Corrected author citation for the combination aster section Biotia (Asteraceae)

The author citation for the combination Aster section Biotia is corrected to: DC. ex Torrey & Gray.     

Phylogenetic relationships and generic delimitation of Eurasian Aster (Asteraceae: Astereae) inferred from ITS, ETS and trnL-F sequence data

Background and Aims

The classification and phylogeny of Eurasian (EA) Aster (Asterinae, Astereae, Asteraceae) remain poorly resolved. Some taxonomists adopt a broad definition of EA Aster, whereas others favour a narrow generic concept. The present study aims to delimit EA Aster sensu stricto (s.s.), elucidate the phylogenetic relationships of EA Aster s.s. and segregate genera.

Methods

The internal and external transcribed spacers of nuclear ribosomal DNA and the plastid DNA trnL-F region were used to reconstruct the phylogeny of EA Aster through maximum parsimony and Bayesian analyses.

Key Results

The analyses strongly support an Aster clade including the genera Sheareria, Rhynchospermum, Kalimeris (excluding Kalimeris longipetiolata), Heteropappus, Miyamayomena, Turczaninowia, Rhinactinidia, eastern Asian Doellingeria, Asterothamnus and Arctogeron. Many well-recognized species of Chinese Aster s.s. lie outside of the Aster clade.

Conclusions

The results reveal that EA Aster s.s. is both paraphyletic and polyphyletic. Sheareria, Rhynchospermum, Kalimeris (excluding K. longipetiolata), Heteropappus, Miyamayomena, Turczaninowia, Rhinactinidia, eastern Asian Doellingeria, Asterothamnus and Arctogeron should be included in Aster, whereas many species of Chinese Aster s.s. should be excluded. The recircumscribed Aster should be divided into two subgenera and nine sections. Kalimeris longipetiolata, Aster batangensis, A. ser. Albescentes, A. series Hersileoides, a two-species group composed of A. senecioides and A. fuscescens, and a six-species group including A. asteroides, should be elevated to generic level. With the Aster clade, they belong to the Australasian lineages. The generic status of Callistephus should be maintained. Whether Galatella (including Crinitina) and Tripolium should remain as genera or be merged into a single genus remains to be determined. In addition, the taxonomic status of A. auriculatus and the A. pycnophyllus–A. panduratus clade remains unresolved, and the systematic position of some segregates of EA Aster requires further study.     

A SYNOPSIS OF NORTH AMERICAN ASTERS: THE SUBGENERA,SECTIONS AND SUBSECTIONS OF ASTER AND LASALLEA

Traditional treatments of Aster were found unsatisfactory in predicting cytological and morphological characteristics of members of various subgenera, sections and subsections of different authors. A realignment of species into these infrageneric categories is presented and is based on sets of shared morphological characteristics of roots or rhizomes, leaves, and phyllaries. Eight subgenera are recognised in Aster. Subgenus Aster is divided into five sections on morphological characteristics: three have X = 9, one has X = 8 and one has X = 7. Subgenus Aster section Dumosi has been redefined and contains all the X = 8 species. Ten X = 5 species and one X = 4 species previously classified in up to six sections of subgenus Aster were found to be morphologically and cytologically related to each other, but unrelated to the other species of Aster. These eleven species have been assigned to the genus Lasallea sensu Semple & Brouillet. Lasallea has been divided in sections and subsections with new combinations presented in this paper.     

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.     

A new species of Ionactis (Asteraceae: Astereae) from Southern Nevada and an overview of the genus

Ionactis caelestis Leary & Nesom is a new species known from a single population that occurs on the Aztec Sandstone near Bridge Mountain in the Spring Mountains of Clark County, Nevada. It is placed in the genus Ionactis (=Aster subg. Ianthe) on the basis of its crowded, multicipital crown, lack of persistent basal leaves and presence of densely arranged cauline ones, strongly carinate phyllaries, blue rays, disc style branches with linear-lanceolate appendages, asymmetric carpopodia, double pappus, and chromosome number of 2n = 9 II. A key to the four species of the genus emphasizes the distinction of the new species in its taproot, the abundant, large, glandular trichomes on its stems and leaves, and disc flowers with sterile ovaries. Ionactis is more similar to the goldenaster (Heterotheca) lineage than to Aster, with which it has been allied formerly. The core of the goldenaster genera differ from Ionactis primarily in their yellow-rayed heads, the crystal complement within cells of their disc corollas, and their primarily multinerved achenes.     

Data on chromosome numbers in Aster (Asteraceae), with comments on the status and relationships of certain North American species

Chromosome counts are presented for more than 250 populations ofAster representing 56 species and eight spontaneous interspecific hybrids. Species are listed according to the infrageneric classification system presented in the preceding paper which utilizes basic chromosome numbers as the pivotal diagnostic character. Fundamental to the classification is an ecological species concept. The list is augmented by a series of comments on field observations and on the examination of types. Six older names previously placed in synonymy or recognized at an inferior rank are reinstated at the species level:A. brachypholis, A.firmus, A. fragilis, A. longifolius, A. simmondsii andA. tardiflorus. Two taxa included as species in current manuals of the eastern North American flora are reduced in status to subspecies rank underA. cordifolius. Putative hybrid status is assigned to four taxa previously treated and originally published as species:A. × coerulescens (= A. praealtus × A. simplex, A. × interior (= A. simplex × A. tradescantii, A. × maccallae (= A. ciliolatus × A. subspicatus) and A. × subgeminatus (= A. ciliolatus × A. tardiflorus). With new data available, the basic chromosome number ofx = 8 has now been confirmed for all species ofAster sectionHeterophylli, except the Mexican A.coahuilensis.     

Lectotypifications and neotypifications in Aster section Biotia (Asteraceae), including a complete annotated synonymy

Thirty-four lectotypes and three neotypes of names of taxa in Aster section Biotia DC. ex Torrey & Gray are designated. In addition, six lectotypes and one neotype that were selected originally in herb. are validated, i.e., published here for the first time. These typifications are presented as part of a complete annotated synonymy of the section, comprising a total of 128 entries.     

EXPERIMENTAL ASSESSMENT OF REPRODUCTIVE INTERACTIONS BETWEEN SYMPATRIC ASTER AND ERIGERON (ASTERACEAE) IN INTERIOR ALASKA

Erigeron glabellus and Aster sibiricus have similar flowers, share pollinators, but bloom sequentially in interior Alaska. Both species depend on insect pollination for seed set: the Erigeron is self-incompatible, and the Aster is apparently self-compatible but allogamous. To test the hypothesis that sequential blooming is maintained by natural selection generated by reproductive interference, we manipulated the flowering time of Erigeron, forcing it to bloom simultaneously with Aster, and measured female fecundity in both species. We found no evidence of reduced female fecundity in either species caused by the presence of the sympatric “competitor” or by artificial pollination with the heterospecific pollen prior to conspecific pollination. Two-species mixtures of simultaneously blooming Aster and Erigeron experienced significant interspecific visitation, which may, under natural conditions, cause loss of pollen to alien stigmas and depressed male fecundity, at least in Erigeron. We found no evidence that sequential blooming in Erigeron and Aster is maintained by depressed female fecundity through pollinator sharing. If sequential blooming is maintained by natural selection, it seems more likely to be the result of selection generated by depressed male fitness through pollen loss to alien stigmas.     

8-Oxo-α-selinen und neue scopoletin-derivate aus Conyza-arten

The investigation of three South African and a North American Conyza species afforded, in addition to known compounds, a new eudesmane ketone and five new scopoletin derivatives. The structures are elucidated by spectroscopic methods and by some chemical transformations. Two of the scopoletin enol ethers have very unusual structures. While most of the Conyza species investigated so far contain typical C₁₀-acetylenes, also present in related genera, there are some with different chemistry. The relationship of the genus to Aster and Erigeron, however, is clearly indicated.     

Beyond the Burgess Shale: Cambrian microfossils track the rise and fall of hallucigeniid lobopodians

Burgess Shale-type deposits are renowned for their exquisite preservation of soft-bodied organisms, representing a range of animal body plans that evolved during the Cambrian ‘explosion’. However, the rarity of these fossil deposits makes it difficult to reconstruct the broader-scale distributions of their constituent organisms. By contrast, microscopic skeletal elements represent an extensive chronicle of early animal evolution—but are difficult to interpret in the absence of corresponding whole-body fossils. Here, we provide new observations on the dorsal spines of the Cambrian lobopodian (panarthropod) worm Hallucigenia sparsa from the Burgess Shale (Cambrian Series 3, Stage 5). These exhibit a distinctive scaly microstructure and layered (cone-in-cone) construction that together identify a hitherto enigmatic suite of carbonaceous and phosphatic Cambrian microfossils—including material attributed to Mongolitubulus, Rushtonites and Rhombocorniculum—as spines of Hallucigenia-type lobopodians. Hallucigeniids are thus revealed as an important and widespread component of disparate Cambrian communities from late in the Terreneuvian (Cambrian Stage 2) through the ‘middle’ Cambrian (Series 3); their apparent decline in the latest Cambrian may be partly taphonomic. The cone-in-cone construction of hallucigeniid sclerites is shared with the sclerotized cuticular structures (jaws and claws) in modern onychophorans. More generally, our results emphasize the reciprocal importance and complementary roles of Burgess Shale-type fossils and isolated microfossils in documenting early animal evolution.     

Bioactive Saponins from Allium and Aster Plants

Plants belonging to the genera Allium and Aster are widely distributed in nature and have been used as food and/or medicine. Their wide use was mainly due to the medicinal properties attributed to these plants since ancient times, recently supported by epidemiological and laboratory studies. Saponin compounds, responsible for many pharmacological activities, are quite abundant in these plants. Thus, a deep knowledge about the saponin composition of these vegetables appears to be essential and could promote the discovery of new potential leads. As part of our continuing research aimed at the identification of bioactive metabolites, we have addressed our attention to several Allium and Aster species. The study resulted in the isolation of over fifty saponins of furostane, spirostane, cholestane, and oleane-type structures. Some of the isolated compounds exhibited promising antiproliferative, antifungal, and antispasmodic activities. Their structure, distribution and bioactivity will be reported here together with a brief overview of the literature on Allium and Aster saponins.     

Inaccessible Biodiversity on Limestone Cliffs: Aster tianmenshanensis (Asteraceae), a New Critically Endangered Species from China

Aster tianmenshanensis G. J. Zhang & T. G. Gao, a new species of Asteraceae from southern China is described and illustrated based on evidence from morphology, micromorphology and molecular phylogeny. The new species is superficially similar to Aster salwinensis Onno in having rosettes of spatulate leaves and a solitary, terminal capitulum, but it differs by its glabrous leaf margins, unequal disc floret lobes and 1-seriate pappus. The molecular phylogenetic analysis, based on nuclear sequences ITS, ETS and chloroplast sequence trnL-F, showed that the new species was nested within the genus Aster and formed a well supported clade with Aster verticillatus (Reinw.) Brouillet et al. The new species differs from the latter in having unbranched stems, much larger capitula, unequal disc floret lobes, beakless achenes and persistent pappus. In particular, A. tianmenshanensis has very short stigmatic lines, only ca. 0.18 mm long and less than 1/3 of the length of sterile style tip appendages, remarkably different from its congeners. This type of stigmatic line, as far as we know, has not been found in any other species of Aster. The very short stigmatic lines plus the unequal disc floret lobes imply that the new species may have a very specialized pollination system, which may be a consequence of habitat specialization. The new species grows only on the limestone cliffs of Mt. Tianmen, Hunan Province, at the elevation of 1400 m. It could only be accessed when a plank walkway was built across the cliffs for tourists. As it is known only from an area estimated at less than 10 km² and a walkway passes through this location, its habitat could be easily disturbed. This species should best be treated as Critically Endangered based on the International Union for Conservation of Nature Red List Categories and Criteria B2a.     

HYBRIDIZATION BETWEEN ASTER AND MACHAERANTHERA AND ITS TAXONOMIC SIGNIFICANCE

A controlled crossing program resulted in the production of four interspecific and three intergeneric hybrids among species of Aster sect. Oxytripolium and Machaeranthera. Meiotic analysis revealed stronger degrees of genome homology in some of the intergeneric hybrids than in some of the interspecific hybrids. The significance of these results to the taxonomy of Aster and Machaeranthera was considered and the result was the transfer of Aster sonorae to Machaeranthera.     

Extraordinary fossils reveal the nature of Cambrian life: a commentary on Whittington (1975) ‘The enigmatic animal Opabinia regalis,Middle Cambrian,Burgess Shale,British Columbia’

Harry Whittington''s 1975 monograph on Opabinia was the first to highlight how some of the Burgess Shale animals differ markedly from those that populate today''s oceans. Categorized by Stephen J. Gould as a ‘weird wonder’ (Wonderful life, 1989) Opabinia, together with other unusual Burgess Shale fossils, stimulated ongoing debates about the early evolution of the major animal groups and the nature of the Cambrian explosion. The subsequent discovery of a number of other exceptionally preserved fossil faunas of Cambrian and early Ordovician age has significantly augmented the information available on this critical interval in the history of life. Although Opabinia initially defied assignment to any group of modern animals, it is now interpreted as lying below anomalocaridids on the stem leading to the living arthropods. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

Cambrian stem-group annelids and a metameric origin of the annelid head

The oldest fossil annelids come from the Early Cambrian Sirius Passet and Guanshan biotas and Middle Cambrian Burgess Shale. While these are among the best preserved polychaete fossils, their relationship to living taxa is contentious, having been interpreted either as members of extant clades or as a grade outside the crown group. New morphological observations from five Cambrian species include the oldest polychaete with head appendages, a new specimen of Pygocirrus from Sirius Passet, and an undescribed form from the Burgess Shale. We propose that the palps of Canadia are on an anterior segment bearing neuropodia and that the head of Phragmochaeta is formed of a segment bearing biramous parapodia and chaetae. The unusual anatomy of these taxa suggests that the head is not differentiated into a prostomium and peristomium, that palps are derived from a modified parapodium and that the annelid head was originally a parapodium-bearing segment. Canadia, Phragmochaeta and the Marble Canyon annelid share the presence of protective notochaetae, interpreted as a primitive character state subsequently lost in Pygocirrus and Burgessochaeta, in which the head is clearly differentiated from the trunk.     

Colour in Burgess Shale animals and the effect of light on evolution in the Cambrian

Diffraction gratings are reported from external surfaces of the hard, protective parts of Wiwaxia corrugata, Canadia spinosa and Marrella splendens from the Burgess Shale (Middle Cambrian (515 million years), British Columbia). As a consequence, these animals would have displayed iridescence in their natural environment: Cambrian animals have previously been accurately reconstructed in black and white only. A diversity of extant marine animals inhabiting a similar depth to the Burgess Shale fauna possess functional diffraction gratings. The Cambrian is a unique period in the history of animal life where predatory lifestyles and eyes capable of producing visual images were evolving rapidly. The discovery of colour in Cambrian animals prompts a new hypothesis on the initiation of the ''Big Bang'' in animal evolution which occurred during the Cambrian: light was introduced into the behavioural systems of metazoan animals for the first time. This introduction, of what was to become generally the most powerful stimulus in metazoan behavioural systems, would have triggered turbulence in metazoan evolution.     

Influence of temperature onOpius dissitus (Hym.: Braconidae), a parasitoid ofLiriomyza trifolii (Dipt.: Agromyzidae)

Rearing experiments at different temperatures (10, 15, 20, 25 and 30°C) were made onOpius dissitus Muesebeck, a parasitoid ofLiriomyza trifolii (Burgess). Temperatures between 20 and 30°C were optimal for the development ofO. dissitus, which was able to adapt to relatively high temperatures despite its holarctic origin.     

Egg-collection equipment for Liriomyza leafminers (Diptera: Agromyzidae)

Egg-collection equipment for Liriomyza leafminers was designed using a small glass tube with a glucose solution supported between two thin films. Adult female Liriomyza trifolii (Burgess), which were previously reared on kidney bean for 2–4 days, laid abundant eggs in 5 or 10% glucose solutions. Hatchability of the eggs in the solutions was greater than 80%. The equipment was used successfully to collect eggs from another leafminer species, Liriomyza sativae Blanchard.     

Decline of the Burgess Shale fauna: ecologic or taphonomic restriction?

Aronson, R. B. 1992 07 15: Decline of the Burgess Shale fauna: ecologic or taphonomic restriction? An important alternative has not been tested in the debate over the origin, significance, and decline of Burgess Shale taxa: their observed stratigraphic distribution could be an artifact of increasing bioturbation. Based on information currently available, comparisons of the observed and expected stratigraphic distributions of Burgess Shale-type faunas falsify bioturbation as the exclusive cause. Ecological factors and/or taphonomic controls other than bioturbation were also responsible for the post-Cambrian decline of Burgess Shale-type faunas. Bioturbation, Burgess Shale fauna, soft-bodied animals, taphonomy.     

A modification of the acetic acid-urea system for use in microslab polyacrylamide gel electrophoresis

A modification of the S. Panyim and R. Chalkley (1969, Arch. Biochem. Biophys.130, 337) acetic acid-urea polyacrylamide gel system to include a stacking gel has been devised. The modification makes possible the use of acetic acid-urea gels in the microslab system of P. T. Matsudaira and D. R. Burgess (1978, Anal. Biochem.87, 386).