Showing papers in "American Journal of Botany in 1998"

The tortoise and the hare: choosing between noncoding plastome and nuclear Adh sequences for phylogeny reconstruction in a recently diverged plant group

Abstract: Phylogenetic resolution is often low within groups of recently diverged taxa due to a paucity of phylogenetically informative characters. We tested the relative utility of seven noncoding cpDNA regions and a pair of homoeologous nuclear genes for resolving recent divergences, using tetraploid cottons ( Gossypium) as a model system. The five tetraploid species of Gossypium are a monophyletic assemblage derived from an allopolyploidization event that probably occurred within the last 0.5‐2 million years. Previous analysis of cpDNA restriction site data provided only partial resolution within this clade despite a large number of enzymes employed. We sequenced three cpDNA introns (rpl16, rpoC1, ndhA) and four cpDNA spacers (accD-psaI, trnL-trnF, trnT-trnL, atpB-rbcL) for a total of over 7 kb of sequence per taxon, yet obtained only four informative nucleotide substitutions (0.05%) resulting in incomplete phylogenetic resolution. In addition, we sequenced a 1.65-kb region of a homoeologous pair of nuclear-encoded alcohol dehydrogenase (Adh) genes. In contrast with the cpDNA sequence data, the Adh homoeologues yielded 25 informative characters (0.76%) and provided a robust and completely resolved topology that is concordant with previous cladistic and phenetic analyses. The enhanced resolution obtained using the nuclear genes reflects an approximately three- to sixfold increase in nucleotide substitution rate relative to the plastome spacers and introns.

Interspecific variation in plant responses to mycorrhizal colonization in tallgrass prairie

Abstract: Symbiotic associations between plants and arbuscular mycorrhizal fungi are ubiquitous and ecologically important in many grasslands. Differences in species responses to mycorrhizal colonization can have a significant influence on plant community structure. The growth responses of 36 species of warm- and cool-season tallgrass prairie grasses and 59 tallgrass prairie forbs to arbuscular mycorrhizal (AM) fungal colonization were assessed in greenhouse studies to examine the extent of interspecific variation in host-plant benefit from the symbiosis and patterns of mycorrhizal dependence among host plant life history (e.g., annual, perennial) and taxonomic (e.g., grass, forb, legume, nonlegume) groups and phenological guilds. There was a strong and significant relationship between phenology of prairie grasses and mycorrhizal responsiveness, however this relationship was less apparent in forbs. Perennial warm-season C(4) grasses and forbs generally benefited significantly from the mycorrhizal symbiosis, whereas biomass production of the cool-season C(3) grasses was not affected. The root systems of the cool-season grasses were also less highly colonized by the AM fungi, as compared to the warm-season grasses or forbs. Unlike the native perennials, annuals were generally not responsive to mycorrhizal colonization and were lower in percentage root colonization than the perennial species. Plant growth responsiveness and AM root colonization were positively correlated for the nonleguminous species, with this relationship being strongest for the cool-season grasses. In contrast, root colonization of prairie legumes showed a significant, but negative, relationship to mycorrhizal growth responsiveness.

Fruit set, nectar reward, and rarity in the Orchidaceae.

Abstract: A review of comparative levels of reproductive success among nectariferous and nectarless orchids worldwide was compiled from a comprehensive survey of fruit set from 117 orchid species in the literature and from our own field studies. It confirms the hypothesis that nectariferous orchids are more successful in setting fruit than are nectarless species. Overall fruit set figures for nectarless and nectariferous orchids were 19.5 and 49.3% for North America, 27.7 and 63.1% for Europe, 41.4 and 74.4% for the temperate southern hemisphere, and 11.5 and 24.9% for the tropics, demonstrating that the dichotomy is consistent across all geographical areas. On average, the provision of nectar doubles the probability of fruit set in both temperate and tropical areas, but tropical orchids are remarkable in that all (whether nectarless or nectariferous, or terrestrial or epiphytic) display low fruit productivity (<50%). Fruiting failure in the tropics may be balanced by higher productivity per capsule, since tropical orchid fruits contain on average 150 times more seeds than temperate ones. Hybridization occurs more frequently among nectarless orchids in Britain and Europe than among nectariferous ones, and there is a significant positive association between orchid rarity and lack of nectar reward in the British Isles. Sexual reproduction in the Orchidaceae is predominantly pollinator dependent, but this can sometimes be successfully circumvented by asexual seed production (agamospermy) or, more frequently, by automatic self-pollination (autogamy). The proportion of highly successful nectarless orchids from all geographic areas is very low and comparable with that of orchids offering rewards other than nectar (∼14% of species in each case) emphasizing that high reproductive success is only associated with nectar reward (53% of species). It is suggested that the evolution of nectar production within the family has been the most frequent means of escaping the reproductive limitations of low pollinator visitation frequencies.

Pollination biology in a lowland dipterocarp forest in Sarawak, Malaysia. I. Characteristics of the plant-pollinator community in a lowland dipterocarp forest

Abstract: Flowerings and flower visitors were observed continuously in alowland dipterocarp forest in Sarawak, Malaysia, for 53 mo in1992-1996. Flower visitors of 270 plant species were observed orcollected, and pollinators were assessed by observing body contact tostigmas and anthers. We recognized 12 categories of pollination systems.Among them, plants pollinated by social bees included the largest numberof species (32%) and were followed by beetle-pollinated species(20%). Pollination systems were significantly related with somefloral characters (flowering time of day, reward, and floral shape), butnot with floral color. Based on the relationships between pollinatorsand floral characters, we described pollination syndromes found in alowland dipterocarp forest. The dominance of social bees and beetlesamong pollinators is discussed in relation to the general floweringobserved in dipterocarp forests of West Malesia. In spite of high plantspecies diversity and consequent low population densities of lowlanddipterocarp forests, long-distance-specific pollinators were uncommoncompared with theNeotropics.

RAPD variation in relation to population size and plant fitness in the rare Gentianella germanica (Gentianaceae)

Abstract: We investigated the distribution of genetic variation and the relationship between population size and genetic variation in the rare plant Gentianella germanicausing RAPD (random amplified polymorphic DNA) profiles. Plants for the analysis were grown from seeds sampled from 72 parent plants in 11 G. germanica populations of different size (40‐5000 fruiting individuals). In large populations, seeds were sampled from parents in two spatially distinct subpopulations comparable in area to the total area covered by small populations. Analysis of molecular variance revealed significant genetic variation among populations (P , 0.001), while genetic variation among subpopulations was marginally significant (P , 0.06). Average molecular variance within subpopulations in large populations did not differ significantly from whole-population values. There was a positive correlation between genetic variation and population size ( P , 0.01). Genetic variation was also positively correlated with the number of seeds per plant in the field (P , 0.02) and the number of flowers per planted seed in a common garden experiment (P , 0.051). We conclude that gene flow among natural populations is very limited and that reduced plant fitness in small populations of G. germanica most likely has genetic causes. Management should aim to increase the size of small populations to minimize further loss of genetic variation. Because a large proportion of genetic variation is among populations, even small populations are worth preserving.

Latitudinal population differentiation in two species of Solidago (Asteraceae) introduced into Europe

Abstract: Solidago altissima and S. gigantea were introduced from North America to Europe ~250 yr ago and are now considered aggressive weeds in abandoned fields and conservation areas. We studied patterns of genetic differentiation in these two species along their present latitudinal range in Europe (44-61 degrees N). Two generations of clonally propagated ramets from randomly selected genets of 24 populations of each species were grown in a common-garden experiment at latitude 47 degrees N from 1991 to 1992. Both species showed significant variation among populations in morphological and life-history characters: at this southern location, plants from northern populations were smaller and flowered earlier than plants from southern populations. The gradient of clinal variation was more pronounced in the second year of cultivation than in the first and was steeper in S. altissima than in S. gigantea. Within-population variation among genotypes was significant tot most characters in the case of S. altissima. Phenological rate (reciprocal of days to flowering) and size at maturity showed a significant negative correlation among populations bot not among genotypes within populations, indicating that genetic trade-offs may occur at one but not another infraspecific level. We suggest that the pattern of among-population variation reflects rapid adaptive population differentiation after introduction of the species to Europe.

The effects of seed size, cotyledon reserves, and herbivory on seedling survival and growth in Quercus rugosa and Q. laurina (Fagaceae).

Abstract: In a greenhouse experiment, seedling survival of two oak species (Quercus rugosa and Q. laurina) was greatly affected by the excision of cotyledons 1 mo after germination, with a greater impact on Q. laurina. The effect of seed size was also significant for both species, with a positive correlation between seed mass and survival and growth. The effect of cotyledon excision on seedling growth persisted throughout the first growing season in Q. rugosa and was not analyzed for Q. laurina due to the low number of seedlings that survived cotyledon excision. Seed size significantly affected seedling height, diameter, leaf area, and biomass at 6 mo. Seed size and cotyledon retention affected the ability of Q. rugosa to recover from herbivory, as both factors had a significant effect on relative growth rates after aerial biomass removal. The results show that seedlings originating from large seeds can better endure loss of cotyledons and aerial biomass and thus are better equipped to confront stress early in their lives.

Phylogeny and radiation of pollination systems in DISA (Orchidaceae).

Abstract: We studied the patterns of adaptive radiation in Disa, a large orchid genus in southern Africa. A cladogram for 27 species was constructed using 44 morphological characters. Pollination systems were then mapped onto the phylogeny in order to analyze pathways of floral evolution. Shifts from one pollination system to another have been a major feature of the evolutionary diversification of Disa. Unlike many plant genera that are pollinated mainly by a single group of insects, radiation in Disa has encompassed nearly all major groups of pollinating insects; in all, 19 different specialized pollination systems have been found in the 27 species included in this analysis. Another striking pattern is the repeated evolution of broadly similar pollination systems in unrelated clades. For example, butterfly-pollinated flowers have evolved twice; showy deceptive flowers pollinated by carpenter bees, twice; long-spurred flowers pollinated by long-tongued flies, four times; night-scented flowers pollinated by moths, three times; and self-pollination, three times. This suggests that a few dominant pollinator species in a region may be sufficient to generate diversification in plants through repeated floral shifts that never retrace the same pathways.

A phylogenetic analysis of Rhamnaceae using rbcL and trnL-F plastid DNA sequences.

Abstract: Previous tribal classifications of Rhamnaceae have been based on fruit characters, resulting in the delimitation of large and otherwise heterogeneous groups. We evaluated the most recent classification with DNA sequences of two regions of the plastid genome, rbcL and trnL-F, from 42 genera of Rhamnaceae and representatives of the related families Elaeagnaceae, Barbeyaceae, Dirachmaceae, Urticaceae, Ulmaceae, Moraceae, and Rosaceae. The trnL-F trees have higher consistency and retention indices than the rbcL trees, and patterns of change in rbcL and trnL-F are compared. The closest relatives of Rhamnaceae are Dirachmaceae and Barbeyaceae, followed by the urticalean families. The plastid trees support the monophyly of the family and provide the basis for a new tribal classification. Three strongly supported clades are identified, but morphological characters could not be found to underpin a formal taxonomic description of these three clades as subfamilies. We therefore only recognize groups that are also defined by morphological characters. The biogeography of Rhamnaceae is discussed with reference to the molecular trees.

Variability in leaf optical properties among 26 species from a broad range of habitats.

Abstract: Leaves from 26 species with growth forms from annual herbs to trees were collected from open, intermediate, and shaded understory habitats in Mississippi and Kansas, USA. Leaf optical properties including reflectance, transmittance, and absorptance in visible and near infrared (NIR) wavelengths were measured along with leaf thickness and specific leaf mass (SLM). These leaf properties and internal light scattering have been reported to vary with light availability in studies that have focused on a limited number of species. Our objective was to determine whether these patterns in leaf optics and light availability were consistent when a greater number of species were evaluated. Leaf thickness and SLM varied by tenfold among species sampled, but within-habitat variance was high. Although there was a strong trend toward thicker leaves in open habitats, only SLM was significantly greater in open vs. understory habitats. In contrast, leaf optical properties were strikingly similar among habitats. Reflectance and reflectance/transmittance in the NIR were used to estimate internal light scattering and there were strong relationships (r 2 . 0.65) between these optical properties and leaf thickness. We concluded that leaf thickness, which did not vary consistently among habitats, was the best predictor of NIR reflectance and internal light scattering. However, because carbon allocation to leaves was lower in understory species (low SLM) yet gross optical properties were similar among all habitats, the energy investment by shade leaves required to achieve optical equivalence with sun leaves was lower. Differences in leaf longevity and growth form within a habitat may help explain the lack of consistent patterns in leaf optics as the number of species sampled increases.

Parallel evolution of glucosinolate biosynthesis inferred from congruent nuclear and plastid gene phylogenies.

Abstract: The phytochemical system of mustard-oil glucosides (glucosinolates) accompanied by the hydrolytic enzyme myrosinase ( b-thioglucosidase), the latter usually compartmented in special myrosin cells, characterizes plants in 16 families of angiosperms. Traditional classifications place these taxa in many separate orders and thus imply multiple convergences in the origin of this chemical defense system. DNA sequencing of the chloroplast rbcL gene for representatives of all 16 families and several putative relatives, with phylogenetic analyses by parsimony and maximum likelihood methods, demonstrated instead a single major clade of mustard-oil plants and one phylogenetic outlier. In a further independent test, DNA sequencing of the nuclear 18S ribosomal RNA gene for all these exemplars has yielded the same result, a major mustard-oil clade of 15 families (Akaniaceae, Bataceae, Brassicaceae, Bretschneideraceae, Capparaceae, Caricaceae, Gyrostemonaceae, Koeberliniaceae, Limnanthaceae, Moringaceae, Pentadiplandraceae, Resedaceae, Salvadoraceae, Tovariaceae, and Tropaeolaceae) and one outlier, the genus Drypetes, traditionally placed in Euphorbiaceae. Concatenating the two gene sequences (for a total of 3254 nucleotides) in a data set for 33 taxa, we obtain robust support for this finding of parallel origins of glucosinolate biosynthesis. From likely cyanogenic ancestors, the ‘‘mustard oil bomb’’ was invented twice.

Molecular systematics of Apiaceae subfamily Apioideae: phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacer and Plastid rpoC1 intron sequences

Abstract: Evolutionary relationships among representatives of Apiaceae (Umbelliferae) subfamily Apioideae have been inferred from phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacer (ITS 1 and ITS 2) and plastid rpoC1 intron sequences. High levels of nucleotide sequence variation preclude the use of the ITS region for examining relationships across subfamilial boundaries in Apiaceae, whereas the rpoC1 intron is more suitably conserved for family-wide phylogenetic study but is too conserved for examining relationships among closely related taxa. In total, 126 ITS sequences from subfamily Apioideae and 100 rpoC1 intron sequences from Apiaceae (all three subfamilies) and outgroups Araliaceae and Pittosporaceae were examined. Phylogenies estimated using parsimony, neighbor-joining, and maximum likelihood methods reveal that: (1) Apiaceae subfamily Apioideae is monophyletic and is sister group to Apiaceae subfamily Saniculoideae; (2) Apiaceae subfamily Hydrocotyloideae is not monophyletic, with some members strongly allied to Araliaceae and others to Apioideae 1 Saniculoideae; and (3) Apiaceae subfamily Apioideae comprises several well-supported subclades, but none of these coincide with previously recognized tribal divisions based largely on morphological and anatomical characters of the fruit. Four major clades in Apioideae are provisionally recognized and provide the framework for future lower level phylogenetic analyses. A putative secondary structure model of the Daucus carota(carrot) rpoC1 group II intron is presented. Of its six major structural domains, domains II and III are the most, and domains V and VI the least, variable.

Cytotype distribution at a diploid–tetraploid contact zone in Chamerion (Epilobium) angustifolium (Onagraceae)

Abstract: In North America, the geographic distributions of diploid and tetraploid Chamerion (formerly Epilobium) angustifolium overlap in a narrow zone along the southern border of the boreal forest and along the Rocky Mountains. We examined the frequency and distribution of diploid and tetraploid cytotypes in a narrow (5 km) zone of sympatry across an elevational gradient and in putatively uniform diploid and tetraploid reference populations on the Beartooth Pass, in the Rocky Mountains of southern Montana-northern Wyoming. All five reference populations sampled were dominated by a single cytotype, but only one was completely uniform. In the zone of sympatry, 27 transects were sampled every 2 m for a total of 238 plants. Reproductive status (vegetative, flower buds, open flowers) was recorded, and the ploidy of each plant was determined by flow cytometry. Diploid and tetraploid plants predominated (36 and 55%, respectively) but were heterogeneously distributed among the transects. Six of the 27 transects were fixed for a single cytotype (four transects, diploid; two transects, tetraploid), and in seven others either diploids or tetraploids predominated (frequency >75%). Triploids represented 9% of the total sample and occurred most frequently in transects containing both diploids and tetraploids (G = 3.4, df = 2, P = 0.07). Diploids were more often reproductive (in bud, flower, or fruit) than either triploids or tetraploids (G = 12.0, df = 2, P < 0.001) and were the only cytotype to have produced open flowers. These results suggest that the zone of sympatry is best characterized as a mosaic rather than a cline, with diploid and tetraploids in close proximity and that the distribution of polyploidy is regulated by ecological sorting in a heterogeneous physical environment.

Phylogenetic relationships of conifers inferred from Partial 28S rRNA gene sequences

Abstract: The conifers, which traditionally comprise seven families, are the largest and most diverse group of living gymnosperms. Efforts to systematize this diversity without a cladistic phylogenetic framework have often resulted in the segregation of certain genera and/or families from the conifers. In order to understand better the relationships between the families, we performed cladistic analyses using a new data set obtained from 28S rRNA gene sequences. These analyses strongly support the monophyly of conifers including Taxaceae. Within the conifers, the Pinaceae are the first to diverge, being the sister group of the rest of conifers. A recently discovered Australian genus Wollemia is confirmed to be a natural member of the Araucariaceae. The Taxaceae are nested within the conifer clade, being the most closely related to the Cephalotaxaceae. The Taxodiaceae and Cupressaceae together form a monophyletic group. Sciadopitys should be considered as constituting a separate family. These relationships are consistent with previous cladistic analyses of morphological and molecular (18S rRNA, rbcL) data. Furthermore, the well-supported clade linking the Araucariaceae and Podocarpaceae, which has not been previously reported, suggests that the common ancestor of these families, both having the greatest diversity in the Southern Hemisphere, inhabited Gondwanaland.

Phylogenetic relationships within Araucariaceae based on rbcL gene sequences.

Abstract: Phylogenetic relationships were determined in the Araucariaceae, which are now found mainly in the Southern Hemisphere. This conifer family was well diversified and widely distributed in both hemispheres during the Mesozoic era. The sequence of 1322 bases of the rbcL gene of cpDNA was determined from 29 species of Araucariaceae, representing almost all the species of the family. Phylogenetic trees determined by the parsimony method indicate that Araucariaceae are well defined by rbcL sequences and also that the monophyly of Agathis or Araucaria is well supported by high bootstrap values. The topology of these trees revealed that Wollemia had derived prior to Agathis and Araucaria. The rbcL phylogeny agrees well with the present recognition of four sections within Araucaria: Araucaria, Bunya, Eutacta, and Intermedia. Morphological characteristics of the number of cotyledons, position of male cone, and cuticular micromorphologies were evaluated as being phylogenetically informative. Section Bunya was found to be derived rather than to be the oldest taxon. Infrageneric relationships of Agathis could not be well elucidated because there are few informative site changes in the rbcL gene, suggesting the more recent differentiation of the species as their fossil records indicate. The New Caledonian Araucaria and Agathis species each formed a monophyletic group with very low differentiation in rbcL sequences among them, indicating rapid adaptive radiation to new edaphic conditions, i.e., ultramafic soils, in the post-Eocene era.

Life history of the long‐lived gynodioecious cushion plant Silene acaulis (Caryophyllaceae), inferred from size‐based population projection matrices

Abstract: Alpine plants often appear to have long life-spans as an adaptation to harsh and unpredictable environmental conditions, yet many lack reliable indicators of age that would make it possible to determine their true longevity Their extended lifespans also pose problems for measuring lifetime reproductive success, a key component of breeding system evolution in species such as the gynodioecious cushion plant Silene acaulis For a population of S acaulis in south-central Alaska, we applied a recently derived analytical approach using size-based population projection matrices that allowed us to estimate: (1) the relationship between cushion diameter and age; and (2) lifetime reproductive success through seed production by females relative to hermaphrodites Because of a combination of slow growth, frequent shrinkage, and extremely high adult survival, we estimate that the largest cushions in our study population exceed 300 yr in age, and some may live substantially longer, despite the seemingly inhospitable alpine environment they inhabit Females are estimated to produce 44 times as many offspring via seed production over the course of their lives as do hermaphrodites, a difference that is more than sufficient to assure the persistence of females despite their inability to transmit genes through pollen These results highlight the utility of size-based projection matrices for studying the life histories of herbaceous perennials whose life-span and lifetime reproductive success cannot be determined easily by any other means Life history theory predicts that perenniality and polycarpic reproduction will be favored where offspring survival is low (Charnov and Schaffer, 1973) or unpredictable (Murphy, 1968), two features that typify arctic and alpine environments While high-altitude and high-latitude plants are often claimed to exhibit a high degree of perenniality (Billings and Mooney, 1968; Bliss, 1971; Walter, 1979), we presently have a poor understanding of the true life-spans of many species, preventing a full quantitative comparison of plant life history strategies in tundra vs other biomes This ignorance reflects the fact that for arctic and alpine plants, as for the dominant plants in many other biomes (eg, grasslands and deserts), it can be exceedingly difficult to obtain direct estimates of plant life-spans Because of their longevity, it is impractical to follow cohorts of tundra plants from birth until death In addition, many characteristic tundra species have herbaceous growth forms, so that standard methods for determining an individual’s age (such as counting annual growth increments as is often done with woody plants) are usually either inappropriate (because many species lack clearly delineated growth increments) or require disturbance or destruction of sampled individ

Specialized phloem parenchyma cells in Norway spruce (Pinaceae) bark are an important site of defense reactions.

Abstract: The bark anatomy of Norway spruce clones that were resistant or susceptible to Ceratocystis polonica, a bark-beetle-vectored fungal pathogen, was compared. The major difference concerned the axial parenchyma cells, called polyphenolic parenchyma (PP cells) because of their vacuolar deposits. The phenolic nature of the deposits was indicated by autofluorescence under blue light, and immunocytochemical studies demonstrating PP cells are enriched in phenylalanine ammonia lyase (EC 4.3.1.5), a key enzyme in phenolic synthesis. Susceptible-clone PP cells occurred as single rows filled with dense deposits. The resistant clone had 40% more PP cells, which occurred in rows two cells thick plus as individual cells scattered among the sieve cells and had lighter deposits. Trees inoculated with fungus were analyzed but a distinct fungal response could not be separated from the general wound response. In the resistant clone, phenolic bodies were reduced in size and density or disappeared completely 12 d after wounding, and PP cell size increased. The susceptible-clone phenolics and cell size changed only slightly. These data show that PP cells are active in synthesis, storage, and modification of phenolics in response to wounding, providing an important site of constitutive and inducible defenses.

Circumscription of the Malvales and relationships to other Rosidae: evidence from rbcL sequence data.

Abstract: The order Malvales remains poorly circumscribed, despite its seemingly indisputable core constituents: Bombacaceae, Malvaceae, Sterculiaceae, and Tiliaceae. We conducted a two-step parsimony analysis on 125 rbcL sequences to clarify the composition of Malvales, to determine the relationships of some controversial families, and to identify the placement of the Malvales within Rosidae. We sampled taxa that have been previously suggested to be within, or close to, Malvales (83 sequences), plus additional rosids (26 sequences) and nonrosid eudicots (16 sequences) to provide a broader framework for the analysis. The resulting trees strongly support the monophyly of the core malvalean families, listed above. In addition, these data serve to identify a broader group of taxa that are closely associated with the core families. This expanded malvalean clade is composed of four major subclades: (1) the core families (Bombacaceae, Malvaceae, Sterculiaceae, Tiliaceae); (2) Bixaceae, Cochlospermaceae, and Sphaerosepalaceae (Rhopalocarpaceae); (3) Thymelaeaceae sensu lato (s.l.); and (4) Cistaceae, Dipterocarpaceae s.l., Sarcolaenaceae (Chlaenaceae), and Muntingia. In addition, Neurada (Neuradaceae or Rosaceae) falls in the expanded malvalean clade but not clearly within any of the four major subclades. This expanded malvalean clade is sister to either the expanded capparalean clade of Rodman et al. or the sapindalean clade of Gadek et al. Members of Elaeocarpaceae, hypothesized by most authors as a sister group to the four core malvalean families, are shown to not fall close to these taxa. Also excluded as members of, or sister groups to, the expanded malvalean clade were the families Aextoxicaceae, Barbeyaceae, Cannabinaceae, Cecropiaceae, Dichapetalaceae, Elaeagnaceae, Euphorbiaceae s.l., Huaceae, Lecythidaceae, Moraceae s.l., Pandaceae, Plagiopteraceae, Rhamnaceae, Scytopetalaceae, Ulmaceae, and Urticaceae. The order Malvales, as traditionally circumscribed, includes four core families, Bombacaceae (;250 spp.), Malvaceae (1500 spp.), Sterculiaceae (1000 spp.), and Tiliaceae (400 spp.), plus from one to eight other families depending upon author (Takhtajan, 1987, 1997; Cronquist, 1988; Dahlgren, 1989; Thorne, 1992). Considering only the core four families, the order comprises predominantly woody and tropical trees, including several economically important genera such as cotton (Gossypium

Cost of reproduction in the pink lady's slipper orchid (Cypripedium acaule, Orchidaceae): an eleven-year experimental study of three populations.

Abstract: An 11-yr experimental study of the cost of reproduction in three wild populations of the perennial orchid Cypripedium acaule contrasted experimental plants that were repeatedly hand-pollinated and often made fruits with control plants that were not hand-pollinated and only rarely made fruits. Repeated flowering without subsequent fruit production resulted in no detectable reduction in either plant size or probability of flowering in subsequent years. A cost of fruit production was evident in experimental plants in all three populations in terms of a reduced probability of flowering and smaller leaf area in subsequent years, but was not evident in terms of mortality rate. Experimental effects of fruit production reached maximum values at 3-7 yr, depending on the population. The probability of remaining dormant below ground in a given year was strongly dependent on plant size in the previous year. Furthermore, the length of the dormancy period (one to several years) was a significant and inverse function of plant size just prior to dormancy. Sample sizes and the consequent ability to detect experimental effects declined over time as more plants died or stopped flowering. Four to seven years appears to be an optimal duration for studies of the cost of reproduction in perennial herbs similar to this species. Studies lasting less than 4 yr may be too brief to reveal experimental effects, whereas those lasting more than 7 yr may fail to reveal new insights.

Fossil Clusiaceae from the late Cretaceous (Turonian) of New Jersey and implications regarding the history of bee pollination.

Abstract: The Turonian flora from Sayreville New Jersey includes one of the world's most diverse assemblages of Cretaceous angiosperm flowers. This flora is made even more interesting by its association with a large insect fauna that is preserved by charcoalification as well as in amber. Floral diversity includes numerous representatives of Magnoliidae, Hamamelididae, Rosidae, Dilleniidae, and Asteridae (Ericales sensu lato). Included are hypogynous, five-merous flowers with uniseriate hairs on the pedicels and stamens in bundles most frequently borne opposite the petals. There is considerable variation in filament length, and some filaments are branched. On some anthers, strands of residue, suggesting the former presence of a liquid of unknown nature, partially occlude the apparent zone of dehiscence. In other cases, open anthers are fully occluded by an amorphous substance. Pollen is rarely found associated with anthers, but is common on stigmatic surfaces. Pollen is prolate and tricolporate with reticulate micromorphology. The superior syncarpous ovary is five-carpellate with axile/intruded parietal placentation and numerous anatropous ovules/carpel. Ovary partitions have closely spaced, parallel ascending channels (secretory canals?), and there are apparent secretory canals/cavities in receptacles, sepals, and petals. Individual stigmas are cuneiform with a central groove and eccentrically peltate. Styles are short and fused. In aggregate, the stigmas form a secondarily peltate stigma. Seeds have a reticulate sculpture pattern, a pronounced raphe, and funicular arils with sculpture similar to the seeds. Phylogenetic analyses of several data matrices of extant taxa place this fossil in a monophyletic group with the modern genera Garcinia and Clusia within the Clusiaceae. As such, these fossils represent the earliest fossil evidence of the family Clusiaceae. Some modern Clusiaceae are notable, in particular, for their close relationship with meliponine and other highly derived bee pollinators; the fossil flowers share several characters that suggest a similar mode of pollination. This possibility is consistent with other floral and insect data from the same locality.

Phylogenetic relationships of Cornaceae and close relatives inferred from matK and rbcL sequences.

Abstract: Phylogenetic relationships were inferred using nucleotide sequences of the chloroplast gene matK for members of Cornales, a well-supported monophyletic group comprising Cornaceae and close relatives. The shortest trees resulting from this analysis were highly concordant with those based on previous phylogenetic analysis of rbcL sequences. Analysis of a combined matK and rbcL sequence data set (a total of 2652 bp [base pairs]) provided greater resolution of relationships and higher internal support for clades compared to the individual data sets. Four major clades (most inclusive monophyletic groups) of Cornales are indicated by both sets of genes: (1) Cornus-Alangium, (2) nyssoids (Nyssa-Davidia-Camptotheca)- mastixioids (Mastixia, Diplopanax), (3) Curtisia, and (4) Hydrangeaceae-Loasaceae. The combined evidence indicates that clades 2 and 3 are sisters, with clade 4 sister to the remainder of Cornales. These relationships are also supported by other lines of evidence, including synapomorphies in fruit and pollen morphology and gynoecial vasculature. Comparisons of matK and rbcL sequences based on one of the most parsimonious rbcL-matK trees indicate that matK has a much higher A-T content (66.9% in matK vs. 55.8% in rbcL) and a lower transition:transversion ratio (1.23 in matK vs. 2.21 in rbcL). The total number of nucleotide substitutions per site for matK is 2.1 times that of rbcL in Cornales. These findings are similar to recent comparisons of matK and rbcL in other dicots. Variable sites of matK are almost evenly distributed among the three codon positions (1.0:1.0:1.3), whereas variable sites of rbcL are mostly at the third position (1.8:1.0 :7.5). Among- lineages rates of nucleotide substitutions in rbcL are basically homogeneous throughout Cornales, but are more heterogeneous in matK.

Ontogenetic changes in size, allometry, and mechanical design of tropical rain forest trees

Abstract: Size, allometry, and mechanical design were measured for trees of three canopy species in a tropical rain forest in French Guiana. Mechanical design was expressed as the safety factor, using the elastic-stability model, and the wind resistance factor, using the constant-stress model. Changes with ontogeny were described as regressions using stem diameter as the independent variable, and they were compared between species. Height, crown size, and the wind resistance factor increased with ontogeny. The safety factor decreased to a minimum and then increased continuously in thicker trees. The crown width/height ratio did not change with ontogeny. Interspecific differences in allometry and mechanical design were related to the adult stature of the species, and not to shade tolerance. The short stature species (Vouacapoua americana) was less slender (height:DBH [stem diameter at 1.3 m] ratio) and had a higher crown width/height ratio than the tall stature species (Goupia glabra and Dicorynia guianensis). Vouacapoua had a higher safety factor, but a similar wind resistance factor. The safety factors of our study species were lower than those of two temperate tree species because of a higher slenderness. Differences in safety factors between tropical and temperate trees may result from unrealistic assumptions of the elastic-stability model, and may also be related to lower light levels and-or wind rates in the tropics.

Fecundity, phenology, and seed dormancy of F1 wild-crop hybrids in Sunflower (Helianthus annuus, Asteraceae).

Abstract: 1 hybrids. In order to assess the ease with which crop-to-wild introgression can proceed, we compared characteristics of F1 wild‐crop progeny with those of purely wild genotypes. Two nontransgenic, cultivated varieties were crossed with wild plants from three different regions— Texas, Kansas, and North Dakota. Seed burial experiments in the region of origin showed that wild‐crop seeds had somewhat higher germination rates (less dormancy) than wild seeds from Kansas and North Dakota, while no differences were seen in seeds from Texas. Progeny from each type of cross were grown in outdoor pots in Ohio and in a weedy field in Kansas to quantify lifetime fecundity and flowering phenology. Flowering periods of hybrid and wild progeny overlapped considerably, especially in plants from North Dakota and Texas, suggesting that these hybrids are very likely to backcross with wild plants. In general, hybrid plants had fewer branches, flower heads, and seeds than wild plants, but in two crosses the fecundity of hybrids was not significantly different from that of purely wild plants. In Ohio, wild‐crop hybrids from North Dakota appeared to be resistant to a rust that infected 53 % of the purely wild progeny, indicating a possible benefit of ‘‘traditional’’ crop genes. In summary, our results suggest that F 1 wild‐crop hybrids had lower fitness than wild genotypes, especially when grown under favorable conditions, but the F1 barrier to the introgression of crop genes is quite permeable.

Effects of soil salt levels on the growth and water use efficiency of Atriplex canescens (Chenopodiaceae) varieties in drying soil

Abstract: The effect of salt stress on the growth and water use efficiency of the xerohalophyte Atriplex canescens (Pursh.) Nutt. in drying soil was determined by growing plants to the wilting point in soils receiving a one-time irrigation of nutrient solution containing low, medium, and high levels of NaCl. The experiment compared three varieties of A. canescens that differed in salt tolerance and capacity for Na and K uptake in previous research. Contrary to expectations, we did not find that water and salt stress were strictly additive in reducing plant performance. Soil salts enhanced the growth performance of the plants in drying soil by increasing their days to wilting, ability to extract water from the soil, organic matter production, and water use efficiency. The variety with the highest salt tolerance also had the highest growth rates and water use efficiency on drying soils. We conclude that tolerances to water and salt stress are linked through a common mechanism of Na uptake for osmotic adjustment in this species.

Independent terrestrial origins of the Halosphaeriales (marine Ascomycota)

Abstract: A phylogenetic study of marine ascomycetes was initiated to test and refine evolutionary hypotheses of marine‐terrestrial transitions among ascomycetes. Taxon sampling focused on the Halosphaeriales, the largest order of marine ascomycetes. Approximately 1050 base pairs (bp) of the gene that codes for the nuclear small subunit (SSU) and 600 bp of the gene that codes for the nuclear large subunit (LSU) ribosomal RNAs (rDNA) were sequenced for 15 halosphaerialean taxa and integrated into a data set of homologous sequences from terrestrial ascomycetes. An initial set of phylogenetic analyses of the SSU rDNA from 38 taxa representing 15 major orders of the phylum Ascomycota confirmed a close phylogenetic relationship of the halosphaerialean species with several other orders of perithecial ascomycetes. A second set of analyses, which involved more intensive taxon sampling of perithecial ascomycetes, was performed using the SSU and LSU rDNA data in combined analyses. These second analyses included 15 halosphaerialean taxa, 26 terrestrial perithecial fungi from eight orders, and five outgroup taxa from the Pezizales. In these analyses the Halosphaeriales were polyphyletic and comprised two distinct lineages. One clade of Halosphaeriales comprised 12 taxa from 11 genera and was most closely related to terrestrial fungi of the Microascales. The second clade of halosphaerialean fungi comprised taxa from the genera Lulworthia and Lindra and was an isolated lineage among the perithecial fungi. Both the main clade of Halosphaeriales and the Lulworthia/Lindra clade are supported by the data as being independently derived from terrestrial ancestors.

Chloroplast DNA characters, phylogeny, and classification of Lathyrus (Fabaceae).

Abstract: Mapped cpDNA restriction site characters were analyzed cladistically and the resulting phylogenetic hypotheses were used to test monophyly and relationships of the infrageneric classification of Lathyrus (Fabaceae) proposed by Kupicha (1983, Notes from the Royal Botanic Garden Edinburgh41: 209‐244). The validity of previously proposed classification systems and questions presented by these classification schemes were explored. Two cpDNA regions, rpo C( rpoC1, its intron, part of rpoC2, and their intergenic spacer) and IR 2 (psbA, trnH-GUG, part of ndhF, and their intergenic spacers), were analyzed for 42 Lathyrus and two Vicia species. PCR (polymerase chain reaction) amplified rpoC and IR 2 products digested with 31 and 27 restriction endonucleases, respectively, resulted in 109 potentially informative characters. The strict consensus tree suggests that several of Kupicha’s sections may be combined in order to constitute clades. The widespread section Orobus and the South American section Notolathyrus should be combined. Section Lathyrus, characterized by a twisted style, should either include sections Orobon and Orobastrum or be redefined as three sections, one of which is characterized by a 100 base pair deletion in the IR 2 region. Finally, a weighted parsimony analysis positions sections Clymenum(excluding L. gloeospermus) and Nissolia, both with phyllodic leaves, as sister sections. The affiliation of Lathyrus gloeospermus (section Clymenum) remains problematic.

Intraspecific variation and phylogeographic patterns of Fagus crenata (Fagaceae) mitochondrial DNA.

Abstract: Mitochondrial (mt) DNA variation in Japanese beech, Fagus crenata (Fagaceae), was studied in 17 populations distributed throughout the species' range. Total genomic DNA of samples from single trees representing each of 12 populations were digested with 18 restriction enzymes and hybridized with three probes containing coxI, coxIII, and atpA gene sequences. Thirty-four of the 54 enzyme/probe combinations showed polymorphisms and all the individuals were subsequently analyzed with six combinations of three probes and two enzymes. Restriction fragment length polymorphisms were evident around all three genes, allowing the identification of eight distinct haplotypes. Haplotype diversity within the populations was found to be very low (HS = 0.031), but population differentiation to be much higher (GST = 0.963). The mtDNA variation was strikingly different from allozyme variation (HS = 0.209; GST = 0.039). Gene flow for maternally inherited mtDNA should be restricted to seed dispersal while nuclear gene flow occurs by both seed and pollen dispersal. Therefore, the difference in the variation between mtDNA and allozymes may be largely a result of the much higher rate of gene flow associated with pollen dispersal than with seed dispersal. The mtDNA variation displayed strong geographic structure, which may reflect the species' distribution in the last glacial maximum and subsequent colonization, and probably also reflects intraspecific phylogeography of the species.

Greater male fitness of a rare invader (Spartina alterniflora, Poaceae) threatens a common native (Spartina foliosa) with hybridization.

Abstract: Hybridization with abundant invaders is a well-known threat to rare native species. Our study addresses mechanisms of hybridization between a rare invader, smooth cordgrass (Spartina alterniflora) and the common native California cordgrass (S. foliosa) in the salt marshes of San Francisco Bay. These species are wind-pollinated and flower in summer. The invader produced 21-fold the viable pollen of the native, and 28% of invader pollen germinated on native stigmas (1.5-fold the rate of the native's own pollen). Invader pollen increased the seed set of native plants almost eightfold over that produced with native pollen, while native pollen failed to increase seed set of the invader. This pollen swamping and superior siring ability by the invader could lead to serial genetic assimilation of a very large native population. Unlike California cordgrass, smooth cordgrass can grow into low intertidal habitats and cover open mud necessary to foraging shorebirds, marine life, navigation, and flood control in channels. To the extent that intertidal range of the hybrids is more similar to the invader than to the native parent, introgression will lead to habitat loss for shore birds and marine life as well to genetic pollution of native California cordgrass.

Seed bank influences on genetic diversity in the rare annual Clarkia springvillensis (Onagraceae).

Abstract: The potential for seed banks to significantly affect the genetic structure of populations was recognized nearly two decades ago. However, there has been little empirical work that examines the problem. In this study, we explore the possibility that the seed bank of a rare annual, Clarkia springvillensis, could act as a buffer against the genetic consequences of small population size. We examined the adult and seed bank cohort in three natural populations. The seed bank was surveyed by collecting soil cores twice during the growing season: postgermination and post seed set. The genetic constitution of the adults and seed bank cohort was determined by examining eight polymorphic isozyme loci via starch gel electrophoresis. The total genetic diversity in the seed bank (Ht = 0.355) was significantly higher than in the adults (Ht = 0.260). Additionally, Fst estimates of genetic differentiation among populations showed significantly less differentiation among population seed banks (Fst = 0.008) than among adults (Fst = 0.045). These results are in agreement with the expectation that seed banks would act to maintain genetic diversity in populations as well as have the effect of slowing differentiation of populations.

Sex ratios, size distributions, and sexual dimorphism in the dioecious tree Ilex aquifolium (Aquifoliaceae).

Abstract: Sex ratio and sexual dimorphism in physiology and growth were studied in the dioecious tree Ilex aquifolium at two localities in northern Spain. Genet sex ratio was significantly male biased in one locality but not in the other. However, ramet and flowering ramet sex ratios were male biased at both study sites. Males had significantly thicker main trunks than females in one locality and produced more ramets in the other. Growth rate, estimated from mean width of annual rings, did not differ between localities, but males produced wider rings than females at both sites. Mean annual growth rates over the last 10, 20, and 30 yr were significantly higher for males. Measurements of chlorophyll fluorescence indicated that the efficiency of photosynthesis of leaves on nonfruiting branches of females was higher than for leaves on branches of male plants under low-light conditions, though not under saturating-light conditions. Efficiency of photosynthesis was significantly lower on fruiting branches of female plants than on nonfruiting branches. We discuss whether the observed between-sex differences are attributable to the higher cost of reproduction in females and/or to pollen competition.