Michael G. Pimenov

Relationships among genera in Saniculoideae and selected Apioideae (Umbelliferae) inferred from nrITS sequences

The internal transcribed spacers (ITS1 and ITS2) of 18S-26S nuclear ribosomal DNA were newly sequenced for eight species of Umbelliferae (six species from subfamily Saniculoideae: Actinolema macrolema, Astrantia minor, Eryngium giganteum, E. coeruleum, Hacquetia epipactis, and Lagoecia cum inoides, two species from subfamily Hydrocotyloideae: Dickinsia hydrocotyloides and Azorella trifurcata), as well as Hohenackeria exscapa, a species of uncertain position in the family. Phylogenetic analyses of new data, plus previously reported sequences of 52 other species using neighbor-joining, maximum parsimony, and maxim um likelihood methods yielded similar results: (1) Actinolema is sister to Astrantia corresponding to Drude’s treatments; (2) in Astrantia, molecular divergence is revealed between sects. Astrantiella (A. minor) and Astrantia (A. major, A. maxima); (3) Eryngium appears to be paraphyletic; (4) Hacquetia might be treated as a part of Sanicula; and (5) Lagoecia is very distant from all other Saniculoideae and close to some genera of Apioideae. Our results correspond to matK data previously published: (1) Hohenackeria forms a clade with Bupleurum, in a position near the base of the Apioideae tree; (2) Azorella is sister to a large cluster uniting all Saniculoideae and Apioideae, being slightly closer to them than to the Hydrocotyle-Araliaceae clade; (3) Dickinsia is very distant from phenetically similar Hydrocotyle, falling within a large cluster of Apioideae, but also including Lagoecia and Naufraga.

nrDNA ITS sequences and affinities of Sino-Himalayan Apioideae (Umbelliferae)

The internal transcribed spacers (ITS1 and ITS2) of 18S-26S nuclear ribosomal DNA were sequenced for 13 species of Apioideae from the Sino-Himalayan flora [Apium ventricosum (= Sium frigidum), Arcuatopterus thalictrioideus, Cyclorhiza peucedanifolia, Meeboldia achilleifolia, Notopterygium forbesii, N. weberbauerianum, Pternopetalum delavayi, P. vulgare, Pterocyclus rivulorum, Sinocarum cruciatum, Sinodielsia delavayi, Tongoloa elata, and Trachydium simplicifolium], relationships of which are controversial or obscure and unresolved on the basis of morphological data. Phylogenetic trees inferred by neighbor-joining, parsimony analysis, and Bayesian inference are topologically congruent, but not identical. The newly obtained data permit testing of several hypotheses regarding these taxa. Some of the early hypotheses treat local species under well-known European genera (e.g., Apium, Carum, Pimpinella, etc.), but these are not supported by phylogenetic analysis of nuclear rDNA spacer sequences. In particular, Sinodielsia, Meeboldia, Tongoloa, and Pternopetalum are distant from Pimpinella; Pternopetalum, Sinocarum and Tongoloa are remote from Carum; Arcuatopterus is separate from Peucedanum and Angelica; Pternopetalum is distinct from Cryptotaenia; and Notopterygium and Pterocyclus are separate from Pleurospermum. Chinese Notopterygium is shown to be closely related to Siberian and Mongolian Hansenia; this, plus similarity in fruit structure, suggests that they are congeneric. Apium ventricosum, more commonly known as Sium frigidum, appears a close relative of Sinocarum. Affinities of most Sino-Himalayan Apioideae, therefore, are found among taxa from the same and adjacent territories. Wide disjunction between presumptively related taxa has been revealed only in Cyclorhiza and Komarovia. Sinodielsia and Meeboldia are also revealed to be clearly distinct genera.

ITS phylogeny of West Asian Heracleum species and related taxa of Umbelliferae–Tordylieae W.D.J.Koch, with notes on evolution of their psbA-trnH sequences

SummaryHeracleum is a large and taxonomically complex genus of the Umbelliferae–Tordylieae. The phylogenetic relationships of West Asian Heracleum species and related taxa were explored using data from sequences of the internal transcribed spacer (ITS1 and ITS2) region of nuclear ribosomal DNA. The data set consists of 56 species, of which 47 were analyzed for the first time; it represents all subdivisions of the genus Heracleum, as well as some representatives of Pastinaca complex. Heracleum was shown to be a polyphyletic genus, as its species fall into two different clades, one of which comprises also Symphyoloma and Mandenovia. Section Pubescentia was confirmed, in contrast to the sections Villosa and Heracleum being polyphyletic. A separate position of the section Wendia was supported. H. marashicum was shown to be a member of a clade comprising Pastinaca and related genera. The sequences of chloroplast psbA-trnH intergenic spacer, the region recently proposed for DNA barcoding in plants, were also analyzed for 33 species, representing all principal clades within Heracleum and its relatives. They have been proven to be very similar and not suitable for DNA barcoding in this group. However, some sequence variation was revealed. This variation could be explained by the combination of such evolutionary events as inversion and duplication. It was shown that these events are rather common in Tordylieae and can occur independently in different lineages. The evolution patterns of psbA-trnH spacer are hypothesized.

A comparison of nrDNA ITS and ETS loci for phylogenetic inference in the Umbelliferae: An example from tribe Tordylieae

The Umbelliferae is a large and taxonomically complex family of flowering plants whose phylogenetic relationships, particularly at low taxonomic levels, are generally obscure based on current and widely used molecular markers. Thus, information on the phylogenetic utility of additional molecular markers at these levels is highly favorable. We investigate the utility of nuclear ribosomal DNA (nrDNA) external transcribed spacer (ETS) sequences for phylogenetic inference in Umbelliferae tribe Tordylieae, a group whose relationships have been previously difficult to resolve owing to low sequence variability, and compare the results to those obtained from the nrDNA internal transcribed spacer (ITS) region. We report that the ETS region evolves at a slightly faster rate and has a higher percentage of parsimony informative characters than that of ITS and all chloroplast DNA loci examined to date. The ETS region is a valuable phylogenetic marker in Umbelliferae for low level analysis, especially when used in combination with ITS.

Towards a standardised terminology for taxonomically important morphological characters in the Umbelliferae

Taxonomically important structural characters and their states are given in an attempt to clarify and standardise the confusing and ambiguous terminology that is currently used in descriptions and identification keys of Umbelliferae taxa. The proposed terms to be used for characters and character states are listed, illustrated and discussed.

Systematic relationships within and between Peucedanum and Angelica (Umbelliferae–Peucedaneae) inferred from immunological studies of seed proteins

Abstract. Immunochemical investigation of the seed storage proteins of some large genera in the tribe Peucedaneae (Umbelliferae) has clearly demonstrated the heterogeneity of Angelica and Peucedanum and the comparative homogeneity of Ferula. Among the taxa included in Angelica, Ostericum appears to be the most distant from the type species A. sylvestris, and should be treated as an independent genus. On the contrary, Archangelica is serologically similar to Angelica s. str. A rather complicated picture has been revealed within Peucedanum s. l. Serotaxonomical studies show a relationship pattern largely corresponding to the infrageneric classification of Thellung (1926). The splitting of satellite genera from Peucedanum s. l. is only partially supported.

Karyotaxonomical analysis in the genusAngelica (Umbelliferae)

Morphometric karyotype characters were studied in 25Angelica spp. (Umbelliferae, Apioideae) and in one species of the related genusTommasinia. For three species the chromosome numbers are new. In our study the majority of the species investigated are diploids with 2n = 22, some are tetraploids with 2n = 44 (for these tetraploids also diploid cytotypes are reported in the literature). Among the diploid species,A. miqueliana has a distinct karyotype consisting of submetacentric and acrocentric chromosomes only, the remaining diploids with 2n = 22 as well as tetraploids with 2n = 44 have rather symmetrical karyotypes, consisting of metacentric and submetacentric chromosomes. The very different chromosome number 2n = 28 has been found inA. gmelinii. Its karyotype includes two distinct groups of chromosomes: 8 pairs of rather large metacentrics and submetacentrics and 6 pairs of very short and asymmetrical chromosomes. Chromosome numbers and structures appear to be useful in the taxonomy of some intrageneric taxa inAngelica.

The tribeSmyrnieae (Umbelliferae) in the light of serotaxonomical analysis

An immunochemical investigation of some Old-WorldUmbelliferae belonging to the tribeSmyrnieae in current classifications has clearly demonstrated that the genera of this tribe do not form a natural monophyletic taxon. The presence of a more or less deep groove on the commissural side of the seeds cannot be regarded as an essential marker of phylogenetic relationships within this family.

Critical taxonomic analysis of Dichoropetalum, Johrenia, Zeravschania and related genera of Umbelliferae-Apioideae-Peucedaneae

Abstract Pimenov, M. G., Kljuykov, E. V. & Ostroumova, T. A.: Critical taxonomic analysis of Dichoropetalum, Johrenia, Zeravschania and related genera of Umbelliferae-Apioideae-Peucedaneae. — Willdenowia 37: 465–502. — ISSN 0511-9618;

An attempt to clarify taxonomic relationships in ‘‘Verwandtschaftskreis der Gattung Ligusticum’’ (Umbelliferae-Apioideae) by molecular analysis

Relationships among the taxa of Umbelliferae, presumably close to Ligusticum and Selinum were investigated by two independent molecular taxonomic methods. 134 ITS 1-2 sequences were analyzed (29 new and 2 reinvestigated species) and immunochemical comparison of storage seed proteins for 38 species of Apioideae of Ligusticum affinity was performed, eight reference systems (antisera) were used. Both approaches yield similar results, showing the extremely polyphyletic nature of this group and some large genera (Ligusticum s.l., Selinum s.l., Pachypleurum) in the Umbelliferae. The independent status of the genera Magadania, Sphaenolobium, Arafoe, Lomatocarpa, Dimorphosciadium and some other segregates of Ligusticum, Cnidium and Selinum have been confirmed, but Cnidium proved to be unnatural even as currently circumscribed. In the group of East Asian taxa the genera Oreocome, Ligusticopsis, Cortia and Cortiella appeared to be closely related. Haplosphaera was shown to be a genus of Hansenia-Notopterygium group.