Begoniaceae) from Mexico

Two new species are described inBegonia sectionGireoudia:B. rafael-torresii andB. mariti.Begonia karwinskyana, known previously only from its type, is described more amply and discussed.     

Molecular evidence for the polyphyly of Olearia (Astereae: Asteraceae)

 Analyses of ITS sequences for 49 species of Olearia, including representatives from all currently recognised intergeneric sections, and 43 species from 23 other genera of Astereae, rooted on eight sequences from Anthemideae, provide no support for the monophyly of this large and morphologically diverse Australasian genus. Eighteen separate lineages of Olearia are recognised, including seven robust groups. Three of these groups and another eight species are placed within a primary clade incorporating representatives of Achnophora, Aster, Brachyscome, Calotis, Camptacra, Erigeron, Felicia, Grangea, Kippistia, Lagenifera, Minuria, Oritrophium, Peripleura, Podocoma, Remya, Solidago, Tetramolopium and Vittadinia. The remaining four groups and three individual species lie within a sister clade that also includes Celmisia, Chiliotrichum, Damnamenia, Pleurophyllum and Pachystegia. Relationships within each primary clade are poorly resolved. There is some congruence between this molecular estimate of the phylogeny and the distribution of types of abaxial leaf-hair, which is the basis of the present sectional classification of Olearia, but all states appear to have arisen more than once within the tribe. It is concluded that those species placed within the second primary clade should be removed from the genus, but the extent to which species placed within the first primary clade constitute a monophyletic group can only be resolved with further sequence data. Received November 12, 2001; accepted April 29, 2002 Published online: November 22, 2002 Addresses of authors: Edward W. Cross, Centre for Plant Biodiversity Research, CSIRO, GPO Box 1600, Canberra, ACT 2601, Australia (E-mail: ed.cross@csiro.au); Christopher J . Quinn, Royal Botanic Gardens, Mrs Macquaries Rd., Sydney, NSW 2000, Australia; Steven J. Wagstaff, Landcare Research, PO Box 69, Lincoln 8152, New Zealand.     

Molecular resolution of the systematics of a problematic group of fishes (Teleostei: Osmeridae) and evidence for morphological homoplasy

Relationships among the species of Northern Hemisphere smelts (family Osmeridae) have long been debated in the fish systematics literature. Eight independent studies based on morphological characters failed to reach any consensus on osmerid interrelationships. We reconstruct the osmerid phylogeny based on DNA sequence data from three mitochondrial (cytb, 16S, 12S) and three nuclear (ITS2, S71, RAG1) gene regions from multiple individuals of the 14 species in 6 genera, using the Japanese ayu (Plecoglossus altivelis) as the outgroup. Analyses with different combinations of nuclear and mitochondrial datasets yielded a generally well-resolved phylogeny of the genera that conflicts with previous hypotheses of osmerid interrelationships, and Shimodaira–Hasegawa tests suggest our topology with the current molecular dataset is significantly better than earlier reconstructions. In addition, mapping 114 morphological characters used in previous studies onto our phylogeny shows widespread homoplasy, which is likely the source of the systematic disagreement produced in earlier works.     

滇东南黄连山秋海棠属一新种

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中国秋海棠属植物的叶表皮特征及其分类学意义

在光学显微镜下,对中国秋海棠属(Begonia)植物7组52种2变种的叶表皮进行观察。结果表明秋海棠属植物叶表皮形态在属内组间具有较大的相似性,表皮细胞为多边形或近多边形,垂周壁平直或弓形,大多数种类表皮细胞内具有晶体,气孔器仅分布于下表皮,且以不等型为主。叶表皮综合特征,例如表皮细胞形状,表皮毛类型,表皮细胞内晶体的类型和形态,气孔器形态以及与一些种类独有特征的组合,在种间,尤其在近缘种之间具有明显的差异。     

Multigene phylogeny of Synura (Synurophyceae) and descriptions of four new species based on morphological and DNA evidence

We used phylogenetic analyses based on multiple gene sequences (partial nr SSU and LSU rDNA, partial pt LSU rDNA, psaA and rbcL) from 148 strains (including three outgroups) and scale ultrastructure to examine phylogenetic relationships among species of the colonial genera Synura and Tessellaria. The phylogenetic tree based on the combined dataset was congruent with ultrastructural characteristics of the scales. Synura was divided into three major clades, two including species in section Synura, and one representing section Peterseniae. One clade, consisting of seven strains of S. uvella (section Synura), diverged at the base of the genus. The second clade consisted of the remaining species belonging to the section Synura. The third clade, containing organisms in the section Peterseniae and characterized by scales possessing a keel, was monophyletic with strong support values. Based on our findings, S. uvella needs to be in a separate section from other spine-bearing species, and we therefore propose new sectional ranks; Synura, Peterseniae, Curtispinae (presence of body scales with slender spines, tubular scales and caudal scales). We further propose four new species based on phylogenetic analyses and unique scale characters: S. longitubularis sp. nov., S. sungminbooi sp. nov., S. soroconopea sp. nov. and S. lanceolata sp. nov. Lastly, we propose a new genus name, Neotessella, to replace the invalid use of the name Tessellaria.     

Molecular phylogeny of Phalaenopsis Blume (Orchidaceae) on the basis of plastid and nuclear DNA

Representatives of the genus Ancistrocladus, woody lianas of tropical Africa and Asia, contain pharmaceutically interesting alkaloids and have been the subjects of intensive phytochemical investigations. In Southeast Asia, Ancistrocladus tectorius, previously regarded as the only species of the genus from this region, is extremely polymorphic with respect to naphthylisoquinoline alkaloids, indicating that this taxon might be comprised of several morphologically similar species. We performed a comparative study of the ITS region of nuclear rDNA and of the trnK intron of cpDNA as well as an ISSR fingerprint analysis. Using 75 samples of A. tectorius from 21 locations in comparison to samples of other Ancistrocladus species from Asia and from West and Central Africa, we investigated patterns of species differentiation within this taxon. We found the high variability of chemical compounds described for A. tectorius to be paralleled by a high genetic variability of the units that have been assigned to this taxon. Samples assigned to A. tectorius were paraphyletic with respect to species from the Indian subcontinent, and intraspecific variability was comparable to interspecific variability among the African taxa. In addition, groups of individuals occurring in sympatry were found to be more similar to those from other locations, suggesting low levels of gene flow between those sympatric groups. This indicates either a considerable number of hybridization events during the evolution of A. tectorius or the existence of several distinguishable species not yet recognized. Our results are a first step in the development of species or population-specific markers for the prediction of the alkaloid spectrum of samples. This will help to improve the reproducibility of phytochemical research on Ancistrocladus.