Polemoniaceae Research Articles

Complex interactions underlie the correlated evolution of floral traits and their association with pollinators in a clade with diverse pollination systems.

Natural selection by pollinators is an important factor in the morphological diversity and adaptive radiation of flowering plants. Selection by similar pollinators in unrelated plants leads to convergence in floral morphology, or "floral syndromes." Previous investigations into floral syndromes have mostly studied relatively small and/or simple systems, emphasizing vertebrate pollination. Despite the importance of multiple floral traits in plant-pollinator interactions, these studies have examined few quantitative traits, so their co-variation and phenotypic integration have been underexplored. To gain better insights into pollinator-trait dynamics, we investigate the model system of the phlox family (Polemoniaceae), a clade of ∼400 species pollinated by a diversity of vectors. Using a comprehensive phylogeny and large dataset of traits and observations of pollinators, we reconstruct ancestral pollination system, accounting for the temporal history of pollinators. We conduct phylogenetically controlled analyses of trait co-variation and association with pollinators, integrating many analyses over phylogenetic uncertainty. Pollinator shifts are more heterogeneous than previously hypothesized. The evolution of floral traits is partially constrained by phylogenetic history and trait co-variation, but traits are convergent and differences are associated with different pollinators. Trait shifts are usually gradual, rather than rapid, suggesting complex genetic and ecological interactions of flowers at macroevolutionary scales.

<i>Puccinia fumosa</i> (Pucciniales) y su hospedante <i>Loeselia glandulosa</i> (Polemoniaceae), nuevos registros para Colombia y Suramérica

Se registra por primera vez para Colombia la roya Puccinia fumosa parasitando a Loeselia glandulosa perteneciente a la familia Polemoniaceae. Es el primer registro de esta familia botánica y del género Loeselia siendo parasitados por Pucciniales en Suramérica. Se describen por primera vez las mesosporas de Puccinia fumosa.

Evolution of floral traits and impact of reproductive mode on diversification in the phlox family (Polemoniaceae)

Pollinator-mediated selection is a major driver of evolution in flowering plants, contributing to the vast diversity of floral features. Despite long-standing interest in floral variation and the evolution of pollination syndromes in Polemoniaceae, the evolution of floral traits and known pollinators has not been investigated in an explicit phylogenetic context. Here we explore macroevolutionary patterns of both pollinator specificity and three floral traits long considered important determinants of pollinator attraction across the most comprehensive species-level phylogenetic tree yet produced for the family. The presence of floral chlorophyll is reconstructed as the ancestral character state of the family, even though the presence of floral anthocyanins is the most prevalent floral pigment in extant taxa. Mean corolla length and width of the opening of the floral tube are correlated, and both appear to vary with pollinator type. The evolution of pollination systems appears labile, with multiple gains and losses of selfing and conflicting implications for patterns of diversification. Explicit testing of diversification models rejects the hypothesis that selfing is an evolutionary dead-end. This study begins to disentangle the individual components that comprise pollination syndromes and lays the foundation for future work on the genetic mechanisms that control each trait.

Towards A Complete Species Level Phylogeny of Leptosiphon (Polemoniaceae)

Abstract Leptosiphon Benth. (Polemoniaceae) has 40 taxa, distributed mainly in western North America, with one species in Chile. Phylogenetic studies of Polemoniaceae have consistently supported recognition of Leptosiphon as a separate genus from Linanthus, and resolved an array of taxonomic problems; however, several issues remained unclear for Leptosiphon. In those studies, sampling of Leptosiphon was incomplete, and certain species having strikingly similar floral morphologies appeared non-monophyletic, while other species groups with divergent floral morphologies formed a clade. Our analyses use greater sampling across taxa in Leptosiphon and, in addition to nrITS, the non-coding chloroplast region trnS–trnG to help resolve the genus phylogeny. These results yielded a more complete phylogeny from which to base further research on floral evolution in this group. However, much work remains to be done in resolving species level relationships. Analyses of morphological character evolution suggest that sev...

Competition for Pollination and the Evolution of Flowering Time.

Competition for Pollination and the Evolution of Flowering Time.

Association of Pigeon pea witches' broom phytoplasma (16Sr IX) infecting Phlox drummondii in India

Phlox is a genus of 67 species of perennial and annual plants in the family Polemoniaceae. Phlox are popular ornamental plants, and traditionally used to treat aches, colds, anaemia, diarrhoea and eczema. During a survey at the Indian Agricultural Research Institute Campus, New Delhi, from 2010 to 2011, symptoms including extensive yellowing and stunting, proliferation of shoots, little leaves and reduced size of flowers were observed in approximately 17% of plants of Phlox drummondii (Fig. 1). To investigate the possibility of a phytoplasma association with the symptoms, total DNA was isolated from leaf midribs and stems of three different infected plants. Healthy-looking plants without symptoms were also collected to use as a negative control using the phytoplasma enrichment procedure (Ahrens & Seemüller, 1). The DNA isolated from periwinkle infected with toria phyllody phytoplasma (group 16SrIX, pigeon pea witches' broom phytoplasma; Azadvar et al., 2) and maintained in greenhouse was used as positive control. Total DNA was used as a template for nested PCR using universal primers that target the phytoplasma 16S rRNA gene: P1/P7 (Deng & Hiruki, 3; Schneider et al., 5) and R16F2n/R16R2 (Gundersen & Lee, 4). Expected size amplicons of ~1.8 kb and ~1.2 kb, respectively, were amplified from all the three symptomatic P. drummondii plants (Fig. 2). However, no PCR products were obtained from the healthy-looking plant samples. The amplicon of ~1.2 kb was sequenced from a representative sample, and the consensus sequence obtained was submitted to GenBank (Accession No. KC178678). BLAST analysis of the partial 16S rDNA sequence of the P. drummondii phytoplasma revealed the highest sequence identity (99%) with that of phytoplasma members of group 16SrIX, ‘Candidatus Phytoplasma phoenicium’, the former Pigeon pea witches' broom group. Phylogenetic analysis (MEGA version 4.0, USA) confirmed the closest phylogenetic relationship of the P. drummondii phytoplasma 16S rDNA sequence with those of the 16SrIX phytoplasma group (Fig. 3). Hence, the P. drummondii phytoplasma was identified as a 16SrIX isolate. Phytoplasmas of group 16SrI ‘Candidatus Phytoplasma asteris’ subgroup M (16SrI-M) have been previously associated with P. drummondii in Lithuania (Samuitienė et al., 6). However, this is the first world report of the association of the 16SrIX group with a phytoplasma disease on P. drummondii. The first author is thankful to Department of Science and Technology, New Delhi, India for providing financial assistance under WOS ‘A’ scheme. The authors wish to express sincere thanks to Head, Division of Plant Pathology and Director, Indian Agricultural Research Institute for providing laboratory facilities.

Allotetraploids in Patagonia with Affinities to Western North American Diploids: Did Dispersal or Genome Doubling Occur First?

Amphitropical disjunct distributions between western North America and western South America have intrigued botanists for over a century. Here, specific examples of migration and speciation are investigated using herbaceous species from the phlox family (Polemoniaceae) as a model for considering the timing of dispersal relative to speciation. Comparative DNA sequencing reveals that, in Collomia and Navarretia, the South American species are allopolyploids, suggesting either two dispersals prior to the allopolyploidization event for each species with subsequent extirpation of the diploid progenitors from South America, or allopolyploid formation prior to dispersal with extirpation of these polyploids from North America. Divergence time estimates support a Pliestocene dispersal hypothesis and sequence data indicate that, at least in Collomia, hybridization of the diploid progenitors occurred in South America.

Nuclear and cpDNA sequences combined provide strong inference of higher phylogenetic relationships in the phlox family (Polemoniaceae)

Members of the phlox family (Polemoniaceae) serve as useful models for studying various evolutionary and biological processes. Despite its biological importance, no family-wide phylogenetic estimate based on multiple DNA regions with complete generic sampling is available. Here, we analyze one nuclear and five chloroplast DNA sequence regions (nuclear ITS, chloroplast matK, trnL intron plus trnL–trnF intergeneric spacer, and the trnS–trnG, trnD–trnT, and psbM–trnD intergenic spacers) using parsimony and Bayesian methods, as well as assessments of congruence and long branch attraction, to explore phylogenetic relationships among 84 ingroup species representing all currently recognized Polemoniaceae genera. Relationships inferred from the ITS and concatenated chloroplast regions are similar overall. A combined analysis provides strong support for the monophyly of Polemoniaceae and subfamilies Acanthogilioideae, Cobaeoideae, and Polemonioideae. Relationships among subfamilies, and thus for the precise root of Polemoniaceae, remain poorly supported. Within the largest subfamily, Polemonioideae, four clades corresponding to tribes Polemonieae, Phlocideae, Gilieae, and Loeselieae receive strong support. The monogeneric Polemonieae appears sister to Phlocideae. Relationships within Polemonieae, Phlocideae, and Gilieae are mostly consistent between analyses and data permutations. Many relationships within Loeselieae remain uncertain. Overall, inferred phylogenetic relationships support a higher-level classification for Polemoniaceae proposed in 2000.

First Report of Tobacco rattle virus and Cucumber mosaic virus in Phlox paniculata in France.

Phlox paniculata L., a perennial plant from the family Polemoniaceae, is cultivated as an ornamental in gardens and for cut-flower production. In spring 2003, two types of symptoms were observed in P. paniculata plants grown for cut flowers on a farm in the Var department, France. Some plants showed a mild leaf mosaic while others showed leaf browning and delayed growth. In plants showing mild mosaic, Cucumber mosaic virus (CMV) was detected on the basis of the symptoms exhibited by a range of inoculated plants, the observation of isometric particles (approximately 30 nm) with the electron microscope in crude sap preparations from the infected plants, and the positive reaction in double-antibody sandwich (DAS)-ELISA to polyclonal antibodies raised against CMV (1). In double-immunodiffusion analysis, the five tested isolates were shown to belong to group II of CMV strains. To determine if CMV was responsible for the symptoms observed, one isolate was multiplied in Nicotiana tabacum cv. Xanthi-nc plants after isolation from local lesions on Vigna unguiculata and mechanically inoculated to 12 1-year-old P. paniculata plants. At 3 months post inoculation (mpi), all plants showed mild mosaic and CMV was detected by DAS-ELISA. In sap preparations from P. paniculata plants showing leaf browning symptoms, rod-shaped particles with two distinct sizes of 190 to 210 and 70 to 90 nm long, typical of those associated with tobraviruses, were revealed using electron microscopy. Local lesions typical of Tobacco rattle virus (TRV) were observed after inoculation of N. tabacum cv. Xanthi-nc, Chenopodium amaranticolor, and C. quinoa. Total nucleic acid preparations were prepared from symptomatic plants, and amplicons of the expected size (463 bp) were generated by reverse-transcription (RT)-PCR using primers specific to TRV RNA 1 (4). The nucleotide sequence of one amplicon was 93.6% identical to the sequence of a reference TRV isolate (GenBank Accession No. AJ586803). Twelve 1-year-old P. paniculata plants were mechanically inoculated with an extract of infected tissues from one symptomatic P. paniculata plant. TRV was detected 2 to 6 mpi in apical leaves of all inoculated plants by RT-PCR, although the plants did not express symptoms. Since no other pathogens were detected in the source plants, it is plausible that the lack of symptoms in back-inoculated plants is either due to a long incubation period or an interaction with particular environmental factors such as cold conditions. The survey of approximately 200 plants revealed that approximately 7, 10, and 1% were infected by TRV, CMV, or by both viruses, respectively. CMV and TRV were previously detected in P. paniculata in Latvian SSR and in Lithuania (2,3). These results show that sanitary selection of P. paniculata prior to vegetative propagation should include a screening for TRV and CMV infections.

Characterization of microsatellite loci in Ipomopsis (Polemoniaceae) wildflowers

AbstractMembers of the Ipomopsis aggregata species complex in the phlox family (Polemoniaceae) often hybridize when they occur in sympatry, and thus have been extensively studied to examine processes involved in plant speciation. I developed 12 microsatellite loci in I. aggregata that are also polymorphic in closely related Ipomopsis tenuituba, producing an average of eight alleles per locus in test populations of 12 individuals per species. Several of these markers also successfully amplified in other Ipomopsis species and more distant members of the Polemoniaceae, suggesting they should prove useful for a broad range of evolutionary studies in this widely distributed system.

Early floral development and androecium organization in Fouquieriaceae (Ericales)

Early floral development with focus on the androecium was studied with the help of scanning electron microscopy and serial microtome sectioning in Fouquieria columnaris and F. splendens. Perianth organs appear in a spiral pattern on the floral apex. The spiral may be a clockwise or anti-clockwise. The androecium is best interpreted as two-whorled with all the stamens arranged in a single series. In F. splendens, two or more of the five epipetalous stamen positions are doubled, i.e. they are occupied by stamen pairs. Unusual features in the floral development of Fouquieriaceae include (1) a strong spiral component even in whorled organ categories and (2) a pronouncedly asymmetric floral apex during an early phase of floral development. From a phylogenetic point of view, it seems plausible that the common ancestor of Fouquieriaceae and its sister family Polemoniaceae was characterized by two alternating, pentamerous stamen-whorls.

(1641) Proposal to conserve the name Gilia splendens (Polemoniaceae) with a conserved type

TAXONVolume 53, Issue 3 p. 842-843 Proposals to Conserve or RejectsFree Access (1641) Proposal to conserve the name Gilia splendens (Polemoniaceae) with a conserved type Verne Grant, Verne Grant Plant Resources Center, The University of Texas at Austin, 1 University Station F0404, Austin, Texas 78712 U.S.A.Search for more papers by this authorTom Wendt, Tom Wendt twendt@mail.utexas.edu Plant Resources Center, The University of Texas at Austin, 1 University Station F0404, Austin, Texas 78712 U.S.A.Search for more papers by this author Verne Grant, Verne Grant Plant Resources Center, The University of Texas at Austin, 1 University Station F0404, Austin, Texas 78712 U.S.A.Search for more papers by this authorTom Wendt, Tom Wendt twendt@mail.utexas.edu Plant Resources Center, The University of Texas at Austin, 1 University Station F0404, Austin, Texas 78712 U.S.A.Search for more papers by this author First published: 01 August 2004 https://doi.org/10.2307/4135466AboutPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinked InRedditWechat No abstract is available for this article. Volume53, Issue3August 2004Pages 842-843 RelatedInformation

The Importance of Species of Dasytinae (Coleoptera: Melyridae) as Pollinators in Western North America

Species of 11 families of Coleoptera are known to pollinate flowering plants. In western North America, the 300 species of the beetle subfamily Dasytinae (Coleoptera: Melyridae) are commonly found on flowers, where adults feed on both nectar and pollen. The dense setae and pubescence of adult dasytines often trap pollen grains, which are transported to other flowers during the course of normal adult feeding. Adults of many dasytine species are abundant, and may form large feeding aggregations. A preliminary list is presented of plant species on whose flowers or pollen cones adult dasytines have been collected, based on literature records and museum specimen labels. These plant species represent 68 genera in 26 families, and include species grown as food crops and species that are important browse or forage plants for livestock and wildlife. At least two plant species, both in the family Polemoniaceae, rely predominantly on species of Dasytinae for pollination.

A Guide to Understanding Recent Classifications of the Family Polemoniaceae

Abstract: Two very different systems of classification of the family Polemoniaceae—one a taxonomic system and the other a molecular cladistic system—have appeared recently. They are compared and reviewed. It is emphasized that taxonomy and cladistics use different basic systematic units (taxa or clades) and have different concepts of monophyly. Furthermore, taxonomy and molecular cladistics differ in the characters used in classification. Conflicts between molecular and reliable morphological characters occur in several parts of the family. The differences in methodology account for most of the incongruences between the two classifications of the family. In general, the methodological differences can be expected to yield incongruences between taxonomic and cladistic systems in any heterogeneous group of medium or large size. New suggestions are made about the early phylogenetic divergences in the Polemoniaceae. The tribes are reviewed from a taxonomic standpoint and a few changes are made.

Synthetic autotetraploid plants of Phlox drummondii exhibit significantly higher photosynthetic rate at C11 generation than teh diploid progenitor

Multiplication of genome has significant roles in the adaptation and evolution of plants and is accompanied by various physiological changes. Synthetic autotetraploids and its diploid progenitor in Phlox drummondii (Polemoniaceae) have given us a unique opportunity to effectively study the effect of gene dosage on biochemical and physiological activities of the plants. The entire genome was doubled by colchicine and now the synthetic autotetraploids are running into 11th generation (C11). They are not only morphologically healthier than their diploid progenitor but are also photosynthetically more efficient on leaf area basis. Moreover, using different generations of tetraploid plants, we are able to study whether the better growth and physiological performance is the result of only genome duplication or the product of genome dosage combined with gradual environmental adaptation over the years. The increase in leaf dry weight is 55% and 80% in C0 and C11 generations, respectively. The increase in chlorophyll content is about 18% in C0 generation while it is 110% in C11 generation over the diploid progenitors. The light saturation point of tetraploids of both the generations is much higher. The stomatal conductance is only little higher at C0 generation while it has increased 2.75 times in C11 generation. Net photosynthetic assimilation is increased by about 43% in C0 generation (12 µmol CO2 m-2s-1) and more than doubled in C11 generation (17 µmol CO2 m-2s-1). We intend to present the anatomical details, quantification of enzymes, DNA and chlorophyll per cell and, assess the response of photosynthesis to CO2 concentration on per cell and per unit DNA basis and discuss its theoretical and practical implications.

A unique and complete polemoniaceous plant from the middle Eocene of Utah, USA

An extinct genus of the Polemoniaceae is described from one complete fossil plant preserved in shale of the Eocene Green River Formation, Utah. Combined vegetative and reproductive characters including the taproot, basal and cauline pinnatifid leaves, primary peduncular leaves, secondary peduncular bracts, pedicel bracts, fruits in groups of three, and persistent calyx, support placement of this plant close to the extant genus Gilia. Gilisenium hueberi gen. et sp. nov. represents a rare record of an herbaceous plant, and the oldest megafossil for the family Polemoniaceae.

Floral Biology: Studies on Floral Evolution in Animal-Pollinated Plants.

Part One : Historical perspective: Discovery of the secret of nature in the structure and fertilization of flowers-- C. K. Sprengel (translated by P. Haase) Christian Konrad Sprengel's theory of the flower: the cradle of floral ecology-- S. Vogel Part Two: Conceptual issues: Floral traits and plant adaptation to insect pollinators: a devil's advocate approach-- C.M. Herrera How do flowers diverge-- P. Wilson and J.D. Thomson Floral longevity: fitness consequences and resource costs-- T.L. Ashman and D.J. Schoen Pollen dispersal and mating patterns in animal-pollinated plants-- L.D. Harder and S.C.H. Barrett The ecology of geitonogamous pollination-- A.A. Snow, T.P. Spira, R. Simpson, and R.A. Klips Flower size dimorphism in plants with unisexual flowers-- L.F. Delph Part Three: Model systems: Evolution of floral morphology and function: an integrative approach to adaptation, constraint and compromise in Dalechampia (Euphorbiaceae)-- W.S. Armbruster The evolution of floral form: insights from an Alpine wildflower, Polemonium viscosum (Polemoniaceae)-- C. Galen Deceit pollination in the monoecious, neotropical herb Begonia oaxacana (Begoniaceae)-- D.W. Schemske, J. Agren, and J. Le Corff Reproductive success and gender variation in deceit-pollinated orchids-- A.L. Fritz and L.A. Nilsson Stylar polymorphisms and the evolution of heterostyly in Narcissus (Amaryllidaceae)-- S.C.H. Barrett, D.G. Lloyd, and J. Arroyo Evolution of Campanula flowers in relation to insect pollinators on islands-- K. Inoue, M. Maki, and M. Masuda Index.

Floral Formula Inconstancy Within and Among Plants and Populations of Ipomopsis aggregata (Polemoniaceae)

The numbers of flower parts are considered among the most constant angiosperm characters However, nearly 33% of the plants in 13 populations of Ipomopsis aggregata in Colorado had at least one anomalous flower, and almost 10% of the flowers displayed abnormal merism (abnormal floral formulas) Floral inconstancy varied significantly among populations The number of abnormal flowers per plant increased with the proportion of plants displaying abnormal merism at a given site Abnormal flowers were nonrandomly distributed; more plants had multiple anomalous flowers than expected by chance The gynoecium was the most variable flower part and varied independently of the other parts, which tended to vary in a coordinated fashion These observations are compared with other examples of abnormal merism in the Polemoniaceae

Checklist of dicotyledons naturalised in New Zealand 10. Polemoniales and Boraginaceae

Abstract Twenty-two taxa of Boraginaceae, 1 of Cobaeaceae, 13 of Convolvulaceae, 1 of Hydrophyllaceae, 1 of Menyanthaceae, 5 of Polemoniaceae, and 47 of Solanaceae are recorded as naturalised in New Zealand. Fourteen other records (6 Boraginaceae, 2 Convolvulaceae, 1 Polemoniaceae, and 5 Solanaceae) are unsubstantiated.

Flavonoid diversification in the polemoniaceae

Analysis of species representing most sections of all the genera in the family Polemoniaceae showed a range of variation in flavonoids comparable to variation already documented for gross morphological features, karyotypes and pollen grains. Three main groups of flavonoids predominate: (A) common flavonols (kaempferol, quercetin, myricetin); (B) 6-methoxyflavonols (patuletin, eupalitin, eupatolitin); and (C) C-glycosylflavones (apigenin and luteolin based). Cobaea, Loeselia], Polemonium, Allophyllum, Collomia and Gymnosteris have predominantly Group A flavonoids; Bonplandia, Ipomopsis and Eriastrum have predominantly Group B flavonoids; Phlox, Microsteris and Leptodactylon have predominantly Group C flavonoids; while the remaining genera (Cantua, Huthia, Gilia, Langloisia, Navarretia and Linanthus) either have flavonoids of all three groups, or some species within a genus have flavonoids of one group, while other species have flavonoids of another group. Linanthus, Gilia and Navarretia (3 of the larger genera in the family) show great flavonoid diversity, while Langloisia (4 species) has 2 species with Group A flavonoids and the other two species have Group B pigments. Two rare hydroxycoumarins, one being daphnetin, were detected in five genera but they proved to be only of limited systematic interest.