A phylogenetic analysis of Diurideae (Orchidaceae) based on plastid DNA sequence data

DNA sequence data from plastid matK and trnL-F regions were used in phylogenetic analyses of Diurideae, which indicate that Diurideae are not monophyletic as currently delimited. However, if Chloraeinae and Pterostylidinae are excluded from Diurideae, the remaining subtribes form a well-supported, monophyletic group that is sister to a "spiranthid" clade. Chloraea, Gavilea, and Megastylis pro parte (Chloraeinae) are all placed among the spiranthid orchids and form a grade with Pterostylis leading to a monophyletic Cranichideae. Codonorchis, previously included among Chloraeinae, is sister to Orchideae. Within the more narrowly delimited Diurideae two major lineages are apparent. One includes Diuridinae, Cryptostylidinae, Thelymitrinae, and an expanded Drakaeinae; the other includes Caladeniinae s.s., Prasophyllinae, and Acianthinae. The achlorophyllous subtribe Rhizanthellinae is a member of Diurideae, but its placement is otherwise uncertain. The sequence-based trees indicate that some morphological characters used in previous classifications, such as subterranean storage organs, anther position, growth habit, fungal symbionts, and pollination syndromes have more complex evolutionary histories than previously hypothesized. Treatments based upon these characters have produced conflicting classifications, and molecular data offer a tool for reevaluating these phylogenetic hypotheses.     

Molecular phylogenetics of Prescottiinae s.l. and their close allies (Orchidaceae, Cranichideae) inferred from plastid and nuclear ribosomal DNA sequences

The Andes are a cradle of orchid evolution, but most phylogenetic studies of Orchidaceae in this biodiversity hotspot have dealt with epiphytic epidendroid lineages. Here we present a study on neotropical, terrestrial, orchidoid taxa of Prescottiinae s.l. (8 genera, ～100 species), which are adapted to some of the highest elevation habitats on earth that support orchids. They are currently included within an expanded concept of Cranichidinae in the tribe Cranichideae, but DNA sequence data show that neither Prescottiinae s.l. nor Cranichidinae s.s. are monophyletic. Prescottiinae s.l. consist of two strongly supported lineages: the Altensteinia and Prescottia clades, which have closer affinities to Spiranthinae than to Cranichidinae. The Prescottia clade comprises two well-supported subclades, one including most sampled species of Prescottia and a second one with Pseudocranichis thysanochila sister to Prescottia tubulosa. As a group, they are sister to Spiranthinae. Sister to this pair is the Altensteinia clade comprised of six genera, whose intergeneric relationships are well resolved. Finally, Cranichidinae s.s. is sister to all three of these clades. Morphological and ecological features distinguishing the major groups are discussed, as are potential synapomorphies to define them. The reconstructed phylogeny indicates that the classification of Cranichideae needs to be reexamined.     

Phylogeny and morphological evolution of tribe Menispermeae (Menispermaceae) inferred from chloroplast and nuclear sequences

The Menispermaceae family contains ca. 72 genera with 450 species that are almost entirely tropical. Its phylogeny at the tribal level has never been examined using molecular data. Here we used DNA sequences of the chloroplast matK gene and trnL-F regions, and the nuclear ITS region to study the delimitation and position of the tribe Menispermeae within the family and its subtribal monophyletic groups. Family-wide phylogenetic analyses of the chloroplast data produced two strongly supported clades. The first clade contains two subclades: Coscinieae including Arcangelisia and Anamirta, and Tinosporeae sensu lato including Fibraureae, supported by morphological characters, such as traits of the cotyledon, stylar scar and embryo. The second clade consists of the tribes Menispermeae sensu DC. and Tiliacoreae Miers. All our analyses surprisingly recognized that tribe Menispermeae is not monophyletic unless tribe Tiliacoreae is included, suggesting that characters of cotyledon and stylar scar are very important for the infrafamilial classification, and that endosperm presence vs. absence was over-emphasized in traditionally tribal division of the family. Our topologies indicate a secondary loss of endosperm. The monophyly of two subtribes of the tribe Menispermeae, Stephaniinae and Cissampelinae, is supported by the cpDNA and ITS data, as well as by morphological characters, including aperture types and shapes, and colpal membrane features of pollen grains, and sepal number of male flowers. The Cocculinae was recognized as a paraphyletic group containing the remaining genera of the tribe Menispermeae.     

Root character evolution and systematics in Cranichidinae, Prescottiinae and Spiranthinae (Orchidaceae, Cranichideae)

Background and Aims: Previous studies have suggested that velamen characteristicsare useful as taxonomic markers in Orchidaceae. Members of tribeCranichideae have been assigned to two velamen types constructedbased on combinations of characters such as the presence ofsecondary cell-wall thickenings and pores. However, such charactershave not been analysed on an individual basis in explicit cladisticanalyses. Methods: The micromorphology of roots of 26 species of Cranichideae wasexamined through scanning electron microscopy and light microscopy,scoring the variation and distribution of four characters: numberof velamen cell layers, velamen cell-wall thickenings, presenceand type of tilosomes, and supraendodermal spaces. The lastthree characters were analysed cladistically in combinationwith DNA sequence data of plastid trnK/matK and nuclear ribosomalinternal transcribed spacer (ITS) regions and optimized on theresulting phylogenetic tree. Key Results: Thickenings of velamen cell walls group Prescottiinae with Spiranthinae,whereas tilosomes, documented here for the first time in Cranichideae,provide an unambiguous synapomorphy for subtribe Spiranthinae.Supraendodermal spaces occur mostly in species dwelling in seasonallydry habitats and appear to have evolved three times. Conclusions: Three of the four structural characters assessed are phylogeneticallyinformative, marking monophyletic groups recovered in the combinedmolecular–morphological analysis. This study highlightsthe need for conducting character-based structural studies toovercome analytical shortcomings of the typological approach.     

A phylogeny of Packera (Senecioneae; asteraceae) based on internal transcribed spacer region sequence data and a broad sampling of outgroups

A phylogeny of Packera is presented based on sequence data from the internal transcribed spacer region of nuclear ribosomal DNA of 26 species (28 populations) of Packera and 23 outgroup taxa, including representatives of all three subtribes of the Senecioneae. The results support a Mexican origin for Packera, with its closest relatives found among Old World taxa in the subtribe Senecioninae, such as Senecio jacobaea and Pericallis. Packera species from the west coast of the United States, previously included in the section Bolanderi of Greenman, are part of a basal assemblage including species of Greenman's Mexican section Sanguisorboidei. The rest of Packera separates into two sister groups, one containing species from the Arizona-New Mexico region and the other containing more geographically diverse taxa. Among the outgroups, New World Senecio species are monophyletic and two Tussilaginoid assemblages are strongly supported; the Tephroseroid group (Tephroseris and Sinosenecio) plus Petasites combine with the Luina complex to form a clade of north temperate taxa, and the four Mexican genera (Psacalium, Robinsonecio, Barkleyanthus, and Pittocaulon) form a monophyletic group.     

Large‐scale molecular phylogeny of metallic wood‐boring beetles (Coleoptera: Buprestoidea) provides new insights into relationships and reveals multiple evolutionary origins of the larval leaf‐mining habit

The family Buprestidae (jewel beetles or metallic wood‐boring beetles), contains nearly 15 000 species in 522 genera. Together with the small family Schizopodidae (seven species, three genera), they form the superfamily Buprestoidea. Adult Buprestoidea feed on flowers or foliage, whereas larvae are mostly internal feeders, boring in roots or stems, or mining the leaves of woody or herbaceous plants. The subfamilial and tribal classification of Buprestoidea remains unsettled, with substantially different schemes proposed by different workers based on morphology. Here we report the first large‐scale molecular phylogenetic study of the superfamily Buprestoidea based on data from four genes for 141 ingroup species. We used these data to reconstruct higher‐level relationships and to assess the current classification and the origins of the larval leaf‐mining habit within Buprestoidea. In our analyses, the monophyly of Buprestoidea was strongly supported, as was the monophyly of Schizopodidae and its placement sister to Buprestidae. Our results are largely consistent with the generally accepted major lineages of buprestoids, including clearly‐defined agrilines, buprestines–chrysochroines and early‐branching julodines–polycestines. In addition to Schizopodidae, three of the six subfamilies were monophyletic in our study: Agrilinae, Julodinae and the monogeneric Galbellinae (Galbella). Polycestinae was monophyletic with the exception of the enigmatic Haplostethini. Chrysochroinae and Buprestinae were not monophyletic, but were recovered together in a large mixed clade along with Galbella. The interrelationships of Chrysochroinae and Buprestinae were not well resolved; however they were clearly polyphyletic, with chrysochroine genera falling into several different well‐supported clades otherwise comprising buprestine genera. All Agrilinae were contained in a single strongly supported clade. Coraebini were dispersed throughout Agrilinae, with strong nodal support for several clades representing subtribes. Neither Agrilini nor Tracheini were monophyletic. The leaf‐mining genus Paratrachys (Paratracheini) was recovered within the Acmaeoderioid clade, consistent with the current classification, and confirming the independent origins of leaf‐mining within Polycestinae and Agrilinae. Additionally, our results strongly suggest that the leaf‐mining agriline tribe Tracheini is polyphyletic, as are several of its constituent subtribes. External root feeding was likely the ancestral larval feeding habit in Buprestoidea. The apparent evolutionary transitions to internal feeding allowed access to a variety of additional plant tissues, including leaves. Interestingly, the several genera of leaf‐mining agrilines do not form a monophyletic group. Many of these genera are diverse and highly specialized, possibly indicating adaptive radiations.     

Phylogenetic relationships among the Ithomiini (Lepidoptera: Nymphalidae) inferred from one mitochondrial and two nuclear gene regions

Abstract.  A phylogenetic hypothesis for the tribe Ithomiini (Lepidoptera: Nymphalidae: Danainae) is presented, based on sequences of the mitochondrial cytochrome oxidase subunits I and II (COI–COII) region and regions of the nuclear genes wingless and Elongation factor 1-alpha . Branch support for each clade is assessed, and a partition congruence index is used to explore conflict among gene regions. The monophyly of the clade is strongly supported, as are many of the traditionally recognized subtribes and genera. The data imply paraphyly of some genera and tribes, but largely support recent classifications and phylogenetic hypotheses based on morphological characters.     

Higher-level phylogeny of Hydrophilinae (Coleoptera: Hydrophilidae) based on larval, pupal and adult characters

Abstract.  A phylogenetic analysis, at a tribal and subtribal level, of the subfamily Hydrophilinae was conducted. The analysis was based on twenty-nine taxa (twenty-three genera) and 148 characters (fifty-eight from immature stages and bionomics, and ninety from adults). According to the present study, Hydrophilinae is monophyletic, and except for the tribe Hydrophilini which appears as polyphyletic (it includes the subtribes Hydrophilina, Hydrobiina, and Acidocerina), the remaining tribes are monophyletic. The tribes Berosini and Chaetarthriini form the basal clade of Hydrophilinae. One unexpected result is the relationship between the subtribe Hydrobiina and the tribe Sperchopsini, which form a well-supported clade. The final tree has the following structure: (((((Sperchopsini Hydrobiina) (Anacaenini Laccobiini)) Acidocerina) Hydrophilina) (Berosini Chaetarthriini)). The results partially disagree with the phylogeny presented by Hansen, in 1991, which was based mostly on adult characters. Several evolutionary trends are briefly discussed: the types of egg case, the morphology of the clypeolabrum, mouthparts, legs, and breathing adaptations in larvae.     

A molecular phylogeny of the grass subfamily Panicoideae (Poaceae) shows multiple origins of C4 photosynthesis

DNA sequence data from the chloroplast gene ndhF were analyzed to estimate the phylogeny of the subfamily Panicoideae, with emphasis on the tribe Paniceae. Our data suggest that the subfamily is divided into three strongly supported clades, corresponding to groups with largely identical base chromosome numbers. Relationships among the three clades are unclear. In unweighted parsimony analyses, the two major clades with x = 10 (Andropogoneae and x = 10 Paniceae) are weakly supported as sister taxa. The third large clade corresponds to x = 9 Paniceae. In analyses under implied weight, the two clades of Paniceae are sisters, making the tribe monophyletic. Neither resolution is strongly supported.Our molecular phylogenies are not congruent with previous classifications of tribes or subtribes. Based on this sample of species, we infer that C(4) photosynthesis has evolved independently several times, although a single origin with multiple reversals and several reacquisitions is only slightly less parsimonious. The phosphoenol pyruvate carboxykinase (PCK) subtype of C(4) photosynthesis has evolved only once, as has the NAD-malic enzyme (ME) subtype; all other origins are NADP-ME. Inflorescence bristles are apparently homologous in the genera Setaria and Pennisetum, contrary to opinions of most previous authors. Some genera, such as Digitaria, Echinochloa, and Homolepis are supported as monophyletic. The large genus Paspalum is shown to be paraphyletic, with Thrasya derived from within it. As expected, Panicum is polyphyletic, with lineages derived from multiple ancestors across the tree. Panicum subg. Panicum is monophyletic. Panicum subg. Dichanthelium, subg. Agrostoides, and subg. Phanopyrum are unrelated to each other, and none is monophyletic. Only Panicum subg. Dichanthelium sect. Dichanthelium, represented by P. sabulorum and P. koolauense, is monophyletic. Panicum subg. Megathyrsus, a monotypic subgenus including only the species P. maximum, is better placed in Urochloa, as suggested by other authors.     

PHYLOGENETIC RELATIONSHIPS AMONG EXTANT BRACHIOPODS

Abstract— The monophyletic status of the Brachiopoda and phylogenetic relationships within the phylum have long been contentious issues for brachiopod systematists. The relationship of brachiopods to other lophophore-bearing taxa is also uncertain; results from recent morphological and molecular studies are in conflict. To test current hypotheses of relationship, a phylogenetic analysis was completed (using PAUP 3.1.1) with 112 morphological and embryological characters that vary among extant representatives of seven brachiopod superfamilies, using bryozoans, phoronids, pterobranchs and sipunculids as outgroups. In the range of analyses performed, brachiopod monophyly is well supported, particularly by characters of soft anatomy. Arguments concerning single or multiple origins of a bivalved shell are not relevant to recognizing brachiopods as a clade. Articulate monophyly is very strongly supported, but inarticulate monophyly receives relatively weak support. Unlike previous studies, the nature of uncertainties about the clade status of Inarticulata are detailed explicitly here, making them easier to test in the future. Calcareous inarticulates appear to share derived characters with the other inarticulates, while sharing many primitive characters with other calcareous brachiopods (the articulates). Experimental manipulation of the data matrix reveals potential sources of bias in previous hypotheses of brachiopod phylogeny. Although not tested explicitly, lophophorate monophyly is very tentatively supported. Molecular systematic studies of a diverse group of brachiopods and other lophophorates will be particularly welcome in providing a test of the conclusions presented here.     

PHYLOGENETIC RELATIONSHIPS IN SEED PLANTS

Abstract— The phylogenetic relationships of nineteen extant and fossil seed plants are considered. Analysis of 31 characters produced ten topologically similar and equally parsimonious cladograms. A strict consensus tree derived from these cladograms places Lyginopteris as the sister taxon to the other seed plants included. Within this clade all the taxa considered, except medullosans and cycads, form a single monophyletic group defined by the presence of flattened seeds and saccate pollen ("platy-sperms"). Relationships between medullosans, cycads, and "platysperms" were not resolved, but within the "platysperm" clade conifers and cordaites ( Cordaixylon, Mesoxylon ) + Ginkgo form a monophyletic group ("coniferophytes"). The "higher platysperms" (glossopterids, Caytonia , corystosperms, Bennettitales, Pentoxylon , Gnetales, and angiosperms) are also monophyletic, but their relationship to "coniferophytes," peltasperms, and Callistophyton is unresolved. Pentoxylon is placed as sister taxon to the Bennettitales, and together they form the sister group to a clade in which Gnetales and angiosperm are sister taxa. The Bennettitales + Pentoxylon + Gnetales + angiosperms ("anthophytes") form a monophyletic sister group to the corystosperms. This analysis is compared with current classifications of seed plants. It does not support a close relationship between Bennettitales and cycads, it provides no evidence for seed plant polyphyly, and it strongly suggests that the current concept of seed ferns has little value in a phylogenetic context.     

A Phylogenetic Study of Tribe Euphorbieae (Euphorbiaceae)

A phylogenetic investigation of a monophyletic lineage of spurge plants, tribe Euphorbieae, was conducted to elucidate evolutionary relationships, to clarify biogeographic patterns, and to reexamine the previous classification of Euphorbieae. Cladistic analyses of the 52 morphological characters of 61 species resulted in 2922 equally most parsimonious trees of 193 steps with a consistency index of 0.34. The strict consensus tree indicates genus Anthostema of subtribe Anthosteminae as a likely sister group to all other members of tribe Euphorbieae. The morphological data support a monophyletic origin of subtribe Euphorbiinae, but the subtribes Anthosteminae and Neoquillauminiinae did not form monophyletic groups. Although the previous taxonomic treatments within tribe Euphorbieae have supported the generic status of Pedilanthus, Monadenium, Synadenium, Chamaesyce, and Elaeophorbia, the results of this analysis do not support generic placement of them based on cladistic principles. Recognition of these groups as genera results in Euphorbia becoming a paraphyletic group. One solution to this problem in Euphorbieae is to divide the largest genus Euphorbia into several monophyletic genera and to keep the generic ranks for previously recognized genera. The distribution of basal endemic genera in Euphorbieae showed African and east Gondwanan affinities and strongly indicated that the ancestor of Euphorbieae originated prior to the breakup of Gondwanaland from an old group in Euphorbiaceae. However, some recent African taxa of Euphorbia should be interpreted by transoceanic dispersal from the New World ancestors.     

Diversification of inflorescence development in the PCK clade (Poaceae: Panicoideae: Paniceae)

In grasses, inflorescence diversification and its correlation with species evolution are intriguing and not well understood. Part of this problem lies in our lack of comprehension about the inflorescence morphological complexity of grasses. We focused our study on the PCK clade (named for phosphoenol pyruvate carboxykinase), a well-supported monophyletic group for which the relationships among its taxa are not well resolved. Interestingly, the PCK clade has an extensive diversity of adult inflorescence forms. A comparative developmental approach can help us to understand the basis of such morphological differences as well as provide characters that can be used in phylogenetic studies of the group. Using SEM studies, we demonstrate that inflorescence morphology in this clade is even more complex than what is typically observed in adult forms. We describe a number of new characters, and some classical features previously used for taxonomic purposes are redefined on the basis of development. We also define four morphological groups combining adult inflorescence form and development, and we discuss some of the evolutionary aspects of inflorescence diversification in the PCK clade. Taxonomic delimitation among genera in the PCK clade remains confusing and unclear where molecular and morphological studies support different classifications.     

Molecular phylogenetic analysis of Passifloraceae sensu lato (Malpighiales) based on plastid and nuclear DNA sequences

A phylogenetic analysis of Passifloraceae sensu lato was performed using rbcL, atpB, matK, and 18S rDNA sequences from 25 genera and 42 species. Parsimony analyses of combined data sets resulted in a single most parsimonious tree, which was very similar to the 50% majority consensus tree from the Bayesian analysis. All nodes except three were supported by more than 50% bootstrap. The monophyly of Passifloraceae s.l. as well as the former families, Malesherbiaceae, Passifloraceae sensu stricto, and Turneraceae were strongly supported. Passifloraceae s.s. and the Turneraceae are sisters, and form a strongly supported clade. Within Passifloraceae s.s., the tribes Passifloreae and Paropsieae are both monophyletic. The intergeneric relationships within Passifloraceae s.s. and Turneraceae are roughly correlated with previous classification systems. The morphological character of an androgynophore/gynophore is better used for characterizing genera grouping within Passifloraceae s.s. Other morphological characters such as the corona and aril are discussed.     

A phylogenetic analysis of tribe Areceae (Arecaceae) using two low-copy nuclear genes

 A phylogenetic study of the largest tribe of palms, the Areceae, was conducted using sequences of two low-copy nuclear genes. Previous morphological and plastid DNA studies have not supported the monophyly of the tribe, but have placed its members in a large clade that includes the monophyletic tribes Geonomeae, Cocoeae, Podococceae, and Hyophorbeae. We analyzed this large clade to test the monophyly of tribe Areceae with nuclear data, to explore relationships among its subtribes, and to identify other monophyletic groups. For 54 palm species, including members of all 17 subtribes of tribe Areceae, we sequenced regions of the malate synthase (MS) and phosphoribulokinase (PRK) genes. Simultaneous analysis of these regions revealed 52 shortest trees, all of which resolved tribe Areceae as polyphyletic. Subtribes Iguanurinae, Dypsidinae, Oncospermatinae, and Arecinae were also resolved as polyphyletic. A clade of Indo-Pacific taxa was resolved with strong support, and would be a suitable target for more focused study. Received February 7, 2001; accepted April 9, 2002 Published online: December 3, 2002     

Phylogenetic relationships of Cranichidinae and Prescottiinae (Orchidaceae, Cranichideae) inferred from plastid and nuclear DNA sequences

Background and Aims

Phylogenetic relationships of subtribes Cranichidinae and Prescottiinae, two diverse groups of neotropical terrestrial orchids, are not satisfactorily understood. A previous molecular phylogenetic study supported monophyly for Cranichidinae, but Prescottiinae consisted of two clades not sister to one another. However, that analysis included only 11 species and eight genera of these subtribes. Here, plastid and nuclear DNA sequences are analysed for an enlarged sample of genera and species of Cranichidinae and Prescottiinae with the aim of clarifying their relationships, evaluating the phylogenetic position of the monospecific genera Exalaria, Ocampoa and Pseudocranichis and examining the value of various structural traits as taxonomic markers.

Methods

Approx. 6000 bp of nucleotide sequences from nuclear ribosomal (ITS) and plastid DNA (rbcL, matK-trnK and trnL-trnF) were analysed with cladistic parsimony and Bayesian inference for 45 species/14 genera of Cranichidinae and Prescottiinae (plus suitable outgroups). The utility of flower orientation, thickenings of velamen cell walls, hamular viscidium and pseudolabellum to mark clades recovered by the molecular analysis was assessed by tracing these characters on the molecular trees.

Key Results

Spiranthinae, Cranichidinae, paraphyletic Prescottia (with Pseudocranichis embedded), and a group of mainly Andean ‘prescottioid’ genera (the ‘Stenoptera clade’) were strongly supported. Relationships among these clades were unresolved by parsimony but the Bayesian tree provided moderately strong support for the resolution (Spiranthinae–(Stenoptera clade-(Prescottia/Pseudocranichis–Cranichidinae))). Three of the four structural characters mark clades on the molecular trees, but the possession of a pseudolabellum is variable in the polyphyletic Ponthieva.

Conclusions

No evidence was found for monophyly of Prescottiinae and the reinstatement of Cranichidinae s.l. (including the genera of ‘Prescottiinae’) is favoured. Cranichidinae s.l. are diagnosed by non-resupinate flowers. Lack of support from parsimony for relationships among the major clades of core spiranthids is suggestive of a rapid morphological radiation or a slow rate of molecular evolution.Key words: Cranichideae, Cranichidinae, matK-trnK, molecular phylogenetics, nrITS, Orchidaceae, Prescottiinae, resupination, trnL-trnF     

Phylogenetic relationships of Early–Middle Ordovician ostracods of Baltoscandia

Phylogenetic analysis of the Early and early Middle Ordovician (Tremadoc and Arenig) ostracod species of Baltoscandia suggests a polyphyletic origin for the suborder Beyrichiocopa. Binodicopes, leiocopes and eridostracans are excluded from the beyrichiocopide clade. An independent origin from the basal ostracods is suggested for the binodicopes and eridostracans. The palaeocopes form a strongly supported monophyletic clade. Within this suborder, the ctenonotellid and the tetradellid families together form a monophyletic clade. The tetradellids are paraphyletic, being a stem-group for the ctenonotellids. Nanopsis nanella , the earliest known ostracod from the Tremadoc, is a basal palaeocope. The early eridostracans Conchoprimitia and Incisua , with their uncomplicated carapace morphology, might be the most primitive ostracods.     

Molecular phylogeny of Nyctaginaceae: taxonomy, biogeography, and characters associated with a radiation of xerophytic genera in North America

The four o'clock family (Nyctaginaceae) has a number of genera with unusual morphological and ecological characters, several of which appear to have a "tendency" to evolve repeatedly in Nyctaginaceae. Despite this, the Nyctaginaceae have attracted little attention from botanists. To produce a phylogeny for the Nyctaginaceae, we sampled 51 species representing 25 genera (of 28-31) for three chloroplast loci (ndhF, rps16, rpl16, and nrITS) and included all genera from North America. Parsimony, likelihood, and Bayesian methods were used to reconstruct the phylogeny for the family. The family is neotropical in origin. A radiation of woody taxa unites Pisonia and Pisoniella with the difficult tropical genera Neea and Guapira, which also form a clade, though neither appears to be monophyletic. This group is sister to a clade containing Bougainvillea, Belemia, and Phaeoptilum. A dramatic radiation of genera occurred in the deserts of North America. The tribe Nyctagineae and its subtribes are paraphyletic, due to over-reliance on a few homoplasious characters, i.e., pollen morphology and involucre presence. Two notable characters associated with the desert radiation are cleistogamy and edaphic endemism on gypsum soils. We discuss evolutionary trends in these traits in light of available data about self-incompatibility and gypsum tolerance in Nyctaginaceae.