CHROMOSOME NUMBERS IN COMPOSITAE. III. SENECIONEAE

Ornduff , Robert (Duke U., Durham, N. C), Peter H. Raven , Donald W. Kyhos , and A. R. Kruckeberg . Chromosome numbers in Compositae. III. Senecioneae. Amer. Jour. Bot. 50(2): 131–139. Illus. 1963.—Chromosome counts are reported for 75 taxa of tribe Senecioneae (Compcsitae) and are listed with a generic summary of previous counts in the tribe. First counts are reported for Bedfordia, Crocidium, Dimeresia, Gamolepis, Lepidospartum, Luina, Peucephyllum, Telradymia, and the first definite count recorded for Euryops. New numbers are added to those previously known in Arnica and Psathyrotes. Intraspecific differences in ploidy-level are reported in 4 North American species of Senecio. Although chromosome numbers are useful as an aid in delimiting some genera of Senecioneae, they are of little use in circumscribing genera peripheral to Senecio, primarily because of the great range of chromosome numbers of that genus. Chromosome numbers support suggestions based on morphological considerations that genera such as Crocidium and Dimeresia do not belong in Senecioneae, whereas chromosome number and morphology of the plants virtually prohibit the removal of such genera as Peucephyllum, Lepidospartum, and Telradymia from Senecioneae, despite the suggestions of several recent authors. It is proposed that the base number for the tribe is 10 and that the tribe originated in the Old World, with subsequent widespread migration and diversification.     

A phylogeny of Packera (Senecioneae; asteraceae) based on internal transcribed spacer region sequence data and a broad sampling of outgroups

A phylogeny of Packera is presented based on sequence data from the internal transcribed spacer region of nuclear ribosomal DNA of 26 species (28 populations) of Packera and 23 outgroup taxa, including representatives of all three subtribes of the Senecioneae. The results support a Mexican origin for Packera, with its closest relatives found among Old World taxa in the subtribe Senecioninae, such as Senecio jacobaea and Pericallis. Packera species from the west coast of the United States, previously included in the section Bolanderi of Greenman, are part of a basal assemblage including species of Greenman's Mexican section Sanguisorboidei. The rest of Packera separates into two sister groups, one containing species from the Arizona-New Mexico region and the other containing more geographically diverse taxa. Among the outgroups, New World Senecio species are monophyletic and two Tussilaginoid assemblages are strongly supported; the Tephroseroid group (Tephroseris and Sinosenecio) plus Petasites combine with the Luina complex to form a clade of north temperate taxa, and the four Mexican genera (Psacalium, Robinsonecio, Barkleyanthus, and Pittocaulon) form a monophyletic group.     

Cytotaxonomy of Senecio tricuspis (Asteraceae–Senecioneae)

The chromosome number of 2n=40 counted for Senecio tricuspis (Asteraceae–Senecioneae) indicates that the species should not be transferred to the genus Parasenecio but be retained in Senecio.     

Tackling speciose genera: species composition and phylogenetic position of Senecio sect. Jacobaea (Asteraceae) based onplastid and nrDNA sequences

The molecular phylogeny of Senecio sect. Jacobaea (Asteraceae; Senecioneae) was studied to clarify species composition and interspecific relationships of Senecio sect. Jacobaea. This information is necessary for studies seeking explanations of the evolutionary success of Senecio, in terms of high species numbers and the evolution of chemical defense mechanisms. Parsimony analyses with 60 species of the tribe Senecioneae, representing 23 genera and 11 sections of Senecio, based on DNA sequence data of the plastid genome (the trnT-L intergenic spacer, the trnL intron, and two parts of the trnK intron, flanking both sides of the matK gene) and nuclear genome (ITS1, 5.8S, and ITS2 gene and spacers) show that sect. Jacobaea is a strongly supported monophyletic group. Fifteen species have been identified as members of section Jacobaea, including three species that have been consistently ascribed to this section in taxonomic literature and 12 species that were either placed in other sections of Senecio or not exclusively ascribed to sect. Jacobaea. This section was traditionally circumscribed as a group of European, biennial, or perennial herbs with pinnately incised leaves, but the results of this study show that one annual species, a species from northeastern Asia, and a species growing in the Himalayas are members of sect. Jacobaea as well. Furthermore, not all species in the section have pinnately incised leaves. The genera Emilia, Packera, and Pseudogynoxys form the sister clade of sect. Jacobaea, but this relationship lacks strong bootstrap support and thus remains provisional.     

(Un)Targeted Metabolomics in Asteraceae: Probing the Applicability of Essential‐Oil Profiles of Senecio L. (Senecioneae) Taxa in Chemotaxonomy

The possible applicability of (un)targeted metabolomics (volatile metabolites) for revealing taxonomic/evolutionary relationships among Senecio L. species (Asteraceae; tribe Senecioneae) was explored. Essential‐oil compositional data of selected Senecio/Senecioneae/Asteraceae taxa (93 samples in total) were mutually compared by means of multivariate statistical analysis (MVA), i.e., agglomerative hierarchical clustering and principal component analysis. The MVA input data set included the very first compositional data on the essential oil extracted from the aerial parts of S. viscosus L. as well as on four different Serbian populations of S. vernalis Waldst . & Kit . (oils from aerial parts and roots; eight samples in total). This metabolomic screening of Senecio/Senecioneae/Asteraceae species (herein presented results and data from the literature) pointed to short‐chain alk‐1‐enes (e.g., oct‐1‐ene, non‐1‐ene, and undec‐1‐ene), with up to now restricted general occurrence in Plantae, as characteristic chemotaxonomic markers/targets for future metabolomic studies of Senecio/Senecioneae taxa. The MVA additionally showed that the evolution of the terpene metabolism (volatile mono‐ and sesquiterpenoids) within the Asteraceae tribe Senecioneae was not genera specific. However, the MVA did confirm plant‐organ specific production/accumulation of volatiles within S. vernalis and suggested the existence of at least two volatile chemotypes for this species.     

CHROMOSOME NUMBERS IN COMPOSITAE. VI. SENECIONEAE. II

Chromosome counts are reported for 76 taxa and 2 natural hybrids of tribe Senecioneae (Compositae). First counts are reported for several species of Senecioneae as well as for the genera Cadiscus and Whitneya. New chromosome numbers are added to those previously known in Arnica, Cacalia, and Senecio. Additional counts from Arnica support our previous suggestion that x = 19 for this genus. It is assumed that observed meiotic irregularities are associated with apomixis in this genus. Basic chromosome numbers for various New World sections of Senecio are proposed, and certain problems of sectional relationships in this genus are discussed. Chromosome numbers and plant morphology of Cadiscus, Hulsea, and Whitneya indicate that these genera should be removed from Helenieae to Senecioneae. The possible affinity of the anomalous genus Adenocaulon with Mutisieae is discussed. Data presented in the paper further support our earlier proposal that the basic chromosome number for Senecioneae is x = 10.     

CHROMOSOME NUMBERS OF THE EAST AFRICAN GIANT SENECIOS AND GIANT LOBELIAS AND THEIR EVOLUTIONARY SIGNIFICANCE

The gametophytic chromosome number for the giant senecios (Asteraceae, Senecioneae, Dendrosenecio) is n = 50, and for the giant lobelias (Lobeliaceae, Lobelia subgenus Tupa section Rhynchopetalum) it is n = 14. Previous sporophytic counts are generally verified, but earlier reports for the giant senecios of 2n = 20 and ca. 80, the bases for claims of intraspecific polyploidy, are unsubstantiated. The 14 new counts for the giant senecios and the ten new counts for the giant lobelias are the first gametophytic records for these plants and include the first reports for six and four taxa, respectively, for the two groups. Only five of the 11 species of giant senecio and three of the 21 species of giant lobelia from eastern Africa remain uncounted. Although both groups are polyploid, the former presumably decaploid and the latter more certainly tetraploid, their adaptive radiations involved no further change in chromosome number. The cytological uniformity within each group, while providing circumstantial evidence of monophyly and simplifying interpretations of cladistic analyses, provides neither positive nor negative support for a possible role of polyploidy in evolving the giant-rosette growth-form.     

Astragalus (Fabaceae): A molecular phylogenetic perspective

Nucleotide sequences of the plastidmatK gene and nuclear rDNA internal transcribed spacer region were sampled fromAstragalus L. (Fabaceae), and its closest relatives within tribe Galegeae, to infer phylogenetic relationships and estimate ages of diversification. Consistent with previous studies that emphasized sampling for nrDNA ITS primarily within either New World or Old World species groups,Astragalus, with the exception of a few morphologically distinct species, is strongly supported as monophyletic based on maximum parsimony and Bayesian analyses ofmatK sequences as well as a combined sequence dataset. ThematK data provides better resolution and stronger clade support for relationships amongAstragalus and traditionally related genera than nrDNA ITS.Astragalus sensu stricto plus the genusOxytropis are strongly supported as sister to a clade composed of strictly Old World (African, Australasian) genera such asColutea. Sutherlandia, Lessertia, Swainsona, andCarmichaelia, plus several morphologically distinct segregates of EurasianAstragalus. Ages of these clades and rates of nucleotide substitution estimated from a fossil-constrained, rate-smoothed, Bayesian analysis ofmatK sequences sampled from Hologalegina indicateAstragalus diverged from its sister group,Oxtropis, 12–16 Ma, with divergence of Neo-Astragalus beginning ca 4.4. Ma. Estimates of absolute rates of nucleotide substitution forAstragalus and sister groups, which range from 8.9 to 10.2×10⁻¹⁰ substitutions per site per year, are not unusual when compared to those estimated for other, mainly temperate groups of papilionoid legumes. The results of previously published work and other recent developments on the phylogenetic relationships and diversification ofAstragalus are reviewed.     

Senecio changii (Asteraceae: Senecioneae), a New Species from Sichuan,China

Senecio changii (Asteraceae: Senecioneae), a new species from Muli, Sichuan, southwestern China, is described. It is distinguished in Chinese Senecio s.s. by having lyrate-pinnatisect to pinnatisect leaves and a single terminal large discoid capitulum which is somewhat nodding. Evidence from floral micromorphology, karyology and molecular phylogenetic analyses based on the nuclear ITS/ETS sequence data all support its membership within Senecio s.s.     

Elucidative studies on the generic concept of Senecio (Asteraceae)

VINCENT, P. L. D. & GETLIFFE, F. M., 1992. Elucidative studies on the generic concept of Senecio (Asteraceae) . This paper presents the results of studies of the generic concept of Senecio sensu stricto. The sample of taxa studied consisted of 93 Natal senecios (including seven varieties and two forms), five Cape heterochromous senecios and nine non-southern African senecios, including the type of the genus, S. vulgaris L. Also included in the study were six species from taxonomically closely related genera in the tribe Senecioneae and one Senecio of uncertain taxonomic position. The phenotype of these taxa was investigated with respect to a large number (122) of morphological and micromorphological characters. Six characters were selected as being taxonomically important with respect to elucidating the generic concept of Senecio sensu stricto. The generic concept of Senecio has been provisionally re-circumscribed and the generic status of each of the senecios and non-senecios studied has been tested according to this concept of Senecio sensu stricto sensu Vincent. In the light of this concept of Senecio , the following species are recommended for exclusion from Senecio sensu stricto sensu Vincent: S. cissampelinus, S. transvaalensis, S. syringifolius and S. hockii. The following species are considered to be peripheral to Senecio sensu stricto sensu Vincent: S. tanacetopsis, S. seminiveus, S. medley-woodii, S. tamoides, S. helminthioides, S. barbertonicus, S. brevilorus, S. viminalis, S. radicans and S.fulgens. Before any taxonomic changes are made to the current composition of Senecio , the concept of Senecio sensu stricto sensu Vincent, is being tested on a worldwide sample of the genus.     

Phylogeny of Mesoamerican freshwater mussels and a revised tribe‐level classification of the Ambleminae

Evolutionary and ecological hypotheses of the freshwater mussel subfamily Ambleminae are intensely geographically biased—a consequence of the complete exclusion of Mesoamerican taxa in phylogenetic reconstructions of the clade. We set out to integrate a portion of the Mesoamerican freshwater mussel assemblage into existing hypotheses of amblemine classification and evolution by generating a molecular phylogeny that includes four previously unsampled Mesoamerican genera and nine species endemic to that region. Given the traditionally hypothesized affinity to Nearctic mussels and the understanding that classification should reflect common ancestry, we predicted that (a) Mesoamerican genera would be recovered as members of the recognized tribes of the Ambleminae, and (b) genera would be supported as monophyletic. The mutilocus phylogeny (COI + 28S + 16S) reported herein does not fully support either of those hypotheses. Neither Cyrtonaias nor Psorula were supported as monophyletic and we predict several other Mesoamerica genera are also non‐monophyletic. The reconstructed phylogeny recovered four independent lineages of Mesoamerican freshwater mussels and these clades are distributed across the phylogeny of the Ambleminae, including the tribe Quadrulini (Megalonaias), Lampsilini (two lineages: Cyrtonaias explicata/Sphenonaias microdon, and Pachynaias), and a previously unrecognized, exclusively Mesoamerican and Rio Grande clade consisting of the genera Psoronaias, Psorula and Popenaias. The latter clade possesses several morphological characteristics that distinguish it from its sister taxon, tribe Lampsilini, and we recognize this newly identified Mesoamerican clade as a fifth tribe of the Ambleminae attributable to the Popenaiadini Heard & Guckert, 1970. This revised classification more completely recognizes the suprageneric diversity of the Ambleminae.     

A SYSTEMATIC REVIEW OF THE SUBTRIBE LAGASCEINAE (COMPOSITAE,HELIANTHEAE)

The subtribe Lagasceinae of the tribe Heliantheae is recognized traditionally to consist of two genera: Lagascea and Coulterella. Although both taxa possess uni-flowered heads aggregated into synflorescences, they are very different in most other morphological and anatomical respects. The six species of Lagascea that were investigated cytologically are n = 17, whereas the monotypic Coulterella is n = 18 with chromosomes one-half the size of those of Lagascea. The dissimilarities suggest that the two genera should not be retained in the same subtribe, and that the subtribe Lagasceinae should be abandoned. Lagascea seems to be related most closely to genera of the subtribe Helianthinae, such as Alvordia. Coulterella, on the other hand, seems closest to Flaveria of the tribe Helenieae or Senecioneae.     

Phylogenetic position of the southern African members of the tribe Psoraleeae based on molecular and morphological data

The tribe Psoraleeae (Leguminosae subfamily Papilionoideae) comprises 185 species in nine genera that have a nearly worldwide distribution, occurring predominantly in Mediterranean regions. About 60% of the species belong to the genera, Otholobium C.H.Stirt. and Psoralea L., which have a centre of diversity in the Cape Floristic Region of South Africa. Since previous molecular studies have sampled only a few species of the tribe from this region, this study sought to determine the phylogenetic position of the southern African genera and to test whether they are monophyletic. Phylogenetic relationships were reconstructed using DNA sequence data (trnL-F, rpoB-trnC and ITS) and seven morphological characters, which diagnose the two southern African genera. The data were analysed using the parsimony method. There was strong support for the Psoraleeae as a clade, but most of the nodes within the large genera were poorly supported. The southern African species of Psoralea and Otholobium together formed a strongly supported clade. This clade was sister to the genus Hoita Rydb., but without support. However, the Psoralea species were nested within the southern African Otholobium. Additionally, some South American species that are currently recognised as Otholobium were resolved in a clade distinct from the southern African species, making Otholobium polyphyletic. Morphological characters that separate Otholobium and Psoralea are discussed. Finally, the southern African genera as currently circumscribed are not monophyletic. However, further investigations using more informative DNA loci are required to validate this observation. Furthermore, the taxonomic placement of the South American species needs to be reviewed.     

Host range of Secusio extensa (Lepidoptera: Arctiidae), and potential for biological control of Senecio madagascariensis (Asteraceae)

Secusio extensa (Lepidoptera: Arctiidae) was evaluated as a potential biological control agent for Madagascar fireweed, Senecio madagascariensis (Asteraceae), which has invaded over 400 000 acres of rangeland in the Hawaiian Islands and is toxic to cattle and horses. The moth was introduced from southeastern Madagascar into containment facilities in Hawaii, and host specificity tests were conducted on 71 endemic and naturalized species (52 genera) in 12 tribes of Asteraceae and 17 species of non‐Asteraceae including six native shrubs and trees considered key components of Hawaiian ecosystems. No‐choice feeding tests indicated that plant species of the tribe Senecioneae were suitable hosts with first instars completing development to adult stage on S. madagascariensis (78.3%), Delairea odorata (66.1%), Senecio vulgaris (57.1%), Crassocephalum crepidioides (41.2%), and at significantly lower rates on Emilia fosbergii (1.8%) and Erechtites hieracifolia (1.3%). A low rate of complete larval development also was observed on sunflower, Helianthus annuus (11.6%), in the tribe Heliantheae. However, sunflower was rejected as a potential host in larval‐feeding and adult oviposition choice tests involving the primary host S. madagascariensis as control. Although larvae died as first instars on most test species, incomplete development and low levels of feeding were observed on nine species in the tribes Heliantheae, Cardueae and Lactuceae. Larvae did not feed on any non‐Asteraceae tested, including species with similar pyrrolizidene alkaloid chemistry, crops, and six ecologically prominent native species. Because all species of Senecioneae are non‐native and weedy in Hawaii, these results indicate that S. extensa is sufficiently host‐specific for introduction for biological control. High levels of feeding damage observed on potted plants indicate that S. extensa can severely impact the target fireweed as well as D. odorata, a noxious weed in native Hawaiian forests.     

Evolution of blindness in scolopendromorph centipedes (Chilopoda: Scolopendromorpha): insight from an expanded sampling of molecular data

Relative to its diversity (34 genera, 700 species), Scolopendromorpha has been undersampled in molecular phylogenetic analyses compared with the other chilopod orders. Previous analyses based on morphology have not resolved several key controversies in systematics and evolutionary morphology unambiguously. Here we apply new molecular and morphological data to scolopendromorph phylogenetics, with a focus on the evolution of blindness. The taxonomic sample includes 19 genera, many lacking previous molecular data, and diverse, cosmopolitan genera of Scolopendridae are sampled by multiple species. Phylogenetic analysis with Direct Optimization used 94 morphological characters and ca. 4.5 kb of sequence data from two nuclear (18S and 28S rRNA) and two mitochondrial (16S rRNA and COI) loci. A single most‐parsimonious cladogram selected after sensitivity analyses resolves Scolopendromorpha as monophyletic, and divides it into a blind clade of three families (Plutoniumidae, Cryptopidae, Scolopocryptopidae) and its ocellate sister group, Scolopendridae. Some species‐rich, cosmopolitan genera (Cormocephalus, Otostigmus, Scolopendra) in Scolopendridae are non‐monophyletic, and in several instances (e.g. New and Old World Scolopendra) relationships are more congruent with geographical distributions than with traditional classifications. The tribe Asanadini is particularly subject to parameter‐sensitivity, nesting in the combined analysis within Scolopendrini but as sister to all other Scolopendrinae for molecular data alone. The total‐evidence tree unambiguously optimizes trunk segmentation: a 23‐segmented trunk has a single origin in the blind clade. © The Willi Hennig Society 2011.     

Molecular phylogenetics of the giant genus Croton and tribe Crotoneae (Euphorbiaceae sensu stricto) using ITS and TRNL-TRNF DNA sequence data

Parsimony, likelihood, and Bayesian analyses of nuclear ITS and plastid trnL-F DNA sequence data are presented for the giant genus Croton (Euphorbiaceae s.s.) and related taxa. Sampling comprises 88 taxa, including 78 of the estimated 1223 species and 29 of the 40 sections previously recognized of Croton. It also includes the satellite genus Moacroton and genera formerly placed in tribe Crotoneae. Croton and all sampled segregate genera form a monophyletic group sister to Brasiliocroton, with the exception of Croton sect. Astraea, which is reinstated to the genus Astraea. A small clade including Moacroton, Croton alabamensis, and C. olivaceus is sister to all other Croton species sampled. The remaining Croton species fall into three major clades. One of these is entirely New World, corresponding to sections Cyclostigma, Cascarilla, and Velamea sensu Webster. The second is entirely Old World and is sister to a third, also entirely New World clade, which is composed of at least 13 of Webster's sections of Croton. This study establishes a phylogenetic framework for future studies in the hyper-diverse genus Croton, indicates a New World origin for the genus, and will soon be used to evaluate wood anatomical, cytological, and morphological data in the Crotoneae tribe.