Galina V. Degtjareva

Inflorescence and Early Flower Development in Loteae (Leguminosae) in a Phylogenetic and Taxonomic Context

Molecular phylogeny shows that the temperate legume tribe Loteae is close to the mostly tropical Robinieae and monogeneric Sesbanieae, but comparative morphological studies of these groups are limited. Unusual patterns of inflorescence symmetry and calyx development have been described in some Loteae, but taxon sampling was low. We studied these features with scanning electron microscopy in 25 species of Loteae plus in three Robinia species. Phylogenetic trees of Loteae based on nuclear ribosomal ITS sequences and 77 morphological characters are constructed. Our data show that whorled flower arrangement is a synapomorphy of Loteae; joint initiation of the two adaxial sepals is a synapomorphy of a clade containing Hippocrepis, Scorpiurus, and Coronilla; floral buds bent backward early in development are a synapomorphy of Coronilla; bilateral umbel symmetry and the presence of a single whorl of flowers are probably primitive within Loteae. Inflorescences of Robinia show no special similarities with those of Loteae. Developmental data support homologies between sterile bracts in all Loteae. Even if the sterile bract is situated at the top of the peduncle, it is morphologically the first leaf on the peduncle. Monosymmetric umbels of Loteae (including the model legume Lotus japonicus) could be useful for investigating genetic control of symmetry in structures of hierarchic levels higher than flowers.

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.

Organization of chloroplast psbA-trnH intergenic spacer in dicotyledonous angiosperms of the family umbelliferae

Chloroplast intergenic psbA-trnH spacer has recently become a popular tool in plant molecular phylogenetic studies at low taxonomic level and as suitable for DNA barcoding studies. In present work, we studied the organization of psbA-trnH in the large family Umbelliferae and its potential as a DNA barcode and phylogenetic marker in this family. Organization of the spacer in Umbelliferae is consistent with a general pattern evident for angiosperms. The 5′-region of the spacer situated directly after the psbA gene is more conserved in length compared to the 3′-region, which has greater sequence variation. This pattern can be attributed to the maintenance of the secondary structural elements in the 5′-region of the spacer needed for posttranscriptional regulation of psbA gene expression. In Umbelliferae only, the conserved region contains a duplication of the fragment corresponding to the loop of the stem-loop structure and an independent appearance of identical sequence complementarities (traits) necessary to stabilize the stem-loop structure in different lineages. The 3′-region of the spacer nearest to trnH ranges greatly in size, mainly due to deletions, and the decrease in spacer length is a general trend in the evolution psbA-trnH in Umbelliferae. The features revealed in spacer organization allow us to use it as phylogenetic marker, and indels seem to be more informative for analyses than nucleotide substitutions. However, high conservation among closely related taxa and occurrence of homoplastic inversions in the stem-loop structure limit its application as DNA barcode.

Redefined generic limits of Carum (Umbelliferae, Apioideae) and new systematic placement of some of its taxa

Abstract Zakharova E. A., Degtjareva G. V. & Pimenov M. G.: Redefined generic limits of Carum (Umbelliferae, Apioideae) and new systematic placement of some of its taxa. — Willdenowia 42: 149–168. December 2012. — Online ISSN 1868-6397;

Phylogenetics of Anthyllis (Leguminosae: Papilionoideae: Loteae): partial incongruence between nuclear and plastid markers, a long branch problem and implications for morphological evolution

Phylogenetic relationships in the genus Anthyllis (Leguminosae: Papilionoideae: Loteae) were investigated using data from the nuclear ribosomal internal transcribed spacer regions (ITS) and three plastid regions (psbA-trnH intergenic spacer, petB-petD region and rps16 intron). Bayesian and maximum parsimony (MP) analysis of a concatenated plastid dataset recovered well-resolved trees that are topologically similar, with many clades supported by unique indels. MP and Bayesian analyses of the ITS sequence data recovered trees that have several well-supported topological differences, both among analyses, and to trees inferred from the plastid data. The most substantial of these concerns A. vulneraria and A. lemanniana, whose placement in the parsimony analysis of the ITS data appears to be due to a strong long-branch effect. Analysis of the secondary structure of the ITS1 spacer showed a strong bias towards transitions in A. vulneraria and A. lemanniana, many of which were also characteristic of certain outgroup taxa. This may contribute to the conflicting placement of this clade in the MP tree for the ITS data. Additional conflicts between the plastid and ITS trees were more taxonomically focused. These differences may reflect the occurrence of reticulate evolution between closely related species, including a possible hybrid origin for A. hystrix. The patterns of incongruence between the plastid and the ITS data seem to correlate with taxon ranks. All of our phylogenetic analyses supported the monophyly of Anthyllis (incl. Hymenocarpos). Although they are often taxonomically associated with Anthyllis, the genera Dorycnopsis and Tripodion are shown here to be more closely related to other genera of Loteae. We infer up to six major clades in Anthyllis that are morphologically well-characterized, and which could be recognized as sections. Four of these agree with various morphology-based classifications, while the other two are novel. We reconstruct the evolution of several morphological characteristics found only in Anthyllis or tribe Loteae. Some of these characters support major clades, while others show evidence of homoplasy within Anthyllis.

Pollination of Vietnamese Aspidistra xuansonensis (Asparagaceae) by female Cecidomyiidi flies: Larvae of pollinator feed on fertile pollen in anthers of anthetic bisexual flowers

UNLABELLED • PREMISE OF THE STUDY Aspidistra is a species-rich, herbaceous monocot genus of tropical Southeast Asia. Most species are recently discovered and apparently endangered, though virtually nothing is known about their biology. Species of the genus are primarily distinguished using flower morphology, which is enormously diverse. However, the pollination process has not been directly observed in the center of diversity of the genus (N Vietnam and S China). Indirect and partly direct data on the only widely cultivated species of the genus (A. elatior) placed it among angiosperms with the most unusual pollination biology, though these data are highly controversial, suggesting pollen transfer by mollusks, crustaceans, flies, or possibly tiny soil invertebrates such as collembolans.• METHODS Pollination of Aspidistra xuansonensis in the center of diversity of the genus was studied using visual observations and videos and light and scanning electron microscopy investigation of flowers and their pollinators. Pollinators and their larvae were molecularly barcoded.• KEY RESULTS Aspidistra xuansonensis is pollinated by female cecidomyiid flies (gall midges). They oviposit on anthers, and larvae develop among the pollen mass. Molecular barcoding proved taxonomic identity of the larvae and the flies. The larvae neither damage floral parts nor cause gall formation, but feed on pollen grains by sucking out their content. The larvae move out of the flowers before decomposition starts. Carebara ants steal developing larvae from flowers but do not contribute to pollination.• CONCLUSIONS More than one kind of myiophily is present in Aspidistra. Brood site pollination was documented for the first time in Aspidistra. The pollination system of A. xuansonensis differs from other kinds of brood site pollination in the exit of the larvae prior to the decomposition of floral parts.

ITS phylogeny of Middle Asian geophilic Umbelliferae-Apioideae genera with comments on their morphology and utility of psbA-trnH sequences

Generic delimitation and relationships within the taxonomically controversial group of geophilic Umbelliferae from Middle Asia are problematic. Geophilic plants in arid regions have a short period of development during spring and early summer and survive unfavorable seasons due to the presence of tuber-like underground storage organs. To investigate the circumscription of recently recognized Middle Asian geophilic genera and their relationships, we conducted phylogenetic analyses using DNA sequence data from rapidly evolving nuclear (nrITS) and chloroplast (psbA-trnH intergenic spacer) regions as well as multivariate analyses of the morphological and ITS data. Our results show that Galagania and Mogoltavia are monophyletic, whereas Bunium, Hyalolaena and Oedibasis are polyphyletic. Elaeosticta, one of the larger genera of geophilic Middle Asian umbellifers, is rendered paraphyletic by the inclusion of the enigmatic Hyalolaenamelanorrhiza. The genus Scaligeria, in which most species of Elaeosticta were initially described, is not grouped with these genera and has other affinities. Our analyses also show that the morphological characters used for the delimitation of genera within this group of Umbelliferae (such as life form, cotyledon number, presence/absence of petiolules of basal leaflets, shape of mericarp ribs, width of mericarp commissure, number of secretory ducts in mericarp furrows) are rather homoplastic. The psbA-trnH sequences were not informative enough to reveal relationships among the genera studied, although some interesting features in the spacer evolution were revealed (maximum reduction in length among Umbelliferae, homoplasious directed substitutions to restore secondary structure).

Are Lotus creticus and Lotus cytisoides (Leguminosae) closely related species? Evidence from nuclear ribosomal ITS sequence data

Lotus creticus is a potentially important perennial legume for soil management in the Mediterranean climate. This plant is in focus of experimental research in different countries. The so-called Lotus creticus group is taxonomically problematic. Although some authors consider all members of the group as a single variable species, others segregate several species distinct from L. creticus, especially L. cytisoides. Recent morphological studies suggested that L. creticus and L. cytisoides are taxonomically distinct at specific level and not even closely related to each other. Two molecular phylogenetic studies published so far and based on one accession of L. creticus each gave conflicting results on relationships of this species. In this paper, sequences of nuclear ribosomal ITS (nrITS) are produced from multiple accessions of both L. creticus and L. cytisoides. Taxonomic identity of each accession has been tested using morphology. The new nrITS sequences are inserted in a data matrix containing a representative set of Lotus species covering all the geographical range and all major taxonomic groups of Lotus. Phylogenetic analysis of nrITS sequence data showed that L. creticus is related to L. assakensis and other species of the section Pedrosia while L. cytisoides is related to L. longisiliquosus and other species of the section Lotea. These results are important for establishing future germplasm collection, breeding and selection programs of perennial Lotus species.

Survey of the genus Diplotaenia (Umbelliferae), with description of two new species from Turkey

Abstract Pimenov M. G, Kljuykov E. V. & Degtjareva G. V.: Survey of the genus Diplotaenia (Umbelliferae), with description of two new species from Turkey. — Willdenowia 41: 67–74. — Online ISSN 1868-6397;

Gradual vs. abrupt reduction of carpels in syncarpous gynoecia: A case study from Polyscias subg. Arthrophyllum (Araliaceae: Apiales)

PREMISE OF THE STUDY Revealing the relative roles of gradual and abrupt transformations of morphological characters is an important topic of evolutionary biology. Gynoecia apparently consisting of one carpel have evolved from pluricarpellate syncarpous gynoecia in several angiosperm clades. The process of reduction can involve intermediate stages, with one fertile and one or more sterile carpels (pseudomonomery). The possible origin of monomery directly via an abrupt change of gynoecium merism has been a matter of dispute. We explore the nature of gynoecium reduction in a clade of Araliaceae. METHODS The anatomy and development of unilocular gynoecia are investigated using light and scanning electron microscopy in two members of Polyscias subg. Arthrophyllum. Gynoecium diversity in the genus is discussed in a phylogenetic framework. KEY RESULTS Unilocular gynoecia with one fertile ovule have evolved at least four times in Polyscias, including one newly discovered case. The two unilocular taxa investigated are unicarpellate, without any traces of reduced sterile carpels. Carpel orientation is unstable, and the ovary roof and style contain numerous vascular bundles without clearly recognizable dorsals or ventrals. In contrast to pluricarpellate Araliaceae and Apiaceae, the cross zone is apparently oblique in the unicarpellate species. CONCLUSIONS No support was found for gradual gynoecium reduction via pseudomonomery. The abrupt origin of monomery via direct change of gynoecium merism and the unstable carpel orientation observed are related to the general lability of the flower groundplan in Polyscias. The apparent occurrence of the unusual oblique cross zone in unicarpellate Araliaceae can be explained by developmental constraints.