The Aristida pansa complex and a key to the Divaricatae group of North America (Gramineae: Aristideae)

Variation in lateral awn length and pulvini development is described for A. pansa and A. gypsophila. The two species are distinguished by lateral awn development. Within each species, forms are based on pulvini development. Three new forms are described (A. pansa f. contracta; A. gypsophila f. gypsophiloides and f. diffusa), A. dissita is reduced to a form of A. pansa, and A. tehuacanensis is considered synonymous with A. pansa f. dissita. Keys, illustrations, and maps are provided.     

New nomenclatural combinations in the North American perennial Triticeae (Gramineae)

Recent generic realignments of the perennial grasses of the tribe Triticeae necessitate some new name combinations for North American species. Three new species combinations and three new subspecies combinations are made to accommodate six long-anthered and usually cross-pollinating species ofAgropyron sensu A. S. Hitchcock in the genusElytrigia sensu N. N. Tzvelev. Three new species combinations are made to accommodate three long-anthered and usually cross-pollinating species ofElymus sensu A. S. Hitchcock in the genusLeymus sensu N. N. Tzvelev. A complete listing is given of the North American species and subspecies ofElytrigia andLeymus as I perceive them.     

Cytotype variation and allopolyploidy in North American species of the Sphagnum subsecundum complex (Sphagnaceae)

Allopolyploid speciation is likely the predominant mode of sympatric speciation in plants. The Sphagnum subsecundum complex includes six species in North America. Three have haploid gametophytes, and three are thought to have diploid gametophytes. Microsatellite analyses indicated that some plants of S. inundatum and S. lescurii are heterozygous at most loci, but others have only one allele at each locus. Flow cytometry and Feulgen staining showed that heterozygous plants have twice the genome size as plants with one allele per locus; thus, microsatellite patterns can be used to survey the distribution and abundance of haploid and diploid gametophytes. Microsatellite analyses also revealed that S. carolinianum is consistently diploid, but S. lescurii and S. inundatum include both haploid and diploid populations. The frequency of diploid plants in S. lescurii increases with latitude. In an analysis of one population of S. lescurii, both cytotypes co-occurred but were genetically differentiated with no evidence of interbreeding. The degree of genetic differentiation showed that the diploids were not derived from simple genome duplication of the local haploids. Heterozygosity appears to be fixed or nearly so in diploids, strongly suggesting that although morphologically indistinguishable from the haploids, they are derived by allopolyploidy.     

Morphometric variation and the classification of Penniseturn section B revivalvula (Gramineae) in tropical Africa

Morphological data for 15 reproductive characters were compiled for 177 African collections of Penniseturn section Brevivalvula (Gramineae). Principal component analyses of the resulting data set revealed the presence of three phenetically distinct groups which are here treated as species. Although extensive morphological variation exists within two of these species, this variation is essentially continuous and is recognized here at the subspecific level.     

Development of Male Flowers in Zizania aquatica (North American Wild-Rice; Gramineae)

We investigated the histology and developmental morphology of flowers of wild-rice (Zizania aquatica), a member of grass subfamily Oryzoideae, to determine whether male flowers in this species develop in a manner similar to those in the subfamily Panicoideae, a group that includes many species with unisexual flowers. Zizania has evolved unisexual flowers from hermaphrodite ancestors and is only distantly related to the Panicoideae; the origins of unisexual flowers are independent in the two taxa. Ontogenetic evidence indicates that many species within the subfamily Panicoideae develop male flowers by a process similar to that already described for maize (Zea mays), a panicoid grass. Unisexual male flowers in maize initiate both the stamen (androecium) and the pistil (gynoecium), but the gynoecium aborts early in development through the death of the subepidermal cells. Cell death in gynoecia of maize is known to be controlled by the product of the gene tasselseed2 (ts2), and an orthologue of ts2 has been shown to have the same effect in the sister genus Tripsacum. It seems likely that ts2 orthologues mediate cell death throughout the Panicoideae, but the phylogenetic range of the cell death mechanism is not known. In this study we show that male flowers of Z. aquatica show neither the distinctive pattern of cell death nor the ontogenetic timing of abortion that are characteristic of male flower formation in studied species of Panicoideae. This indicates that these unisexual flowers may be produced by an entirely different mechanism from that employed by the Panicoideae. Either ts2 does not control sex expression in Zizania, or it is deployed at a different time, and possibly in different tissues, with a different histological result. Our results indicate that the independent origins of male flowers in Gramineae apparently do not have a common system of genetic control.     

A worldwide phylogenetic classification of the Poaceae (Gramineae)

Based on recent molecular and morphological studies we present a modern worldwide phylogenetic classification of the ± 12074 grasses and place the 771 grass genera into 12 subfamilies (Anomochlooideae, Aristidoideae, Arundinoideae, Bambusoideae, Chloridoideae, Danthonioideae, Micraioideae, Oryzoideae, Panicoideae, Pharoideae, Puelioideae, and Pooideae), 6 supertribes (Andropogonodae, Arundinarodae, Bambusodae, Panicodae, Poodae, Triticodae), 51 tribes (Ampelodesmeae, Andropogoneae, Anomochloeae, Aristideae, Arundinarieae, Arundineae, Arundinelleae, Atractocarpeae, Bambuseae, Brachyelytreae, Brachypodieae, Bromeae, Brylkinieae, Centotheceae, Centropodieae, Chasmanthieae, Cynodonteae, Cyperochloeae, Danthonieae, Diarrheneae, Ehrharteae, Eragrostideae, Eriachneae, Guaduellieae, Gynerieae, Hubbardieae, Isachneae, Littledaleeae, Lygeeae, Meliceae, Micraireae, Molinieae, Nardeae, Olyreae, Oryzeae, Paniceae, Paspaleae, Phaenospermateae, Phareae, Phyllorachideae, Poeae, Steyermarkochloeae, Stipeae, Streptochaeteae, Streptogyneae, Thysanolaeneae, Triraphideae, Tristachyideae, Triticeae, Zeugiteae, and Zoysieae), and 80 subtribes (Aeluropodinae, Agrostidinae, Airinae, Ammochloinae, Andropogoninae, Anthephorinae, Anthistiriinae, Anthoxanthinae, Arthraxoninae, Arthropogoninae, Arthrostylidiinae, Arundinariinae, Aveninae, Bambusinae, Boivinellinae, Boutelouinae, Brizinae, Buergersiochloinae, Calothecinae, Cenchrinae, Chionachninae, Chusqueinae, Coicinae, Coleanthinae, Cotteinae, Cteniinae, Cynosurinae, Dactylidinae, Dichantheliinae, Dimeriinae, Duthieinae, Eleusininae, Eragrostidinae, Farragininae, Germainiinae, Gouiniinae, Guaduinae, Gymnopogoninae, Hickeliinae, Hilariinae, Holcinae, Hordeinae, Ischaeminae, Loliinae, Melinidinae, Melocanninae, Miliinae, Monanthochloinae, Muhlenbergiinae, Neurachninae, Olyrinae, Orcuttiinae, Oryzinae, Otachyriinae, Panicinae, Pappophorinae, Parapholiinae, Parianinae, Paspalinae, Perotidinae, Phalaridinae, Poinae, Racemobambosinae, Rottboelliinae, Saccharinae, Scleropogoninae, Scolochloinae, Sesleriinae, Sorghinae, Sporobolinae, Torreyochloinae, Traginae, Trichoneurinae, Triodiinae, Tripogoninae, Tripsacinae, Triticinae, Unioliinae, Zizaniinae, and Zoysiinae). In addition, we include a radial tree illustrating the hierarchical relationships among the subtribes, tribes, and subfamilies. We use the subfamilial name, Oryzoideae, over Ehrhartoideae because the latter was initially published as a misplaced rank, and we circumscribe Molinieae to include 13 Arundinoideae genera. The subtribe Calothecinae is newly described and the tribe Littledaleeae is new at that rank.     

Fecundity and egg size variation in North American Pacific salmon (Oncorhynchus)

We examined regional and latitudinal variation in fecundity and egg weight for five species of Pacific salmon ( Oncorhynchus ) along the Pacific coast of North America. Data were examined for 24 chum salmon, 15 pink salmon, 34 sockeye salmon, 44 chinook salmon, and 40 coho salmon populations from published sources, unpublished Canadian hatchery records, our own laboratory investigations, and other unpublished sources. Substantial regional variation in fecundity and egg weight was observed, with salmon on the Queen Charlotte Islands and Vancouver Island in British Columbia generally having lower fecundity and larger egg size than nearby mainland populations. The relative distance of freshwater migration to the spawning grounds generally had a marked effect on both fecundity and egg size, with populations spawning in the upper portions in the drainages of large rivers like the Fraser River in British Columbia having reduced fecundity and egg size compared with coastal spawning populations. Fecundity was generally higher and egg size generally lower in more northern populations of sockeye, chinook, and coho salmon compared with southern ones. We suggest that egg size tends to be lower in northern populations of some species as a result of increased fecundity due to their older ages at maturity and a limited amount of energy that can be expended on egg production.     

Genetic variation and reproductive system among North American species of Nuttallanthus (Plantaginaceae)

We examined the effect of reproductive and life history strategies on the amount and partitioning of genetic variation in three annual species of Nuttallanthus. The North American species N. canadensis, N. floridanus, and N. texanus have regional to widespread ranges that overlap in the southeastern USA, are characterized by homogeneous populations and high fecundity, and possess showy, fragrant flowers seemingly adapted for insect pollination and outbreeding. Field and greenhouse studies on plants from 25 populations indicated that reproductive strategies were similar among species and showed predominant self-fertilization via cleistogamy and self-pollination prior to anthesis in chasmogamous flowers. Species were reproductively isolated and demonstrated complete cross-incompatibility after experimental crosses and no evidence for hybridization in mixed populations. Genetic variation was assessed using starch gel electrophoresis to resolve 15 isozyme loci in 50 populations. Conspecific genetic identity (I) values were high (0.819-0.936), but interspecific comparisons indicated many qualitative allelic differences and correspondingly low I values (0.516-0.623). Low levels of polymorphism and observed heterozygosity within populations and the disproportionate amount of gene diversity distributed among populations were concordant with reproductive data. The pattern of genetic differentiation was most similar to that observed in species with a predominantly inbreeding mating system.     

Habitat shift and geographical variation in North American tanagers (Thraupinae: Piranga)

Summary Relationships of habitat structure among North American tanagers were studied in large areas of their breeding ranges in the USA. In the large areas sampled, the four species differ significantly in multivariate scores of habitat structure. In Scarlet and Summer Tanagers (for which the most data was collected) there is intraspecific geographical variation in habitat features. Variation in habitats of Summer Tanagers is more pronounced than that of Scarlet Tanagers, and is related to changes of macrohabitats in low elevations from oak-pine and mixed woods in the southeast to riparian vegetation in the dry southwest. In the southeast these two species often breed in the same localities, and establish territories next to each other. In six localities in which I found both of them, the difference between the two species in habitat structure is significantly greater than in allopatric areas in the southeast. These results, and those of another study on song variation and behavioral responses to song suggest that coexistence of Scarlet and Summer Tanagers in sympatry can be explained both by partial habitat shift, and by interspecific aggressive division of space. It is concluded that quantitative sampling of habitats in large areas of distributional ranges of species is needed before any conclusions on species-specific preference and interspecific competition can be made.     

A worldwide phylogenetic classification of the Poaceae (Gramineae) III: An update

We present an updated worldwide phylogenetic classification of Poaceae with 11 783 species in 12 subfamilies, 7 supertribes, 54 tribes, 5 super subtribes, 109 subtribes, and 789 accepted genera. The subfamilies (in descending order based on the number of species) are Pooideae with 4126 species in 219 genera, 15 tribes, and 34 subtribes; Panicoideae with 3325 species in 242 genera, 14 tribes, and 24 subtribes; Bambusoideae with 1698 species in 136 genera, 3 tribes, and 19 subtribes; Chloridoideae with 1603 species in 121 genera, 5 tribes, and 30 subtribes; Aristidoideae with 367 species in three generaand one tribe; Danthonioideae with 292 species in 19 generaand 1 tribe; Micrairoideae with 192 species in nine generaand three tribes; Oryzoideae with 117 species in 19 genera, 4 tribes, and 2 subtribes; Arundinoideae with 36 species in 14 genera and 3 tribes; Pharoideae with 12 species in three generaand one tribe; Puelioideae with 11 species in two generaand two tribes; and the Anomochlooideae with four species in two generaand two tribes. Two new tribes and 22 new or resurrected subtribes are recognized. Forty-five new (28) and resurrected (17) genera are accepted, and 24 previously accepted genera are placed in synonymy. We also provide an updated list of all accepted genera including common synonyms, genus authors, number of species in each accepted genus, and subfamily affiliation. We propose Locajonoa, a new name and rank with a new combination, L. coerulescens. The following seven new combinations are made in Lorenzochloa: L. bomanii, L. henrardiana, L. mucronata, L. obtusa, L. orurensis, L. rigidiseta, and L. venusta.     

A biosystematic study of the Panicum amarum-P. Amarulum complex (Gramineae)

Panicum amarum andP. amarulum exhibit north-south clinal variation and are virtually indistinguishable in the southern part of their range. Cytological evidence indicates thatP. amarum (2n = 54) is probably an autohexaploid derivative ofP. amarulum (2n = 36). The minor morphological differences, the apparent close genetic relationship, and the close ecological and chemical affinities between these two taxa indicate thatP. amarum andP. amarulum should have varietal rather than specific rank.     

A new species of Agrostis (Gramineae) in the A. alpina complex

A new species of Agrostis L. (A. barceloi ) is described from the northern mountains of Mallorca (Balearic Islands). The new taxon belongs to sect. Agtush and is mainly related to A. alpina Scop. and A. schleicheri Jord. & Verl. on morphological grounds. However, A. barceloi differs from A. alpina by its narrowly-lanceolate inflorescence, with panicle branches erect during and after anthesis, non-scabrid leaves, and by smaller lemmas and others. The new species is distinguished from A. schleicheri by the smaller spikelets, lemma, palea, and anthers. In addition, A. barceloi is tetraploid (2n = 28), and differs cytologically from the diploid A. alpina (2n= 14) and the hexaploid A. schleicheri (2n = 56).     

Genetic assessment of species boundaries in the North American Daphnia longispina complex (Crustacea: Daphniidae)

Species boundaries in the North American Daphnia longispina group have proved difficult to establish on the basis of morphology alone. This confusion may be due to hydridization, phenotypic plasticity or the existence of sibling species. We therefore used genetic analysis to delineate species boundaries by examining 27 North American populations belonging to the longispina complex for variation at 15–26 allozyme loci. The populations consisted of Daphnia thorata from two western sites and two eastern sites, Daphnia galeata mendotae from its type location and seven sites across its range, and Daphnia rosea from eight temperate and seven arctic sites. Two populations from the Eurasian longispina complex were also included for reference. Populations assigned to D. galeata mendotae formed a genetically cohesive group, whereas a genetic dichotomy was found between temperate and arctic D. rosea , suggesting that this taxon includes two species. Genetic analysis also confirmed the distinctness of western D. thorata from other members of the longispina group. Unexpectedly, eastern populations resembling D. thorata were genetically more similar to temperate D. rosea than to any helmeted species ( D. galeata, Daphnia hyalina or D. thorata ). Our results suggest that the helmet character is a poor indicator of phylogenetic relationships, as the genetic ability to produce this feature has been lost or acquired several times in the evolution of the longispina group.     

Molecular Phylogeny,Diversity, and Bioprospecting of Endophytic Fungi Associated with wild Ethnomedicinal North American Plant Echinacea purpurea (Asteraceae)

The endophytic fungal community associated with the ethnomedicinal plant Echinacea purpurea was investigated as well as its potential for providing antifungal compounds against plant pathogenic fungi. A total of 233 endophytic fungal isolates were obtained and classified into 42 different taxa of 16 genera, of which Alternaria alternata, Colletotrichum dematium, and Stagonosporopsis sp. 2 are the most frequent colonizers. The extracts of 29 endophytic fungi displayed activities against important phytopathogenic fungi. Eight antifungal extracts were selected for chemical analysis. Forty fatty acids were identified by gas chromatography‐flame‐ionization detection (GC‐FID) analysis. The compounds (–)‐5‐methylmellein and (–)‐(3R)‐8‐hydroxy‐6‐methoxy‐3,5‐dimethyl‐3,4‐dihydroisocoumarin were isolated from Biscogniauxia mediterraneaEPU38CA crude extract. (–)‐5‐Methylmellein showed weak activity against Phomopsis obscurans, P. viticola, and Fusarium oxysporum, and caused growth stimulation of C. fragariae, C. acutatum, C. gloeosporioides, and Botrytis cinerea. (–)‐(3R)‐8‐Hydroxy‐6‐methoxy‐3,5‐dimethyl‐3,4‐dihydroisocoumarin appeared slightly more active in the microtiter environment than 5‐methylmellein. Our results indicate that E. purpurea lives symbiotically with different endophytic fungi, which are able to produce bioactive fatty acids and aromatic compounds active against important phytopathogenic fungi. The detection of the different fatty acids and aromatic compounds produced by the endophytic community associated with wild E. purpurea suggests that it may have intrinsic mutualistic resistance against phytopathogen attacks in its natural environment.     

No evidence for niche segregation in a North American Cattail (Typha) species complex

Interspecific hybridization can lead to a breakdown of species boundaries, and is of particular concern in cases in which one of the parental species is invasive. Cattails (Typha spp.) have increased their abundance in the Great Lakes region of North America over the past 150 years. This increase in the distribution of cattails is associated with hybridization between broad-leaved (Typha latifolia) and narrow-leaved cattails (T. angustifolia). The resulting hybrids occur predominantly as F(1)s, which are known as T. × glauca, although later-generation hybrids have also been documented. It has been proposed that in sympatric populations, the parental species and hybrids are often spatially segregated according to growth in contrasting water depths, and that this should promote the maintenance of parental species. In this study, we tested the hypothesis that the two species and their hybrids segregate along a water-depth gradient at sites where they are sympatric. We identified the two parental species and their hybrids using molecular genetic markers (SSR), and measured shoot elevations (a proxy for water depth) at 18 sites in Southern Ontario, Canada. We found no evidence for niche segregation among species based on elevation. Our data indicate that all three lineages compete for similar habitat where they co-occur suggesting that there is potential for an overall loss of biodiversity in the species complex, particularly if the hybrid lineage is more vigorous compared to the parental species, as has been suggested by other authors.     

Pollen morphology of the Chloridoideae (Gramineae)

Pollen grains of 57 species (representing 42 genera) of the Chloridoideae have been investigated using light, scanning and transmission electron microscopy. Two aperture patterns and nine exine patterns are distinguished using SEM and TEM. These are categorized into five pollen types. Two pollen types are previously not recorded in Gramineae. A key for the identification of these pollen types is presented. Generally, pollen characters have limited systematic value in recognizing taxa at generic level or above in the Chloridoideae. An evolutionary trend is proposed that awaits verification by further systematic study. Pollen characters can be used as indicators for the areas of origin and distribution of the Chloridoideae.     

Patterns of range-wide genetic variation in six North American bumble bee (Apidae: Bombus) species

The increasing evidence for population declines in bumble bee (Bombus) species worldwide has accelerated research efforts to explain losses in these important pollinators. In North America, a number of once widespread Bombus species have suffered serious reductions in range and abundance, although other species remain healthy. To examine whether declining and stable species exhibit different levels of genetic diversity or population fragmentation, we used microsatellite markers to genotype populations sampled across the geographic distributions of two declining (Bombus occidentalis and Bombus pensylvanicus) and four stable (Bombus bifarius; Bombus vosnesenskii; Bombus impatiens and Bombus bimaculatus) Bombus species. Populations of declining species generally have reduced levels of genetic diversity throughout their range compared to codistributed stable species. Genetic diversity can be affected by overall range size and degree of isolation of local populations, potentially confounding comparisons among species in some cases. We find no evidence for consistent differences in gene flow among stable and declining species, with all species exhibiting weak genetic differentiation over large distances (e.g. >1000 km). Populations on islands and at high elevations experience relatively strong genetic drift, suggesting that some conditions lead to genetic isolation in otherwise weakly differentiated species. B. occidentalis and B. bifarius exhibit stronger genetic differentiation than the other species, indicating greater phylogeographic structure consistent with their broader geographic distributions across topographically complex regions of western North America. Screening genetic diversity in North American Bombus should prove useful for identifying species that warrant monitoring, and developing management strategies that promote high levels of gene flow will be a key component in efforts to maintain healthy populations.     

Competition effects on wiregrass (Aristida beyrichana) growth and survival

We investigated neighboring plant effects (competition and facilitation)on wiregrass establishment in two frequently encountered restoration situationsof former longleaf pine-wiregrass habitats in the Southeastern United States:longleaf pine plantations and previously cultivated fields. In the plantationexperiment, we specifically examined canopy removal, neighboring wiregrassdensity, and aboveground and belowground effects on establishment and growth ofwiregrass seedlings at two different ages (3 weeks and 6 months) with 3competitive exclusion treatments (aboveground exclusion, belowground exclusion,or no exclusion). Competition treatment effects were age-dependent forsurvivorship and growth. Survival of 6 month old seedlings was unaffected bycompetition treatment; whereas, three week old seedling survival was greatestwhere roots were excluded. Seedling size increased with root exclusion for 6month seedlings, but not for 3 week old seedlings in plots lacking neighboringwiregrass. Where wiregrass was present, both 3 week and 6 month old seedlingsincreased growth with root exclusion. Furthermore, where neighboring wiregrassplants were absent, increasing canopy density resulted in decreased seedlingsize, but did not affect survivorship. In old fields, fertilizer treatments andweeding effects were also assessed using 3 week and 6 month old seedlings.Fertilizer application did not benefit seedling survival or growth and reducedsurvivorship of 3 week old seedlings. Seedlings were smaller where neighboringold field, weedy vegetation was present regardless of wiregrass seedling age;whereas, survival was dependent on seedling age at time of planting. Six monthold seedling survivorship remained high regardless of weeding treatment. It isstill unclear whether fertilizer application is beneficial to the successfulestablishment of wiregrass.