Taxonomic importance of seed macro‐ and micro‐morphology in Abelmoschus (Malvaceae)

Seed morphology of Abelmoschus is known to be variable, but patterns of variation have never been critically studied. We studied seed macro‐ and micro‐morphological characters, including seed shape/size, seed coat pattern and trichome density/structure in multiple samples to evaluate the taxonomic significance of seed characters. Among the studied characters, seed shape and trichome structure were found to have major taxonomic importance and proved to be valuable characters for separating taxa. Two main seed types were present: seeds with deciduous trichomes and seeds with persistent trichomes. These characters offer significant evidence to the distinctness of certain species (A. esculentus, A. moschatus subsp. moschatus, A. moschatus subsp. tuberosus, A. crinitus and A. angulosus). Further, our results indicate that A. moschatus subsp. tuberosus should be maintained as a separate subspecies while A. manihot subsp. tetraphyllus var. pungens may be merged in A. angulosus. No significant intraspecific variation was observed, except in A. esculentus. We conclude that seed coat sculpturing and seed trichomes do indeed provide stable and diagnostic characters for many morphologically closely related taxa of Abelmoschus and that LM/SEM techniques can be useful in solving systematic problems and management of Abelmoschus genetic resources.     

Floral sex ratios and gynomonoecy in Aster (Asteraceae)

Gynomonoecy is the sexual system in which female and bisexual flowers occur on the same plant. This system has received little attention despite the considerable work on other plant sexual systems in the past few decades. Our study examines one hypothesized advantage of having two flower types on a plant, namely that this arrangement permits flexibility in allocation of resources to male and female reproductive functions. We examined 16 species of Aster (Asteraceae), a genus of gynomonoecious, perennial herbs. Plants in this genus produce heads consisting of a whorl of female flowers around a cluster of bisexual flowers. Among field-grown plants we found no evidence that plant size, date, position of heads, rainfall, or shade influenced the proportion of female flowers. A series of greenhouse experiments likewise revealed no large or consistent effects of light, nutrients, or position of heads on the proportion of ray flowers. While floral ratios proved very stable in the face of environmental and physiological variables, they exhibited significant variation among plants and among sibships in most species. We conclude that the presence of two flower types in gynomonoecious asters is not advantageous in permitting flexibility in allocation of resources to male and female functions. We believe that the advantage of the female flowers in aster heads lies either in reducing pollen-pistil interference or in attracting pollinators.     

Chromosome number determinations in Aster section Conyzopsis (Asteraceae)

Chromosome numbers forAster brachyactis, A. frondosus, andA. laurentianus were determined to be 2n=14. The latter two are the first documented records for these taxa. The basic chromosome number for sectionConyzopsis is confirmed asx=7.     

Galatella anatolica sp. nov. (Asteraceae: Astereae) from Osmaniye,Turkey

Galatella anatolica Hamzao?lu & Budak sp. nov. (Asteraceae), collected from Osmaniye (Turkey) is here described as a new species. It is similar to G. angustissima (Tausch) Novopokr. in general habit. Both have stems with few branches and 1‐veined middle leaves, but are distinguished by involucral features, series of phyllaries, and lengths of disc florets, achenes and pappus.     

Taxonomic status of the subspecies of Teucrium lamiifolium in Turkey: reevaluation based on macro‐ and micro‐morphology,anatomy and chemistry

Teucrium stachyophyllum P. H. Davis was first described as a separate species in the ‘Syrian, Lebanon and Palestine Flora’, but was reduced to a subspecies of T. lamiifolium d’Urv. in the ‘Turkish Flora’. Studies on the macro‐ and micro‐morphology as well as the anatomy of populations of T. lamiifolium subsp. stachyophyllum (P. H. Davis) Hedge & Ekim and T. lamiifolium subsp. lamiifolium in Turkey have been carried out based on herbarium specimens and field observations. The results from the anatomical studies on stems and leaves show that although the two taxa are similar in stem anatomy, they differ in terms of mesophyll thickness and occurrence of crystals in the upper epidermal cells of the leaf. Light and scanning electron microscope analyzes revealed striking differences in colour, size, sculpture and gland density between nutlets of the two taxa. Morphological studies also showed that there were important differences between the two taxa in terms of inflorescence, indumentum, and stem length and thickness. In addition, the essential oils from aerial parts of T. lamiifolium subsp. stachyophyllum and T. lamiifolium subsp. lamiifolium were isolated by steam distillation and analysed by gas chromatography (GC) and gas chromatography–mass spectrometry (GC–MS). The major components were characterized as β‐caryophyllene (23.5–44.8%), trans‐β‐bergamotene (22.4–26.4%), germacrene D (22.2%) and (Z)‐β‐farnesene (14.0%) for subp. lamiifolium and trans‐β‐bergamotene (38.1–41.1%) and β‐caryophyllene (8.7–8.9%) for subsp. stachyophyllum, respectively. As a result of the studies, it is proposed that the taxon treated as T. lamiifolium subsp. stachyophyllum in the Turkish flora should be raised to specific rank as T. stachyophyllum.     

Aster saxicola sp. nov. (Asteraceae,Astereae), a new species from Guizhou,China: evidence from morphological,molecular and cytological studies

Aster saxicola W. P. Li & Z. Li, a new species of Asteraceae from southeastern Guizhou province, China, is described and illustrated based on morphological, molecular and cytological data. Morphological comparisons showed that A. saxicola is similar to A. oliganthus, but can be distinguished from the latter by its purple abaxial surface of the lower and middle cauline leaves, cordate‐ovate basal leaves with strigose hairs, outer three‐seriate phyllaries with purple or purplish apex, 9–14 ray florets and 10–18 disk florets. Phylogenetic analyses based on nrDNA ITS, ETS and plastid trnL–F DNA sequence data support that A. saxicola is a distinct species and belongs to Aster subgen. Aster sect. Ageratoides. Cytological observation shows that the new species is diploid with a karyotype formula 2n = 2x = 18 = 16m + 2sm. The new species should be considered endangered (EN) based on the IUCN red list criteria because of its restricted geographic range.     

Rib and vertebral micro‐anatomical characteristics of hydropelvic mosasauroids

Houssaye, A. & Bardet, N. 2011: Rib and vertebral micro‐anatomical characteristics of hydropelvic mosasauroids. Lethaia, Vol. 45, pp. 200–209. Mosasauroids are squamates secondarily adapted to an aquatic life that dominated the sea during the Late Cretaceous. Mosasauroids display distinct types of morphologies illustrating steps in the adaptation of this lineage to increasing obligatory habits. Hydropelvic mosasauroids (sensu Caldwell & Palci) were the most highly adapted to an open‐sea environment. Contrary to plesiopelvic forms, they are considered to have relied on a hydrodynamic, rather than hydrostatic buoyancy and body trim control strategies. This led previous authors to consider that these taxa would favour bone lightening (osteoporotic‐like condition) rather than bone mass increase. Although an osteoporotic‐like state was indeed described in Clidastes and Tylosaurus, bone mass increase was reported in Platecarpus. As a matter of fact, the new analysis of vertebral thin sections of various taxa combined with the reanalysis of the rib sections available in Sheldon’s PhD thesis in a micro‐anatomical perspective reveals the absence of both bone mass increase and bone lightening in these organisms. These taxa in fact display a vertebral micro‐anatomy much peculiar within squamates. It characteristically corresponds to a true network of thin trabeculae whose tightness varies between taxa, probably as a result of both species and individual size differences, particularly the latter. In addition, analysis of the pattern of vascularization as observed in hydropelvic mosasauroids, which is unique amongst squamates, suggests that large size in hydropelvic mosasauroids would mainly rely on protracted rather than faster growth rates. □Histology, hydropelvic mosasauroids, micro‐anatomy, ribs, vascularization, vertebrae.     

Bottleneck-induced dissolution of self-incompatibility and breeding system consequences in Aster furcatus (Asteraceae)

Aster furcatus is a rare species with extremely limited genetic variation at isozyme loci. We utilized crossing experiments and seed set data obtained from natural populations to verify that there is also little allelic variation at the self-incompatibility (S) locus. Seed set in several populations was limited by a low number of S-alleles. Associated with a low number of S-alleles in populations was the dissolution of the incompatibility system, manifest by individual variation in self-compatibility, and by complex dominance relationships among S-alleles. Plant self-compatibility was correlated with mean number of ovules per inflorescence. Thus, self-compatibility appeared to be under partial environmental influence. Computer simulations revealed that the shapes of seed set distribution curves of modeled self-incompatible plant populations depend on the number of incompatibility alleles in the populations. By varying the number of S-alleles in modeled populations, we generated seed set distribution curves similar to those of natural populations. Genetic bottlenecks reduce the number of S-alleles and the proportion of compatible matings in populations of multiallelic self-incompatible species. Self-compatible genotypes are at a selective advantage in populations that lack a sufficient number of S-alleles to produce compatible crosses. Aster furcatus appears to be evolving self-compatibility as a result of bottleneck-induced losses of S-alleles.     

A classification of the New World species of Aster (Asteraceae)

In order effectively to list and comment on the results of cytological investigations inAster in a companion paper, a scheme of infrageneric classification is presented which utilizes the basic chromosome number as a pivotal diagnostic character. Reasons are stated as to why, with the exception ofUnamia Greene which is transferred toSolidago, and the commonly recognized generaLeucelene Greene,Machaeranthera Nees andXylorhiza Nutt., none of the segregate genera previously proposed or recorded in the literature is upheld. Instead, these taxa are being given subgeneric or sectional rank. Two additional subgenera are established to accommodate the species groups traditionally placed in “Aster proper,” which are characterized by having basic chromosome numbers ofx = 5 andx = 8, respectively. Altogether ten subgenera of the genusAster, five of them subdivided further into a total of 24 sections, are recognized as having representative species in the New World. All basionyms and type species are listed, and a number of new combinations and status changes are validated in accordance with the International Rules of Botanical Nomenclature. Where known (from literature and personal research), chromosome numbers are recorded for the species.     

Extensive chromosome number variation in Aster ageratoides var. pendulus (Asteraceae)

Aster ageratoides var. pendulus, a recently described taxon, is endemic to Mt Hupingshan of north‐western Hunan, China. Field observations and collections were made from the only known population. Root‐tip squashes were used to determine the chromosome numbers of 96 plants and 61 seedlings from the achenes of eight sample plants. The results show that var. pedulus is a swarm of 30 cytotypes with nearly continuous chromosome numbers from 2n = 60 to 2n = 92. Chromosome numbers of 61 seedlings vary from 2n = 61 to 2n = 91, belonging to 18 cytotypes. The chromosome number variation of var. pendulus is highly unusual not only in the A. ageratoides polyploid complex but also in angiosperms. Such an enormous continuous variation of chromosome numbers could have arisen by the combined effect of hybridization, recent origin and high levels of polyploidy. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 378–387.     

Taxonomic notes on Senecio tomentosus and its allies (Asteraceae)

Distribution maps, specimen citations and taxonomic commentary are provided for 14 species ofSenecio in temperate North America allied withS. tomentosus (the “Tomentosi”). These data support the author’s treatment of the group inNorth American Flora.     

Identification and characterization of eight microsatellite loci in Aster amellus L. (Asteraceae)

Eight polymorphic microsatellites were developed in the perennial herbaceous Aster amellus L. (Asteraceae) and characterized on three populations from France and Switzerland. The number of alleles ranged between four and 30 depending on the locus, and the mean number of effective alleles was 5.8. The average gene diversity equalled 0.744 (range: 0.419–0.957) and the overall differentiation was found significant (θ = 0.092, P < 0.01). Three loci displayed significant heterozygote deficiencies, which might indicate the presence of null alleles. Amplifications were detected on eight loci in Aster alpinus L.     

Ploidy, geographical distribution and morphological differentiation of Parasenecio auriculata (Senecioneae; Asteraceae) in Japan

Parasenecio auriculata is a woodland perennial herb widely distributed in Northeastern Asia, constituted by a poorly understood polyploidy with a diploid (2n=2x=60) and a tetraploid (2n=4x=120). In this study, for a better understanding of the polyploidal evolution, cytogeography and morphological variation were analyzed in Japanese P. auriculata, including two varieties; var. bulbifera endemic to central Hokkaido and var. kamtschatica widely distributed in northern Honshu and Hokkaido. The occurrence of two polyploidal levels was reconfirmed. While var. bulbifera is predominantly tetraploid, var. kamtschatica is comprised of diploid and tetraploid. Morphological variation among 22 quantitative characteristics is continuous and not distinctive among cytotypes or varieties, but plant size tended to be larger in the order, diploid of var. kamtschatica, var. bulbifera, and tetraploid of var. kamtschatica. The cytotype distribution showed a conspicuous geographical pattern. Besides var. bulbifera endemic to the central Hokkaido, the diploid of var. kamtschatica is mainly found in Southern Hokkaido, and the tetraploid has a disjunct distribution in eastern and northern Hokkaido and northern Honshu. Such a geographical pattern is possibly attributable to the differentiation of climatic preference among cytotypes and varieties, and may have been established in association with the climatic cline along the Japanese archipelago.     

Genetic,cytogenetic and morphological diversity in Helicrysum leucocephalum (Asteraceae) populations

Helichrysum (family Asteraceae) is a large, heterogeneous and polyphyletic genus. Most of the Helichrysum species have ornamental and medicinal values. Helicrysum leucocephalum Boiss., has wide geographical distribution in Iran and forms several local populations. The present investigation was performed to study inter populations, genetic, cytogenetic and morphological diversity of this medicinal species. The AMOVA test showed significant genetic difference among the studied populations but the Mantel test did not show correlation between the genetic distance and geographical distance of these populations. STRUCTURE and reticulation analyses showed some degree of genetic admixture and gene exchange among the studied populations. These populations had different chromosome numbers and also differed significantly in the size of their chromosomes.     

Genetic diversity and hierarchical population structure of a rare autotetraploid plant,Aster kantoensis (Asteraceae)

We sampled 17 populations of a rare autotetraploid Aster kantoensis (Asteraceae) from three river systems located in central Japan, and studied them for allelic variation at 22 enzyme loci. There was no significant correlation between the actual population size and three genetic diversity parameters, suggesting that the effective population size was very small even for the large populations, i.e., even large populations may still have a high probability of being of recent origin and remain influenced by the founder effect. Compared to other autotetraploid species, the total genetic variation of A. kantoensis is small. The number of alleles and gene diversity of a population were not significantly different among the river systems, although the percentage of polymorphic loci was different. Genetic differentiation among river systems was larger than between populations within the river systems, thereby indicating that gene flow between river systems is small, especially between the Kinu River system and Tama or Sagami River systems.