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Smithsonian Plant Collections, Guiana Shield: 2002–2012, Karen M. Redden and Kenneth J. Wurdack. Part I provides the collector’s notes on trips with maps by date. Part II lists collection localities, with collection number ranges, habitat descriptions, geographic coordinates, and assisting collectors. Part III lists collections in numerical order with identifications and authors. Part IV lists collections ordered by determined name.

German-born Charles Mohr (1824–1901) spent most of his adult life working as a pharmacist in Mobile, Alabama. In addition, he contributed multiple scientific and popular articles on the botany of the Southeastern United States. Suffering from many physical ailments, he sought treatment in thermal baths and cool climates, including Asheville, North Carolina. There he was able to work with botanists at the newly established Biltmore Herbarium. In early 1900, he and his family moved to Asheville, where he completed reading the proof of his magnum opus, Plant Life of Alabama. He died in Asheville 17 July 1901—two weeks before his book was published—and is buried there.

We report on the discovery and characterization of a novel Fusarium species that produced yellow-orange pseudoflowers on Xyris spp. (yellow-eyed grass; Xyridaceae) growing in the savannas of the Pakaraima Mountains of western Guyana. The petaloid fungal structures produced on infected plants mimic host flowers in gross morphology. Molecular phylogenetic analyses of full-length RPB1 (RNA polymerase largest subunit), RPB2 (RNA polymerase second largest subunit), and TEF1 (elongation factor 1-α) DNA sequences mined from genome sequences resolved the fungus, described herein as F. xyrophilum, sp. nov., as sister to F. pseudocircinatum within the African clade of the F. fujikuroi species complex. Results of a polymerase chain reaction (PCR) assay for mating type idiomorph revealed that single-conidial isolates of F. xyrophilum had only one of the MAT idiomorphs (MAT1-1 or MAT1-2), which suggests that the fungus may have a heterothallic sexual reproductive mode. BLASTn searches of whole-genome sequence of three strains of F. xyrophilum indicated that it has the genetic potential to produce secondary metabolites, including phytohormones, pigments, and mycotoxins. However, a polyketide-derived pigment, 8-O-methylbostrycoidin, was the only metabolite detected in cracked maize kernel cultures. When grown on carnation leaf agar, F. xyrophilum is phenotypically distinct from other described Fusarium species in that it produces aseptate microconidia on erect indeterminate synnemata that are up to 2 mm tall and it does not produce multiseptate macroconidia.

Many aspects of the dynamics of tropical fungal endophyte communities are poorly known, including the influence of host taxonomy, host life stage, host defence, and host geographical distance on community assembly and composition. Recent fungal endophyte research has focused on Hevea brasiliensis due to its global importance as the main source of natural rubber. However, almost no data exist on the fungal community harboured within other Hevea species or its sister genus Micrandra. In this study, we expanded sampling to include four additional Hevea spp. and two Micrandra spp., as well as two host developmental stages. Through culture-dependent and -independent (metagenomic) approaches, a total of 381 seedlings and 144 adults distributed across three remote areas within the Peruvian Amazon were sampled. Results from both sampling methodologies indicate that host developmental stage had a greater influence in community assemblage than host taxonomy or locality. Based on FunGuild ecological guild assignments, saprotrophic and mycotrophic endophytes were more frequent in adults, while plant pathogens were dominant in seedlings. Trichoderma was the most abundant genus recovered from adult trees while Diaporthe prevailed in seedlings. Potential explanations for that disparity of abundance are discussed in relation to plant physiological traits and community ecology hypotheses.

During the environmental impact study for a proposed nickel mine near Weda Bay on Halmahera in North Moluccas (Maluku Utara Province), Indonesia, two unknown Euphorbiaceae were discovered. Morphological comparisons and molecular phylogenetic analyses using four markers (plastid trnL-F and rbcL, and nuclear ribosomal internal transcribed spacer and external transcribed spacer) indicated that they should be recognized as constituting a new, distinct genus of two species, which are described and illustrated here as Weda fragarioides and Weda lutea. The new taxa are members of the Australasian tribe Ricinocarpeae in subfamily Crotonoideae, and they are most closely related to Alphandia. In contrast with the otherwise mostly sclerophyllous Ricinocarpeae, Weda possesses stellate to dendritic hairs, large, long-petiolate, glandular leaves, and inflorescences with a pair of large, leafy, subopposite bracts. The two narrowly distributed species are distinguished from each other by vegetative and floral features, molecular data, and elevational preferences. Leaf elemental analysis of Weda indicated manganese, but not nickel, accumulation. Newly resolved generic relationships and potential morphological synapomorphies within Crotonoideae are discussed, and the circumscription of Ricinocarpeae is expanded from 7 to 11 genera.

Recent systematic treatments for Jamaica have shown that the island is home to seven endemic genera of seed-plants [viz., Dendrocousinsia (Euphorbiaceae), Jacmaia (Asteraceae), Odontocline (Asteraceae), Portlandia (Rubiaceae), Salpixantha (Acanthaceae), Tetrasiphon (Celastraceae), and Zemisia (Asteraceae)]. These taxa account for over 23 species, with three genera (Jacmaia, Tetrasiphon, and Zemisia) being monotypic. Further study of these taxa revealed that five names need typification: (1) Tetrasiphon jamaicensis Urb.; (2) Senecio fadyenii Griseb.; (3) S. fadyenii var. dolichanthus Krug & Urb.; (4) S. hollickii Britton ex Greenm. [all the three Senecio taxa included in Odontocline]; and (5) Gymnanthes elliptica Sw. Lectotypes for these names are designated here. Furthermore, G. elliptica is transferred to Dendrocousinsia and the new combination is made here: D. elliptica (Sw.) Commock & K. Wurdack.

Systematics is the science of discovering, organizing and interpreting the diversity of all living organisms. Recent developments in genomics and biodiversity informatics are transforming systematics and have opened up many new opportunities. Major digitization efforts and developments in biodiversity informatics have helped the systematics community explore ways to enhance the efficiency in organizing, publishing, and utilizing systematic information. At the same time, genomics is rapidly facilitating construction of the tree of life, improving taxonomic classification, and disentangling complex evolutionary histories. In the informatics and genomics era, systematics has an incredible capacity to integrate with computational and exploratory platforms for discovery as well as with other, related disciplines while maintaining its core strengths in biological collections and morphology. We call for the establishment of a new global cyberinfrastructure or Biodiversity Cyberbank that will function as the main repository of many types of biodiversity data to ensure the long-term sustainability of the vast and growing amount of systematic data. This Biodiversity Cyberbank will contain new and efficient analytical pipelines for systematics research, especially for efficiently generating taxonomic treatments (revisions, e-monographs and floras). Integrative systematics requires the training of the next-generation systematists with taxonomic, phylogenomic and informatics skills to address grand questions about biodiversity and its assembly and continue to develop the Biodiversity Cyberbank. Integrative systematics must also proactively educate the public and policy makers on the importance of systematics and collections for addressing the biodiversity crisis of the Anthropocene, and a Biodiversity Cyberbank may represent one powerful tool for outreach.

Background Plukenetia is a small pantropical genus of lianas and vines with variably sized edible oil-rich seeds that presents an ideal system to investigate neotropical and pantropical diversification patterns and seed size evolution. We assessed the biogeography and seed evolution of Plukenetia through phylogenetic analyses of a 5069 character molecular dataset comprising five nuclear and two plastid markers for 86 terminals in subtribe Plukenetiinae (representing 20 of ~ 23 Plukenetia species). Two nuclear genes, KEA1 and TEB, were used for phylogenetic reconstruction for the first time. Our goals were: (1) produce a robust, time-dependent evolutionary framework for Plukenetia using BEAST; (2) reconstruct its biogeographical history with ancestral range estimation in BioGeoBEARS; (3) define seed size categories; (4) identify patterns of seed size evolution using ancestral state estimation; and (5) conduct regression analyses with putative drivers of seed size using the threshold model. Results Plukenetia was resolved into two major groups, which we refer to as the pinnately- and palmately-veined clades. Our analyses suggest Plukenetia originated in the Amazon or Atlantic Forest of Brazil during the Oligocene (28.7 Mya) and migrated/dispersed between those regions and Central America/Mexico throughout the Miocene. Trans-oceanic dispersals explain the pantropical distribution of Plukenetia, including from the Amazon to Africa in the Early Miocene (17.4 Mya), followed by Africa to Madagascar and Africa to Southeast Asia in the Late Miocene (9.4 Mya) and Pliocene (4.5 Mya), respectively. We infer a single origin of large seeds in the ancestor of Plukenetia. Seed size fits a Brownian motion model of trait evolution and is moderately to strongly associated with plant size, fruit type/dispersal syndrome, and seedling ecology. Biome shifts were not drivers of seed size, although there was a weak association with a transition to fire prone semi-arid savannas. Conclusions The major relationships among the species of Plukenetia are now well-resolved. Our biogeographical analyses support growing evidence that many pantropical distributions developed by periodic trans-oceanic dispersals throughout the Miocene and Pliocene. Selection on a combination of traits contributed to seed size variation, while movement between forest edge/light gap and canopy niches likely contributed to the seed size extremes in Plukenetia. Electronic supplementary material The online version of this article (10.1186/s12862-018-1308-9) contains supplementary material, which is available to authorized users.

Pseudoflower formation is arguably the rarest outcome of a plant-fungus interaction. Here we report on a novel putative floral mimicry system in which the pseudoflowers are composed entirely of fungal tissues in contrast to modified leaves documented in previous mimicry systems. Pseudoflowers on two perennial Xyris species (yellow-eyed grass, X. setigera and X. surinamensis) collected from savannas in Guyana were produced by Fusarium xyrophilum, a novel Fusarium species. These pseudoflowers mimic Xyris flowers in gross morphology and are ultraviolet reflective. Axenic cultures of F. xyrophilum produced two pigments that had fluorescence emission maxima in light ranges that trichromatic insects are sensitive to and volatiles known to attract insect pollinators. One of the volatiles emitted by F. xyrophilum cultures (i.e., 2-ethylhexanol) was also detected in the head space of X. laxifolia var. iridifolia flowers, a perennial species native to the New World. Results of microscopic and PCR analyses, combined with examination of gross morphology of the pseudoflowers, provide evidence that the fungus had established a systemic infection in both Xyris species, sterilized them and formed fungal pseudoflowers containing both mating type idiomorphs. Fusarium xyrophilum cultures also produced the auxin indole-3-acetic acid (IAA) and the cytokinin isopentenyl adenosine (iPR). Field observations revealed that pseudoflowers and Xyris flowers were both visited by bees. Together, the results suggest that F. xyrophilum pseudoflowers are a novel floral mimicry system that attracts insect pollinators, via visual and olfactory cues, into vectoring its conidia, which might facilitate outcrossing of this putatively heterothallic fungus and infection of previously uninfected plants.

Andeimalvaperuviana Dorr & C.Romero, sp. nov., the third Peruvian endemic in a small genus of five species, is described and illustrated from a single collection made at high elevation on the eastern slopes of the Andes. Molecular phylogenetic analyses of nuclear ribosomal ITS sequence data resolve a group of northern species of Andeimalva found in Bolivia and Peru from the morphologically very different southern A.chilensis. The new species bears the largest flowers of any Andeimalva and is compared with Bolivian A.mandonii. A revised key to the genus is presented.

The mid-Cenozoic decline of atmospheric CO2 levels that promoted global climate change was critical to shaping contemporary arid ecosystems. Within angiosperms, two CO2 -concentrating mechanisms (CCMs)-crassulacean acid metabolism (CAM) and C4 -evolved from the C3 photosynthetic pathway, enabling more efficient whole-plant function in such environments. Many angiosperm clades with CCMs are thought to have diversified rapidly due to Miocene aridification, but links between this climate change, CCM evolution, and increased net diversification rates (r) remain to be further understood. Euphorbia (∼2000 species) includes a diversity of CAM-using stem succulents, plus a single species-rich C4 subclade. We used ancestral state reconstructions with a dated molecular phylogeny to reveal that CCMs independently evolved 17-22 times in Euphorbia, principally from the Miocene onwards. Analyses assessing among-lineage variation in r identified eight Euphorbia subclades with significantly increased r, six of which have a close temporal relationship with a lineage-corresponding CCM origin. Our trait-dependent diversification analysis indicated that r of Euphorbia CCM lineages is approximately threefold greater than C3 lineages. Overall, these results suggest that CCM evolution in Euphorbia was likely an adaptive strategy that enabled the occupation of increased arid niche space accompanying Miocene expansion of arid ecosystems. These opportunities evidently facilitated recent, replicated bursts of diversification in Euphorbia.

The current study examines all genera of Sapindaceae tribe Paullinieae sensu Acevedo-Rodriguez et al. (2011). Based on molecular phylogenetic analyses of trnL intron and ITS sequence data along with critical evaluation of morphology, supertribe Paulliniodae is newly recognized and morphologically characterized by zygomorphic flowers, thyrses with lateral cincinni, corollas of 4 petals, and alternate leaves with a well-developed distal leaflet. Paulliniodae contains four successively nested subclades designated as tribes Athyaneae, Bridgesieae, Thouinieae, and Paullinieae. Athyaneae contains Athyana and Diatenopteryx, and is composed of trees with exstipulate pinnately compound leaves, and isopolar, spherical, colporate pollen grains. Bridgesieae contains the monospecific shrub genus Bridgesia with exstipulate, simple leaves, and isopolar, spherical, tricolporate pollen grains. Thouinieae is resurrected and amended to include three genera of trees or shrubs with exstipulate trifoliolate or unifoli...

The introduction of Next Generation Sequencing into the disciplines of plant systematics, ecology, and metagenomics, among others, has resulted in a phenomenal increase in the collecting and storing of tissue samples and their respective vouchers. This manual suggests standard practices that will insure the quality and preservation of the tissue and vouchers and their respective data. Although written for use by the Smithsonian Institution botanists it suggests a framework for collecting tissues and vouchers that other research programs can adapt to their own needs. It includes information on collecting voucher specimens, collecting plant tissue intended for genomic analysis, how to manage these collections, and how to incorporate the data into a database management system. It also includes many useful references for collecting and processing collections. We hope it will be useful for a variety of botanists but especially those who know how to collect plants and want to collect tissue samples that will be useful for genomic research, and those who are skilled in lab work and want to know how to properly voucher and record their tissue collections.

Although much attention has been paid to the role of stabilizing selection, empirical analyses testing the role of developmental constraints in evolutionary stasis remain rare, particularly for plants. This topic is studied here with a focus on the evolution of a pollen ontogenetic feature, the last points of callose deposition (LPCD) pattern, involved in the determination of an adaptive morphological pollen character (aperture pattern). The LPCD pattern exhibits a low level of evolution in eudicots, as compared to the evolution observed in monocots. Stasis in this pattern might be explained by developmental constraints expressed during male meiosis (microsporogenesis) or by selective pressures expressed through the adaptive role of the aperture pattern. Here, we demonstrate that the LPCD pattern is conserved in Euphorbiaceae s.s. and that this conservatism is primarily due to selective pressures. A phylogenetic association was found between the putative removal of selective pressures on pollen morphology after the origin of inaperturate pollen, and the appearance of variation in microsporogenesis and in the resulting LPCD pattern, suggesting that stasis was due to these selective pressures. However, even in a neutral context, variation in microsporogenesis was biased. This should therefore favour the appearance of some developmental and morphological phenotypes rather than others.

The legume subfamily Caesalpinioideae is a paraphyletic grade, within which are nested the monophyletic subfamilies Mimosoideae and Papilionoideae. Although higher level relationships within Caesalpinioideae are now better understood, few studies have examined generic and species level relationships. A morphological and molecular phylogenetic analysis of selected members of the Brownea clade (Detarieae: Caesalpinioideae) is presented here, focusing on relationships within and among the genera Elizabetha, Heterostemon, Paloveopsis, and Paloue. Morphological characters (125) and DNA sequence data from the plastid trnL intron and nuclear ITS were used to reconstruct phlogenetic relationships. These results indicate that (1) Heterostemon is monophyletic; (2) the majority of Elizabetha species form a poorly supported, monophyletic group sister to Paloue; and (3) Paloveopsis is nested within Paloue. Intergeneric hybridization between species of Paloue and Elizabetha has been identified and traditional generic, species, and intraspecies boundaries are assessed and reevaluated using the results of the combined phylogenetic analysis.

Two unispecific genera, Eccremis and Pasithea, are the only representatives of Hemerocallidaceae (Asparagales) native to the Western Hemisphere. The affinities of the former genus have been recently questioned. Phylogenetic analyses of DNA sequence data (plastid atpB, ndhF, rbcL, trnL-F) spanning the Asparagales recover a monophyletic Hemerocallidaceae and strongly support the inclusion of Eccremis within that family. Eccremis is sister to Dianella and not closely associated with Pasithea, which indicates two introductions of Hemerocallidaceae to the New World. A taxonomic synopsis of Eccremis and Pasithea is presented and morphological character evolution and biogeography are discussed.

Enriquebeltrania, a genus of trees and shrubs endemic to Mexico, traditionally has been included in Euphorbiaceae (tribe Adelieae, subfamily Acalyphoideae). It was previously thought to be monotypic with a disjunct distribution on the Yucatan Peninsula and along the Pacific coast in Jalisco and Sinaloa. There is some question about whether Enriquebeltrania should be treated as distinct from Adelia. Morphological and molecular characters (trnL-F spacer and rbcL DNA sequences) were utilized in phylogenetic analyses to determine if Enriquebeltrania belongs to the tribe Adelieae, to test whether it should be recognized as a genus separate from Adelia, and to determine the number of species that should be recognized. Results indicate that Enriquebeltrania does not belong to tribe Adelieae, that it should be recognized as a genus separate from Adelia, and that it consists of two species. The second species, Enriquebeltrania disjuncta, is named and described here.

Phylogenetic relationships within tribe Phyllantheae, the largest tribe of the family Phyllanthaceae, were examined with special emphasis on the large genus Phyllanthus. Nuclear ribosomal ITS and plastid matK DNA sequence data for 95 species of tribe Phyllantheae, including representatives of all subgenera of Phyllanthus (except Cyclanthera) and several hitherto unplaced infrageneric groups, were analyzed. Results for ITS and matK are generally concordant, although some species are placed differently in the plastid and ITS trees, indicating that hybridization/paralogy is involved. Results confirm paraphyly of Phyllanthus in its traditional circumscription with embedded Breynia, Glochidion, Reverchonia, and Sauropus. We favor the inclusion of the embedded taxa in Phyllanthus over further generic segregation. Monophyletic Phyllanthus comprises an estimated 1269 species, making it one of the "giant" genera. Phyllanthus maderaspatensis is sister to all other species of Phyllanthus, and the genus appears to be of paleotropical origin. Subgenera Isocladus, Kirganelia, and Phyllanthus are polyphyletic, whereas other subgenera appear to be monophyletic. Monotypic Reverchonia is sister to P. abnormis, arborescent section Emblica to herbaceous Urinaria, free-floating aquatic P. fluitans to the weed P. caroliniensis, and the phyllocladous section Choretropsis to the delicate leafy P. claussenii. The unique branching architecture known as "phyllanthoid branching" found in most Phyllanthus taxa has been lost (and/or has been derived) repeatedly. Taxonomic divisions within Phyllantheae based on similar pollen morphology are confirmed, and related taxa share similar distributions. We recommend recognition of six clades at generic level: Flueggea s.l. (including Richeriella), Lingelsheimia, Margaritaria, Phyllanthus s.l. (including Breynia, Glochidion, Reverchonia, and Sauropus), P. diandrus, and Savia section Heterosavia.

The rattlesnake fern ( Botrychium virginianum (L.) Sw.) is obligately mycotrophic and widely distributed across the northern hemisphere. Three mitochondrial gene regions place this species with other ferns in Ophioglossaceae, while two regions place it as a member of the largely parasitic angiosperm order Santalales (sandalwoods and mistletoes). These discordant phylogenetic placements suggest that part of the genome in B. virginianum was acquired by horizontal gene transfer (HGT), perhaps from root-parasitic Loranthaceae. These transgenes are restricted to B. virginianum and occur across the range of the species. Molecular and life-history traits indicate that the transfer preceded the global expansion of B. virginianum , and that the latter may have happened very rapidly. This is the first report of HGT from an angiosperm to a fern, through either direct parasitism or the mediation of interconnecting fungal symbionts.

Parsimony and Bayesian analyses of plastid rbcL and trnL-F DNA sequence data of the pantropical family Euphorbiaceae sensu stricto (s.s.) are presented. Sampling includes representatives of all three subfamilies (Acalyphoideae, Crotonoideae, and Euphorbioideae), 35 of 37 tribes and 179 of the 247 genera of uniovulate Euphorbiaceae sensu lato (s.l.). Euphorbiaceae s.s. were recovered as a monophyletic group with no new adjustments in circumscription. Two clades containing taxa previously placed in Acalyphoideae are found to be successive sisters to all other Euphorbiaceae s.s. and are proposed here at subfamilial rank as Peroideae and Cheilosoideae. The remainder of the family fall into seven major lineages including Erismantheae and Acalyphoideae s.s. (parts of Acalyphoideae), Adenoclineae s.l., Gelonieae, articulated crotonoids and inaperturate crotonoids (parts of Crotonoideae), and Euphorbioideae. Potential synapomorphies and biogeographical trends are suggested for these clades. Acalyphoideae s.s., inaperturate crotonoids, and Euphorbioideae tribe Hippomaneae each have two major subclades that represent novel groupings without apparent morphological synapomorphies. Two subfamilies, 14 tribes, and 10 genera were found to be para- or polyphyletic. Noteworthy among these, Omphaleae are embedded in Adenoclineae, Hureae + Pachystromateae in Hippomaneae, Ditta in Tetrorchidium, and Sapium s.s. in Stillingia.

Fossil data have been interpreted as indicating that Late Cretaceous tropical forests were open and dry adapted and that mod- ern closed-canopy rain forest did not originate until after the Cretaceous-Tertiary (K/T) boundary. However, some mid-Cretaceous leaf floras have been interpreted as rain forest. Molecular divergence- time estimates within the clade Malpighiales, which constitute a large percentage of species in the shaded, shrub, and small tree layer in tropical rain forests worldwide, provide new tests of these hypotheses. We estimate that all 28 major lineages (i.e., traditionally recognized families) within this clade originated in tropical rain forest well before the Tertiary, mostly during the Albian and Cenomanian (112-94 Ma). Their rapid rise in the mid-Cretaceous may have resulted from the origin of adaptations to survive and reproduce under a closed forest canopy. This pattern may also be paralleled by other similarly diverse lineages and supports fossil indications that closed-canopy tropical rain forests existed well before the K/T boundary. This case illustrates that dated phylogenies can provide an important new source of ev- idence bearing on the timing of major environmental changes, which may be especially useful when fossil evidence is limited or controversial.

The classification of the Catesbaeeae and Chiococceae tribes, along with that of the entire Rubiaceae, has long been debated. The Catesbaeeae-Chiococceae complex (CCC) includes approximately 28 genera and 190 species primarily concentrated in the Greater Antilles (nearly 70% of the species), Central and South America, and in the western Pacific (three genera). Previous molecular studies, with broad sampling of the Rubiaceae, have shown the CCC to be a monophyletic group. The present study is a more detailed examination of the generic relationships within the CCC using two data sets, the nuclear ribosomal ITS regions and the trnL-F chloroplast intron and spacer. Maximum parsimony analyses lend further support to the previous hypotheses that the CCC is monophyletic and sister to Strumpfia maritima. However, within the complex several genera do not form monophyletic groups. Previous studies of the Rubiaceae suggest that the ancestral fruit type in the CCC is a multiseeded capsule. Indehiscent, fleshy fruits appear to have evolved three to four times within this lineage. Changes in floral morphologies within the complex tend to correspond to cladogenesis among and within genera. Finally, molecular analyses suggest one or possibly two long-distance dispersals from the Americas to the western Pacific.

Analysis of plastid rbcL DNA sequence data of the pantropical family Phyllanthaceae (Malpighiales) and related biovulate lineages of Euphorbiaceae sensu lato is presented. Sampling for this study includes representatives of all 10 tribes and 51 of the 60 genera attributed to Euphorbiaceae-Phyllanthoideae. Centroplacus and Putranjivaceae (Phyllanthoideae-Drypeteae) containing a paraphyletic Drypetes are excluded from Phyllanthaceae. Croizatia, previously thought to be a "basal" member of Euphorbiaceae-Oldfieldioideae (Picrodendraceae), falls within Phyllanthaceae. Phyllanthaceae with the mentioned adjustments form a monophyletic group consisting of two sister clades that mostly correspond to the distribution of tanniniferous leaf epidermal cells and inflorescence structure. With the exception of bigeneric Hymenocardieae and monotypic Bischofieae, none of the current Phyllanthoideae (Phyllanthaceae) tribal circumscriptions are supported by rbcL. Antidesma, Bischofia, Hymenocardia, Martretia, and Uapaca, all of which have previously been placed in monogeneric families, are confirmed as members of Phyllanthaceae. Savia is polyphyletic, and Cleistanthus appears paraphyletic. Paraphyly of Phyllanthus is also indicated, but this pattern lacks bootstrap support. Morphological characters are discussed and mapped for inflorescence structure, tanniniferous epidermal cells, breeding system, and fruit and embryo type. A table summarizes the main characters of six euphorbiaceous lineages.

The mid-Cenozoic decline of atmospheric CO2 levels that promoted global climate change was critical to shaping contemporary arid ecosystems. Within angiosperms, two CO2 -concentrating mechanisms (CCMs)-crassulacean acid metabolism (CAM) and C4 -evolved from the C3 photosynthetic pathway, enabling more efficient whole-plant function in such environments. Many angiosperm clades with CCMs are thought to have diversified rapidly due to Miocene aridification, but links between this climate change, CCM evolution, and increased net diversification rates (r) remain to be further understood. Euphorbia (∼2000 species) includes a diversity of CAM-using stem succulents, plus a single species-rich C4 subclade. We used ancestral state reconstructions with a dated molecular phylogeny to reveal that CCMs independently evolved 17-22 times in Euphorbia, principally from the Miocene onwards. Analyses assessing among-lineage variation in r identified eight Euphorbia subclades with significantly increased r, six of which have a close temporal relationship with a lineage-corresponding CCM origin. Our trait-dependent diversification analysis indicated that r of Euphorbia CCM lineages is approximately threefold greater than C3 lineages. Overall, these results suggest that CCM evolution in Euphorbia was likely an adaptive strategy that enabled the occupation of increased arid niche space accompanying Miocene expansion of arid ecosystems. These opportunities evidently facilitated recent, replicated bursts of diversification in Euphorbia.

The angiosperm order Malpighiales includes ∼16,000 species and constitutes up to 40% of the understory tree diversity in tropical rain forests. Despite remarkable progress in angiosperm systematics during the last 20 y, relationships within Malpighiales remain poorly resolved, possibly owing to its rapid rise during the mid-Cretaceous. Using phylogenomic approaches, including analyses of 82 plastid genes from 58 species, we identified 12 additional clades in Malpighiales and substantially increased resolution along the backbone. This greatly improved phylogeny revealed a dynamic history of shifts in net diversification rates across Malpighiales, with bursts of diversification noted in the Barbados cherries (Malpighiaceae), cocas (Erythroxylaceae), and passion flowers (Passifloraceae). We found that commonly used a priori approaches for partitioning concatenated data in maximum likelihood analyses, by gene or by codon position, performed poorly relative to the use of partitions identified a posteriori using a Bayesian mixture model. We also found better branch support in trees inferred from a taxon-rich, data-sparse matrix, which deeply sampled only the phylogenetically critical placeholders, than in trees inferred from a taxon-sparse matrix with little missing data. Although this matrix has more missing data, our a posteriori partitioning strategy reduced the possibility of producing multiple distinct but equally optimal topologies and increased phylogenetic decisiveness, compared with the strategy of partitioning by gene. These approaches are likely to help improve phylogenetic resolution in other poorly resolved major clades of angiosperms and to be more broadly useful in studies across the Tree of Life.

Euphorbia is among the largest genera of angiosperms, with about 2000 species that are renowned for their remarkably diverse growth forms. To clarify phylogenetic relationships in the genus, we used maximum likelihood, Bayesian, and parsimony analyses of DNA sequence data from 10 markers representing all three plant genomes, averaging more than 16. kbp for each accession. Taxon sampling included 176 representatives from Euphorbioideae (including 161 of Euphorbia). Analyses of these data robustly resolve a backbone topology of four major, subgeneric clades- Esula, Rhizanthium, Euphorbia, and Chamaesyce-that are successively sister lineages. Ancestral state reconstructions of six reproductive and growth form characters indicate that the earliest Euphorbia species were likely woody, non-succulent plants with helically arranged leaves and 5-glanded cyathia in terminal inflorescences. The highly modified growth forms and reproductive features in Euphorbia have independent origins within the subgeneric clades. Examples of extreme parallelism in trait evolution include at least 14 origins of xeromorphic growth forms and at least 13 origins of seed caruncles. The evolution of growth form and inflorescence position are significantly correlated, and a pathway of evolutionary transitions is supported that has implications for the evolution of Euphorbia xerophytes of large stature. Such xerophytes total more than 400 species and are dominants of vegetation types throughout much of arid Africa and Madagascar. © 2012., Support for this study came from a National Science Foundation PBI Grant (DEB 0616533) and the Smithsonian Institution.

The snap-trap leaves of the aquatic waterwheel plant (Aldrovanda) resemble those of Venus' flytrap (Dionaea), its distribution and habit are reminiscent of bladderworts (Utricularia), but it shares many reproductive characters with sundews (Drosera). Moreover, Aldrovanda has never been included in molecular phylogenetic studies, so it has been unclear whether snap-traps evolved only once or more than once among angiosperms. Using sequences from nuclear 18S and plastid rbcL, atpB, and matK genes, we show that Aldrovanda is sister to Dionaea, and this pair is sister to Drosera. Our results indicate that snap-traps are derived from flypaper-traps and have a common ancestry among flowering plants, despite the fact that this mechanism is used by both a terrestrial species and an aquatic one. Genetic and fossil evidence for the close relationship between these unique and threatened organisms indicate that carnivory evolved from a common ancestor within this caryophyllid clade at least 65 million years ago.

Parsimony, likelihood, and Bayesian analyses of nuclear ITS and plastid trnL-F DNA sequence data are presented for the giant genus Croton (Euphorbiaceae s.s.) and related taxa. Sampling comprises 88 taxa, including 78 of the estimated 1223 species and 29 of the 40 sections previously recognized of Croton. It also includes the satellite genus Moacroton and genera formerly placed in tribe Crotoneae. Croton and all sampled segregate genera form a monophyletic group sister to Brasiliocroton, with the exception of Croton sect. Astraea, which is reinstated to the genus Astraea. A small clade including Moacroton, Croton alabamensis, and C. olivaceus is sister to all other Croton species sampled. The remaining Croton species fall into three major clades. One of these is entirely New World, corresponding to sections Cyclostigma, Cascarilla, and Velamea sensu Webster. The second is entirely Old World and is sister to a third, also entirely New World clade, which is composed of at least 13 of Webster’s sections of Croton. This study establishes a phylogenetic framework for future studies in the hyper-diverse genus Croton, indicates a New World origin for the genus, and will soon be used to evaluate wood anatomical, cytological, and morphological data in the Crotoneae tribe.

The eudicot order Malpighiales contains ∼16000 species and is the most poorly resolved large rosid clade. To clarify phylogenetic relationships in the order, we used maximum likelihood, Bayesian, and parsimony analyses of DNA sequence data from 13 gene regions, totaling 15604 bp, and representing all three genomic compartments (i.e., plastid: atpB, matK, ndhF, and rbcL; mitochondrial: ccmB, cob, matR, nad1B-C, nad6, and rps3; and nuclear: 18S rDNA, PHYC, and newly developed low-copy EMB2765). Our sampling of 190 taxa includes representatives from all families of Malpighiales. These data provide greatly increased support for the recent additions of Aneulophus, Bhesa, Centroplacus, Ploiarium, and Rafflesiaceae to Malpighiales; sister relations of Phyllanthaceae + Picrodendraceae, monophyly of Hypericaceae, and polyphyly of Clusiaceae. Oxalidales + Huaceae, followed by Celastrales are successive sisters to Malpighiales. Parasitic Rafflesiaceae, which produce the world's largest flowers, are confirmed as embedded within a paraphyletic Euphorbiaceae. Novel findings show a well-supported placement of Ctenolophonaceae with Erythroxylaceae + Rhizophoraceae, sister-group relationships of Bhesa + Centroplacus, and the exclusion of Medusandra from Malpighiales. New taxonomic circumscriptions include the addition of Bhesa to Centroplacaceae, Medusandra to Peridiscaceae (Saxifragales), Calophyllaceae applied to Clusiaceae subfamily Kielmeyeroideae, Peraceae applied to Euphorbiaceae subfamily Peroideae, and Huaceae included in Oxalidales.

PREMISE OF THE STUDY Recent analyses employing up to five genes have provided numerous insights into angiosperm phylogeny, but many relationships have remained unresolved or poorly supported. In the hope of improving our understanding of angiosperm phylogeny, we expanded sampling of taxa and genes beyond previous analyses. METHODS We conducted two primary analyses based on 640 species representing 330 families. The first included 25260 aligned base pairs (bp) from 17 genes (representing all three plant genomes, i.e., nucleus, plastid, and mitochondrion). The second included 19846 aligned bp from 13 genes (representing only the nucleus and plastid). KEY RESULTS Many important questions of deep-level relationships in the nonmonocot angiosperms have now been resolved with strong support. Amborellaceae, Nymphaeales, and Austrobaileyales are successive sisters to the remaining angiosperms (Mesangiospermae), which are resolved into Chloranthales + Magnoliidae as sister to Monocotyledoneae + [Ceratophyllaceae + Eudicotyledoneae]. Eudicotyledoneae contains a basal grade subtending Gunneridae. Within Gunneridae, Gunnerales are sister to the remainder (Pentapetalae), which comprises (1) Superrosidae, consisting of Rosidae (including Vitaceae) and Saxifragales; and (2) Superasteridae, comprising Berberidopsidales, Santalales, Caryophyllales, Asteridae, and, based on this study, Dilleniaceae (although other recent analyses disagree with this placement). Within the major subclades of Pentapetalae, most deep-level relationships are resolved with strong support. CONCLUSIONS Our analyses confirm that with large amounts of sequence data, most deep-level relationships within the angiosperms can be resolved. We anticipate that this well-resolved angiosperm tree will be of broad utility for many areas of biology, including physiology, ecology, paleobiology, and genomics.