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Revised taxonomy of the Arctotis Annual Clade (Arctotideae, Asteraceae) from Southern Africa: integration of molecular phylogenetic and morphological evidence
<p>Previous phylogenetic analysis of ITS nrDNA sequence data for Arctotidinae species resolved a highly supported clade containing all but one of the showy annual <i>Arctotis </i>species (informally designated the '<i>Arctotis</i> Annual Clade')<i>.</i> In the present study, phylogenetic relationships in the <i>Arctotis </i>Annual<i> </i>Clade were investigated by Bayesian inference and maximum parsimony analyses of cpDNA (<i>trnT-trnL-trnF</i> and <i>trnH-psbA</i>) and nrDNA (ITS) sequence data. The cpDNA and nrDNA phylogenies were notably incongruent. <i>Arctotis venusta </i>and a putative unnamed species<i> </i>('sp. B') were highly supported as monophyletic by both datasets. The monophyly of <i>A. leiocarpa </i>was strongly supported by the ITS dataset, whereas the remaining accessions formed a poorly resolved complex (the '<i>A. fastuosa </i>complex'). Within the <i>A. fastuosa </i>complex, <i>A. hirsuta </i>was monophyletic with high support in the ITS phylogeny. A statistical parsimony-derived cpDNA haplotype network resolved five broad groups of haplotypes and showed no consistent geographical structure, but species-specific haplotype lineages<i> </i>for<i> A. venusta </i>and sp. B were resolved. <i>Arctotis fastuosa </i>accessions were distributed among four haplotype groups. Incongruence between the datasets and poor resolution within the <i>A. fastuosa </i>complex may reflect reticulate evolution, ancestral polymorphism, and incomplete lineage sorting, in tandem with the low information content of the datasets. The greatest phenotypic diversification in the clade is in cypsela morphology. Comparison of cypsela morphology with the phylogenies suggests a general trend for reduction in the sizes of the cypsela, abaxial wings, and pappus scales, and loss of pubescence during diversification. A revised taxonomy, integrating currently available evidence, accompanied by full descriptive accounts and a key to the taxa are presented. Eight species are recognized, including the nomenclatural novelties <span><b><i><span>Arctotis chrysantha</span></i></b></span> (sp. nov.) and <span><b><i><span>Arctotis namibiensis</span></i></b></span><i> </i>(sp. nov.). The names <i>Arctotis karasmontana</i>, <i>Venidium fugax</i>, and <i>Venidium macrocephalum</i> are lectotypified.</p>
FIGURE 4 in Revision of the genus Pelecocera Meigen, 1822 (Diptera: Syrphidae) from France taxonomy, ecology and distribution
FIGURE 4. Consensus NJ tree of COI Pelecocera sequences based on 1000 bootstraps. Only clusters found in more than 50 % of the trees are shown.
FIGURE 7 in Revision of the genus Pelecocera Meigen, 1822 (Diptera: Syrphidae) from France taxonomy, ecology and distribution
FIGURE 7. Photographs and schematic dorsal views of Pelecocera heads, showing the shape of the swollen part of the vertex in females of A: P. scaevoides, B: P. pruinosomaculata, C: P. lugubris, D: P. garrigae sp. nov. and E: P. caledonica.
FIGURE 10 in Revision of the genus Pelecocera Meigen, 1822 (Diptera: Syrphidae) from France taxonomy, ecology and distribution
FIGURE 10. Head semi-lateral view to show the lunules (arrow) of: A. Pelecocera lugubris, B: Pelecocera garrigae sp. nov., C: Pelecocera scaevoides, D: Pelecocera pruinosomaculata.
FIGURE 9 in Revision of the genus Pelecocera Meigen, 1822 (Diptera: Syrphidae) from France taxonomy, ecology and distribution
FIGURE 9. Head, frontal view to show parafaces (arrow) of different Pelecocera species from France. A. Pelecocera lugubris, B: Pelecocera garrigae sp. nov., C: Pelecocera scaevoides, D: Pelecocera pruinosomaculata.
FIGURE 6 in Revision of the genus Pelecocera Meigen, 1822 (Diptera: Syrphidae) from France taxonomy, ecology and distribution
FIGURE 6. Plates from Perris (1839) with illustrations of Pelecocera lugubris (photo Bibliothèque nationale de France).
FIGURE 2 in Revision of the genus Pelecocera Meigen, 1822 (Diptera: Syrphidae) from France taxonomy, ecology and distribution
FIGURE 2. Pelecocera garrigae sp. nov. paratype female: A: head and abdomen: lateral view, B head frontal view, C: head, dorso-lateral view, D and E: abdomen: dorsal views. Scale bars = 1 mm.
FIGURE 1 in Revision of the genus Pelecocera Meigen, 1822 (Diptera: Syrphidae) from France taxonomy, ecology and distribution
FIGURE 1. Pelecocera garrigae sp. nov. holotype male (A, C, D) and paratype male (B, E, F): A and C: habitus lateral views, B: abdomen, ventral view, C head, frontal view, E: genitalia, ventral view, F genitalia lateral view. Scale bars (A, B, C, D) = 1 mm, scale bars (E, F) = 0.5 mm.
FIGURE 5 in Revision of the genus Pelecocera Meigen, 1822 (Diptera: Syrphidae) from France taxonomy, ecology and distribution
FIGURE 5. Habitus of the female lectotype of Pelecocera lugubris Perris, 1839: specimen 2731a, Perris collection, CBGP, with the original label of the syntype series (photos J.-C. Streito & É. Pierre, CBGP, INRAE, CIRAD, IRD, Institut Agro Montpellier, Univ. Montpellier).
FIGURE 8 in Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa
FIGURE 8. Dorsal, lateral, and ventral views of the body of the holotype of Nototriton vereh sp. nov. in preservation. Photograph by BK.
FIGURE 10 in Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa
FIGURE 10. Map of Costa Rica showing the distribution range of the eight species within the subgenus Nototriton.
FIGURE 13 in Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa
FIGURE 13. Dorsal views of the hand and foot and detail of the face of the holotype of Nototriton kenorum sp. nov. in preservation. Photograph by BK.
FIGURE 5 in Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa
FIGURE 5. Linear discriminant analysis showing the morphological separation among the eight species within the subgenus Nototriton.
FIGURE 16 in Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa
FIGURE 16. Dorsal views of the hand and foot and detail of the face of the holotype of Nototriton lateomuscus sp. nov. in preservation. Photograph by BK.
FIGURE 4 in Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa
FIGURE 4. Linear discriminant analysis showing the morphological separation among the three species within the subgenus Taylorotriton.
FIGURE 3 in Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa
FIGURE 3. Time-calibrated phylogeny from BEAST analysis of concatenated data set including all species of the genus Nototriton with sequence data available. The map insert shows the distribution range of the three subgenera within the genus Nototriton.
FIGURE 1. A in Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa
FIGURE 1. A) Map showing the historical distribution of the genus Nototriton in Central America, the colors indicate the two know subgenera. B) The insert map shows the localities for the genetic sequences from Costa Rica. The solid diamonds indicate the historical sequences, the open squares indicate the newly created sequences resulting from this study.
FIGURE 2 in Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa
FIGURE 2. Maximum likelihood phylogeny (log likelihood = -10741.341147) showing the relationships of the subgenera Nototriton and Taylorotriton based on the 16S, cyt b, and COI mitochondrial DNA genes fragment. Bootstraps proportions are shown above branches, below the branches are shown posterior probabilities (multiplied by 100) from BEAST analysis (left) and posterior probabilities (multiplied by 100) from MrBayes analysis (right). The scale bar refers to the estimated substitutions per site. The support values of any node within species are not shown. The asterisks represent support of 100.
FIGURE 9 in Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa
FIGURE 9. Dorsal views of the hand and foot and detail of the face of the holotype of Nototriton vereh sp. nov. in preservation. Photograph by BK.
FIGURE 12 in Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa
FIGURE 12. Dorsal, lateral, and ventral views of the body of the holotype of Nototriton kenorum sp. nov. in preservation. Photograph by BK.
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