The major opsin in bees (Insecta: Hymenoptera): A promising nuclear gene for higher level phylogenetics.

We report the phylogenetic utility of the nuclear gene encoding the long-wavelength opsin (LW Rh) for tribes of bees. Aligned nucleotide sequences were examined in multiple taxa from the four tribes comprising the corbiculate bees within the subfamily Apinae. Phylogenetic analyses of sequence variation in a 502-bp fragment (approx 40% of the coding region) strongly supported the monophyly of each of the four tribes, which are well established from previous studies of morphology and DNA. Trees estimated from parsimony and maximum likelihood analyses of LW Rh sequences show a strongly supported relationship between the tribes Meliponini and Bombini, a relationship that has been found uniformly in studies of other genes (28S, 16S, and cytochrome b). All of the tribal clades as well as relationships among the tribes are supported by high bootstrap values, suggesting the utility of LW Rh in estimating tribal and subfamily rank for these bees. The sequences exhibit minimal base composition bias. Both 1st + 2nd and 3rd position sites provide information for estimating a reliable tree topology. These results suggest that LW Rh, which has not been reported previously in studies of organismal phylogenetics, could provide important new data from the nuclear genome for phylogeny reconstruction.     

Single-copy nuclear genes recover cretaceous-age divergences in bees

We analyzed the higher level phylogeny of the bee family Halictidae based on the coding regions of three single-copy nuclear genes (long-wavelength [LW] opsin, wingless, and elongation factor 1-alpha [EF-1 alpha]). Our combined data set consisted of 2,234 aligned nucleotide sites (702 base pairs [bp] for LW opsin, 405 bp for wingless, and 1,127 bp for EF-1 alpha) and 779 parsimony-informative sites. We included 58 species of halictid bees from 33 genera, representing all subfamilies and tribes, and rooted the trees using seven outgroups from other bee families: Colletidae, Andrenidae, Melittidae, and Apidae. We analyzed the separate and combined data sets by a variety of methods, including equal weights parsimony, maximum likelihood, and Bayesian methods. Analysis of the combined data set produced a strong phylogenetic signal with high bootstrap and Bremer support and high posterior probability well into the base of the tree. The phylogeny recovered the monophyly of the Halictidae and of all four subfamilies and both tribes, recovered relationships among the subfamilies and tribes congruent with morphology, and provided robust support for the relationships among the numerous genera in the tribe Halictini, sensu Michener (2000). Using our combined nucleotide data set, several recently described halictid fossils from the Oligocene and Eocene, and recently developed Bayesian methods, we estimated the antiquity of major clades within the family. Our results indicate that each of the four subfamilies arose well before the Cretaceous-Tertiary boundary and suggest that the early radiation of halictid bees involved substantial African-South American interchange roughly coincident with the separation of these two continents in the late Cretaceous. This combination of single-copy nuclear genes is capable of recovering Cretaceous-age divergences in bees with high levels of support. We propose that LW opsin, wingless, and EF-1 alpha(F2 copy) may be useful in resolving relationships among bee families and other Cretaceous-age insect lineages.     

Behavioral Phylogeny of Corbiculate Apidae (Hymenoptera; Apinae), with Special Reference to Social Behavior

The phylogenetic relationships among the four tribes of corbiculate bees (Euglossini, Bombini, Meliponini, and Apini) are controversial. There is substantial incongruence between morphological and molecular data, and the single origin of eusociality is questionable. The use of behavioral characters by previous workers has been restricted to some typological definitions, such as solitary and eusocial. Here, I expand the term "social" to 42 characters and present a tree based only on behavioral characters. The reconstructed relationships were similar to those observed in morphological and "total evidence" analyses, i.e., Euglossini + (Bombini + (Meliponini + Apini)), all of which support a single origin of eusociality.     

Analysis of family-level relationships in bees (Hymenoptera: Apiformes) using 28S and two previously unexplored nuclear genes: CAD and RNA polymerase II

We analyzed a combined data set of two protein-coding nuclear genes (CAD and RNA polymerase II) and a nuclear ribosomal gene (28S D2-D4 region) for 68 bee species and 11 wasp outgroups. Our taxon sampling included all seven extant bee families, 17 of 20 subfamilies, and diverse tribes. Wasp outgroups included the two families most closely related to bees: Crabronidae and Sphecidae. We analyzed the combined and single gene data sets using parsimony and Bayesian methods, which yielded largely congruent results. Our results provide reasonably strong support for family and subfamily-level relationships among bees. Our data set strongly supports the sister-group relationship of the Colletidae and Stenotritidae, and places Halictidae as sister to this clade combined. Our analyses place the Melittidae and the long-tongued (LT) bee clade (Apidae+Megachilidae) near the base of the tree with Colletidae (and Stenotritidae) in a fairly highly derived position. This topology ("Melittidae-LT basal") was obtained in previous morphological studies under certain methods of character coding. A more widely accepted tree topology that places Colletidae (and/or Stenotritidae) as sister to all other bees ("Colletidae basal") is not supported by our data. The "Melittidae-LT basal" hypothesis may better explain patterns in the bee fossil record as well as historical biogeography of certain bee groups. Our results provide new insights into higher-level bee phylogeny and indicate that CAD, RNA polymerase II, and 28S are useful data sets for resolving Cretaceous-age divergences in bees and other Hymenoptera.     

Australian Lasioglossum + Homalictus form a monophyletic group: resolving the "Australian enigma"

The bee genus Lasioglossum includes > 1,000 species of bees distributed on all continents except Antarctica. Lasioglossum is a major component of the bee fauna in the Holarctic, Ethiopian, and Asian regions and is an important group for investigating the evolution of social behavior in bees. Given its cosmopolitan distribution, the historical biogeography of the genus is of considerable interest. We reconstructed phylogenetic relationships among the subgenera and species within Lasioglossum s.s., using DNA sequence data from a slowly evolving nuclear gene, elongation factor-1 alpha. The entire data set includes > 1,604 aligned nucleotide sites (including three exons plus two introns) for 89 species (17 outgroups plus 72 ingroups). Parsimony and maximum likelihood analyses provide strong evidence that the primarily Indoaustralian subgenera (Homalictus, Chilalictus, Parasphecodes) form a monophyletic group. Bootstrap support for the Australian clade ranged from 73% to 77%, depending on the method of analysis. Monophyly of the Australian Lasioglossum suggests that a single colonization event (by way of Southeast Asia and New Guinea) gave rise to a lineage of > 350 native Indoaustralian bees. We discuss the implications of Australian monophyly for resolving the "Australian enigma"--the similarity in social behavior among the Australian halictine bees relative to that of Holarctic groups.     

Phylogenetic analysis of the corbiculate Apinae based on morphology of the sting apparatus (Hymenoptera: Apidae)

This study aimed to test the various competing hypotheses regarding the relationships among the four tribes of corbiculate apine bees (Euglossini “orchid bees”, Bombini “bumble bees”, Meliponini “stingless bees”, and Apini “honey bees”) with a completely new set of previously unstudied morphological characters derived from the sting apparatus. The result was one most parsimonious tree of 49 steps, CI = 89, RI = 93 that is perfectly congruent with most studies based on morphological and combined morphological/molecular data, i.e., Euglossini + (Bombini + (Meliponini + Apini)), supporting a well accepted scenario of social evolution for these bees. This data matrix was then combined with other published matrices for this group in order to perform simultaneous analyses. The problem of how to best combine the multiple matrices that did not use the same exemplars was investigated. © The Willi Hennig Society 2007.     

Task-partitioned nectar transfer in stingless bees: work organisation in a phylogenetic context

Abstract 1. The eusocial corbiculate bee tribes comprise the Apini (honey bees), Bombini (bumble bees), and Meliponini (stingless bees). Honey bee foragers ( Apis ) transfer nectar to receiver bees within the nest. This is an example of task partitioning, in which a task is split into sub-tasks connected by material transfer. Nectar transfer does not occur in Bombini. Although it is reported in some species of Meliponini, it has not been subject to detailed study.
2. Nectar transfer was investigated in five genera of Meliponini from Yucatan, Mexico ( Melipona , Trigona , Scaptotrigona , Nannotrigona , and Plebeia ). Nectar transfer occurred in all species and for > 99% of foragers. Multiple transfer, in which a forager unloads nectar to more than one receiver, occurred but at a lower level than in Apis . In M. beecheii , multiple transfer was associated strongly with putative recruitment dances.
3. The data provide some support for the hypothesis that task partitioning is favoured by large colony size, in that the Meliponini never have small colonies because colonies are swarm founded. This ensures that colonies are always large enough to prevent delays in finding a transfer partner imposing high costs. Further tests of this hypothesis are suggested.
4. Viewed in a phylogenetic context, the most parsimonious interpretation is that nectar transfer evolved once in the clade (Apini + Meliponini).     

Evolution of green lacewings (Neuroptera: Chrysopidae): a molecular supermatrix approach

We present a time‐calibrated phylogeny of the charismatic green lacewings (Neuroptera: Chrysopidae). Previous phylogenetic studies on the family using DNA sequences have suffered from sparse taxon sampling and/or limited amounts of data. Here we combine all available previously published DNA sequence data and add to it new DNA sequences generated for this study. We analysed these data in a supermatrix using Bayesian and maximum likelihood methods and provide a phylogenetic hypothesis for the family that recovers strong support for the monophyly of all subfamilies and resolves relationships among a large proportion of chrysopine genera. Chrysopinae tribes Leucochrysini and Belonopterygini were recovered as monophyletic sister clades, while the species‐rich tribe Chrysopini was rendered paraphyletic by Ankylopterygini. Relationships among the subfamilies were resolved, although with relatively low statistical support, and the topology varied based on the method of analysis. Greatest support was found for Apochrysinae as sister to Nothochrysinae and Chrysopinae, which is in contrast to traditional concepts that place Nothochrysinae as sister to the rest of the family. Divergence estimates suggest that the stem groups to the various subfamilies diverged during the Triassic‐Jurassic, and that stem groups of the chrysopine tribes diverged during the Cretaceous.     

Bees diversified in the age of eudicots

Reliable estimates on the ages of the major bee clades are needed to further understand the evolutionary history of bees and their close association with flowering plants. Divergence times have been estimated for a few groups of bees, but no study has yet provided estimates for all major bee lineages. To date the origin of bees and their major clades, we first perform a phylogenetic analysis of bees including representatives from every extant family, subfamily and almost all tribes, using sequence data from seven genes. We then use this phylogeny to place 14 time calibration points based on information from the fossil record for an uncorrelated relaxed clock divergence time analysis taking into account uncertainties in phylogenetic relationships and the fossil record. We explore the effect of placing a hard upper age bound near the root of the tree and the effect of different topologies on our divergence time estimates. We estimate that crown bees originated approximately 123 Ma (million years ago) (113–132 Ma), concurrently with the origin or diversification of the eudicots, a group comprising 75 per cent of angiosperm species. All of the major bee clades are estimated to have originated during the Middle to Late Cretaceous, which is when angiosperms became the dominant group of land plants.     

Partitioned Gene-Tree Analyses and Gene-Based Topology Testing Help Resolve Incongruence in a Phylogenomic Study of Host-Specialist Bees (Apidae: Eucerinae)

Incongruence among phylogenetic results has become a common occurrence in analyses of genome-scale data sets. Incongruence originates from uncertainty in underlying evolutionary processes (e.g., incomplete lineage sorting) and from difficulties in determining the best analytical approaches for each situation. To overcome these difficulties, more studies are needed that identify incongruences and demonstrate practical ways to confidently resolve them. Here, we present results of a phylogenomic study based on the analysis 197 taxa and 2,526 ultraconserved element (UCE) loci. We investigate evolutionary relationships of Eucerinae, a diverse subfamily of apid bees (relatives of honey bees and bumble bees) with >1,200 species. We sampled representatives of all tribes within the group and >80% of genera, including two mysterious South American genera, Chilimalopsis and Teratognatha. Initial analysis of the UCE data revealed two conflicting hypotheses for relationships among tribes. To resolve the incongruence, we tested concatenation and species tree approaches and used a variety of additional strategies including locus filtering, partitioned gene-trees searches, and gene-based topological tests. We show that within-locus partitioning improves gene tree and subsequent species-tree estimation, and that this approach, confidently resolves the incongruence observed in our data set. After exploring our proposed analytical strategy on eucerine bees, we validated its efficacy to resolve hard phylogenetic problems by implementing it on a published UCE data set of Adephaga (Insecta: Coleoptera). Our results provide a robust phylogenetic hypothesis for Eucerinae and demonstrate a practical strategy for resolving incongruence in other phylogenomic data sets.     

Insights into the phylogenetic relationships within Cixiidae (Hemiptera: Fulgoromorpha): cladistic analysis of a morphological dataset

Abstract.  According to the most recent classifications proposed, the planthopper family Cixiidae comprises three subfamilies, namely Borystheninae, Bothriocerinae and Cixiinae, the latter with 16 tribes. Here we examine morphological characters to present the first phylogenetic reconstructions within Cixiidae derived from a cladistic analysis. We scored 85 characters of the head, thorax, and male and female genitalia for 50 taxa representative of all cixiid subfamilies and tribes and for six outgroup taxa. Analyses were based on maximum parsimony – using both equally weighted and successive weighting procedures – and Bayesian inferences. The monophyly of most currently accepted tribes and subfamilies was investigated through Templeton statistical tests of alternative phylogenetic hypotheses. The cladistic analyses recover the monophyly of Cixiidae, the subfamily Bothriocerinae, and the tribes Pentastirini, Mnemosynini, and Eucarpiini. Successive weighting and Bayesian inference recover the monophyly of the tribe Gelastocephalini, but only Bayesian inference supports the monophyly of Semoniini. The relationships recovered support the groups [Stenophlepsini (Borystheninae + Bothriocerinae)] arising from the tribe Oecleini, and [Andini + Brixiidini + Brixiini (polyphyletic) + Bennini]. Templeton tests reject the alternative hypothesis of a monophyletic condition for the tribe Pintaliini as presently defined.     

Phylogenetic relationships and host-plant evolution within the basal clade of Halictidae (Hymenoptera, Apoidea)

Bees are among the most important pollinators of angiosperm plants. Many bee species show narrow host‐plant preferences, reflected both in behavioral and morphological adaptations to particular attributes of host‐plant pollen or floral morphology. Whether bee host‐plant associations reflect co‐cladogenesis of bees and their host plants or host‐switches to unrelated host plants is not clear. Rophitinae is a basal subfamily of Halictidae in which most species show narrow host‐plant preferences (oligolecty). We reconstructed the phylogenetic relationships among the rophitine genera using a combination of adult morphology (24 characters) and DNA sequence data (EF‐1α, LW rhodopsin, wingless; 2700 bp total). The data set was analyzed by parsimony, maximum likelihood and Bayesian methods. All methods yielded highly congruent results. Using the phylogeny, we investigated the pattern of host‐plant association as well as the historical biogeography of Rophitinae. Our biogeographical analysis suggests a number of dispersal/vicariance events: (1) a basal split between North America and South America (most likely a dispersal from South America to North America), and (2) at least two subsequent interchanges between North America and Eurasia (presumably via the northern hemisphere land bridges). Our analysis of host‐plant associations indicates that Rophitinae specialized on a closely related group of angiosperm orders in the Euasterid I clade (mainly Gentianales, Lamiales and Solanales). However, there is little evidence of cocladogenesis between bees and plants and strong evidence of host switches to unrelated host plants. Based on our phylogenetic results we describe two new tribes of Rophitinae: Conanthalictini new tribe (including the genus Conanthalictus) and Xeralictini new tribe (including Xeralictus and Protodufourea). © The Willi Hennig Society 2007.     

Phylogeny of the orchid bees (Hymenoptera: Apinae: Euglossini): DNA and morphology yield equivalent patterns

Orchid bees (Euglossini) are spectacular long-tongued Neotropical bees important in the pollination of Neotropical long-corolla flowers, particularly some orchids. Besides remarkably long tongues, males in particular exhibit other flower-related adaptations, including setal brushes on the foretarsi used for rasping the petals of orchids while collecting aromatic compounds. These compounds are stored in large swollen tibiae and are thought to play an important role in courtship behavior. Euglossini are also unusual in lacking sociality; they are the only tribe among the corbiculate bees that are not eusocial, and two of the genera are cleptoparasitic. Each genus exhibits distinct behavioral traits including nest architecture and host-parasite interactions, yet their evolution is unknown. Despite previous phylogenetic studies of on morphological characters, the relationships among the five euglossine genera remain under debate. We investigate euglossine generic relationships using DNA sequence data from four genes and new morphological characters. The morphological and molecular data yield congruent evolutionary patterns, and combining the data gives a fully resolved and well supported phylogeny of Euglossini.     

Phylogeny and a new tribal classification of the Panicoideae s.l. (Poaceae) based on plastid and nuclear sequence data and structural data

? Premise of the study: The subfamily Panicoideae (Poaceae) encompasses nearly one-third of the diversity of grass species, including important crops such as maize and sugarcane. Previous analyses recovered strong support for a Panicoideae+Centothecoideae lineage within the diverse Panicoideae+Arundinoideae+Chloridoideae+Micrairoideae+Aristidoideae+Danthonioideae (PACMAD) clade, although support for internal relationships was inconsistent. The objectives of this research were to (1) further test the monophyly of each subfamily and previously recovered clades within the Panicoideae+Centothecoideae lineage, (2) establish phylogenetic relationships among these groups, and (3) propose a new tribal classification for this lineage based explicitly on the phylogeny. ? Methods: Maximum parsimony and Bayesian inference analyses of 37 taxa were based on previously published sequences (ndhF and rpl16 intron) and on new plastid and nuclear (rbcL and granule-bound starch synthase I) sequence data as well as structural data. ? Key results. The Panicoideae+Centothecoideae lineage and a majority of the clades identified in previous analyses continue to be robustly supported, but resolution along the backbone of the topology remains elusive. Support for the monophyly of both subfamilies was lacking although support values for some clades increased. The tribes Centotheceae and Arundinelleae were confirmed as polyphyletic. ? Conclusions: Subfamily Centothecoideae is formally submerged into the Panicoideae, and a new tribal classification for the expanded Panicoideae is proposed based explicitly on the phylogeny. This classification includes 12 tribes of which Chasmanthieae and Zeugiteae are segretated from the Centotheceae; Tristachyideae is segregated from Arundinelleae, and a new tribe, Cyperochloeae, is validated to accommodate two isolated genera. A key to the tribes is provided.     

Molecular phylogeny of a circum-global, diverse gastropod superfamily (Cerithioidea: Mollusca: Caenogastropoda): pushing the deepest phylogenetic limits of mitochondrial LSU rDNA sequences

The Cerithioidea is a very diverse group of gastropods with ca. 14 extant families and more than 200 genera occupying, and often dominating, marine, estuarine, and freshwater habitats. While the composition of Cerithioidea is now better understood due to recent anatomical and ultrastructural studies, the phylogenetic relationships among families remain chaotic. Morphology-based studies have provided conflicting views of relationships among families. We generated a phylogeny of cerithioideans based on mitochondrial large subunit rRNA and flanking tRNA gene sequences (total aligned data set 1873 bp). Nucleotide evidence and the presence of a unique pair of tRNA genes (i.e., threonine + glycine) between valine-mtLSU and the mtSSU rRNA gene support conclusions based on ultrastructural data that Vermetidae and Campanilidae are not Cerithioidea, certain anatomical similarities being due to convergent evolution. The molecular phylogeny shows support for the monophyly of the marine families Cerithiidae [corrected], Turritellidae, Batillariidae, Potamididae, and Scaliolidae as currently recognized. The phylogenetic data reveal that freshwater taxa evolved on three separate occasions; however, all three recognized freshwater families (Pleuroceridae, Melanopsidae, and Thiaridae) are polyphyletic. Mitochondrial rDNA sequences provide valuable data for testing the monophyly of cerithioidean [corrected] families and relationships within families, but fail to provide strong evidence for resolving relationships among families. It appears that the deepest phylogenetic limits for resolving caenogastropod relationships is less than about 245--241 mya, based on estimates of divergence derived from the fossil record.     

Evaluating alternative hypotheses for the origin of eusociality in corbiculate bees

We use a likelihood-based statistical test to evaluate the extent to which the available molecular data sets can be used to falsify alternative phylogenetic hypotheses describing the inter-relationship among corbiculate bee tribes. Based on the results of this test, we explore three alternative models of behavioural character state evolution and evaluate the support each model has for single-origin versus dual-origin hypotheses for 'highly' eusocial behaviour. We show that only one of four data sets could statistically reject any of the 15 possible outgroup-rooted phylogenetic hypotheses. However, a cytochrome b data set rejected all but three alternative topologies. Using this information, a simple model of behavioural character state evolution, in which transitions between solitary/communal, 'primitively' eusocial, and 'highly' eusocial are unconstrained, supports single-origin hypotheses for 'highly' eusocial behaviour, in spite of phylogenetic uncertainty. By contrast, an ordered model, in which 'highly' eusocial is constrained to be an evolutionarily terminal state, supports a dual-origins hypothesis. Our results show that the molecular phylogenetic evidence favouring a dual-origins hypothesis for 'highly' eusocial behaviour is, at present, conditional on information from one gene (cyt b) and on specific, though likely realistic, assumptions regarding the nature of eusocial evolution.     

Phylogeny of colletid bees (Hymenoptera: Colletidae) inferred from four nuclear genes

Colletidae comprise approximately 2500 species of bees primarily distributed in the southern continents (only two colletid genera are widely distributed: Colletes and Hylaeus). Previously published studies have failed to resolve phylogenetic relationships on a worldwide basis and this has been a major barrier to the progress of research regarding systematics and evolution of colletid bees. For this study, data from four nuclear gene loci: elongation factor-1alpha (F2 copy), opsin, wingless, and 28S rRNA were analyzed for 122 species of colletid bees, representing all subfamilies and tribes currently recognized; 22 species belonging to three other bee families were used as outgroups. Bayesian, maximum likelihood, and parsimony methods were employed to investigate the phylogenetic relationships within Colletidae and resulted in highly congruent and well-resolved trees. The phylogenetic results show that Colletidae are monophyletic and that all traditionally recognized subfamilies (except Paracolletinae) are also strongly supported as monophyletic. Our phylogenetic hypothesis provides a framework within which broad questions related to the taxonomy, biogeography, morphology, evolution, and ecology of colletid bees can be addressed.     

Multiple molecular data sets suggest independent origins of highly eusocial behavior in bees (Hymenoptera:Apinae)

Different views of the pattern of social evolution among the highly eusocial bees have arisen as a result of discordance between past molecular and morphology-based phylogenies. Here we present new data and taxa for four molecular data sets and reassess the morphological characters available to date. We show there is no significant character incongruence between four molecular data sets (two nuclear and two mitochondrial), but highly significant character incongruence leads to topological incongruence between the molecular and morphological data. We investigate the effects of using different outgroup combinations to root the estimated tree. We also consider various ways in which biases in the sequence data could be misleading, using several maximum likelihood models, LogDet corrections, and spectral analyses. Ultimately, we concede there is strong discordance between the molecular and morphological data partitions and appropriately apply the conditional combination approach in this case. We also find two equally well supported placements of the root for the molecular trees, one supported by 16S and 28S sequences, the other supported by cytochrome b and opsin. The strength of the evidence leads us to accept two equally well supported hypotheses based on analyses of the molecular data sets. These are the most rigorously supported hypotheses of corbiculate bee relationships at this time, and frame our argument that highly eusocial behavior within the corbiculate bees evolved twice independently.     

Phylogeny and biogeography of the flowering plant genus Styrax (Styracaceae) based on chloroplast DNA restriction sites and DNA sequences of the internal transcribed spacer region

Phylogenetic relationships within the flowering plant genus Styrax were investigated with DNA sequence data from the internal transcribed spacer (ITS) region of nuclear ribosomal DNA (nrDNA) and with chloroplast DNA restriction site data from the genes trnK, rpoC1, and rpoC2. The data sets from each genome were analyzed separately and in combination with parsimony methods. The results strongly support the monophyly of each of the four series of the genus but provide little phylogenetic resolution among them. Reticulate evolution may at least partly explain discordance between the molecular phylogenetic estimates and a prior morphological estimate within series Cyrta. The historical biogeography of the genus was inferred with unweighted parsimony character optimization of trees recovered from a combined ITS and morphological data set, after a series of combinability tests for data set congruence was conducted. The results are consistent with the fossil record in supporting a Eurasian origin of Styrax. The nested phylogenetic position of the South American members of the genus within those from southern North America and Eurasia suggests that the boreotropics hypothesis best explains the amphi-Pacific tropical disjunct distribution occurring within section Valvatae. The pattern of relationship recovered among the species of section Styrax ((western North America + western Eurasia) (eastern North America + eastern Eurasia)) is rare among north-temperate Tertiary forest relicts. The monophyly of the group of species from western North America and western Eurasia provides qualified support for the Madrean-Tethyan hypothesis, which posits a Tertiary floristic connection among the semiarid regions in which these taxa occur. A single vicariance event between eastern Asia and eastern North America accounts for the pattern of relationship among intercontinental disjuncts in series Cyrta.