A molecular phylogeny of the wild onions (Allium; Alliaceae) with a focus on the western North American center of diversity

Nuclear ribosomal DNA (ITS and ETS) sequences from 39 native Californian (USA) Allium species and congeners were combined with 154 ITS sequences available on GenBank to develop a global Allium phylogeny with the simultaneous goals of investigating the evolutionary history (monophyly) of Allium in the Californian center of diversity and exploring patterns of adaptation to serpentine soils. Phylogenies constructed with ITS alone or ITS in combination with ETS provided sufficient resolution for investigating evolutionary relationships among species. The ITS region alone was sufficient to resolve the deeper relationships in North American species. Addition of a second marker (ETS) further supports the phylogenetic placements of the North American species and adds resolution within subgenus Amerallium, a clade containing many Californian endemics. Within the global phylogeny, the native North American species were found to be monophyletic, with the exception of Allium tricoccum and Allium schoenoprasum. All native Californian species included in the analysis fell into a monophyletic subgenus Amerallium section Lophioprason, although endemic Californian species were not monophyletic due to the inclusion of species with ranges extending beyond the California Floristic Province. The molecular phylogeny strongly supports previous morphology-based taxonomic groupings. Based on our results, serpentine adaptation appears to have occurred multiple times within section Lophioprason, while the ancestor of the Californian center of diversity may not have been serpentine-adapted.     

Phylogenetic relationships of North American nymphophiline gastropods based on mitochondrial DNA sequences

Phylogenetic relationships of 36 nymphophiline species representing 10 genera were inferred from mtCOI sequence data and compared to recent morphology-based classifications of this group. Parsimony and maximum likelihood analyses of the molecular data set suggested monophyly of the North American nymphophilines and a sister or otherwise close relationship between this fauna and a European species assigned to the subfamily. Results also supported a previously hypothesized close relationship between the predominantly freshwater nymphophilines and the brackish-water genus Hydrobia . Our analyses resolved a North American nymphophiline subclade composed of Floridobia , Nymphophilus , and Pyrgulopsis , and depicted the remaining North American genera ( Cincinnatia , Marstonia , Notogillia , Rhapinema , Spilochlamys , Stiobia ) as either a monophyletic or paraphyletic group. Two of the large North American genera ( Floridobia , Marstonia ) were supported as monophyletic groups while monophyly of Pyrgulopsis , a western North American group containing > 100 species, was equivocal. North American nymphophiline phylogeny implies that vicariance of eastern and western North American groups was followed by a secondary invasion of eastern coastal areas from the west. We attribute this to dispersal of salt-tolerant progenitors along the Gulf of Mexico coast     

Molecular phylogeny of the speciose vole genus Microtus (Arvicolinae, Rodentia) inferred from mitochondrial DNA sequences

Voles of the genus Microtus represent one of the most speciose mammalian genera in the Holarctic. We established a molecular phylogeny for Microtus to resolve contentious issues of systematic relationships and evolutionary history in this genus. A total of 81 specimens representing ten Microtus species endemic to Europe as well as eight Eurasian, six Asian and one Holarctic species were sequenced for the entire cytochrome b gene (1140 bp). A further 25 sequences were retrieved from GenBank, providing data on an additional 23, mainly Nearctic, Microtus species. Phylogenetic analysis of these 48 species generated four well-supported monophyletic lineages. The genus Chionomys, snow voles, formed a distinct and well-supported lineage separate from the genus Microtus. The subgenus Microtus formed the strongest supported lineage with two sublineages displaying a close relationship between the arvalis species group (common voles) and the socialis species group (social voles). Monophyly of the Palearctic pitymyid voles, subgenus Terricola, was supported, and this subgenus was also subdivided into two monophyletic species groups. Together, these groupings clarify long-standing taxonomic uncertainties in Microtus. In addition, the "Asian" and the Nearctic lineages reported previously were identified although the latter group was not supported. However, relationships among the main Microtus branches were not resolved, suggesting a rapid and potentially simultaneous radiation of a widespread ancestor early in the history of the genus. This and subsequent radiations discernible in the cytochrome b phylogeny, show the considerable potential of Microtus for analysis of historical and ecological determinants of speciation in small mammals. It is evident that speciation is an ongoing process in the genus and that the molecular data provides a vital insight into current species limits as well as cladogenic events of the past.     

Phylogenetics and evolution of the aphid genus Uroleucon based on mitochondrial and nuclear DNA sequences

The genus Uroleucon, and the related genus Macrosiphoniella, represent a large Tertiary radiation of aphids, with a total of about 300 species distributed throughout the world, primarily on host plant species in the family Asteraceae. A molecular phylogenetic study was conducted to identify major clades within Uroleucon and to address the cladistic validity of current subgeneric categories, the evolution of host plant associations, the age of origin, and intercontinental movements in this genus. The seventeen study species included members of the three major subgenera of Uroleucon, species from Europe and North America, one member of Macrosiphoniella, and two outgroups. Data consisted of DNA sequences for three mitochondrial regions and the nuclear gene EF1alpha, for a total of 4287 sites. Nodes supported strongly in both parsimony and maximum likelihood analyses suggest that: (1) Nearctic Uromelan are a monophyletic group branching near the base of the genus and not related to European Uromelan, (2) the New World subgenus Lambersius is possibly monophyletic but is not a tightly related group and is not closely related to other North American species, and (3) Nearctic members of subgenus Uroleucon are a closely related monophyletic group not allied with Nearctic Uromelan or Lambersius. Instead they represent a separate colonization by an Old World ancestor, as they are nested within a strongly supported clade containing European members of both subgenera Uroleucon and Uromelan. Neither of these subgenera is monophyletic. Molecular clock calculations, based on calibrations of mitochondrial divergences from other insects, suggest that Uroleucon + Macrosiphoniella is a relatively recent radiation, probably no more than 5–10 million years old. Although largely confined to Asteraceae, this clade did not radiate in parallel with its host plants. Rather, lateral movement between lineages of Asteraceae must have occurred repeatedly.     

Phylogeny of North American amblemines (Bivalvia, Unionoida): prodigious polyphyly proves pervasive across genera

Abstract. The subfamily Ambleminae is the most diverse subfamily of fresh‐water mussels (order Unionoida), a globally diverse and ecologically prominent group of bivalves. About 250 amblemine species occur in North America; however, this diversity is highly imperiled, with the majority of species at risk. Assessing and protecting this diversity has been hampered by the uncertain systematics of this group. This study sought to provide an improved phylogenetic framework for the Ambleminae. Currently, 37 North American genera are recognized in Ambleminae. Previous phylogenetic studies of amblemines highlighted the need for more extensive sampling due to the uncertainties arising from polyphyly of many currently recognized taxa. The present study incorporated all amblemine genera occurring in North America north of the Rio Grande, with multiple species of most genera, including the type species for all but seven genera. A total of 192 new DNA sequences were obtained for three mitochondrial gene regions: COI, 16S, and ND1. In combination with published data, this produced a data matrix incorporating 357 gene sequences for 143 operational taxonomic units, representing 107 currently recognized species. Inclusion of published data provides additional taxa and a summary of present molecular evidence on amblemine phylogeny, if at the cost of increasing the amount of missing data. Parsimony and Bayesian analyses suggest that most amblemine genera, as currently defined, are polyphyletic. At higher taxonomic levels, the tribes Quadrulini, Lampsilini, and Pleurobemini were supported; the extent of Amblemini and the relationships of some genera previously assigned to that tribe remain unclear. The eastern North American amblemines appear monophyletic. Gonidea and some Eurasian taxa place as probable sister taxa for the eastern North American Ambleminae. The results also highlight problematic taxa of particular interest for further work.     

Molecular phylogenetic systematics and biogeography of tribe Neillieae (Rosaceae) using DNA sequences of cpDNA, rDNA, and LEAFY

A phylogeny of the tribe Neillieae (Rosaceae), which comprises Neillia, Stephanandra, and Physocarpus, was reconstructed based on nucleotide sequences of several regions of cpDNA, the ITS and ETS regions of rDNA, and the second intron of LEAFY, to elucidate relationships among genera and species in Neillieae and to assess the historical biogeography of the tribe. Phylogenetic analyses indicated that Physocarpus and Neillia-Stephanandra were strongly supported as monophyletic and suggested that Stephanandra may have originated by hybridization between two lineages of Neillia. Dispersal-vicariance analyses suggested that the most recent common ancestor of Neillieae may have occupied eastern Asia and western North America and that Physocarpus and Neillia-Stephanandra may have been split by an intercontinental vicariance event in the early Miocene. The biogeographic analyses also suggested that species of Neillia and Stephanandra diversified in eastern Asia, whereas in Physocarpus one dispersal event from western North America to eastern Asia occurred. Two divergent types of LEAFY sequences were found in the eastern North American species, P. opulifolius, but only one type was present in each plant. The two types of sequences may represent homeologous genes that originated by hybridization between P. capitatus and P. monogynus, both western North American species.     

Testing whether ecological factors promote cladogenesis in a group of tiger beetles (Coleoptera: Cicindelidae)

We investigate the role of ecological differentiation in cladogenesis of a monophyletic group of North American tiger beetles, the subgenus Ellipsoptera (genus: Cicindela), by reconstructing their species-level phylogeny from mitochondrial DNA sequences. Observed reconstructions of ecological characters on the phylogeny are compared to those expected under simple null models of no association with cladogenesis. We find no evidence that ecological disparity is associated with either species coexistence, speciation or long-term persistence and/or radiation of lineages. Ecomorphological traits have evolved in response to differences in habitat occupied by species, but without detectable relationship with cladogenesis.     

Aphis (Hemiptera: Aphididae) species groups found in the Midwestern United States and their contribution to the phylogenetic knowledge of the genus

A phylogeny of the genus Aphis Linnaeus, 1 758 was built primarily from specimens collected in the Midwest of the United States. A data matrix was constructed with 68 species and 41 morphological characters with respective character states of alate and apterous viviparous females. Dendrogram topologies of analyses performed using UPGMA (Unweighted Pair Group Method with Arithmetic Mean), Maximum Parsimony and Bayesian analysis of Cytochrome Oxidase I, Elongation Factor 1‐α and primary endosymbiont Buchnera aphidicola 16S sequences were not congruent. Bayesian analysis strongly supported most terminal nodes of the phylogenetic trees. The phylogeny was strongly supported by EF1‐α, and analysis of COI and EF1‐α molecular data combined with morphological characters. It was not supported by single analysis of COI or Buchnera aphidicola 16S. Results from the Bayesian phylogeny show 4 main species groups: asclepiadis, fabae, gossypii, and middletonii. Results place Aphis and species of the genera Protaphis Börner, 1952, Toxoptera Koch, 1856 and Xerobion Nevsky, 1928 in a monophyletic clade. Morphological characters support this monophyly as well. The phylogeny shows that the monophyletic clade of the North American middletonii species group belong to the genus Protaphis: P. debilicornis (Gillette & Palmer, 1929 ), comb. nov., P. echinaceae (Lagos and Voegtlin, 2009 ), comb. nov., and P. middletonii (Thomas, 1879 ). The genus Toxoptera should be considered a subgenus of Aphis (stat. nov.). The analysis also indicates that the current genus Iowana Frison, 1954 should be considered a subgenus of Aphis (stat. nov.).     

Phylogenetic reassessment of the Teloschistaceae (lichen-forming Ascomycota, Lecanoromycetes)

The Teloschistaceae is a widespread family with considerable morphological and ecological heterogeneity across genera and species groups. In order to provide a comprehensive molecular phylogeny for this family, phylogenetic analyses were carried out on sequences from the nuclear ribosomal ITS region obtained from 114 individuals that represent virtually all main lineages of Teloschistaceae. Our study confirmed the polyphyly of Caloplaca, Fulgensia and Xanthoria, and revealed that Teloschistes is probably non-monophyletic. We also confirm here that species traditionally included in Caloplaca subgenus Gasparrinia do not form a monophyletic entity. Caloplaca aurantia, C. carphinea and C. saxicola s. str. groups were recovered as monophyletic. The subgenera Caloplaca and Pyrenodesmia were also polyphyletic. In the subgenus Caloplaca, the traditionally recognized C. cerina group was recovered as monophyletic. Because this study is based solely on ITS, to maximize taxon sampling, the inclusion of phylogenetic signal from ambiguously aligned regions in MP (recoded INAASE and arc characters) resulted in the most highly supported phylogenetic reconstruction, compared with Bayesian inference restricted to alignable sites.     

Phylogenetic analysis of Phlebotomus species belonging to the subgenus Larroussius (Diptera, psychodidae) by ITS2 rDNA sequences

In the genealogy of Phlebotomus (Diptera: Psychodidae), morphological analyses have indicated that the subgenus Larroussius is a monophyletic group which is most closely related to the subgenera Transphlebotomus and Adlerius. We conducted a phylogenetic analysis of the relationships among six representative species of the subgenus Larroussius and one species representatitive of the Phlebotomus subgenus, assessing sequences of the Second Internal Transcribed Spacer (ITS2) of the ribosomal RNA (rRNA). Three of the species (P. perniciosus, P. ariasi and P. perfiliewi perfiliewi) were collected in different parts of the Mediterranean area. The trees estimated from parsimony and neighbour-joining analyses supported the monophyly of the Larroussius subgenus inferred from the morphological analysis. According to our data, P. ariasi may be a sister group to the rest of the Larroussius subgenus, although additional sequence data are needed to confirm this observation. Our results suggest that P. perniciosus and P. longicuspis are distinct species, in spite of the fact that there are only slight morphological differences. The strict congruence between the phylogeny of the Larroussius subgenus inferred from the ITS2 sequences and that based on morphological studies further confirmed the ability of the spacer sequence to identify recently-derived affiliations.     

牦牛的分类学地位及起源研究:mtDNA D-loop序列的分析

牦牛的起源与属级分类学地位至今仍然存在一定的争议。我们测定了家养牦牛和野生牦牛线粒体控制区(D-loop)序列,并以此构建牦牛和牛属、野牛属、水牛属以及非洲水牛属相关种的系统发育树。研究结果表明线粒体D-loop区与Cytb基因序列在构建牛族的系统发育具有同样重要的价值。系统发育关系显示野牛属的灭绝种草原野牛与现存种美洲野牛先聚合为一单系群,然后再和牦牛形成一单系分支,表明牦牛与野牛属的草原野牛、美洲野牛亲缘关系最近,具有最近的共同祖先,而与牛属的其它亚洲物种亲缘关系较远。因此,本研究不支持将牦牛独立为牦牛属———Poephagus,牛属与野牛属在分类上也应合并为一个属。基于上述研究结果和化石证据,我们进一步对牦牛起源的历史背景进行了讨论,认为牦牛与野牛属的分化是由于第四纪气候变化在欧亚大陆发生的,野牛通过白令陆桥进入北美;冰期结束后,由于欧亚大陆其它地区温度升高,牦牛只能局限分布在较为寒冷的青藏高原;而野牛属在北美先后分化为草原野牛和美洲野牛,前者可能是后者的直接祖先。     

Taxonomy and biogeography of the fiddler crabs (Ocypodidae: Genus Uca) of the Atlantic and Gulf coasts of eastern North America

Fifteen species of fiddler crabs are reported for eastern North America between Massachusetts and Quintana Roo, Mexico. Thirteen occur in the United States and 11 in Mexico, with eight in common to the two countries. Of 13 species in the Gulf of Mexico, five are endemic and a sixth is restricted largely to the peninsulas of Florida and Yucatan. The status of U. rapax in the northern Gulf remains to be resolved. Range limits of most species approximate one or the other of two sets of intersecting thermal and geological boundaries that subdivide the Gulf of Mexico along north-south and east-west axes. Species belonging to subgenus Minuca tend to replace one another at the thermally-controlled Carolinian-Caribbean marine biotic boundary across the Florida peninsula and northern Gulf. However, only U. minax of all the North American fiddler crabs exhibits the classical disjunct Carolinian distribution, and this appears basically to reflect the discontinuous distribution of temperate salt marshes that are the habitat of the species. Distributions of species belonging to subgenus Celuca adhere for the most part to the subdivision of the Gulf into western terrigenous and eastern carbonate sedimentary provinces. The northern transition occurs in the vicinity of Apalachee Bay and the southern at Laguna de Terminos. A third distributional pattern is shown by U. subcylindrica , a specialized endemic species of the hypersaline Laguna Madre system of the western Gulf. The level of endemism in the fiddler crabs is relatively high in comparison with that of other marine groups within the Gulf of Mexico. This may be a consequence of the adaptations of fiddler crabs as specialized deposit feeders to regional differences in climatic and edaphic characteristics of a marginally marine upper shore habitat. The distributional patterns of the endemics could prove useful in reconstructing palaeoecological events of evolutionary significance within the Gulf of Mexico.     

Delimitation of major lineages within Cuscuta subgenus Grammica (Convolvulaceae) using plastid and nuclear DNA sequences

Subgenus Grammica, the largest and most diverse group in the parasitic genus Cuscuta, includes 130 species distributed primarily throughout the New World, with Mexico as its center of diversity. To circumscribe the subgenus and assess the relationships among its major lineages, we conducted the first phylogenetic study of Grammica using plastid trnL-F and nrITS sequences from a wide taxonomic sampling covering its morphological, physiological, and geographical diversity. With the exception of one species belonging elsewhere, the subgenus was found to be monophyletic. The results further indicate the presence of 15 well-supported major clades within Grammica. Some of those lineages correspond partially to earlier taxonomic treatments, but the majority of groups are identified in this study for the first time. The backbone relationships among major clades, however, remain weakly supported or unresolved in some cases. The phylogenetic results indicate that the fruit dehiscence character is homoplastic, thus compromising its value as a major taxonomic and evolutionary feature. While several striking cases of long-distance dispersal are inferred, vicariance emerges as the most dominant biogeographical pattern for Cuscuta. Species placed within one of the clades with a predominantly South American distribution are hypothesized to have substantially altered plastid genomes.     

Phylogeny of the New World diploid cottons (Gossypium L., Malvaceae) based on sequences of three low-copy nuclear genes

American diploid cottons (Gossypium L., subgenus Houzingenia Fryxell) form a monophyletic group of 13 species distributed mainly in western Mexico, extending into Arizona, Baja California, and with one disjunct species each in the Galapagos Islands and Peru. Prior phylogenetic analyses based on an alcohol dehydrogenase gene (AdhA) and nuclear ribosomal DNA indicated the need for additional data from other molecular markers to resolve phylogenetic relationships within this subgenus. Toward this end, we sequenced three nuclear genes, the anonymous locus A1341, an alcohol dehydrogenase gene (AdhC), and a cellulose synthase gene (CesA1b). Independent and combined analyses resolved clades that are congruent with current taxonomy and previous phylogenies. Our analyses diagnose at least two long distance dispersal events from the Mexican mainland to Baja California, following a rapid radiation of the primary lineages early in the diversification of the subgenus. Molecular data support the proposed recognition of a new species closely related to Gossypium laxum that was recently collected in Mexico.     

Molecular evolution of Holarctic martens (genus Martes,Mammalia: Carnivora: Mustelidae)

The Bering Land Bridge has served as a major corridor of interchange between the northern continents for many organisms. We investigated the phylogeny of all extant species of Martes (except for Martes gwatkinsi from India) to infer evolutionary relationships and characterize the extent of trans-Beringian movements. Analyses of complete sequences of the mitochondrial cytochrome b gene and partial sequences of the nuclear aldolase C gene (241bp) suggested that the genus Martes may be paraphyletic with respect to Gulo gulo. These data supported the fossil record's indication that early radiations gave rise to two subgenera (Pekania and Charronia) and that a more recent, possibly rapid, radiation gave rise to species of the third subgenus (Martes). Two colonizations of North America are evident, one by members of the subgenus Pekania and another by member of the subgenus Martes. Contrary to hypotheses based on morphological evidence, the "americana" and "caurina" subspecies groups of Martes americana are not the result of independent colonizations of North America. The phylogenetic analyses of cytochrome b data were consistent with the recognition of these subspecies groups as monophyletic clades; however, variation in the aldolase C sequences indicated that these generally parapatric groups may interbreed in a region of limited geographic overlap.     

A phylogeographic study of the cosmopolitan genus Phragmites (Poaceae) based on AFLPs

Within the genus Phragmites (Poaceae), the species P. australis (the common reed) is virtually cosmopolitan, and shows considerable variation in ploidy level and morphology. Genetic variation in Phragmites was studied using AFLPs, and analysed with parsimony and distance methods. Groups of P. australis strongly supported in the analyses include one that comprises all South American clones, a distinct group from the US Gulf Coast, and a group of E. Asian and Australian octoploids. Among the other species, the paleotropical P. vallatoria is supported as monophyletic and most closely related to the paraphyletic P. mauritianus and to the Gulf Coast and S. American groups. The E. Asian species P. japonicus is closely related to a group of P. australis clones mostly from central North America. Tetraploidy predominates in the genus, and optimisation of chromosome numbers onto the phylogeny shows that higher ploidy levels have evolved many times.     

Mid-Pleistocene divergence of Cuban and North American ivory-billed woodpeckers

We used ancient DNA analysis of seven museum specimens of the endangered North American ivory-billed woodpecker (Campephilus principalis) and three specimens of the species from Cuba to document their degree of differentiation and their relationships to other Campephilus woodpeckers. Analysis of these mtDNA sequences reveals that the Cuban and North American ivory bills, along with the imperial woodpecker (Campephilus imperialis) of Mexico, are a monophyletic group and are roughly equidistant genetically, suggesting each lineage may be a separate species. Application of both internal and external rate calibrations indicates that the three lineages split more than one million years ago, in the Mid-Pleistocene. We thus can exclude the hypothesis that Native Americans introduced North American ivory-billed woodpeckers to Cuba. Our sequences of all three woodpeckers also provide an important DNA barcoding resource for identification of non-invasive samples or remains of these critically endangered and charismatic woodpeckers.     

Molecular phylogeny and historical biogeography of the Holarctic wetland leaf beetle of the genus Plateumaris

Leaf beetles of the genus Plateumaris inhabit wetlands across the temperate zone of the Holarctic region. To explore the phylogeographic relationships among North American, East Asian, and European members of this genus and the origin of the species endemic to Japan, we studied the molecular phylogeny of 20 of the 27 species in this genus using partial sequences of mitochondrial cytochrome oxidase subunit I (COI) and the 16S and nuclear 28S rRNA genes. The molecular phylogeny revealed that three species endemic to Europe are monophyletic and sister to the remaining 11 North American and six Asian species. Within the latter clade, North American and Asian species did not show reciprocal monophyly. Dispersal-vicariance analysis and divergence time estimation revealed that the European and North America-Asian lineages diverged during the Eocene. Moreover, subsequent differentiation occurred repeatedly between North American and Asian species, which was facilitated by three dispersal events from North America to Asia and one in the opposite direction during the late Eocene through the late Miocene. Two Japanese endemics originated from different divergence events; one differentiated from the mainland lineage after differentiation from the North American lineage, whereas the other showed a deep coalescence from the North American lineage with no present-day sister species on the East Asian mainland. This study of extant insects provides molecular phylogenetic evidence for ancient vicariance between Europe and East Asia-North America, and for more recent (but pre-Pleistocene) faunal exchanges between East Asia and North America.     

Systematics of the subgenus Oligocephalus (Teleostei: Percidae: Etheostoma) with complete subgeneric sampling of the genus Etheostoma

The genus Etheostoma is the most diverse clade of freshwater fishes in North America. While studies have been performed with complete sampling of a single subgenus, none have included representatives of all remaining subgenera. The subgenus Oligocephalus is the largest, consisting of 25-27 species in four species groups, and its monophyly has never been clearly demonstrated. The monophyly of this subgenus and its constituent groups was tested using parsimony and Bayesian analyses of ND2 (mtDNA) and the first intron of S7 (nDNA) with complete species sampling from Oligocephalus and complete subgeneric sampling from Etheostoma. Although the subgenus Oligocephalus was not recovered as a monophyletic group in any analyses, monophyletic E. whipplei, Southwestern Darter, and E. spectabile (in part) species groups were recovered in all analyses. All analyses agree that E. okaloosae and both subspecies of E. hopkinsi are not closely related to other members of the subgenus Oligocephalus. E. exile is, however, presenting the strongest evidence yet that recognition of the subgenus Boleichthys is unwarranted.     

Phylogeny of Eastern North American Coreopsis (Asteraceae-Coreopsideae): insights from nuclear and plastid sequences, and comments on character evolution

A molecular phylogenetic study of eastern North American Coreopsis and representatives of other genera of tribe Coreopsideae was conducted using combined sequences from nuclear ITS and two plastid regions (matK, rpl16). A total of 25-30 species has been recognized in five sections of Coreopsis in eastern North America. Based on morphological characters, these taxa have generally been considered a monophyletic group. Our well-resolved phylogeny supports the monophyly of sections that have been recognized in Coreopsis, but the sections collectively do not comprise a monophyletic group because species of north temperate Bidens occur within one of the two major Coreopsis clades. The most notable departure of present results from prior views of relationships among sections is the lack of a sister group relationship between sections Calliopsis and Eublepharis; the shared presence of four-lobed disk floret corollas had been used to support a close relationship between these two sections. Relationships within sections show both similarities and differences with the results of previous studies based primarily on morphological characters. Mapping of morphological characters used taxonomically in Coreopsis and related genera onto the phylogeny indicates that the evolution of these characters has been complex, and this compromises their value for defining monophyletic groups. Examples include the annual habit, alternate leaves, winged fruits, red or brown basal spots on the yellow ligules, and four-lobed disk floret corollas.