Heliantheae sensu lato (Asteraceae), clades and classification

The interrelationships within theHeliantheae s. lato and the closely relatedEupatorieae are analyzed and discussed. The basis to this discussion is a cladistic analysis of 141 morphological characters (172 apomorphic states) scored for 97 genera. TheHeleniae s. lato, a subgroup of theHeliantheae s. lato, are paraphyletic, and a monophyletic group corresponding largely to theHeliantheae s. str. is recognized. TheEcliptinae sensuRobinson are polyphyletic. TheCoreopsidinae form an ingroup in theHeliantheae s. str. TheTageteae (Pectidinae) and theMadieae (Madiinae) are two separate branches within the helenioid assemblage of taxa.     

A cladistic analysis of the tribeAstereae (Asteraceae) with notes on their evolution and subtribal classification

TheAstereae were surveyed and the genera arranged in 23 informal groups. The generic groups were used to sample representative genera for a cladistic analysis based on morphological characters. The resulting cladogram was used for discussion of evolution and subtribal classification within the tribe. The lower basic chromosome numbers x = 4, 5, 6, and 8 are interpreted as reductions from a primitive x = 9. The subtribeGrangeinae occupies a phylogenetically basal position as sister group to the rest of the tribe. This may be divided into two large groups, largely corresponding to the homochromousSolidagininae and to the heterochromousAsterinae sensu lato, i.e. including theBellidinae, Hinterhuberinae, Conyzinae, andBaccharidinae. The latter four subtribes are derived within theAsterinae, and hence reduced to synonymy. Several intercontinental relationships indicate that a geographical subdivision of the tribe should be avoided, although in our analysis most of the groups proved to be restricted to one of five major regions.     

A phylogenetic analysis of the lichen familySphaerophoraceae (Caliciales); a new generic classification and notes on character evolution

A phylogenetic analysis of the familySphaerophoraceae (Caliciales, lichenized ascomycetes) has resulted in a new generic classification. Notes on character evolution are given. The generaSphaerophorus s. str.,Bunodophoron andLeifidium, gen. nov., are accepted.Pleurocybe andPseudosphaerophorus are considered synonyms ofBunodophoron andThysanophoron is considered synonym toSphaerophorus. The following new combinations are proposed:Bunodophoron coomerense (Ohlsson)Wedin,B. diplotypum (Vain.)Wedin,B. dodgei (Ohlsson)Wedin,B. flaccidum (Kantvilas & Wedin)Wedin,B. formosanum (Zahlbr.)Wedin,B. imshaugii (Ohlsson)Wedin,B. insigne (Laurer)Wedin,B. kinabaluense (M. Satô)Wedin,B. macrocarpum (Ohlsson)Wedin,B. madagascareum (Nyl.)Wedin,B. microsporum (Ohlsson)Wedin,B. murrayi (Ohlsson)Wedin,B. notatum (Tibell)Wedin,B. ohlssonii (Wedin)Wedin,B. patagonicum (C. W. Dodge)Wedin,B. ramuliferum (I. M. Lamb)Wedin,B. scrobiculatum (C. Bab.)Wedin,B. tibellii (Wedin)Wedin,B. whakapapaense (Wedin)Wedin, andLeifidium tenerum (Laurer)Wedin.     

Morphometric differentiation,patterns of interfertility,and the genetic basis of character evolution inCrepis tectorum (Asteraceae)

Morphometric analyses of populations ofCrepis tectorum raised under uniform conditions support the earlier finding based on herbarium studies that extensive divergence has taken place in western Europe with a centre of diversity in the Baltic lowland area. A crossing experiment showed that subsp.pumila on the island of Öland has acquired a wide range of derived characters without the establishment of strong reproductive barriers, the only exception being a weak tendency for crosses with other populations on Öland to yield more fertile F₁ offspring than crosses with weedy and non-weedy populations in adjacent regions. Canadian weed populations showed a more uniform response in crosses with subsp.pumila than did morphologically similar weed populations scattered throughout Central Europe. While F₂ data suggest a monogenic basis of differences in the extent of leaf dissection, other distinctive traits appear to be governed by multiple genetic factors with individually slight effects.     

Karyological notes onCentaurea sect.Acrocentron (Asteraceae)

The karyology ofCentaurea sect.Acrocentron is surveyed. 19 chromosome counts on 8 species are reported; those onC. acaulis, C. crocata, C. galianoi, C. pubescens, andC. malinvaldiana are new. The basic chromosome numbers of the section are x = 11 and x = 10. Karyological arguments have been used to show that evolution was from x = 11 to x = 10. This is supported by biogeographical data. Two main centres of diversification of sect.Acrocentron were studied from that point of view: the East and the Southwest Mediterranean region.     

Systematic study of the genusDelilia (Asteraceae,Heliantheae)

A taxonomic study of the genusDelilia is presented. Two species are recognized:D. biflora, a widespread weed of tropical and subtropical regions of the New World, extending from Mexico to northern Argentina and recently introduced in the Cape Verde Islands; andD. repens, endemic to the Galapagos islands. A key to the species, illustrations, distribution maps, and complete synonymy are presented.     

Taxonomy ofCarminatia (Asteraceae,Eupatorieae)

A taxonomic study of the largely Mexican genusCarminatia is rendered. It is comprised of three closely related species:C. tenuiflora, C. recondita andC. anomala spec. nova. Illustrations, dot maps, keys to species and complete synonymy are presented.     

Fourfold polyphyly of the genus formerly known as Upucerthia, with notes on the systematics and evolution of the avian subfamily Furnariinae

The traditional avian subfamily Furnariinae, a group of terrestrial ovenbirds typical of the Andean and Patagonian arid zones, consists of the genera Furnarius, Cinclodes, Geositta, Upucerthia, Chilia, and Eremobius. We investigated phylogenetic relationships within the Furnariinae, with particular attention to the nine species of the genus Upucerthia, using nuclear and mitochondrial DNA sequences from all genera in the subfamily. Upucerthia was found to be highly polyphyletic, its constituent species forming four non-sister clades: (1) a basal lineage consisting of two Upucerthia species, U. ruficaudus and U. andaecola, as well as the monotypic genera Eremobius and Chilia; (2) a lineage consisting of U. harterti and U. certhioides, two species behaviorally divergent from other Upucerthia species; (3) a lineage consisting of U. serrana, which is not closely related to any other Upucerthia species; and (4) a lineage, sister to Cinclodes, consisting of the four Upucerthia species U. dumetaria, U. albigula, U. validirostris, and U. jelskii. The larger Furnariinae was also found to be highly polyphyletic; the terrestrial open country ecotype characteristic of this subfamily occurs in four unrelated clades in the family Furnariidae, including a basal lineage as well as derived lineages. Although the large degree of divergence among Upucerthia clades was not previously recognized, owing to ecological, behavioral, and morphological similarities, the groupings correspond closely to relationships suggested by plumage. This is in contrast to studies of other avian genera in which plumage patterns have been shown to be extensively convergent. The generic names Upucerthia and Ochetorhynchus are available for two of the former Upucerthia clades; new generic names may be warranted for the other two.     

Taxonomy and phylogeny of the tribePlucheeae (Asteraceae)

The tribePlucheeae (Benth.)A. Anderb., has been analysed cladistically by means of a computerized parsimony program (Hennig 86), using theArctotideae as outgroup. The results of the analysis are presented in a consensus tree and one cladogram. Four major monophyletic subgroups can be recognized: TheColeocoma group (3 genera), thePterocaulon group (3 genera), theLaggera group (6 genera), and thePluchea group (12 genera). All recognized genera are described and most genera are supplied with taxonomical notes including comments on their taxonomic status. Genera such asBlumea, Pluchea, andEpaltes are demonstrated to be unnatural assemblages.Monarrhenus andTessaria are both closely related to thePluchea complex. The old generic nameLitogyne Harv. has been taken up for one species ofEpaltes, the genusRhodogeron is reduced to a synonym ofSachsia, and the following new combinations are made;Litogyne gariepina (DC.)A. Anderb., andSachsia coronopifolia (Griseb.)A. Anderb.     

Invasive tapetum and tricelled pollen inAmbrosia trifida (Asteraceae,tribeHeliantheae)

Staminate flowers of giant ragweed,Ambrosia trifida L. (Asteraceae, tribeHeliantheae, subtribeAmbrosiinae) were processed into resin and sectioned 1–2 µm thick. The invasive (amoeboid) anther tapetum remains parietal until microspores are released from tetrads, then it swells and invades the locule, merging gradually into a single protoplast that flows among the microspores. After the tapetal membrane ruptures at late microspore stage, tapetal debris fills the locule, then disappears as pollen matures. Pollen becomes tricelled before anthesis. The two sperm cell nuclei are slender and wormlike. The present report supports the two generalizations that invasive tapetum and tricelled pollen are attributes of theAsteraceae.     

Comparative seed morphology and character evolution in the genus Lysimachia (Myrsinaceae) and related taxa

Summary We investigated seed morphology in 34 species of the genus Lysimachia and in 14 species and two subspecies of six additional genera (Anagallis, Ardisiandra, Asterolinon, Glaux, Pelletiera, Trientalis), which have been shown to be closely related to, or are placed within Lysimachia in previous molecular studies. We studied seed shape, seed coat structure, and seed coat surface patterns. Three major types of seed shape were identified: (1) sectoroid, (2) polyhedral, and (3) coarsely rugose with concave hilar area. In addition, seeds may be keeled or winged. The outer layer of the seed coat is either sponge-like and adhering only loosely to the inner seed coat or it is thin and tightly adhering to the underlying tissue. Seed surface patterns can be divided into six main types: (1) reticulate, (2) tuberculate, (3) vesiculose, (4) colliculate, (5) undulate, or (6) poroid-alveolate. Seed surface patterns are mostly congruent with molecular phylogenetic relationships. A reticulate surface pattern is diagnostic of, e.g. Lysimachia subgenera Palladia and Hawaiian Lysimachiopsis. Mapping seed characters onto a recent phylogenetic tree, reveals that they provide potentially synapomorphic character states for various subclades of Lysimachia. Salient examples include a rugose seed shape, which turns out to be synapomorphic for the clade comprising the genus Pelletiera plus Asterolinon linum-stellatum and a sponge-like outer seed coat layer, which characterizes a clade with Lysimachia vulgaris, L. thyrsiflora, and L. terrestris, with an analogue that apparently evolved in parallel in Trientalis europaea. We also discuss possible habitat factors that may have favored the independent evolution of particular seed types such as winged seeds in various lineages.     

Early contributions of molecular phylogenetics to understanding the evolution of Rotifera

The past decade has seen the application of DNA sequence data to phylogenetic investigations of Rotifera, both expanding and challenging our understanding of the evolution of the phylum. Evidence that Acanthocephala, long regarded as a separate but closely related phylum, is a highly derived class of Rotifera demonstrates the potential of molecular analyses to suggest relationships not obvious from morphological analysis. Phylogenies based on the sequence of the gene for the small ribosomal RNA suggest that rotifers and acanthocephalans are associated with Platyhelminthes and Gastrotricha, perhaps in a clade with Gnathostomula and Cycliophora; at present, this group lacks a clear morphological synapomorphy. A more complete resolution of the molecular phylogeny of Rotifera will require surveying multiple genes and several species from each clade under investigation.     

New insights to the molecular phylogenetics and generic assessment in the Rhacophoridae (Amphibia: Anura) based on five nuclear and three mitochondrial genes, with comments on the evolution of reproduction

The phylogenetic relationships among 12 genera of treefrogs (Family, Rhacophoridae), were investigated based on a large sequence data set, including five nuclear (brain-derived neurotrophic factor, proopiomelanocortin, recombination activating gene 1, tyrosinase, rhodopsin) and three mitochondrial (partial 12S and 16S ribosomal RNA and the complete valine t-RNA) genes. Phylogenetic analysis of the nuclear gene sequences resolved three major clades. The first group included Philautus, Pseudophilautus, Kurixalus, Gracixalus, and Theloderma moloch; Pseudophilautus and Kurixalus were sister taxa. The second group consisted of Nyctixalus and Theloderma. The third group contained Feihyla, Polypedates, Rhacophorus, and Chiromantis vittatus; Polypedates and Feihyla were sister taxa. Analyses of the nuclear and mitochondrial genes supported the following results: (1) Genus Liuixalus formed the sister group of all other rhacophorines. (2) Philautus, Theloderma, and Chiromantis were not resolved as monophyletic genera. Four groups, including Philautus ocellatus and P. hainanus, P. longchuanensis and P. gryllus, P. banaensis, and P. quyeti nested well within the genera Liuixalus, Pseudophilautus, Kurixalus, and Gracixalus, respectively. (3) Theloderma moloch and Chiromantis vittatus did not cluster with other species of Theloderma and Chiromantis, respectively. Foam nesting evolved only once, as did laying eggs in a jelly-like matrix containing some bubbles. Terrestrial direct development evolved twice in the Rhacophoridae.     

The systematics of Porphyra: character evolution in closely related species

Isozymes, vegetative and reproductive morphology, seasonality, vertical and geographic distributions and chromosomes were compared for six pairs of putative sibling species of Porphyra (P. abbottae/P. torta, P. fallax subsp. fallax/P. fallax subsp. conwayae, P. amplissima/P. cuneiformis, P. fucicola/P. leucostica, P. miniata/P. variegata, P. umbilicalis/P. umbilicalis) and among five species in a complex (P. brumalis, P. kurogii, P. linearis, P. pseudolinearis, and P. purpurea.) Geographic distribution and zymograms for certain proteins showed the greatest change between species pairs: only one pair of species had identical distributions, and most species pairs were disjunct; every species had a different allozyme for GOT-1, whereas all species had apparently identical proteins for phycoerythrin. Seasonality and habitat exhibited moderate differentiation: Northeast Pacific sibling species were characterized by a high intertidal winter species pairing with a mid intertidal spring species, whereas all but one of the other species pairs exhibited nearly identical vertical distributions and seasonalities. There were few changes in morphology: most species pairs had essentially identical morphologies and coloration and the same arrangement of reproductive cells. Chromosome numbers and karyotypes were identical for species pairs and in the species complex. These results provide evidence for different rates of evolution of different characters in the genus Porphyra.     

Synopsis ofVerbesina sect.Ochractinia (Asteraceae)

A taxonomic revision ofVerbesina sect.Ochractinia is presented. 40 species plus 7 additional infraspecific taxa are recognized. A table of the distribution of the species, key to the species and synonymy are included. Relationships to other sections ofVerbesina and species relationships within sect.Ochractinia are discussed.     

Giemsa banded karyotypes,systematics, and evolution inAnacyclus (Asteraceae-Anthemideae)

Giemsa C-banding allows the differentiation of six, otherwise very similar karyotypes from the small genusAnacyclus. Banding style—with stable centromeric and nucleolar bands, and diverse specific banding patterns in distal chromosome segments—contributes significantly to generic demarcation and systematic grouping. The amount of banding corresponds to heterochromatic chromocentres and increases from perennials to annuals. Relationships with other nucleotype parameters and evolutionary mechanisms are discussed.First contribution of a series on Giemsa Banded Karyotypes, Systematics, and Evolution inAnthemideae (Asteraceae).