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Showing papers in "American Journal of Botany in 2009"


Journal ArticleDOI
TL;DR: Comparisons of diversification rates suggest that genome doubling may have led to a dramatic increase in species richness in several angiosperm lineages, including Poaceae, Solanaceae, Fabaceae, and Brassicaceae, but additional genomic studies are needed to pinpoint the exact phylogenetic placement of the ancient polyploidy events within these lineages.
Abstract: Polyploidy has long been recognized as a major force in angiosperm evolution. Recent genomic investigations not only indicate that polyploidy is ubiquitous among angiosperms, but also suggest several ancient genome-doubling events. These include ancient whole genome duplication (WGD) events in basal angiosperm lineages, as well as a proposed paleohexaploid event that may have occurred close to the eudicot divergence. However, there is currently no evidence for WGD in Amborella, the putative sister species to other extant angiosperms. The question is no longer "What proportion of angiosperms are polyploid?", but "How many episodes of polyploidy characterize any given lineage?" New algorithms provide promise that ancestral genomes can be reconstructed for deep divergences (e.g., it may be possible to reconstruct the ancestral eudicot or even the ancestral angiosperm genome). Comparisons of diversification rates suggest that genome doubling may have led to a dramatic increase in species richness in several angiosperm lineages, including Poaceae, Solanaceae, Fabaceae, and Brassicaceae. However, additional genomic studies are needed to pinpoint the exact phylogenetic placement of the ancient polyploidy events within these lineages and to determine when novel genes resulting from polyploidy have enabled adaptive radiations.

1,040 citations


Journal ArticleDOI
TL;DR: Research from the social sciences on how the public makes sense of and participates in societal decisions about science and technology is reviewed, offering a set of detailed recommendations for improved public engagement efforts on the part of scientists and their organizations.
Abstract: In this essay, we review research from the social sciences on how the public makes sense of and participates in societal decisions about science and technology. We specifically highlight the role of the media and public communication in this process, challeng ing the still dominant assumption that science literacy is both the problem and the solution to societal conflicts. After reviewing the cases of evolution, climate change, food biotechnology, and nanotechnology, we offer a set of detailed recommendations for improved public engagement efforts on the part of scientists and their organizations. We emphasize the need for science commu nication initiatives that are guided by careful formative research; that span a diversity of media platforms and audiences; and that facilitate conversations with the public that recognize, respect, and incorporate differences in knowledge, values, perspectives, and goals. Over the past few years, there have been signs of a gradual shift in how the scientific community in the United States views public engagement. One can detect a growing recognition that effective communication requires initiatives that sponsor dialogue, trust, re lationships, and public participation across a diversity of social settings and media platforms. Yet despite notable new directions, many communication efforts continue to be based on ad-hoc, intu ition-driven approaches, paying little attention to several decades of interdisciplinary research on what makes for effective public engagement. Many of these initiatives start with the false premise that deficits in public knowledge are the central culprit driving so cietal conflict over science, when in fact, science literacy has only a limited role in shaping public perceptions and decisions. In this article, we describe what we know from social science research on how members of the public from diverse back grounds are likely to use information and reach decisions about science. We then offer recommendations on how this research should inform effective public engagement and communica tion. To clearly demonstrate these principles, we highlight no table successes and mistakes specific to the cases of climate change, evolution, plant biotechnology, and nanotechnology. Across each of our recommendations, we emphasize the fol lowing basic premise: any science communication efforts need to be based on a systematic empirical understanding of an in tended audience's existing values, knowledge, and attitudes,

769 citations


Journal ArticleDOI
TL;DR: Angiosperm diversity is found to have mixed origins: slightly less than half of the living species belong to lineages with low to moderate diversification rates, which appeared between 130 and 102 Mya (Barremian-uppermost Albian; Lower Cretaceous).
Abstract: The extraordinary diversity of angiosperms is the ultimate outcome of the interplay of speciation and extinction, which determine the net diversification of different lineages. We document the temporal trends of angiosperm diversification rates during their early history. Absolute diversification rates were estimated for order-level clades using ages derived from relaxed molecular clock analyses that included or excluded a maximal constraint to angiosperm age. Diversification rates for angiosperms as a whole ranged from 0.0781 to 0.0909 net speciation events per million years, with dates from the constrained analysis. Diversification through time plots show an inverse relationship between clade age and rate, where the younger clades tend to have the highest rates. Angiosperm diversity is found to have mixed origins: slightly less than half of the living species belong to lineages with low to moderate diversification rates, which appeared between 130 and 102 Mya (Barremian-uppermost Albian; Lower Cretaceous). Slightly over half of the living species belong to lineages with moderate to high diversification rates, which appeared between 102 and 77 Mya (Cenomanian-mid Campanian; Upper Cretaceous). Terminal lineages leading to living angiosperm species, however, may have originated soon or long after the phylogenetic differentiation of the clade to which they belong.

389 citations


Journal ArticleDOI
TL;DR: 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.
Abstract: 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.

301 citations


Journal ArticleDOI
TL;DR: This work evaluates early floral evolution in angiosperms by parsimony optimization of morphological characters on phylogenetic trees derived from morphological and molecular data and indicates that the simple flowers of the newly recognized basal groups are reduced rather than primitively simple.
Abstract: Increasingly robust understanding of angiosperm phylogeny allows more secure reconstruction of the flower in the most recent common ancestor of extant angiosperms and its early evolution. The surprising emergence of several extant and fossil taxa with simple flowers near the base of the angiosperms-Chloranthaceae, Ceratophyllum, Hydatellaceae, and the Early Cretaceous fossil Archaefructus (the last three are water plants)-has brought a new twist to this problem. We evaluate early floral evolution in angiosperms by parsimony optimization of morphological characters on phylogenetic trees derived from morphological and molecular data. Our analyses imply that Ceratophyllum may be related to Chloranthaceae, and Archaefructus to either Hydatellaceae or Ceratophyllum. Inferred ancestral features include more than two whorls (or series) of tepals and stamens, stamens with protruding adaxial or lateral pollen sacs, several free, ascidiate carpels closed by secretion, extended stigma, extragynoecial compitum, and one or several ventral pendent ovule(s). The ancestral state in other characters is equivocal: e.g., bisexual vs. unisexual flowers, whorled vs. spiral floral phyllotaxis, presence vs. absence of tepal differentiation, anatropous vs. orthotropous ovules. Our results indicate that the simple flowers of the newly recognized basal groups are reduced rather than primitively simple.

293 citations


Journal ArticleDOI
TL;DR: The data suggest that SEM images of intact pit membranes underestimate the porosity of pit membranes in situ, and pit membrane porosity based on SEM offers a relative estimate of air-seeding thresholds, but absolute pore diameters must be treated with caution.
Abstract: Pit membranes between xylem vessels have been suggested to have functional adaptive traits because of their influence on hydraulic resistance and vulnerability to embolism in plants. Observations of intervessel pit membranes in 26 hardwood species using electron microscopy showed significant variation in their structure, with a more than 25-fold difference in thickness (70-1892 nm) and observed maximum pore diameter (10-225 nm). In some SEM images, pit membrane porosity was affected by sample preparation, although pores were resolvable in intact pit membranes of many species. A significant relationship (r(2) = 0.7, P = 0.002) was found between pit membrane thickness and maximum pore diameter, indicating that the thinner membranes are usually more porous. In a subset of nine species, maximum pore diameter determined from SEM was correlated with pore diameter calculated from air-seeding thresholds (r(2) = 0.8, P < 0.001). Our data suggest that SEM images of intact pit membranes underestimate the porosity of pit membranes in situ. Pit membrane porosity based on SEM offers a relative estimate of air-seeding thresholds, but absolute pore diameters must be treated with caution. The implications of variation in pit membrane thickness and porosity to plant function are discussed.

282 citations


Journal ArticleDOI
TL;DR: Evidence that, in response to increasing CO(2) concentration, anthropic perturbation and/ or increasing forest fragmentation, lianas are increasing relative to tree species is cited.
Abstract: We provide an overview of research on climbing plants from Charles Darwin to the present day. Following Darwin's interests, this review will focus on functional perspectives including attachment mechanisms and stem structure and function. We draw attention to a number of unsolved problems inviting future research. These include the mechanism for establishment of the twining habit, a quantitative description following the development of a tissue element through space and time, the chemistry of sticky exudates, the microstructure of xylem and the capacity for water storage, the vulnerability to embolism, and the mechanism for embolism repair. In conclusion we cite evidence that, in response to increasing CO(2) concentration, anthropic perturbation and/ or increasing forest fragmentation, lianas are increasing relative to tree species. In the 21st century, we are returning to the multiscale, multidisciplinary approach taken by Darwin to understand natural history.

254 citations


Journal ArticleDOI
TL;DR: The survey of the literature suggests that no single vegetative, reproductive, or ecological feature taken in isolation can account for the evolutionary success of the angiosperms, and it is believed that the answer to Darwin's second "abominable mystery" lies in a confluence of features that collectively make the angosperms unique among the land plants.
Abstract: The rapid diversification and ecological dominance of the flowering plants beg the question "Why are there so many angiosperm species and why are they so successful?" A number of equally plausible hypotheses have been advanced in response to this question, among which the most widely accepted highlights the mutually beneficial animal-plant relationships that are nowhere better developed nor more widespread than among angiosperm species and their biotic vectors for pollination and dispersal. Nevertheless, consensus acknowledges that there are many other attributes unique to or characteristic of the flowering plants. In addition, the remarkable coevolution of the angiosperms and pollination/dispersal animal agents could be an effect of the intrinsic adaptability of the flowering plants rather than a primary cause of their success, suggesting that the search for underlying causes should focus on an exploration of the genetic and epigenetic mechanisms that might facilitate adaptive evolution and speciation. Here, we explore angiosperm diversity promoting attributes in their general form and draw particular attention to those that, either individually or collectively, have been shown empirically to favor high speciation rates, low extinction rates, or broad ecological tolerances. Among these are the annual growth form, homeotic gene effects, asexual/sexual reproduction, a propensity for hybrid polyploidy, and apparent "resistance" to extinction. Our survey of the literature suggests that no single vegetative, reproductive, or ecological feature taken in isolation can account for the evolutionary success of the angiosperms. Rather, we believe that the answer to Darwin's second "abominable mystery" lies in a confluence of features that collectively make the angiosperms unique among the land plants.

194 citations


Journal ArticleDOI
TL;DR: Darwin's thoughts and frustrations with the fossil record of flowering plants as revealed in correspondence with Joseph Hooker, Gaston de Saporta, and Oswald Heer between 1875 and 1881 are explicated.
Abstract: Charles Darwin's "abominable mystery" has come to symbolize just about all aspects of the origin and early evolution of flowering plants. Yet, there has never been an analysis of precisely what Darwin thought was so abominably mysterious. Here I explicate Darwin's thoughts and frustrations with the fossil record of flowering plants as revealed in correspondence with Joseph Hooker, Gaston de Saporta, and Oswald Heer between 1875 and 1881. I also examine the essay by John Ball that prompted Darwin to write his "abominable mystery" letter to Hooker in July of 1879. Contrary to what is generally believed, Darwin's abominable mystery has little if anything to do with the fossil prehistory of angiosperms, identification of the closest relatives of flowering plants, questions of the homologies (and character transformations) of defining features of flowering plants, or the phylogeny of flowering plants themselves. Darwin's abominable mystery and his abiding interest in the radiation of angiosperms were never driven primarily by a need to understand the literal text of the evolutionary history of flowering plants. Rather, Darwin was deeply bothered by what he perceived to be an abrupt origin and highly accelerated rate of diversification of flowering plants in the mid-Cretaceous. This led Darwin to create speculative arguments for a long, gradual, and undiscovered pre-Cretaceous history of flowering plants on a lost island or continent. Darwin also took refuge in the possibility that a rapid diversification of flowering plants in the mid-Cretaceous might, if real, have a biological explanation involving coevolutionary interactions between pollinating insects and angiosperms. Nevertheless, although generations of plant biologists have seized upon Darwin's abominable mystery as a metaphor for their struggle to understand angiosperm history, the evidence strongly suggests that the abominable mystery is not about angiosperms per se. On the contrary, Darwin's abominable mystery is about his abhorrence that evolution could be both rapid and potentially even saltational. Throughout the last years of his life, it just so happens that flowering plants, among all groups of organisms, presented Darwin with the most extreme exception to his strongly held notion natura non facit saltum, nature does not make a leap.

186 citations


Journal ArticleDOI
TL;DR: The proportion of fibers in direct contact with vessels decreased with and was independent of wood density, indicating that the number of fiber-vessel contacts does not explain the previously observed correlation between wood density and implosion resistance
Abstract: Wood density plays a key role in ecological strategies and life history variation in woody plants, but little is known about its anatomical basis in shrubs. We quantified the relationships between wood density, anatomy, and climate in 61 shrub species from eight field sites along latitudinal belts between 31° and 35° in North and South America. Measurements included cell dimensions, transverse areas of each xylem cell type and percentage contact between different cell types and vessels. Wood density was more significantly correlated with precipitation and aridity than with temperature. High wood density was achieved through reductions in cell size and increases in the proportion of wall relative to lumen. Wood density was independent of vessel traits, suggesting that this trait does not impose conduction limitations in shrubs. The proportion of fibers in direct contact with vessels decreased with and was independent of wood density, indicating that the number of fiber-vessel contacts does not explain the previously observed correlation between wood density and implosion resistance. Axial and radial parenchyma each had a significant but opposite association with wood density. Fiber size and wall thickness link wood density, life history, and ecological strategies by controlling the proportion of carbon invested per unit stem volume.

180 citations


Journal ArticleDOI
TL;DR: The breeding of new, high-quality citrus cultivars depends on dependable information about the relationships of taxa within the tribe Citreae; therefore, it is important to have a well-supported phylogeny of the relationships between species not only to advance breeding strategies, but also to advance conservation strategies for the wild taxa.
Abstract: The breeding of new, high-quality citrus cultivars depends on dependable information about the relationships of taxa within the tribe Citreae; therefore, it is important to have a well-supported phylogeny of the relationships between species not only to advance breeding strategies, but also to advance conservation strategies for the wild taxa. The recent history of the systematics of Citrus (Rutaceae: Aurantioideae) and its allies, in the context of Rutaceae taxonomy as a whole, is reviewed. The most recent classifi cation is tested using nine cpDNA sequence regions in representatives of all genera of the subfam. Aurantioideae (save Limnocitrus ) and numerous species and hybrids referred to Citrus s.l. Aurantioideae are confi rmed as monophyletic. Within Aurantioideae, tribe Clauseneae are not monophyletic unless Murraya s.s. and Merrillia are removed to Aurantieae. Within tribe Aurantieae, the three traditionally recognized subtribes are not monophyletic. Triphasiinae is not monophyletic unless Oxanthera is returned to Citrus (Citrinae). Balsamocitrinae is polyphyletic. Feroniella , traditionally considered allied closely to Limonia (= Feronia ), is shown to be nested in Citrus. The proposed congenericity of Severinia and Atalantia is confi rmed. The most recent circumscription of Citrus is strongly supported by this analysis, with hybrids appearing with their putative maternal parents. The genus was resolved into two clades, one comprising wild species from New Guinea, Australia, and New Caledonia (formerly Clymenia , Eremocitrus , Microcitrus , Oxanthera ), but surprisingly also Citrus medica , traditionally believed to be native in India. The second clade is largely from the Asian mainland (including species formerly referred to Fortunella and Poncirus ).

Journal ArticleDOI
TL;DR: Barcoding is a retroactive procedure that relies on well-defined species to function, is based solely on a limited number of DNA sequences that are often inappropriate at the species level, has been poorly tested with geographically well-dispersed replicate samples from difficult taxonomic groups, and discounts substantial practical and theoretical problems in defining species.
Abstract: DNA barcoding ("barcoding") has been proposed as a rapid and practical molecular method to identify species via diagnostic variation in short orthologous DNA sequences from one or a few universal genomic regions. It seeks to address in a rapid and simple way the "taxonomic impediment" of a greater need for taxonomic identifications than can be supplied by taxonomists. Using a complicated plant group, Solanum sect. Petota (wild potatoes), I tested barcoding with the most variable and frequently suggested plant barcoding regions: the internal nontranscribed spacer of nuclear ribosomal DNA (ITS) and the plastid markers trnH-psbA intergenic spacer and matK. These DNA regions fail to provide species-specific markers in sect. Petota because the ITS has too much intraspecific variation and the plastid markers lack sufficient polymorphism. The complications seen in wild potatoes are common in many plant groups, but they have not been assessed with barcoding. Barcoding is a retroactive procedure that relies on well-defined species to function, is based solely on a limited number of DNA sequences that are often inappropriate at the species level, has been poorly tested with geographically well-dispersed replicate samples from difficult taxonomic groups, and discounts substantial practical and theoretical problems in defining species.

Journal ArticleDOI
TL;DR: Several traits responded to relatedness in shared pots, including increased leaf to root allocation with strangers and increased stem elongation and branchiness in response to kin, potentially indicating both increased competition toward strangers and reduced interference (cooperation) toward kin.
Abstract: The ability to recognize kin is an important element in social behavior and can lead to the evolution of altruism. Recently, it has been shown that plants are capable of kin recognition through root interactions. Here we tested for kin recognition in a North American species of Impatiens that has a high opportunity of growing with kin and responds strongly to aboveground competition. We measured how the plants responded to the aboveground light quality cues of competition and to the presence of root neighbors and determined whether the responses depended on whether the neighbors were siblings or strangers. The study families were identified by DNA sequencing as members of the same species, provisionally identified as Impatiens pallida (hereafter I. cf. pallida). We found that I. cf. pallida plants were capable of kin recognition, but only in the presence of another plant's roots. Several traits responded to relatedness in shared pots, including increased leaf to root allocation with strangers and increased stem elongation and branchiness in response to kin, potentially indicating both increased competition toward strangers and reduced interference (cooperation) toward kin. Impatiens cf. pallida responded to both competition cues simultaneously, with the responses to the aboveground competition cue dependent on the presence of the belowground competition cue.

Journal ArticleDOI
TL;DR: This study tested the performance and accuracy of several nondestructive, reflectance-based techniques for estimating anthocyanin and chlorophyll contents in leaves of four unrelated species, European hazel, Siberian dogwood, Norwegian maple, Norway maple, and Virginia creeper.
Abstract: The anthocyanin and chlorophyll contents in leaves provide valuable information about the physiological status of plants. Thus, there is a need for accurate, efficient, and practical methodologies to estimate these biochemical parameters of vegetation. In this study, we tested the performance and accuracy of several nondestructive, reflectance-based techniques for estimating anthocyanin and chlorophyll contents in leaves of four unrelated species, European hazel (Corylus avellana), Siberian dogwood (Cornus alba =Swida alba), Norway maple (Acer platanoides), and Virginia creeper (Parthenocissus quinquefolia), with widely variable pigment content and composition. An anthocyanin reflectance index, which uses reflectances in the green and red edge spectral bands, and a modified anthocyanin reflectance index, employing, in addition, the near-infrared (NIR) band, were able to accurately estimate leaf anthocyanin for all species taken together with no reparameterization of algorithms. Total chlorophyll content was accurately estimated by a red edge chlorophyll index that uses spectral bands in the red edge and the NIR. These approaches can be used to estimate anthocyanin and chlorophyll nondestructively and allow the development of simple handheld field instrumentation.

Journal ArticleDOI
TL;DR: The present sampling of loci did not conclusively resolve the number and location of domestications of C. annuum, but several lines of evidence suggest multiple independent domestications from widely distributed progenitor populations.
Abstract: The chile of Mesoamerica, Capsicum annuum, is one of five domesticated chiles in the Americas. Among the chiles, it varies the most in size, form, and color of its fruits. Together with maize, C. annuum is one of the principal elements of the neotropical diets of Mesoamerican civilizations. Despite the great economic and cultural importance of C. annuum both worldwide and in Mexico, however, very little is known about its geographic origin and number of domestications. Here we sampled a total of 80 accessions from Mexico (58 semiwild and 22 domesticated) and examined nucleotide sequence diversity at three single- or low-copy nuclear loci, Dhn, G3pdh, and Waxy. Across the three loci, we found an average reduction of ca. 10% in the diversity of domesticates relative to semiwild chiles and geographic structure within Mexican populations. The Yucatan Peninsula contained a large number of haplotypes, many of which were unique, suggesting an important region of chile domestication and center of diversity. The present sampling of loci did not conclusively resolve the number and location of domestications, but several lines of evidence suggest multiple independent domestications from widely distributed progenitor populations.

Journal ArticleDOI
TL;DR: Parsimony inference suggests that the family originated in the neotropics, with at least five dispersal events leading to the Old World representatives.
Abstract: Bignoniaceae are woody, trees, shrubs, and lianas found in all tropical fl oras of the world with lesser representation in temperate regions. Phylogenetic analyses of chloroplast sequences ( rbcL , ndhF , trnL-F ) were undertaken to infer evolutionary relationships in Bignoniaceae and to revise its classifi cation. Eight clades are recognized as tribes (Bignonieae, Catalpeae, Coleeae, Crescentieae, Jacarandeae, Oroxyleae, Tecomeae, Tourrettieae); additional inclusive clades are named informally. Jacarandeae and Catalpeae are resurrected; the former is sister to the rest of the family, and the latter occupies an unresolved position within the “ core ” Bignoniaceae. Tribe Eccremocarpeae is included in Tourrettieae. Past classifi cations recognized a large Tecomeae, but this tribe is paraphyletic with respect to all other tribes. Here Tecomeae are reduced to a clade of approximately 12 genera with a worldwide distribution in both temperate and tropical ecosystems. Two large clades, Bignonieae and Crescentiina, account for over 80% of the species in the family. Coleeae and Crescentieae are each included in larger clades, the Paleotropical alliance and Tabebuia alliance, respectively; each alliance includes a grade of taxa assigned to the traditional Tecomeae. Parsimony inference suggests that the family originated in the neotropics, with at least fi ve dispersal events leading to the Old World representatives.

Journal ArticleDOI
TL;DR: It is hypothesized that large flowers in Nymphaeaceae are the result of the interaction of heat, floral odors, and colored tepals to trap insects to increase fitness.
Abstract: The fi rst three branches of the angiosperm phylogenetic tree consist of eight families with ~201 species of plants (the ANITA grade). The oldest fl ower fossil for the group is dated to the Early Cretaceous (115 - 125 Mya) and identifi ed to the Nymphaeales. The fl owers of extant plants in the ANITA grade are small, and pollen is the edible reward (rarely nectar or starch bodies). Unlike many gymnosperms that secrete " pollination drops, " ANITA-grade members examined thus far have a dry-type stigma. Copious secretions of stigmatic fl uid are restricted to the Nymphaeales, but this is not nectar. Floral odors, fl oral thermogenesis (a re- source), and colored tepals attract insects in deceit-based pollination syndromes throughout the fi rst three branches of the phylo- genetic tree. Self-incompatibility and an extragynoecial compitum occur in some species in the Austrobaileyales. Flies are primary pollinators in six families (10 genera). Beetles are pollinators in fi families varying in importance as primary (exclusive) to secondary vectors of pollen. Bees are major pollinators only in the Nymphaeaceae. It is hypothesized that large fl owers in Nympha- eaceae are the result of the interaction of heat, fl oral odors, and colored tepals to trap insects to increase fi tness.

Journal ArticleDOI
TL;DR: The flexibility of the aerial system is found to be critical to reducing the resistance to the airflow and thus to minimizing the risk of failure, and slender trees are shown to be relatively more vulnerable to stem breakage than uprooting.
Abstract: Wind can alter plant growth and cause extensive, irreversible damage in forested areas. To hotter understand how to mitigate the effects of wind action, we investigated the sensitivity of tree aerodynamic behavior to the material and geometrical factors characterizing the aerial system. The mechanical response of a 35 yr-old maritime pine (Pinas /thoser. Pinaceae) submitted to static and dynamic wind loads is simulated with a finite element model. The branching structure is represented in three dimensions. Factor effects are evaluated using, a fractional experimental design. Results show that material properties play only a limited role in tree dynamics. In contrast, small morphological variations can produce extreme behaviors such as either very little or nearly critical dissipation of stem oscillations. Slender trees are shown to be relatively more vulnerable to stem breakage than uprooting. Dynamic loading, leads to deflections and forces up to 20% higher near the base of the tree than those calculated for a static loading of similar magnitude. Effects of branch geometry on dynamic amplification are substantial yet not linear. The flexibility of the aerial system is found to be critical to reducing the resistance to the airflow and thus to minimizing the risk of failure.

Journal ArticleDOI
Sarah Mathews1
TL;DR: It is likely that improved sampling of gymnosperm diversity in nucleotide data sets will help alleviate some of the analytical issues encountered in the estimation of seed plant phylogeny, providing a more definitive test of morphological trees.
Abstract: Trees inferred from DNA sequence data provide only limited insight into the phylogeny of seed plants because the living lineages (cycads, Ginkgo, conifers, gnetophytes, and angiosperms) represent fewer than half of the major lineages that have been detected in the fossil record. Nevertheless, phylogenetic trees of living seed plants inferred from sequence data can provide a test of relationships inferred in analyses that include fossils. So far, however, significant uncertainty persists because nucleotide data support several conflicting hypotheses. It is likely that improved sampling of gymnosperm diversity in nucleotide data sets will help alleviate some of the analytical issues encountered in the estimation of seed plant phylogeny, providing a more definitive test of morphological trees. Still, rigorous morphological analyses will be required to answer certain fundamental questions, such as the identity of the angiosperm sister group and the rooting of crown seed plants. Moreover, it will be important to identify approaches for incorporating insights from data that may be accurate but less likely than sequence data to generate results supported by high bootstrap values. How best to weigh evidence and distinguish among hypotheses when some types of data give high support values and others do not remains an important problem.

Journal ArticleDOI
TL;DR: This analysis goes beyond many phylogenies in exploring how phylogenetic structure imposed by morphology, ecology, and geography reveals useful evolutionary data, suggesting niche conservation and dispersal limitation are important processes rendering signature shapes to the Indigofereae phylogeny in different biomes.
Abstract: This analysis goes beyond many phylogenies in exploring how phylogenetic structure imposed by morphology, ecology, and geography reveals useful evolutionary data. A comprehensive range of such diversity is evaluated within tribe Indigofereae and outgroups from sister tribes. A combined data set of 321 taxa (over one-third of the tribe) by 80 morphological characters, 833 aligned nuclear ribosomal ITS/5.8S sites, and an indel data set of 33 characters was subjected to parsimony analysis. Notable results include the Madagascan dry forest Disynstemon resolved as sister to tribe Indigofereae, and all species of the large genus Indigofera comprise just four main clades, each diagnosable by morphological synapomorphies and ecological and geographical predilections. These results suggest niche conservation (ecology) and dispersal limitation (geography) are important processes rendering signature shapes to the Indigofereae phylogeny in different biomes. Clades confined to temperate and succulent-rich biomes are more dispersal limited and have more geographical phylogenetic structure than those inhabiting tropical grass-rich vegetation. The African arid corridor, particularly the Namib center of endemism, harbors many of the oldest Indigofera lineages. A rates analysis of nucleotide substitutions confirms that the ages of the oldest crown clades are mostly younger than 16 Ma, implicating dispersal in explaining the worldwide distribution of the tribe.

Journal ArticleDOI
TL;DR: Data support the concept that coiling and twining in vines is caused by the presence of G fibers, which occurred concomitantly with the ability to coil in redvine.
Abstract: Although the coiling of tendrils and the twining of vines has been investigated since Darwin's time, a full understanding of the mechanism(s) of this coiling and twining ability has not yet been obtained. In a previous study (Planta 225: 485-498), gelatinous (G) fibers in tendrils of redvine occurred concomitantly with the ability to coil, strongly indicating their role in the coiling process. In this study, tendrils and twining vines of a number of species were examined using microscopic and immunocytochemical techniques to determine if a similar presence and distribution of these fibers exists in other plant species. Tendrils that coiled in many different directions had a cylinder of cortical G fibers, similar to redvine. However, tendrils that coiled only in a single direction had gelatinous fibers only along the inner surface of the coil. In tendrils with adhesive tips, the gelatinous fibers occurred in the central/core region of the tendril. Coiling occurred later in development in these tendrils, after the adhesive pad had attached. In twining stems, G fibers were not observed during the rapid circumnutation stage, but were found at later stages when the vine's position was fixed, generally one or two nodes below the node still circumnutating. The number and extent of fiber development correlated roughly with the amount of torsion required for the vine to ascend a support. In contrast, species that use adventitious roots for climbing or were trailing/scrambling-type vines did not have G fibers. These data strongly support the concept that coiling and twining in vines is caused by the presence of G fibers.

Journal ArticleDOI
TL;DR: While it is confirmed that filamentous and unicellular photobiont morphotypes belong to a single clade, this clade does not cluster with Scytonema but represents a novel, previously unrecognized, highly diverse, exclusively lichenized lineage, for which the name Rhizonema is available.
Abstract: Phylogenetic diversity of lichen photobionts is low compared to that of fungal counterparts. Most lichen fungi are thought to be associated with just four photobiont genera, among them the cyanobacteria Nostoc and Scytonema , two of the most important nitrogen fi xers in humid ecosystems. Although many Nostoc photobionts have been identifi ed using isolated cultures and sequences, the identity of Scytonema photobionts has never been confi rmed by culturing or sequencing. We investigated the phylogenetic placement of presumed Scytonema photobionts and unicellular morphotypes previously assigned to Chroococcus , from tropical Dictyonema , Acantholichen , Coccocarpia , and Stereocaulon lichens. While we confi rm that fi lamentous and unicellular photobiont morphotypes belong to a single clade, this clade does not cluster with Scytonema but represents a novel, previously unrecognized, highly diverse, exclusively lichenized lineage, for which the name Rhizonema is available. The phylogenetic structure observed in this novel lineage suggests absence of coevolution with associated mycobionts at the species or clade level. Instead, highly effi cient photobiont strains appear to have evolved through photobiont sharing between unrelated, but ecologically similar, coexisting lineages of lichenized fungi ( “ lichen guilds ” ), via the selection of particular photobiont strains through and subsequent horizontal transfer among unrelated mycobionts, a phenomenon not unlike crop domestication.

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TL;DR: The results show that the reported differences between early- and late-flowering species may be widespread, but they are not ubiquitous.
Abstract: Variability in plant phenological responses to climate change is likely to lead to changes in many ecological relationships as the climate continues to change. We used a 34-yr record of flowering times and flower abundance for four species (two Delphinium [Ranunculaceae] species and two Mertensia [Boraginaceae] species) from a subalpine plant community near the Rocky Mountain Biological Laboratory to test the hypothesis that the phenologies of early-flowering species change more rapidly in response to climatological and other abiotic cues than do late-flowering species, a pattern previously found in plant communities in North America and Europe. We also explored a related hypothesis, that flower abundance of late-flowering species is more responsive to changes in climate than that of early-flowering species. The Delphinium species did not support these hypotheses, but the Mertensia species did. The difference between the peak flowering times of the early and late Mertensia species is expanding, leading to a period of diminished resources for pollinators that specialize on this genus. Mertensia ciliata populations are already severely declining in our study area, possibly as a result of earlier snowmelt. Together, these results show that the reported differences between early- and late-flowering species may be widespread, but they are not ubiquitous.

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TL;DR: The data support earlier hypotheses that divergence in flowering time causes assortative mating, allowing these ecologically distinct sister species to occur in sympatry and help to explain the overdispersion of oak species in local communities.
Abstract: The genus Quercus (the oaks) is notorious for interspecific hybrization, generating questions about the mechanisms that permit coexistence of closely related species. Two sister oak species, Quercus virginiana and Q. geminata, occur in sympatry in Florida and throughout the southeastern United States. In 11 sites from northern and southeastern regions of Florida, we used a leaf-based morphological index to identify individuals to species. Eleven nuclear microsatellite markers significantly differentiated between the species with a high correspondence between molecular and morphological typing of specimens. Nevertheless, Bayesian clustering analysis indicates interspecific gene flow, and six of 109 individuals had mixed ancestry. The identity of several individuals also was mismatched using molecular markers and morphological characters. In a common environment, the two species performed differently in terms of photosynthetic performance and growth, corresponding to their divergent ecological niches with respect to soil moisture and other edaphic properties. Our data support earlier hypotheses that divergence in flowering time causes assortative mating, allowing these ecologically distinct sister species to occur in sympatry. Limited gene flow that permits ecological differentiation helps to explain the overdispersion of oak species in local communities.

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TL;DR: Four new genera and six new species of fossil seed are described from five Early Cretaceous mesofossil floras from Portugal and eastern North America, suggesting that this newly recognized complex of extinct plants, together with Bennettitales, Erdtmanithecales, and Gnetales (the BEG group), is phylogenetically closely related.
Abstract: Four new genera and six new species of fossil seed (Buarcospermum tetragonium, Lignierispermum maroneae, Lobospermum glabrum, L. rugosum, L. stampanonii, Rugonella trigonospermum) are described from five Early Cretaceous mesofossil floras from Portugal and eastern North America. The four genera are distinguished by differences in size, shape, and details of seed anatomy, but all are unusual in having an outer seed envelope with a distinctive anatomical structure that surrounds the nucellus and the integument. The integument is extended apically into a long, narrow micropylar tube. The four new genera are part of a diverse, but previously unrecognized, complex of extinct plants that was widespread in Early Cretaceous vegetation and that coexisted in similar habitats with early angiosperms. The distinctive structure of these seeds, and the strong similarities to other fossil seeds (Ephedra, Ephedripites, Erdtmanispermum, Raunsgaardispermum, and some Bennettitales) already known from the Early Cretaceous, suggests that this newly recognized complex of extinct plants, together with Bennettitales, Erdtmanithecales, and Gnetales (the BEG group), is phylogenetically closely related.

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TL;DR: Combined evidence indicates a biome similar to extant subtropical, or tropical montane, rainforests that persisted for at least 4.4 Myr, linking elevated floral richness to abundant rainfall.
Abstract: The 51.9 Ma Laguna del Hunco (LH) and 47.5 Ma Rio Pichileufu (RP) floras from Patagonia, Argentina are unusually rich, angiosperm-dominated assemblages with living relatives in the low-latitude West Pacific, neotropics, and temperate southern latitudes. The diverse gymnosperms in these floras are important for Gondwanan biogeographic history and paleoclimatic interpretations. "Libocedrus" prechilensis Berry 1938 (Cupressaceae), previously known only from the holotype (RP), a vegetative branch, is revised here based on new material from both localities, including a seed cone attached to a shoot with cuticle (LH). Characters of these fossils are diagnostic of monotypic Papuacedrus (highlands of New Guinea and Moluccas). Living P. papuana is most abundant in cloud forests receiving up to 4 m rainfall annually, whereas Austrocedrus (Libocedrus) chilensis, the basis of comparison when the fossil species was named, inhabits dry, cold steppe margins to mediterranean climates in southern South America. We establish Papuacedrus prechilensis comb. nov., which simultaneously invalidates a southern South American connection for the fossil floras and reveals a link to West Pacific montane rainforests. Combined evidence indicates a biome similar to extant subtropical, or tropical montane, rainforests that persisted for at least 4.4 Myr, linking elevated floral richness to abundant rainfall.

Journal ArticleDOI
TL;DR: Characters of bennettitalean plants are summarized and new evidence for the structure of cones and seeds is presented that help clarify relationships of Bennettitales to flowering plants, Gnetales, and other potential angiosperm sister groups.
Abstract: Bennettitales is an extinct group of seed plants with reproductive structures that are similar in some respects to both Gnetales and angiosperms, but systematic relationships among the three clades remain controversial. This study summarizes characters of bennettitalean plants and presents new evidence for the structure of cones and seeds that help clarify relationships of Bennettitales to flowering plants, Gnetales, and other potential angiosperm sister groups. Bennettitales have simple mono- or bisporangiate cones. Seeds are borne terminally on sporophylls. They have a unique structure that includes a nucellus with a solid apex, no pollen chamber, and a single integument, and they are clearly not enclosed by a cupule or other specialized structures. Such features differ substantially from Gnetales, flowering plants, and the seed fern Caytonia, providing no compelling evidence for the origin of the angiospermous carpel. Cladistic tests were performed to assess the strength of the "anthophyte hypothesis" and possible relationships of Bennettitales, Gnetales, and Caytonia to flowering plants. Our results do not support the anthophyte hypothesis for the origin of angiosperms by a transformation of fertile organs that were already aggregated into a cone or flower-like structure. However, the anthophyte topology of the seed plant tree continues to be supported by morphological analyses of living and extinct taxa.

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TL;DR: This work explored the effects of invasive Lythrum salicaria on the reproductive success of Mimulus ringens, a wetland plant native to eastern North America and found that grooming by bees as they forage on a competitor may also significantly reduce outcross pollen export and seed set in Mimulusringens.
Abstract: Movement of pollinators between cofl owering plant species may infl uence conspecifi c pollen deposition and seed set. Interspecifi c pollinator movements between native and showy invasive plants may be particularly detrimental to the pollination and reproductive success of native species. We explored the effects of invasive Lythrum salicaria on the reproductive success of Mimulus ringens , a wetland plant native to eastern North America. Pollinator fl ights between these species signifi cantly reduced the amount of conspecifi c pollen deposited on Mimulus stigmas and the number of seeds in Mimulus fruits, suggesting that pollen loss is an important mechanism of competition for pollination. Although pollen loss is often attributed to pollen wastage on heterospecifi c fl oral structures, our novel fi ndings suggest that grooming by bees as they forage on a competitor may also signifi cantly reduce outcross pollen export and seed set in Mimulus ringens .

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TL;DR: The distinctive, raised tetra- to cyclocytic stomatal subsidiary cells of Earina and characteristic papilla-like absorbing glands and "ringed" guard cells of Dendrobium support the placement of the fossils into these genera, and demonstrate expansion of epiphytic orchids into Zealandia by the mid-Cenozoic and an important role for southern continents in the diversification of Orchidaceae.
Abstract: Fossil leaves of two Early Miocene orchids (Dendrobium and Earina) are reported from New Zealand. The distinctive, raised tetra- to cyclocytic stomatal subsidiary cells of Earina and characteristic papilla-like absorbing glands and "ringed" guard cells of Dendrobium support the placement of the fossils into these genera. These therefore represent the first Orchidaceae macrofossils with cuticular preservation, the oldest records for subfamily Epidendroideae, as well as the first New Zealand and southern hemisphere fossil records for Orchidaceae. These taxa belong in basal clades to the Vandeae/Cymbideae or Epidendreae (Earina) and the Australasian clade of Dendrobium sensu lato. This phylogenetic placement demonstrates expansion of epiphytic orchids into Zealandia by the mid-Cenozoic and an important role for southern continents in the diversification of Orchidaceae.

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TL;DR: It is found here that the widespread apomictic triploid, Cheilanthes lindheimeri, is an autopolyploid derived from a rare, previously undetected sexual diploid.
Abstract: Deciphering species relationships and hybrid origins in polyploid agamic species complexes is notoriously diffi cult. In this study of cheilanthoid ferns, we demonstrate increased resolving power for clarifying the origins of polyploid lineages by integrating evidence from a diverse selection of biosystematic methods. The prevalence of polyploidy, hybridization, and apomixis in ferns suggests that these processes play a signifi cant role in their evolution and diversifi cation. Using a combination of systematic approaches, we investigated the origins of apomictic polyploids belonging to the Cheilanthes yavapensis complex. Spore studies allowed us to assess ploidy levels; plastid and nuclear DNA sequencing revealed evolutionary relationships and confi rmed the putative progenitors (both maternal and paternal) of taxa of hybrid origin; enzyme electrophoretic evidence provided information on genome dosage in allopolyploids. We fi nd here that the widespread apomictic triploid, Cheilanthes lindheimeri , is an autopolyploid derived from a rare, previously undetected sexual diploid. The apomictic triploid Cheilanthes wootonii is shown to be an interspecifi c hybrid between C. fendleri and C. lindheimeri , whereas the apomictic tetraploid C. yavapensis is comprised of two cryptic and geographically distinct lineages. We show that earlier morphology-based hypotheses of species relationships, while not altogether incorrect, only partially explain the complicated evolutionary history of these ferns.