Tribe Cynoglosseae Research Articles

Phylogenetic relationships within the subtribe Cynoglossinae (Cynoglossoideae: Boraginaceae): new insights from nuclear and plastid DNA sequence data

Cynoglossinae with ca. 200 spp. is one of the taxonomically most challenging subtribes of tribe Cynoglosseae with regard to generic delimitations. Here, we present a phylogenetic analysis based on nuclear ITS and cpDNA trnL-F and rps16 sequences of 270 accessions including 102 newly sequenced ones, representing all currently recognized genera of Cynoglossinae to clarify relationships among Cynoglossum and allied genera. We conducted maximum likelihood, maximum parsimony, and Bayesian inference analyses on separate and combined datasets. Cynoglossinae is recovered as a monophylum of closely related genera. Subtribe Cynoglossinae falls into three major groups: (1) Clade I including Microparacaryum spp., (2) Clade II comprising two species of Lindelofia, several Cynoglossum and Paracynoglossum spp., and (3) a larger clade (Clades III + IV) consisting of Solenanthus, Rindera, Trachelanthus, Paracaryum, Pardoglossum, Mattiastrum, and the remainder of Cynoglossum. None of the genera were monophyletic in the combined nuclear-plastid and nuclear datasets, except for Microparacaryum. A close relationship between Solenanthus apenninus and Cynoglossum officinale is shown, which suggests transferring S. apenninus to Cynoglossum. Moreover, Pardoglossum atlanticum, the type of the genus, is placed within Mediterranean Cynoglossum corroborating its synonymy under Cynoglossum. Lindelofia olgae is distantly related to the type of the genus (L. longiflora), but more closely connected with Solenanthus core-group. The majority of species currently recognized as members of Rindera do not form a clade with the type of the genus (R. tetraspis). The genus Paracaryum as currently circumscribed is paraphyletic and includes some species of Mattiastrum and Rindera.

Foliar and stem epidermal anatomy of the tribe Cynoglosseae (Boraginaceae) and their taxonomic significance

Foliar and stem epidermal anatomical features of the tribe Cynoglosseae have been studied in detail for the taxonomic identification using light microscopy (LM) and scanning electron microscopic (SEM) techniques. A comparative study was conducted on different plant parts (leaf and stem epidermal anatomy) of 25 species belonging to eight genera of the tribe Cynoglosseae (Boraginaceae) collected from different phytogeographical regions of Iran for the first time. Different qualitative and quantitative characteristics were observed in detail using LM and SEM. Results showed that although generally the stem and leaf anatomical traits were similar, but some diagnostic features were examined for distinguishing the closely related genera in the tribe. The ratio of cortex/diameter of stem and phloem/xylem, the average row number of collenchyma, palisade and spongy cells, structure of trichomes, type of indumentum and palisade arrangement were found taxonomically important. The anatomical characters were statistically analyzed using cluster analysis and principal component analysis. The study found that stem and leaf eccentrics are variable in the genus but constant within species of the same genus. Most species had typical isobilateral leaves, but some showed an incipient dorsoventrally symmetry with a layer of abaxial palisade tissue. Eglandular trichomes were observed found in all the studied species, which were recognized based on structure and function. In present study some novel characters have been observed which are of great interest to the taxonomist for the correct identification some genera delimitations. The characters studied here are of less taxonomic value and delimitating at species level.

Taxonomic identification in the tribe Cynoglosseae (Boraginaceae) using palynological characteristics

The palynomorphological characteristics of 31 species belonging to 8 genera (Paracaryum, Mattiastrum, Microparacaryum, Rindera, Cynoglossum, Solenanthus, Trachelanthus and Lindelofia) of the tribe Cynoglosseae (Boraginaceae) were studied in detail. Palynological observation of the tribe revealed that shape of pollen grains was frequently prolate, subprolate, prolate spheroidal and preprolate. The similarity in some features of the pollen shape and apertural type has been found in several genera such as Cynoglossum, Trachelanthus and Lindelofia. The main ornamentation type was psilate-punctate, psilate-perforate, microgranulate, punctate-microreticulate, scabrate, reticulate, microechinate, punctate-perforate. The species were classified into two pollen types: (i) Rindera tetraspis type and (ii) Cynoglossum officinale type on the basis of aperture characters and surface ornamentation. In present research both genera and species were delimited in the tribe Cynoglosseae. Main objective of the present research is to study different palynological characters for the identification and species delimitation in the tribe Cynoglosseae (Borginaceae). The present findings of the palynological characters are of great interest for the taxonomists for the accurate identification of genera and species in tribe Cynoglosseae. The micro-morphological characteristics of pollen show many novel features for the correct taxonomic identification and species delimitations and this study provide many valuable information to plant taxonomists for further studies.

<div class="page" title="Page 1"><div class="layoutArea"><div class="column">Dwarves on the roof of the world: A taxonomic revision of the Himalayan Lasiocaryeae Weigend (Boraginaceae)</div></div></div>

Recent molecular studies have contributed significantly to our understanding of the systematics and phylogeny of Boraginaceae. A morphologically interesting placement within this plant family is the sister-group relationship between two Himalayan genera of dwarf annuals (Lasiocaryum and Microcaryum) and the genera Trichodesma and Caccinia, a clade that is sister to the remainder of tribe Cynoglosseae. This study investigates the systematics, taxonomy and relationships of Lasiocaryum and Microcaryum and compares it with other, morphologically similar genera, Setulocarya and Chionocharis. Together, Lasiocaryum, Microcaryum, Setulocarya and Chionocharis include seven species as currently recognized. However, morphological studies show that the monotypic Setulocarya should be treated as a synonym of Lasiocaryum munroi while L. ludlowii is best placed within L. trichocarpum, leading to a total of five species within this clade. Sequences from three chloroplast markers and one nuclear region generated for four of the five species here recognized confirm the monophyly of the clade Chionocharis [Microcaryum + (Lasiocaryum trichocarpum + L. munroi)]. This exclusively Himalayan lineage is morphologically homogeneous, and consists of tiny annual herbs and perennial cushion plants with small, turbinate to oblong-ovoid, pubescent or rugulate nutlets.

Chemotaxonomic perspectives of the Paracaryum (Cynoglosseae, Boraginaceae) taxa based on fruit fatty acid composition

Paracaryum is a medium sized genus in Cynoglosseae. This study represents the most comprehensive phytochemical investigation of Paracaryum to date. The fatty acid compositions of the fruits of ten Paracaryum taxa belonging to three different subgenera were investigated for chemotaxonomic allocation using gas chromatography. The fatty acid profiles of seven Paracaryum taxa, five of which are endemic to Turkey, were recorded for the first time. Among the twenty-two analysed fatty acids, oleic, linoleic and α-linolenic acids were the major fatty acids represented. The oleic acid content ranged from 22.1% in P. hirsutum to 51.7% in P. lithospermifolium subsp. cariense var. erectum; linolenic acid content ranged from 8.6% in P. lithospermifolium subsp. cariense var. erectum to 20.7% in P. erysimifolium; α-linolenic acid content ranged from 7.5% in P. lithospermifolium subsp. cariense var. erectum to 13.5% in P. cristatum subsp. cristatum; gamma linolenic acid content ranged from 2.8% in P. erysimifolium to 6.0% in P. hirsutum. Additional fatty acids also displayed varying levels in different species; palmitic acid content accounted for 17.7% in P. erysimifolium, erucic acid content was 8.73% in P. strictum, eicosenoic acid content was 6.0% in P. cristatum subsp. cristatum, eicosadienoic acid content was 4.4% in P. hirsutum, and stearic acid content was 4.3% in P. erysimifolium. The classification of the tribe Cynoglosseae remains controversial despite the many intensive morphological and phylogenetic investigations that have been carried out. Our fatty acid data from Paracaryum were analysed together with previously recorded fatty acid data from Cynoglosseae s.l. taxa to examine the chemotaxonomic contribution to the classification among taxa in Cynoglosseae by multivariate methods, including the unweighted pair group method with arithmetic mean and principal component analysis. An assessment of these chemometrics data supported the chemotaxonomic position of the genus Paracaryum in the tribe Cynoglosseae. While the principal component graphic did not depict clear separation of the three subgenera of Paracaryum, the principal component analysis revealed the chemotaxonomic significance of palmitic, linoleic, capric, and oleic acids.

Molecular phylogenetics and morphology support two new genera (Memoremea and Nihon) of Boraginaceae s.s.

Omphalodes (Boraginaceae s.s., Cynoglosseae s.l.) comprises ca. 29 species of annual and perennial plants distributed in three main disjunct areas: Western Palearctic, Japan and SW of North America. This paper uses micromorphological and DNA data to re-assess the monophyly and taxonomy of the genus. Morphological characters of 15 species of Omphalodes and four closely-related genera were analysed using SEM. A total of 82 ITS and 68 trnL–trnF sequences were newly sequenced, including 14 species of Omphalodes and three genera of the tribe Cynoglosseae. Phylogenetic analyses of 57 genera (186 species) of Boraginaceae indicated that Omphalodes as currently circumscribed is formed by three independent lineages, which were supported by morphological characters of the fruit. As a result, and in the interest of a more natural classification, two new genera are described to accommodate Omphalodes scorpioides (Memoremea) from Europe and the Japanese species (Nihon). Memoremea is distinguished from all the other species previously included in Omphalodes by the apical attachment scar and the hollow nutlet margin. Nihon is easily discriminated by the abrupt change of margin ornamentation towards the nutlet aperture. We also provided a taxonomic treatment that proposes the lowest number of nomenclature changes, although six new combinations are required.

A NEW SUBSPECIES OF OMPHALODES LUCILIAE (BORAGINACEAE) FROM TURKEY

Omphalodes luciliae Boiss. subsp. pisidica R.R.Mill subsp. nov. (Boraginaceae, tribe Cynoglosseae) is described from Isparta and Antalya vilayets, Turkey. Its relations with the other subspecies of Omphalodes luciliae are discussed.

Oncaglossum, a New Genus of the Boraginaceae, Tribe Cynoglosseae, from Mexico

A new monotypic genus, Oncaglossum Sutorý, endemic to central Mexico, is described based on Cynoglossum pringlei Greenm. from the state of Mexico and belongs within the tribe Cynoglosseae of the Boraginaceae. The new genus is related to Cynoglossum L. subg. Eleutherostylum (Brand) Riedl and can be distinguished from the North American species currently classified in Cynoglossum by its calyx with imbricate, colored, and broadly ovate sepals, the corolla lobes reduced to narrow strips, anthers on short filaments inserted on the corolla margin, the stigma in the form of a lenticular disc, and tricolporate pollen grains with an additional aperture in their polar region.

Palynological contribution to the systematics ofRinderaand the allied generaParacaryumandSolenanthus (Boraginaceae-Cynoglosseae)

Bigazzi, M., Nardi, E. & Selvi, F.: Palynological contribution to the systematics of Rindera and the allied genera Paracaryum and Solenanthus (Boraginaceae-Cynoglosseae). — Willdenowia 36 (Special Issue): 37–46. — ISSN 0511-9618; © 2006 BGBM Berlin-Dahlem. doi:10.3372/wi.36.36103 (available via http//dx.doi.org/)The pollen of thirteen species of Rindera, ten of Solenanthus and eight of Paracaryum was examined by light and scanning electron microscopy. Two main pollen types are recognised, one typical of most Rindera species (Rindera tetraspis type) and one occurring in all but one species examined of Paracaryum and Solenanthus (Cynoglossum officinale type). Deviations from the Rindera tetraspis type in Rindera were observed in R. tianschanica, which is characterised by the Cynoglossum officinale pollen type, and R. gymnandra, which shows the Pardoglossum atlanticum type as already published. Within Paracaryum and Solenanthus the only remarkable deviation from the Cynoglossum officinale type was found in Solenanthus hupehensis from China. In the constricted shape and the lack of an ectocingulus, the pollen of this species approaches the Myosotis arvensis type, which is rarely found in the tribe Cynoglosseae. Combined with other morphological peculiarities, pollen characters support the placement of S. hupehensis in a subgenus of its own, S. subg. Silvestria, named after the discoverer of this species.

Host Specificity of Longitarsus quadriguttatus Pont., a Below-Ground Herbivore for the Biological Control of Houndstongue

Abstract Cynoglossum officinale is a Eurasian plant that has become an invasive weed of rangelands in the United States and Canada. It is a herbaceous, monocarpic, short-lived perennial. Since there are no satisfactory means of control of houndstongue, a biological control project was initiated in 1988. The first species introduced for the control of houndstongue is a root-mining weevil, Mogulones cruciger, which was first released in Canada in 1997. The flea beetle Longitarsus quadriguttatus is another below-ground herbivore species considered for the biological control of C. officinale. Previous host-specificity investigations with 49 European test-plant species demonstrated that larval development of the flea beetle is limited largely to plant species within the tribe Cynoglosseae in Boraginaceae. However, investigations neither included any native North American Boraginaceae species nor examined results of earlier tests for plant species that were susceptible to attack by the flea beetle. Therefore, additional host-range investigations were conducted between 1994 and 1996. Different no-choice and choice tests were conducted with 7 plant species that showed susceptibility to attack in previous tests and 5 native North American plant species. Experiments were carried out under laboratory and open-field conditions. L. quadriguttatus was host specific in these tests. Two European test-plant species outside the genus Cynoglossum and 2 native North American Cynoglossum species supported complete development of the flea beetle to some extent, but in all cases the test plants were significantly less suitable hosts than houndstongue. The remaining 5 European and 3 native North American test-plant species did not support development and/or survival of L. quadriguttatus. The introduction of L. quadriguttatus into Canada was approved in 1998.

Host specificity of Longitarsus quadriguttatus (Pont., 1765) (Col., Chrysomelidae), an agent for the biological control of hound's tongue (Cynoglossum officinale L., Boraginaceae) in North America

AbstractThe host specificity of the European flea‐beetle Longitarsus quadriguttatus was investigated between 1991 and 1993 as a potential agent for the biological control of hound's‐tongue (Cynoglossum officinale), a weed of European origin that has become naturalized in North America. Life‐history studies showed a good synchronization of the univoltine flea‐beetle with its host plant. Adults feed on leaves and deposit eggs on host plant rosettes. The larvae develop inside the roots. Host specificity screening tests comprising adult feeding‐, oviposition‐ and larval development tests were carried out with 49 European plant species. Adults or larvae exclusively attacked plants in the family Boraginaceae. The 18 test plant species in nine other families remained free from feeding and oviposition and supported no larval development. Adults nibbled on most of the 31 Boraginaceae species tested. Normal adult feeding was mainly found on plants in the tribe Cynoglosseae, which were preferrably attacked in oviposition tests. Larvae developed mainly in roots of Cynoglosseae species. Occasional attack of other Boraginoideae species in laboratory tests could not be verified under field conditions. Host acceptance tests showed normal feeding, oviposition and development on the North American biotype of hound's‐tongue. L. quadriguttatus has a narrow host range and a good potential for controlling the target plant in North America.

The generic position of C.B.Clarke's species of Paracaryum (Boraginaceae)

The classification of three subtropical, primarily Indian, members of the tribe Cynoglosseae (Boraginaceae) which C. B. Clarke included in Paracaryum P. coelestinum (Lindley) Bentham, P. lambertianum C. B. Clarke and P.malabaricum C. B. Clarke – is discussed. The conclusion reached is that Clarke was correct to assign them all to one genus, but that none has any affinity with the mainly Irano-Turanian genus Paracaryum (DC.) Boiss. Cynoglossum L. sect. Paracaryopsis H. Riedl, to which Riedl transferred P. coelestinum, is elevated to generic rank as Paracaryopsis (H. Riedl) R. Mill, with three new combinations at specific rank. A systematic treatment of Paracaryopsis is given.

Pollen morphology in the genus Pardoglossum (Boraginacea) with some observations on heterocolpate pollen

The pollen morphology of the genus Pardoglossum Barbier et Mathez (Boraginaceae) has been described using light and scanning electron microscopy. All five species in this genus have heterocolpate pollen, as do other genera in the tribe Cynoglosseae to which Pardoglossum belongs. The Pardoglossum pollen type is most unusual in having alternating colpi and pori round the equator, the pori bearing the endoapertures. A new term ‘poro-colpate’ has been coined for this condition. Evidence is presented to suggest that heterocolpate grains are more advanced than grains with a singe apertures type, and that poro-colpate grains are more advanced than heterocolpate grains where all the apertures are colpi. Pardoglossm should therefore be considered an advanced rather than primitive genus in the Cynoglosseae.