A review of the phylogeny and classification of the Asteraceae

The Asteraceae are commonly divided into two large subfamilies, the Cichorioideae (syn. Lactucoideae; Mutisieae, Cardueae, Lactuceae, Vernonieae, Liabeae, Arctoteae) and the Asteroideae (Inuleae, Astereae, Anthemideae, Senecioneae, Calenduleae, Heliantheae, Eupatorieae). Recent phylogenetic analyses based on morphological and chloroplast DNA data conclusively show that the Mutisieae-Barnadesiinae are the sister group to the rest of the family and that the Asteroideae tribes form a monophyletic group. The Vernonieae and Liabeae are sister tribes and the Eupatorieae are nested within a paraphyletic Heliantheae; otherwise tribal interrelationships are still largely uncertain. The Mutisieae-Barnadesiinae are excluded from the Mutisieae and elevated to the new subfamily Barnadesioideae. The two subfamilies Barnadesioideae and Asteroideae are monophyletic, whereas the status of the Cichorioideae remains uncertain. Analyses of chloroplast DNA data support the monophyly of the Cichorioideae; however, morphological data indicate that the subfamily is paraphyletic. Further studies are needed to test the monophyly of the Cichorioideae, as well as to further resolve tribal interrelationships in the two larger subfamilies.     

TRIBAL PLACEMENT OF MARSHALLIA (ASTERACEAE) USING CHLOROPLAST DNA RESTRICTION SITE MAPPING

The examination of morphological traits has failed to resolve the tribal placement of Marshallia. Suggested relationships for this anomalous genus have, at various times, included Eupatorieae, Heliantheae, Vemonieae, and Inuleae. Chloroplast DNA restriction site mapping, using Bamadesiinae (Mutisieae) as the outgroup, revealed 981 restriction site mutations, 332 of which were phylogenetically informative, for 60 genera representing 15 tribes of Asteraceae. Wagner parsimony produced 36 equally parsimonious tress of 729 steps, and Dollo parsimony produced 34 equally parsimonious trees of 759 steps. Monophyletic groups, resulting from the Wagner analysis, were further tested with the bootstrap method. The placement of Marshallia in the Heliantheae-Tageteae-Coreopsideae-Eupatorieae complex was consistent for all trees produced. Tageteae and Coreopsideae form the sister group to paraphyletic Heliantheae, with Marshallia sharing its most recent common ancestor with Galinsoga, Palafoxia, and Bahia. The Eupatorieae form a monophyletic clade that is derived from helianthoid ancestors.     

Chemodiversity of exudate flavonoids in seven tribes of Cichorioideae and Asteroideae (Asteraceae)

Members of several genera of Asteraceae, belonging to the tribes Mutisieae, Cardueae, Lactuceae (all subfamily Cichorioideae), and of Astereae, Senecioneae, Helenieae and Heliantheae (all subfamily Asteroideae) have been analyzed for chemodiversity of their exudate flavonoid profiles. The majority of structures found were flavones and flavonols, sometimes with 6- and/or 8-substitution, and with a varying degree of oxidation and methylation. Flavanones were observed in exudates of some genera, and, in some cases, also flavonol- and flavone glycosides were detected. This was mostly the case when exudates were poor both in yield and chemical complexity. Structurally diverse profiles are found particularly within Astereae and Heliantheae. The tribes in the subfamily Cichorioideae exhibited less complex flavonoid profiles. Current results are compared to literature data, and botanical information is included on the studied taxa.     

Implications of morphological phylogenetics for the placement of the generaAdenocaulon andEriachaenium (Asteraceae)

Adenocaulon andEriachaenium are two problematic genera because their tribal and subfamilial placement in Asteraceae is uncertain. Previous cladistic analyses based on molecular data undertaken to analyze the relationships within Asteraceae, placeAdenocaulon in the tribe Mutisieae (Cichorioideae). This paper investigates cladistic relationships amongAdenocaulon andEriachaenium and tribes of subfamilies Cichorioideae and Asteroideae using morphological data. Thirty-eight characters were scored across 52 genera selected as exemplar taxa to represent the current classification system. In the analysis (one tree, length = 86, c.i. = 0.55, r.i. = 0.64)Adenocaulon andEriachaenium are sister taxa and appear as an isolated clade nested in Cichorioideae. A new, tentative position among the tribes of the paraphyletic Cichorioideae is proposed for these two isolated genera.     

A general survey and some taxonomic implications of diterpenes in the Asteraceae

This paper provides a general survey of the occurrence of diterpenes in the Asteraceae. Data on 4351 botanical occurrences were obtained from the literature. These were grouped by skeleton for each genus. Then, the genera were grouped by subtribes, which, in turn, were gathered in tribes, followed by subfamilies. In spite of the low number of species containing diterpenes, it was possible to describe some structural features of these compounds, i.e. the skeletal types in various taxa and the positions in some skeletons that are always oxidized or never undergo oxidation in some genera. Thus, it was verified that: in the subfamily Cichorioideae, only a few of the studied species possess diterpenes, wherein kaurane is the most frequent diterpene skeleton. In the Asteroideae, the presence of diterpenes is much greater than that in the Cichorioideae and Carduoideae. At tribal taxonomic level, for example, the Astereae produce labdanes and clerodanes; Heliantheae and Eupatorieae produce kauranes and labdanes, respectively; and Calenduleae produce pimaranes. Some taxonomic implications are presented.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 291–308.     

A comprehensive generic-level phylogeny of the sunflower family: Implications for the systematics of Chinese Asteraceae

The sunflower family (Asteraceae) is the largest and the most diverse flowering plant family, comprising 24 000–30 000 species and 1600–1700 genera. In China, Asteraceae are also the largest family, with approximately 2336 indigenous species in 248 genera. In the past two decades, molecular phylogenetic analyses has contributed greatly to our understanding of the systematics of Asteraceae. Nevertheless, the large-scale analyses and knowledge about the relationships of Chinese Asteraceae at the generic level as a whole are far from complete due to difficulties in sampling. In this study, we presented a three-marker (rbcL, ndhF, and matK) phylogeny of Asteraceae, including 506 genera (i.e., approximately one-third of Asteraceae genera). The study sampled 200 Chinese genera (i.e., approximately 80% of Chinese Asteraceae genera). The backbones of the new phylogeny were largely congruent with earlier studies, with 13 subfamilies and 45 tribes recognized. Chinese Asteraceae were distributed in 7 subfamilies (Mutisioideae, Wunderlichioideae, Carduoideae, Pertyoideae, Gymnarrhenoideae, Cichorioideae, and Asteroideae) and 22 tribes (Mutiseae, Hyalideae, Cardueae, Pertyeae, Gymnarrheneae, Vernonieae, Cichorieae, Doroniceae, Senecioneae, Astereae, Anthemideae, Gnaphalieae, Calenduleae, Inuleae, Athroismeae, Helenieae, Coreopsideae, Neurolaeneae, Tageteae, Millieae, Eupatorieae, and Heliantheae). Chinese Asteraceae lacked 6 basal subfamilies and 23 tribes. Several previously ambiguous relationships were clarified. Our analyses also resolved some unplaced genera within Chinese Asteraceae. Finally, our phylogenetic tree was used to revise the classification for all genera of Chinese Asteraceae. In total, 255 genera, 22 tribes, and 7 subfamilies in China are recognized.     

Evolutionary relationships in the Asteraceae tribe Inuleae (incl. Plucheeae) evidenced by DNA sequences of ndhF; with notes on the systematic positions of some aberrant genera

The phylogenetic relationships between the tribes Inuleae sensu stricto and Plucheeae are investigated by analysis of sequence data from the cpDNA gene ndhF. The delimitation between the two tribes is elucidated, and the systematic positions of a number of genera associated with these groups, i.e. genera with either aberrant morphological characters or a debated systematic position, are clarified. Together, the Inuleae and Plucheeae form a monophyletic group in which the majority of genera of Inuleae s.str. form one clade, and all the taxa from the Plucheeae together with the genera Antiphiona, Calostephane, Geigeria, Ondetia, Pechuel-loeschea, Pegolettia, and Iphionopsis from Inuleae s.str. form another. Members of the Plucheeae are nested with genera of the Inuleae s.str., and support for the Plucheeae clade is weak. Consequently, the latter cannot be maintained and the two groups are treated as one tribe, Inuleae, with the two subtribes Inulinae and Plucheinae. The genera Asteriscus, Chrysophthalmum, Inula, Laggera, Pentanema, Pluchea, and Pulicaria are demonstrated to be non-monophyletic. Cratystylis and Iphionopsis are found to belong to the same clade as the taxa of the former Plucheeae. Caesulia is shown to be a close relative of Duhaldea and Blumea of the Inuleae-Inulinae. The genera Callilepis and Zoutpansbergia belong to the major clade of the family that includes the tribes Heliantheae sensu lato and Inuleae (incl. Plucheeae), but their exact position remains unresolved. The genus Gymnarrhena is not part of the Inuleae, but is either part of the unresolved basal complex of the paraphyletic Cichorioideae, or sister to the entire Asteroideae.     

A reassessment of tribal affinities of Cratystylis and Haegiela (Asteraceae) based on three chloroplast DNA sequences

 The tribal affinities of Cratystylis and Haegiela were assessed using three chloroplast DNA sequences, the trnL/F spacer, the trnL intron and the matK coding region. The outgroup was represented by two species of the subfamily Barnadesioideae, whereas one to seven genera (45 species including Cratystylis and Haegiela) of the tribes of the Asteroideae [Anthemideae (6 genera), Astereae (7), Calenduleae (2), Gnaphalieae (7), Heliantheae s.l. (5), Inuleae s.str. (3), Plucheeae (3), Senecioneae (4)] and Cichorioideae, [Arctotideae (1), Cardueae (2), Lactuceae (2), Liabeae (1), Mutisieae (1) and Vernonieae (1)] comprise the ingroup. Phylogenetic analysis indicates that Cratystylis has strong support as a member of the tribe Plucheeae, whereas Haegiela is a member of Gnaphalieae. At some point in their taxonomic history, both genera have been placed in these tribes and there are good morphological and chemical characters that justify these placements. The monotypic Haegiela was once included in Epaltes (Plucheeae) and this study has confirmed the need for the separation of the two genera. The genus Cratystylis appears to be monophyletic. Received August 26, 2002; accepted September 19, 2002 Published online: February 7, 2003     

A survey of autofluorescent patterns in the staminal connective base epidermis in 60 species of Asteraceae

All 60 Asteraceae species sampled, i.e., two species of Bamadesioideae, 21 species in six tribes in Cichorioideae, and 37 species in nine tribes of Asteroideae, show specific patterns of autofluorescence in abaxial connective base epidermal cells. Two species of Cichorioideae and 15 species of Asteroideae have autofluorescence in adaxial epidermal cells of the connective base. Several taxonomically useful features of the connective base are identified; they include shapes of autofluorescent cells, patterns of distribution of cells with autofluorescent walls, and patterns of distribution of autofluorescence in walls. Cellular features of many species coincide with recognized generic, tribal, and subtribal classifications. The results extend Cassini's characterization of the article anthérifere and indicate that the variable features of the connective bases of stamens of Asteraceae could contribute to phylogenetic understanding.     

ITS secondary structure derived from comparative analysis: implications for sequence alignment and phylogeny of the Asteraceae

An RNA secondary structure model is presented for the nuclear ribosomal internal transcribed spacers (ITS) based on comparative analysis of 340 sequences from the angiosperm family Asteraceae. The model based on covariation analysis agrees with structural features proposed in previous studies using mainly thermodynamic criteria and provides evidence for additional structural motifs within ITS1 and ITS2. The minimum structure model suggests that at least 20% of ITS1 and 38% of ITS2 nucleotide positions are involved in base pairing to form helices. The sequence alignment enabled by conserved structural features provides a framework for broadscale molecular evolutionary studies and the first family-level phylogeny of the Asteraceae based on nuclear DNA data. The phylogeny based on ITS sequence data is very well resolved and shows considerable congruence with relationships among major lineages of the family suggested by chloroplast DNA studies, including a monophyletic subfamily Asteroideae and a paraphyletic subfamily Cichorioideae. Combined analyses of ndhF and ITS sequences provide additional resolution and support for relationships in the family.     

The importance of the Mexican taxa of Asteraceae in the family phylogeny

Asteraceae is the largest plant family in México with about 417 genera and 3113 species, and with more than 60% of them being endemic. Phylogenetic relationships at subfamily and tribal levels have been previously resolved employing both nuclear and plastid molecular markers. However, Asteraceae species native to Mexico have been underrepresented in such phylogenies. To tackle this issue, the taxon sampling of this study included 90 Asteraceae species native to México, four species from the Caribbean, 119 previously sequenced species, and six outgroups. With this sampling, all the Asteraceae subfamilies and all of the tribes recognized to date are represented. The analyzed dataset consisted of eleven chloroplast markers (atpB, matK, ndhC, ndhD, ndhF, ndhI, ndhJ, ndhK, rbcL, trnL-trnF, and 23S-trnA). We present two phylogenetic reconstructions obtained by maximum likelihood and pseudocoalescent methods. Besides, we present a time-calibrated phylogeny, which is used to infer the best configuration of diversification rate shifts. Our results show that Mexican species are distributed mainly in the subfamily Asteroideae (80 species), followed by Cichorioideae (6 species), Carduoideae (2 species), and Mutisioideae (2 species). Four net diversification rate shifts were found: One near the base of the tree and four within Asteroideae subfamily. Our extended sampling of the family with the representation of native species to Mexico allowed us to identify important events in the evolutionary history of the family.     

Contribution to the genome size knowledge of New World species from the Heliantheae alliance (Asteraceae)

This article contributes first genome size assessments by flow cytometry for 16 species, 12 genera, and 3 tribes from family Asteraceae, mostly belonging to the Heliantheae alliance, an assembly of 13 tribes from subfamily Asteroideae with a large majority of its species in the New World. Most genome sizes are accompanied by their own chromosome counts, confirming in most cases, although not all, previous counts for the species, and revealing possible cases of unknown dysploidy or polyploidy for certain taxa. The data contribute to the pool of knowledge on genome size and chromosome numbers in the family Asteraceae and will further allow deeper studies and a better understanding on the role of dysploidy in the evolution of the Heliantheae alliance. However, we still lack data for tribes Chaenactideae, Neurolaeneae, Polymnieae, and Feddeeae (the latter, monospecific) to complete the alliance representation.     

Sesquiterpene lactone-based classification of three Asteraceae tribes: a study based on self-organizing neural networks applied to chemosystematics

This work describes an application of artificial neural networks on a small data set of sesquiterpene lactones (STLs) of three tribes of the family Asteraceae. Structurally different types of representative STLs from seven subtribes of the tribes Eupatorieae, Heliantheae and Vernonieae were selected as input data for self-organizing neural networks. Encoding the 3D molecular structures of STLs and their projection onto Kohonen maps allowed the classification of Asteraceae into tribes and subtribes. This approach allowed the evaluation of structural similarities among different sets of 3D structures of sesquiterpene lactones and their correlation with the current taxonomic classification of the family. Predictions of the occurrence of STLs from a plant species according to the taxa they belong to were also performed by the networks. The methodology used in this work can be applied to chemosystematic or chemotaxonomic studies of Asteraceae.     

The value of sampling anomalous taxa in phylogenetic studies: major clades of the Asteraceae revealed

The largest family of flowering plants Asteraceae (Compositae) is found to contain 12 major lineages rather than five as previously suggested. Five of these lineages heretofore had been circumscribed in tribe Mutisieae (Cichorioideae), a taxon shown by earlier molecular studies to be paraphyletic and to include some of the deepest divergences of the family. Combined analyses of 10 chloroplast DNA loci by different phylogenetic methods yielded highly congruent well-resolved trees with 95% of the branches receiving moderate to strong statistical support. Our strategy of sampling genera identified by morphological studies as anomalous, supported by broader character sampling than previous studies, resulted in identification of several novel clades. The generic compositions of subfamilies Carduoideae, Gochnatioideae, Hecastocleidoideae, Mutisioideae, Pertyoideae, Stifftioideae, and Wunderlichioideae are novel in Asteraceae systematics and the taxonomy of the family has been revised to reflect only monophyletic groups. Our results contradict earlier hypotheses that early divergences in the family took place on and spread from the Guayana Highlands (Pantepui Province of northern South America) and raise new hypotheses about how Asteraceae dispersed out of the continent of their origin. Several nodes of this new phylogeny illustrate the vast differential in success of sister lineages suggesting focal points for future study of species diversification. Our results also provide a backbone exemplar of Asteraceae for supertree construction.     

Phylogeny of the bee family Megachilidae (Hymenoptera: Apoidea) based on adult morphology

Phylogenetic relationships within the bee family Megachilidae are poorly understood. The monophyly of the subfamily Fideliinae is questionable, the relationships among the tribes and subtribes in the subfamily Megachilinae are unknown, and some extant genera cannot be placed with certainty at the tribal level. Using a cladistic analysis of adult external morphological characters, we explore the relationships of the eight tribes and two subtribes currently recognised in Megachilidae. Our dataset included 80% of the extant generic‐level diversity, representatives of all fossil taxa, and was analysed using parsimony. We employed 200 characters and selected 7 outgroups and 72 ingroup species of 60 genera, plus 7 species of 4 extinct genera from Baltic amber. Our analysis shows that Fideliinae and the tribes Anthidiini and Osmiini of Megachilinae are paraphyletic; it supports the monophyly of Megachilinae, including the extinct taxa, and the sister group relationship of Lithurgini to the remaining megachilines. The Sub‐Saharan genus Aspidosmia, a rare group with a mixture of osmiine and anthidiine features, is herein removed from Anthidiini and placed in its own tribe, Aspidosmiini, new tribe . Protolithurgini is the sister of Lithurgini, both placed herein in the subfamily Lithurginae; the other extinct taxa, Glyptapina and Ctenoplectrellina, are more basally related among Megachilinae than Osmiini, near Aspidosmia, and are herein treated at the tribal level. Noteriades, a genus presently in the Osmiini, is herein transferred to the Megachilini. Thus, we recognise four subfamilies (Fideliinae, Pararhophitinae, Lithurginae and Megachilinae) and nine tribes in Megachilidae. We briefly discuss the evolutionary history and biogeography of the family, present alternative classifications, and provide a revised key to the extant tribes of Megachilinae.     

PHYLOGENETIC IMPLICATIONS OF CHLOROPLAST DNA RESTRICTION SITE VARIATION IN THE MUTISIEAE (ASTERACEAE)

Phylogenetic relationships among 13 species in the tribe Mutisieae and a single species from each of three other tribes in the Asteraceae were assessed by chloroplast DNA restriction site mapping. Initially, 211 restriction site mutations were detected among 16 species using 10 restriction enzymes. Examination of 12 of these species using nine more enzymes revealed 179 additional restriction site mutations. Phylogenetic analyses of restriction site mutations were performed using both Dolio and Wagner parsimony, and the resulting monophyletic groups were statistically tested by the bootstrap method. The phylogenetic trees confirm an ancient evolutionary split in the Asteraceae that was previously suggested by the distribution of a chloroplast DNA inversion. The subtribe Barnadesiinae of the tribe Mutisieae is shown to be the ancestral group within the Asteraceae. The molecular phylogenies also confirm the paraphyly of the Mutisieae and provide statistical support for the monophyly of three of its four currently recognized subtribes (Barnadesiinae, Mutisiinae, and Nassauviinae). The fourth subtribe, Gochnatiinae, is shown to be paraphyletic. Within the subtribes, several closely related generic pairs are identified. Chloroplast DNA sequence divergence among genera of the Asteraceae ranges between 0.7 and 5.4%, which is relatively low in comparison to other angiosperm groups. This suggests that the Asteraceae is either a relatively young family or that its chloroplast DNA has evolved at a slower rate than in other families.     

Phylogenetic relationships of subfamilies and circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea)

A comprehensive tribal‐level classification for the world’s subfamilies of Hesperiidae, the skipper butterflies, is proposed for the first time. Phylogenetic relationships between tribes and subfamilies are inferred using DNA sequence data from three gene regions (cytochrome oxidase subunit I‐subunit II, elongation factor‐1α and wingless). Monophyly of the family is strongly supported, as are some of the traditionally recognized subfamilies, with the following relationships: (Coeliadinae + (“Pyrginae” + (Heteropterinae + (Trapezitinae + Hesperiinae)))). The subfamily Pyrginae of contemporary authors was recovered as a paraphyletic grade of taxa. The formerly recognized subfamily Pyrrhopyginae, although monophyletic, is downgraded to a tribe of the “Pyrginae”. The former subfamily Megathyminae is an infra‐tribal group of the Hesperiinae. The Australian endemic Euschemon rafflesia is a hesperiid, possibly related to “Pyrginae” (Eudamini). Most of the traditionally recognized groups and subgroups of genera currently employed to partition the subfamilies of the Hesperiidae are not monophyletic. We recognize eight pyrgine and six hesperiine tribes, including the new tribe Moncini. © The Willi Hennig Society 2008.     

An ITS phylogeny of Balsamorhiza and Wyethia (Asteraceae: Heliantheae)

The relationships among the species of Balsamorhiza and Wyethia (Asteraceae: Heliantheae) were examined using data from the internal transcribed spacer (ITS) region of the nuclear ribosomal DNA. The ITS sequences were obtained from nine species of Balsamorhiza and 14 species of Wyethia as well as seven outgroup genera. Five of the outgroup genera were members of the subtribe Engelmanniinae of the tribe Heliantheae, the subtribe that includes Balsamorhiza and Wyethia. The resulting trees show that Balsamorhiza and Wyethia together form a monophyletic group. Balsamorhiza alone is monophyletic, but neither of its two sections is monophyletic. Wyethia is paraphyletic. One group of Wyethia species, including all members of sections Alarconia and Wyethia as well as W. bolanderi from section Agnorhiza, is monophyletic and sister to Balsamorhiza. The other species of Wyethia (all placed in section Agnorhiza) are part of a polytomy along with the clade composed of Balsamorhiza plus the rest of Wyethia.     

First phylogeny of predatory flower flies (Diptera, Syrphidae, Syrphinae) using mitochondrial COI and nuclear 28S rRNA genes: conflict and congruence with the current tribal classification

The family Syrphidae (Diptera) is traditionally divided into three subfamilies. The aim of this study was to address the monophyly of the tribes within the subfamily Syrphinae (virtually all with predaceous habits), as well as the phylogenetic placement of particular genera using molecular characters. Sequence data from the mitochondrial protein-coding gene cytochrome c oxidase subunit I ( COI ) and the nuclear 28S ribosomal RNA gene of 98 Syrphinae taxa were analyzed using optimization alignment to explore phylogenetic relationships among included taxa. Volucella pellucens was used as outgroup, and representatives of the tribe Pipizini (Eristalinae), with similar larval feeding mode, were also included. Congruence of our results with current tribal classification of Syrphinae is discussed. Our results include the tribe Toxomerini resolved as monophyletic but placed in a clade with genera Ocyptamus and Eosalpingogaster . Some genera traditionally placed into Syrphini were resolved outside of this tribe, as the sister groups to other tribes or genera. The tribe Bacchini was resolved into several different clades. We recovered Paragini as a monophyletic group, and sister group of the genus Allobaccha . The present results highlight the need of a reclassification of Syrphinae.
© The Willi Hennig Society 2008.