Tragopogon Species Research Articles

Molecular (ISSR, cp DNA, ITS) and morphological study of the genus Tragopogon L. (Asteraceae)

Tragopogon is an Old World genus with 150 species. Mediterranean, Middle East, and Eastern Europe are the distribution centers of this genus. This genus has 26 species in Iran, of which 11 are endemic. The morphology studies of 32 species and Molecular studies (ISSR, ITS, cp DNA) of 22 species of the genus Tragopogon were investigated. Despite the anatomical and molecular studies done around the world, the exact classification of this genus is not clear due to the high number of secret species, hybridization, polyploidy, and rapid diversification. The purpose of these studies is to classify, determine interspecific relationship in this genus, and determine the important morphological characteristics in taxon differentiation. We conclude that sections of Rubriflori, Sosnowskya, Chromopappus, Majores, Angustissimi, and Krascheninnikovia introduced by the flora of Iranica are confirmed by our morphometry and molecular studies. Section of Profundisulcati in flora Iranica is confirmed based on morphometry data. The Species of T. jesdianus, T. porphyrocephalus, T. rezaiyensis, and T. Stroterocarpus in the flora of Iranica are not classified in any section which we classified in the Rubriflori section. Cp DNA dendrogram is not useful for classification in this genus and Chloroplast sequences are very similar among Tragopogon species. Therefore, the use of cp DNA markers in the classification of this genus is not recommended. The use of ISSR and ITS molecular markers are useful for classifying the genus Tragopogon.

Pollen Morphology of the Genus Tragopogon (Asteraceae)

Tragopogon L. (Cichorioideae, Lactuceae, Scorzonerinae) is an Old World genus with 150 species. Pollen morphology has proved useful in the systematics of some genera and species of Asteraceae as well as in that of some of its genera and species. The pollen morphology of 24 taxa of the genus Tragopogon was investigated in detail by scanning electron microscopy (SEM). The pollen grain type ranged from suboblate, oblate-spheroidal to prolate-spheroidal in equatorial view and hexagonal, obtuse-hexagonal to hexagonal-angular in polar view. In this study separation of the species of the sections Majores, Profundisulcati, Sosnovsky, Chromopappus, Rubriflori according to Flora Iranica is presented from the other species of Tragopogon. T. jezdianus, T. porphyrocephalus, T. rezaiyensis are suggested to belong to Rubriflori section. The results indicate that the palynological characters of the genus Tragopogon are valuable for taxonomic applications and are useful for classification.

Comparison of sand burial-dependent growth responses of two Tragopogon species from different habitats

Wind-driven sand movement leading to sand burial is an essential environmental factor in coastal sand dunes; therefore, plants native to sand dunes need to possess specific adaptations. The present study aimed to compare responses to variable sand burial intensity of Tragopogon heterospermus, rare plant species native to coastal dunes, and Tragopogon pratensis taxonomically related grassland species. Plants in culture were established from seeds collected in natural habitats, cultivated in an automated greenhouse, and individuals of different age were buried by sand in the different depths. Both plants had basic sand burial tolerance, seen as an elongation of adult plants’ leaf bases under moderate sand burial conditions. However, only dune-adapted T. heterospermus plants exhibited efficient resource allocation from roots to shoots with increased sand accretion intensity. T. heterospermus plants had pronounced age dependence of burial tolerance, with higher sensitivity to sand burial at an earlier stage due to small shoot height and, possibly, certain physiological factors.

The genus Tragopogon (Asteraceae): A review of its traditional uses, phytochemistry, and pharmacological properties.

Species of Tragopogon are used in traditional medicine, and consumed as vegetables across the world. In terms of the medicinal uses of Tragopogon, different species have found use in traditional medicine, including uses for wound-healing, treatment of gastrointestinal and hepatic complaints, cancer, kidney and liver dysfunction, inflamed skin and certain cutaneous diseases, as well as constipation, fatigue and anoxia. The aim of this review is to highlight and critically summarize those species of the genus that have been studied as a source of interesting lead compounds, and their traditional uses and bioactivities. A comprehensive and systematic review of literature on traditional uses, phytochemicals and pharmacological properties of the genus Tragopogon was carried out. Information was retrieved from secondary databases such as Scopus, Chemical Abstracts Services (Scifinder), Pubmed, Google Scholar and ScienceDirect, in addition to primary sources including books, PhD and MSc dissertations, and official websites. Species names were validated using "The Plant List" (www.theplantlist.org). The taxa of the genus Tragopogon are known for their local and traditional uses as medicine for treatment of various diseases, and have been consumed as vegetables and snacks for generations in Eurasia, the Mediterranean, Caucasus, Europe and North America. From the approximately 110 species of Tragopogon, only twelve species have been scientifically evaluated for their bioactivity and/or phytochemical composition. Tragopogon species are a rich source of phytochemical constituents and among those that have been identified are 19 flavonoids, 35 terpenoids, seven bibenzyl derivatives, five benzylphtalides, six stilbenes, nine dihydroisocoumarin derivatives, nine phenylmethane derivatives, three hydroxyphenylacetic acid derivatives, four phenylpropane derivatives, four esters of phenylpropanoic acids, a coumarin derivative, and a spermine derivative. Various extracts of the taxa, in addition to the isolated compounds, demonstrated pharmacological properties such as antitumor, antimicrobial, anti-inflammatory and enzyme inhibitory activities, in addition to hepatoprotective, antihyperlipidaemic and wound-healing properties. This review highlights the traditional uses, phytochemistry and pharmacological properties of the few studied taxa of the genus Tragopogon. Some of the reviewed papers were not of an appropriate methodological standard. For instance, phytochemical profiles were not determined, and the fundamental requirements of the pharmacological properties were not defined such as including appropriate positive and negative controls, and calculating the MIC values. Furthermore, these studies did not provide an in depth evaluation of bioactivity of the extracts and the isolated compounds or in vivo experiments which could indicate therapeutic relevance. From a phytopharmacological point of view, this review recommends more high quality evidence-based research on Tragopogon species for further development of plant-derived remedies and compounds.

Ecological-anatomical characteristics of some Tragopogon (Asteraceae) species of the flora of Ukraine

The results of the study of the anatomical structure of assimilating organs of the three species of the genus Tragopogon are described. Nowadays the anatomical structure of vegetative organs of representatives of many taxa have already been researched. The in-depth study of the anatomical structure of species of the genus Tragopogon which was conducted revealed the presence of different ways of structural adaptation of plants to growing on sands, on the one hand, and, on the other hand, revealed that the investigated taxa have structurally different leaves and stems, whose main features do not depend on the influence of conditions of existence. Almost all species of the genus Tragopogon are mesoxerophytes, the purpose of this work was to investigate the species of the Tragopogon genus which inhabit sandy soil. Such researches will be useful for solving problems of genus taxonomy and for detection of adaptive features of plants. The specimens for investigation were collected from a natural habitat. The plants were in the generative phase of development. Leaves and stems in the middle of their length were conserved for investigation. As a result, we detected features which distinguish species but they can not be used for taxonomy, as they depend on ecology. Thus, the studied species differ in the length of leaf lamina. T. ucrainicus has the thinnest lamina and T. orientalis has the thickest one. In addition, T. orientalis has the most laminaled mesophyll (it is distinctly multilayered). The type of mesophyll varied from isolateral (T. borysthenicus and T. ucrainicus) to pseudo-isolateral (T. orientalis). The numbers of layers of palisade parenchyma (both adaxial as well as abaxial) hardly varied at all between species, except that a third layer of cells was rarely represented in T. orientalis. The species differ in the degree of development of spongy parenchyma. It is most developed in T. orientalis (five or six layers of cells) and it is less developed in T. borysthenicus (two layers of cells). There are differences between the species in the anatomical structure of the stem. Thus, T. borysthenicus has the most developed chlorenchyma (four or five layers) and in T. orientalis it is less developed (two or three layers of cells). Cholophylless parenchyma is represented by the greatest number of layers in T. borysthenicus and by a smaller number of layers in T. ucrainicus, it is less developed in T. orientalis. The conduction system of the three investigated species is developed approximately equally. As the result of the analysis, it is possible to conclude that in the anatomical structure of all investigated species of the genus Tragopogon there is a combination of xeromorphic and mesomorphic features . T. borysthenicus from the Black Sea Biosphere Reserve exhibits the most xeromorphic features. The least xeromorphic characteristics are shown by T. orientalis from the sandy terraces of the Desna river (Chernihiv Region). We singled out a complex of anatomical features which can be additional diagnostic ones for the investigated species. T. borysthenicus is characterised by the presence of cortical conducting vascular bundles and by the placement of laticifers in the perimedullar region of the pith; in T. ucrainicus laticifers are also in the perimedullar region but cortical vascular bundles are absent; cortical vascular bundles are observed in T. orientalis but laticifers are placed all over the pith.

Tragopogon erostris Boiss. & Hausskn. (Asteraceae: Cichorieae, Scorzonerinae), the correct name for T. maroofii Mahmoodi & Safavi and T. kurdicus Safavi & Maroofi

The Eurasian genus Tragopogon Linnaeus (1753: 789) comprises approximately 150 species (Mavrodiev et al. 2012, Bell et al. 2012). Twenty six species of Tragopogon were treated in Flora Iranica (Rechinger 1977) as occurring in Iran, of which 10 species were considered endemic to Iran. The same number of species was retained in the new Flora of Iran (Safavi et al. 2013).

Carpology of the genus Tragopogon L. (Asteraceae)

67 species of Tragopogon were investigated with regard to fruit anatomy. The outer achenes, especially the beak and the central part of the seed-containing body, provide the most valuable features (diameter and outlines of the body and the beak, and arrangement of the mechanical elements in the body parenchyma). Some specimens of widely distributed taxa (e.g. T. capitatus, T. dubius, T. pratensis, T. pseudomajor) show variation in the character set and require more investigation prior to further taxonomic treatment. The species studied are classified into informal groups to demonstrate the diversity of carpological traits within the genus, and a comparison is made with the existing molecular phylogeny. The separation of the genus Geropogon from Tragopogon is supported by the achene anatomy.

A cryptic taxon rather than a hybrid species of Tragopogon (Asteraceae) from the Czech Republic

Tragopogon ×mirabilis Rouy is described as a diploid hybrid between T. porrifolius and T. pratensis. A population of T. ×mirabilis from Central Bohemia, Czech Republic, was recently investigated and, unlike previous reports of T. ×mirabilis, was found to be highly fertile. This fertile diploid hybrid population was considered to represent an alternative evolutionary pathway to polyploidy in Tragopogon. To determine the parentage of the plants from Bohemia, we investigated 12 samples of T. ×mirabilis with ITS, ETS, LFY and plastid (rpL16 gene, intron 1, tRNA-Leu (trnL) gene, intron, trnL-trnF intergenic spacer, psbA-trnH intergenic spacer, and trnG-trnT intergenic spacer) sequence data. None of the Bohemian plants have sequences that are consistent with a hybrid origin between T. porrifolius (incl. T. australis) and T. pratensis. Our data suggest that this fertile population of “T. ×mirabilis” may represent an unrecognised diploid species from the Angustissimi clade sensu Mavrodiev et al. (Int. J. Pl. Sci. 164: 1 – 19, 2005), a clade with a centre of distribution in the Caucasus, and hybrids of this unknown species with T. orientalis or T. hayekii (= T. orientalis L. var. hayekii Soo), a species closely related to T. pratensis and native to Bohemia. The Bohemian population of “Tragopogon ×mirabilis” clearly requires more investigation, but based on our data it does not appear to represent T. porrifolius × T. pratensis.

Tragopogon lainzii a New Species of Tragopogon (Asteraceae) Segregated from T. dubius: Evidence from Morphological and Molecular Data

Abstract Tragopogon dubius is one of the most widespread species of Tragopogon, extending across much of Eurasia. Traditionally, T. dubius has been considered a morphologically homogeneous species that includes all Tragopogon collections with yellow flowers and swollen peduncles under capitula. Here we describe a new species of Tragopogon from the Iberian Peninsula, T. lainzii, which has heretofore been included in T. dubius. To this end, we performed comparative morphological, cytogenetic, and molecular analyses on many populations of both species. Our results show that T. dubius is not a homogeneous species and that different lineages exist across its broad geographic distribution. Moreover, we show that hybridization has occurred in the wild between sympatric populations of T. dubius, T. lainzii, and T. porrifolius.

Tomato spotted wilt virus Found in Five Species of the genus Tragopogon in a Florida Greenhouse.

An obviously unhealthy plant identified as Tragopogon mirus Ownbey (remarkable goatsbeard) was sent for diagnosis to the Division of Plant Industry (DPI), Gainesville, FL in May of 2008. T. mirus is a recently formed allotetraploid that has T. dubius Scop. and T. porrifolius L. (goatsbeard or salsify) as parents. The parents (family Asteraceae) are diploid and originate from Eurasia. They were introduced to the northwest United States in the early 1900s. The allotetraploid T. mirus, which does not occur in Eurasia, was discovered in 1949 and named in 1950. It has been found in the northwest states of Washington and Idaho. It has also been found in Arizona (4). The plant sent to the DPI was grown in a greenhouse for research purposes at the Botany Department of the University of Florida (Alachua County). Symptoms exhibited on the leaves included mottling, chlorotic and necrotic spots, and mild distortion. Epidermal leaf strips from a mottled leaf were stained with the Orange-Green protein stain and Azure A nucleic acid stain (1). With a light microscope, granular inclusions typical for Tomato spotted wilt virus (TSWV) (1) were seen in leaf strips from both stains. The remainder of the leaf was ground in buffer and tested serologically for TSWV by TSWV-specific ImmunoStrips (Agdia, Elkhart, IN). The ImmunoStrip was positive for the presence of TSWV. This test was confirmed by double-antibody sandwich-ELISA using antiserum and conjugate for TSWV (Agdia). Further serological testing of other Tragopogon species with similar symptoms growing in the same greenhouse revealed that T. miscellus (another recently formed allotetraploid found in the northwestern United States; parents T. dubius and T. pratensis), T. dubius, T. porrifolius, and T. pratensis were also infected with TSWV. Total RNA was extracted from symptomatic leaves of T. mirus, T. dubius, T. porrifolius, and T. miscellus. Reverse transcription-PCR was performed with universal tospovirus primers BR60 and BR65 that amplify part of the nucleocapsid protein gene (2). Target amplicons of 454 bp were produced for all four samples. The PCR product from T. porrifolius was cloned and sequenced. The resulting sequence (GenBank Accession No. FJ655913) shows high homology, 98%, to several isolates of the Tomato spotted wilt virus deposited in the GenBank (Accession Nos. AY870391, AY744477, and AF020659). T. porrifolius has been reported to be naturally infected with TSWV in Italy (3); however, to our knowledge, this is the first report of this virus in the allotetraploids T. mirus and T. miscellus and in the diploids T. dubius and T. pratensis. This report adds five new Asteraceae weeds to the list of possible reservoirs of TSWV in the United States.

Polyphyly of Tragopogon porrifolius L. (Asteraceae), a European Native with Intercontinental Disjuncts

Tragopogon L. comprises 150 species from Eurasia, with three diploids well established in North America (T. porrifolius, T. dubius, and T. pratensis). ITS, external transcribed spacer, and plastid sequence data indicate that T. porrifolius is polyphyletic. One lineage is phylogenetically well separated from other collections identified as T. porrifolius. This lineage is widespread in North America and Europe, includes the cultivated salsify, and has been considered T. porrifolius ssp. porrifolius. It is distinct from other T. porrifolius in having longer ligules and an additional set of chromosomal satellites. This “salsify” lineage is one parent (with T. dubius) of the North American allotetraploid T. mirus. Collections identified as T. porrifolius ssp. australis and ssp. cupani appear together in a clade well removed from ssp. porrifolius. These subspecies do not appear to have been introduced into North America. Available data suggest that ssp. cupani comprises both diploid and putative allotetraploid populations; hence, as currently described, this subspecies may encompass a previously unrecognized species. Additional samples of T. porrifolius not classified to subspecies, from both Europe and North America, form a third lineage and appear with collections of several species from Europe, including specimens identified as T. longirostris (which itself may not be monophyletic), another purple‐flowered Tragopogon. This lineage appears to represent an unrecognized species and indicates that an additional species of Tragopogon has been introduced into North America.

The biology of Canadian weeds. 110. Tragopogon dubius Scop., Tragopogon pratensis L., and Tragopogon porrifolius L.

Several types of salsify or goat's-beard, Tragopogon species (Asteraceae), are found in Canada as biennial or monocarpic perennial herbs with yellow or purple flowers. Introduced from Eurasia, T. pratensis and T. dubius have become established in all provinces in Canada except Newfoundland and the territories. Tragopogon porrifolius tends to be more local, but is also found in southern areas of most provinces. Tragopogon species hybridize readily, and the tetraploid species T. mirus and T. miscellus have resulted from crosses among the three diploid Tragopogon species in North America. Salsify species produce relatively large umbrella-like pappuses which promote long-distance dispersal. These species also invade rangeland, and are considered noxious weeds in some parts of Canada. Key words: Tragopogon dubius, Tragopogon pratensis, Tragopogon porrifolius, Tragopogon mirus, Tragopogon miscellus, salsify, goat's-beard, Asteraceae, Compositae, hybridization, weed biology

Fruit morphology in Tragopogon L. (Compositae: Lactuceae) from the Iberian Peninsula

Abstract Achenes of all the Tragopogon species from the Iberian Peninsula were examined by means of scanning–electron microscopy and stereomicroscopy. The achenes of the eight species are described, illustrated and compared. The results are contrasted with the systematics of this genus. The isolated position of T. lamottei with regard to the other seven species is noted. A key is provided to enable the different species to be distinguished.

Lectotypification of five Linnaean species of Tragopogon L. (Compositae)

SummaryDíaz de la Guardia, C. & Bianca, G.: Lectotypification of five Linnaean species of Tragopogon L. (Compositae). — Taxon 41: 548–551. 1992. — ISSN 0040‐0262.Lectotypes are designated for Tragopogon pratensis L., T. orientalis L., T. porrifolius L., T. crocifolius L. and T. villosus L. The latter is considered to be a name of uncertain application, at least as far as the Iberian Peninsula is concerned.

OWNBEY'S TRAGOPOGONS: 40 YEARS LATER

Two of the classic examples of recent allopolyploid speciation are Tragopogon mirus and T. miscellus. Previous studies have documented that both allotetraploids originated within the past 50–60 years; the diploid parents of T. minis are T. dubius and T. porrifolius; those of T. miscellus are T. dubius and T. pratensis. It has now been 40 years since these allotetraploids were first described by Ownbey in 1950. To assess whether population size (the absolute number of plants) and population number of these species have changed during the past 40 years, we determined their distribution and numbers in the Palouse region of eastern Washington and northern Idaho in the spring of 1990. We visited 90 localities, 60 in Washington and 30 in Idaho. Tragopogon minis is found in nine locations, with eight of these sites located in eastern Whitman County, Washington. Tragopogon miscellus is much more widespread, occurring in 38 of 90 locations. The latter species is now one of the most common weeds in the vicinity of Spokane, Washington. Comparison of the current distributions of these species with historical records indicates that both allotetraploids, especially T. miscellus, have increased substantially in both geographic range and numbers. These Tragopogon species offer an excellent model system for testing questions concerning the ecology and population biology of recently derived allotetraploid species and their diploid progenitors.

Electrophoretic Evidence for Hybridization Between Tragopogon mirus and T. miscellus (Compositae)

A mixed population of the two recently derived allopolyploid species of Tragopogon, T. mirus and T. miscellus, was discovered in Pullman, Washington. The presence of morphologically intermediate plants suggested that interspecific hybridization had occurred. Evidence from enzyme electrophoresis confirmed the occurrence of hybridization between T. mirus and T. miscellus. Each species possessed unique enzyme bands at three of the eleven enzyme loci examined. Seedlings obtained from the presumed F1 hybrid plants combined the enzyme bands of the two parental species, and displayed additive enzyme phenotypes. Because the seedlings were F1 or later generation plants, or the result of backcrossing, individual seedlings did not display additivity at all three loci. Although diploid species of Tragopogon have provided classical, thoroughly documented examples of interspecific hybridization since 1759, this is the first report of naturally occurring hybridization between the newly derived tetraploid species. Interspecific hybrids have been observed in Tragopogon (Compositae) for nearly two centu- ries and the genus has furnished unusually fa- vorable material for the study of hybridization (Mertens 1972; Ownbey 1950) and polyploidy (Roose and Gottlieb 1976). Although hybrid- ization occurs frequently between diploid species of Tragopogon, natural hybridization be- tween the recently derived amphidiploid species, T. mirus Ownbey and T. miscellus Own- bey, has not been reported. Based on morphol- ogy, interspecific hybridization appeared evi- dent in a mixed population of T. miscellus and T. mirus in Pullman, Washington. Tragopogon is an old-world genus represented in North America by three introduced diploid species, T. dubius Scop., T. porrifolius L., and T. pratensis L. The three species are widespread on this continent and highly sterile F1 hybrids are formed in areas of sympatry (Ownbey 1950). The two tetraploid species, T. mirus and T. mis- cellus, originated near Pullman, Washington and Moscow, Idaho, and are now common weeds in this area. Ownbey (1950) clearly documented their parentage utilizing evidence from mor- phology, meiotic analysis, and genetic analysis of floral color: T. dubius x T. porrifolius gave rise to T. mirus, and T. dubius x T. pratensis gave rise to T. miscellus. Ownbey and McCollum (1953, 1954) later provided evidence for cytoplasmic inheritance and repeated independent origins for the two tetraploid species. Finally, Roose and Gottlieb (1976) utilized enzyme electro- phoresis to document isozyme multiplicity in

Karyotypes of Tragopogon (Compositae: Lactuceae)

Karyotypes of 24 diploid (2n=12)Tragopogon species are similar with one long pair of chromosomes (A), two medium-length pairs (B and C), and three short pairs (D, E, and F). These species may be divided into three karyotypic groups: 1) seven species with a satellite on A and on D; 2) 14 species with a satellite on A only; and 3) three species with a satellite on D only. Most species within karyotype groups may be separated from each other either by distinctive features of certain chromosomes or by statistical differences in length of chromosome arms or long arm: short arm ratios of chromosome A. Three tetraploid (2n=24) species had two long pairs (A,A′), four medium-length pairs (B,B′;C,C′), and six short pairs (D,D′;E,E′;F,F′). Suggestions are made as to the putative diploid parents of these presumed allotetraploids.

Windkan�le f�r die Untersuchung anemochorer Verbreitungseinheiten

New constructions of a vertical and a horizontal wind tunnel are described. Their function is demonstrated by comparative and quantitative analyses of anemochorous dispersal units. The results for 6 Angiosperm genera with pogonochorous, lophochorous and pterochorous dispersal units, and for 2 species ofTragopogon indicate remarkable differences and a broad range of dispersal effectiveness.

Chromatographic Comparison of Tragopogon Species and Hybrids

Approximately 1,700 plants representing five species of Tragopogon (Compositae) and their F1 and F2 hybrids were analyzed by two-dimensional descending paper chromatography. Each species, or population within a species, was chromatographically distinct. Often, however, the differences were more quantitative than qualitative. The chromatographic data generally supported the species relationships which had been determined from previous morphologic, hybridization, and fertility studies. Inheritance of the flavonoid compounds was usually additive in the F1's. Segregation and recombination of the genes controlling the synthesis of these compounds sometimes approximated 3:1 or 9:7 ratios in the F2's. Occasionally parental compounds were missing from some of the hybrids. “Hybrid” compounds which had not been found in either parent were absent from the F1 but did occur in several F2 populations. Two linkage groups were present. The first contains genes controlling the synthesis of three compounds and the second, four compounds.

CHROMATOGRAPHIC COMPARISON OF TRAGOPOGON SPECIES AND HYBRIDS

Approximately 1,700 plants representing five species of Tragopogon (Compositae) and their F1 and F2 hybrids were analyzed by two‐dimensional descending paper chromatography. Each species, or population within a species, was chromatographically distinct. Often, however, the differences were more quantitative than qualitative. The chromatographic data generally supported the species relationships which had been determined from previous morphologic, hybridization, and fertility studies. Inheritance of the flavonoid compounds was usually additive in the F1's. Segregation and recombination of the genes controlling the synthesis of these compounds sometimes approximated 3:1 or 9:7 ratios in the F2's. Occasionally parental compounds were missing from some of the hybrids. “Hybrid” compounds which had not been found in either parent were absent from the F1 but did occur in several F2 populations. Two linkage groups were present. The first contains genes controlling the synthesis of three compounds and the second, four compounds.