Effects of virus infection on demographic traits of an agamospermous population of Eupatorium chinense (Asteraceae)

There are few studies of the interaction between wild plants and viruses. In this paper, the incidence of a geminivirus (tobacco leaf curl virus, TLCV) infection, and its effects on mortality, growth and reproduction of its host-plant, Eupatorium chinense, are reported. A total of 221 plants of an agamospermous population of E. chinense were chosen and their demographic behaviour followed over 2 years (1991–1992). The proportion of infected plants differed between years, with fewer plants infected in 1991 than in 1992. Under low virus incidence (35.3% in 1991), infection was significantly associated with taller plants (>80 cm). However, when the incidence of infected plants increased by almost two times (69.1%) in 1992, this tendency disappeared and small plants were also infected. Virus infection had significant effects on mortality of agamospermous plants. Almost half of the initial number of marked plants (n=221) died after 1 year of observations. Of those dead plants (n=105), 86 plants (82%) were infected in 1991, indicating that virus infection was an important, but not the sole cause of mortality. In 1992, 116 plants were alive, and of these, 40% were infected in 1991, indicating that some infected plants survived 1 year. Agamospermous plants were classified in three groups according to the extent of virus infection (plants infected in 2 years, infected in 1 year and uninfected plants) to detect the effect of virus infection on growth of plants of E. chinense. Infected plants had significantly lower growth rates than healthy plants. Infected plants also produced significantly fewer seeds than uninfected plants. Virus infection, however, had no significant effect on the probability of reproduction in plants of E. chinense, suggesting that infected plants may reproduce but with a lower seed output. In this study, we showed that virus infection may have a strong effect on demographic traits and, as a consequence, on fitness components of plants of E. chinense. These effects were higher than those sometimes observed in other plant-herbivore or plant-pathogen interactions.     

Genotypic variation in agamospermous Erigeron compositus (Asteraceae)

Few integrative analyses of the structure of agamospermous plant populations have been conducted. Erigeron compositus occurs in montane western North America and comprises both sexual and agamospermous populations. Sexual E. compositus has previously been characterized as outcrossing and predominantly diploid (2n = 18). Agamic E. compositus is usually hexaploid (2n = 54), though counts herein range from 2n = 36 to 2n = 80. Starch-gel electrophoresis, cytology, and analysis of pollen production were used to evaluate variation within and among agamospermous populations. Fifteen enzyme loci were used to identify 24 unique multilocus genotypes in seven populations, an average of 3.4 genotypes per population. Proportion of distinct genotypes per population sample size (GIN) and measures of genetic diversity (D) and evenness (E) are 0.10, 0.48, and 0.61, respectively, which indicate that E. compositus maintains levels of diversity similar to other agamospermous taxa. Most agamospermous populations are mosaics comprising groups of genetically distinct individuals that are frequently distinguished by cytotype and capacity for pollen production. The geographical and ecological separation of sexual and agamospermous populations make it unlikely that gene flow from sexual populations is a direct source of genetic variation in agamospermous populations. Instead, crossing between genetically distinct facultative agamosperms probably accounts for most variation. Genetic and morphological evidence document one such putative crossing event. Agamospermous E. compositus is very similar genetically to sexual E. compositus. Allozyme analysis further shows that genetic variation in agamospermous populations is partitioned among a few highly heterozygous genotypes, whereas sexual populations maintain numerous genotypes of relatively low heterozygosity.     

DNA evidence for multiple origins of intergeneric allopolyploids in annualMicroseris (Asteraceae)

Seventy populations of North American annualMicroseris, Stebbinsoseris, andUropappus species were examined for chloroplast and nuclear ribosomal DNA restriction site variability to determine the origin of the allotetraploid speciesS. heterocarpa andS. decipiens. Previously identified chloroplast DNA restriction site variants were used in concert with restriction site variation forNco I in the nuclear-encoded ribosomal DNA repeat. The presence of two, mutually exclusive restriction site gains were observed in diploid populations ofM. douglasii; these same variants were also found in populations of allotetraploidS. heterocarpa, indicating mutiple origins of this species from different maternal diploid populations ofM. douglasii. Variation in the rDNA repeat between the diploid annual species and the putative paternal genome ofU. lindleyi was found to be additive inS. heterocarpa. A similar relationship was observed for the origin ofS. decipiens; cpDNA restriction site variants found inM. bigelovii andM. douglasii were present inS. decipiens. The rDNANco I variants also were additive in this purported allotetraploid. These results confirm the reticulate evolutionary pattern inStebbinsoseris and provide another example of multiple origins of intergeneric allopolyploids.     

Pollen quantity and quality as explanations for low seed set in small populations exemplified by Eupatorium (Asteraceae)

As plant populations decrease in size, reduced seed set may contribute to their ultimate extirpation. In this study, effects of pollen quantity and compatibility relationships (quality) on seed set were investigated in a rare species (Eupatorium resinosum) and a closely related common species (E. perfoliatum). The impact of pollen quantity was studied through pollen supplementation experiments in two populations of each species. Addition of pollen increased seed set only in the smaller population of E. resinosum. Compatibility relationships (pollen quality) were investigated in a diallel crossing experiment using ten genotypes from the same populations. Plants from the smaller population of E. resinosum were found to be 40% cross-incompatible, which was higher than the larger population of E. resinosum and the two populations of E. perfoliatum, the latter showing signs of self-compatibility in some individuals. In addition the variance in number of compatible matings per individual was higher in the smaller population of E. resinosum. These results are consistent with a computer simulation model that investigated the effect of small population size on S-allele diversity. Sufficient pollination accompanied by a partial breakdown of the incompatibility system may account, in part, for the relative success of E. perfoliatum.     

Evidence for multiple introductions of Centaurea stoebe micranthos (spotted knapweed,Asteraceae) to North America

Invasive species’ success may depend strongly on the genetic resources they maintain through the invasion process. We ask how many introductions have occurred in the North American weed Centaurea stoebe micranthos (Asteraceae), and explore whether genetic diversity and population structure have changed as a result of introduction. We surveyed individuals from 15 European native range sites and 11 North American introduced range sites at six polymorphic microsatellite loci. No significant difference existed in the total number of alleles or in the number of private alleles found in each range. Shannon–Weaver diversity of phenotype frequencies was also not significantly different between the ranges, while expected heterozygosity was significantly higher in the invasive range. Population structure was similar between the native range and the invasive range, and isolation by distance was not significant in either range. Traditional assignment methods did not allocate any North American individuals to the sampled European populations, while Bayesian assignment methods grouped individuals into nine genetic clusters, with three of them shared between North America and Europe. Invasive individuals tended to have genetically admixed profiles, while natives tended to assign more strongly to a single cluster. Many North American individuals share assignment with Romania and Bulgaria, suggesting two separate invasions that have undergone gene flow in North America. Samples from three other invasive range sites were genetically distinct, possibly representing three other unique introductions. Multiple introductions and the maintenance of high genetic diversity through the introduction process may be partially responsible for the invasive success of C. stoebe micranthos.     

A new species of Eupatorium (Asteraceae) from the Philippines

Eupatorium leonardii, presently known only from Mt. Mingan in the border region between Aurora and Nueva Ecija Provinces in Luzon, Philippines, is described and illustrated. The new species is remarkable because it is the only woody representative of the genus with a peculiar spiral phyllotaxy. Previously, Eupatorium s. str. consisted of herbs with opposite or verticillate phyllotaxy.     

Two new species of Eupatorium (Asteraceae) from northeastern Mexico

Two closely related new species ofEupatorium,E. tamaulipanum from southeastern Tamaulipas andE. xalapanum from central Veracruz, are described herein. The latter is diploid with 2n=20. Their relationship appears to be with the genusEupatoriastrum (subtribe Critoniinae, sensu Robinson and King), but an equally good case might be made for inclusion in the generaChromolaena, Bartlettina, orTamaulipa of subtribes Praxelinae, Hebecliniinae, and Gyptidinae respectively, as circumscribed by King and Robinson. While their splintered treatment ofEupatorium, and the Eupatorieae generally, often provides unequivocal “generic” placement of known or newly described species, many others appear to be rather arbitrarily placed, including some whose possible generic positions might include three or more of their subtribal groupings.     

Molecular confirmation of the hybrid origin of Eupatorium godfreyanum (Asteraceae)

Analysis of nuclear ribosomal ITS sequence data was used to assess the relationships of Eupatorium godfreyanum, an agamospermous polyploid species of putative hybrid origin. A data set of ITS sequences that included representatives of all but two of the North American species of Eupatorium was compiled from a combination of previously published and newly obtained results. Assessment of the data showed that each species was relatively distinctive, although the results from parsimony analysis suggested that there was little phylogenetic structure within the data beyond a basal split between members of the dog fennel group ("Traganthes") and the remainder of the genus. Cloning was required to obtain readable ITS sequence from E. godfreyanum, and analysis of individual clones produced sequences that matched closely those of either E. rotundifolium or E. sessilifolium. The ITS sequence data thus supported the hypothesis that Eupatorium godfreyanum is of hybrid origin from a combination of E. rotundifolium and E. sessilifolium.     

入侵植物紫茎泽兰化感作用及其途径研究

紫茎泽兰广泛入侵中国西南地区,研究结果表明化感作用是其入侵的重要武器,但其化感作用的途径并不十分清楚。本研究中,我们发现紫茎泽兰可以通过多种途径对两种栽培植物大麦和玉米的生长产生化感作用,这些途径包括了叶挥发物、叶淋溶物以及根分泌物。并且在紫茎泽兰幼苗早期就可以检测到这些化感作用。然而,没有实验证据表明紫茎泽兰落叶的微生物降解物对两种测试植物具有化感作用。     

入侵植物紫茎泽兰化感作用及其途径研究

紫茎泽兰广泛入侵中国西南地区,研究结果表明化感作用是其入侵的重要武器,但其化感作用的途径并不十分清楚。本研究中,我们发现紫茎泽兰可以通过多种途径对两种栽培植物大麦和玉米的生长产生化感作用,这些途径包括了叶挥发物、叶淋溶物以及根分泌物。并且在紫茎泽兰幼苗早期就可以检测到这些化感作用。然而,没有实验证据表明紫茎泽兰落叶的微生物降解物对两种测试植物具有化感作用。     

Genetic,cytogenetic and morphological diversity in Helicrysum leucocephalum (Asteraceae) populations

Helichrysum (family Asteraceae) is a large, heterogeneous and polyphyletic genus. Most of the Helichrysum species have ornamental and medicinal values. Helicrysum leucocephalum Boiss., has wide geographical distribution in Iran and forms several local populations. The present investigation was performed to study inter populations, genetic, cytogenetic and morphological diversity of this medicinal species. The AMOVA test showed significant genetic difference among the studied populations but the Mantel test did not show correlation between the genetic distance and geographical distance of these populations. STRUCTURE and reticulation analyses showed some degree of genetic admixture and gene exchange among the studied populations. These populations had different chromosome numbers and also differed significantly in the size of their chromosomes.     

Phytochemical variation within populations of Echinacea angustifolia (Asteraceae)

Quantitative evaluation of phytochemical diversity in Echinacea angustifolia DC. populations from different natural geographic areas supports the existence of distinct natural chemotypes within the species. Consumers, growers and manufacturers of phytomedicines are interested in chemotype identification for prediction of phytochemical content in cultivar development. Six month old E. angustifolia roots, grown from nine different wild seed sources in a controlled environment, were extracted into 70% ethanol and 28 reported phytochemicals were measured by HPLC separation. Two-way ANOVA between the nine populations revealed quantitative differences (p<0.05) in the caffeic acid derivatives 2,3-O-dicaffeoyl tartaric acid (cichoric acid), 2-O-caffeoyl tartaric acid (caftaric acid), 1,3-dicaffeoyl-quinic acid (cynarin), echinacoside and ten reported alkamides. Canonical discriminant analysis determined the phytochemical variables which contributed the most towards chemotype distinction for five of the nine populations: undeca-2E,4Z-diene-8,10-diynoic acid-2-methylbutylamide¹, dodeca-2E,4E-dienoic acid isobutylamide¹, dodeca-2E-ene-8,10-diynoic acid isobutylamide⁷, hexadeca-2E,9Z-diene-12,14-diynoic acid isobutylamide¹, cichoric acid⁷, caftaric acid¹, and echinacoside⁷ (¹p<0.0001, ⁷p<0.05). Five of those compounds were also significantly associated with latitudinal variation by regression analyses (p<0.05).     

Evidence for autoploid evolution in the artemisia ludoviciana complex of the Pacific Northwest

TheArtemisia ludoviciana complex of the Northwest is considered to be an intervarietal autoploid complex on the basis of evidence obtained from cytogenetic analysis. The evidence includes the occurrence of chromosomal races within all but two of the inclusive taxa, the degree and constancy of multivalent formation in the polyploid races, and the high degree of homology among the genomes of the various taxa as demonstrated by the pairing relationships in the F₁ progeny. Both triploid and tetraploid progeny were produced in diploid-tetraploid crosses, and the tetraploid offspring were fully as fertile as the natural tetraploids. The triploids, on the other hand, produced very few viable pollen grains. The production of tetraploid offspring in interracial crosses could provide a mechanism for gene flow from the diploid to the tetraploid population. With the observation of both diploid and tetraploid populations ofA. douglasiana, in addition to the well-known hexaploid, a reasonable doubt is cast upon the putative amphidiploid origin of the hexaploid via hybridization betweenA. suksdorfii andA. ludoviciana.     

Phenotypic differentiation of peripheral populations of Santolina rosmarinifolia (Asteraceae)

Phenotypic differentiation of two tetraploid (2n = 4x = 36, 36+1B, 36+2B) populations of Santolina rosmarinifolia geographically isolated from diploid populations was investigated. The karyotype was relatively homogeneous, meiosis was regular and pollen was fertile in both cytotypes. An autopolyploid or allopolyploid origin for tetraploid cytotypes is discussed. Overall, 80.82% of all variance in achene weight, time t₀, t₅₀ and t₉₀ of germination and accumulated germination rate was due to achene age at each ploidy level. Partition of the total phenotypic variance showed that there was extensive variation between ploidy levels. The mean of morphological characters was generally higher in polyploids. For diploid cytotypes, flower number, achene production and fruiting percentage were significantly higher than for tetraploid cytotypes. Cluster analysis indicated that the patterns of seedling morphology and development were similar in three diploid individuals and several tetraploids; the same analysis showed high similarity between diploid individuals of the natural populations, whereas tetraploid individuals showed high dissimilarity among themselves and with diploid individuals. Multiple correspondence analysis and logistic regression analysis indicated that qualitative characters contribute strongly to cytotype differentiation. The results support recognition of the tetraploid cytotypes at the subspecies level. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 650–668.     

Deep sequencing of amplicons reveals widespread intraspecific hybridization and multiple origins of polyploidy in big sagebrush (Artemisia tridentata; Asteraceae)

• Premise of the study: Hybridization has played an important role in the evolution and ecological adaptation of diploid and polyploid plants. Artemisia tridentata (Asteraceae) tetraploids are extremely widespread and of great ecological importance. These tetraploids are often taxonomically identified as A. tridentata subsp. wyomingensis or as autotetraploids of diploid subspecies tridentata and vaseyana. Few details are available as to how these tetraploids are formed or how they are related to diploid subspecies. • Methods: We used amplicon sequencing to assess phylogenetic relationships among three recognized subspecies: tridentata, vaseyana, and wyomingensis. DNA sequence data from putative genes were pyrosequenced and assembled from 329 samples. Nucleotide diversity and putative haplotypes were estimated from the high-read coverage. Phylogenies were constructed from Bayesian coalescence and neighbor-net network analyses. • Key results: Analyses support distinct diploid subspecies of tridentata and vaseyana in spite of known hybridization in ecotones. Nucleotide diversity estimates of populations compared to the total diversity indicate the relationships are predominately driven by a small proportion of the amplicons. Tetraploids, including subspecies wyomingensis, are polyphyletic occurring within and between diploid subspecies groups. • Conclusions: Artemisia tridentata is a species comprising phylogenetically distinct diploid progenitors and a tetraploid complex with varying degrees of phylogenetic and morphological affinities to the diploid subspecies. These analyses suggest tetraploids are formed locally or regionally from diploid tridentata and vaseyana populations via autotetraploidy, followed by introgression between tetraploid groups. Understanding the phylogenetic vs. ecological relationships of A. tridentata subspecies will have bearing on how to restore these desert ecosystems.     

Effects of virus infection and light environment on population dynamics of Eupatorium makinoi (Asteraceae)

We studied the effects of virus infection on dynamics of three Eupatorium makinoi populations in contrasting light environments, Gora-dani (a shaded population) and Minou 1 and Minou 2 (open-site populations). Censuses of the plants were taken for 8 yr in Gora-dani and 4 yr in Minou 1 and Minou 2. After the epidemics of virus infection, most plants were virus infected at both sites. The number of plants and the proportion of flowering individuals decreased rapidly and simultaneously in the shaded population in Gora-dani. By contrast, in the open-site populations of Minou, the proportion of flowering plants decreased first, and then the number of plants decreased gradually. Growth analysis of the plants in the Gora-dani population revealed that stem growth was significantly suppressed by infection and that flowering and survivorship of the infected plants decreased with reducing plant height. Since light availability affected plant growth and thereby flowering and survivorship, the differences in population dynamics between the two field sites could be caused by the differences in light environments. Although populations in open sites may persist for considerable periods after virus epidemics, the individual local populations of E. makinoi would eventually become extinct irrespective of light environments.     

Phylogenetics of Darwiniothamnus (Asteraceae: Astereae) – molecular evidence for multiple origins in the endemic flora of the Galápagos Islands

Aim The aims of this study were (1) to investigate whether the two growth forms of Darwiniothamnus Harling (Asteraceae) originated from the colonization of a single ancestor, (2) to identify the closest relative(s) of Darwiniothamnus, and (3) to review molecular phylogenies from other plant groups to infer the origin of Galápagos endemics. Location Darwiniothamnus is endemic to the Galápagos Islands. Methods All putative relatives of Darwiniothamnus plus 38 additional species were included. Nucleotide sequences of the internal transcribed spacers of the nuclear ribosomal DNA were used for Bayesian and parsimony analyses. Results Darwiniothamnus is polyphyletic. Two species (D. lancifolius (Hook. f.) Harling and D. tenuifolius (Hook. f.) Harling) are woody shrubs that usually grow to 1–2 m in height; they belong to a clade composed of species otherwise restricted to the Caribbean. These two species are sister to Erigeron bellidiastroides Griseb., a herbaceous species endemic to Cuba. The third species (D. alternifolius Lawesson & Adsersen) is a perennial herbaceous plant, woody at the base and reaching only up to 50 cm in height. It is sister to two Chilean (Coquimbo–Valparaiso region) species that also have a perennial herbaceous habit: E. fasciculatus Colla and E. luxurians (Skottsb.) Solbrig. They are placed in an assemblage restricted to South America. The review of previous molecular phylogenetic studies revealed that two of the endemic genera and endemic species of three non‐endemic genera have their closest relatives in South America. Endemic species belonging to three non‐endemic genera have sister species in North America or the West Indies. One endemic genus and endemic species in three non‐endemic genera have sister taxa with a widespread continental distribution, or their molecular phylogenies yielded equivocal results. Main conclusions The flora of Galápagos has affinities with both North America (including the Antilles) and South America. Darwiniothamnus exhibits both patterns: two species of this genus are sister to a taxon endemic to Cuba, supporting a connection between the Cocos plate and the West Indies; the third species, D. alternifolius, provides a link with the Coquimbo–Valparaiso region, suggesting a biogeographical connection between the Nazca plate and southern South America.     

Unreduced gametes and neopolyploids in natural populations of Achillea borealis (Asteraceae)

Polyploidy is a major mechanism of speciation and adaptation, yet little is known about the origins of polyploids in natural species. I investigated gametic nonreduction and neopolyploid formation in natural tetraploid populations of Achillea borealis (Asteraceae), an autopolyploid complex consisting of tetraploid and hexaploid cytotypes. Cytological analyses of tetraploid populations revealed the occurrence of reduced (n=2x) as well as unreduced 'big' (2n=4x) and 'jumbo' (4n=8x) pollen grains, which were clearly distinguished by size. Production of unreduced pollen was monitored in two tetraploid populations in 1997 and 1998. Mean population-level frequencies of unreduced pollen ranged from 0.030 to 0.538%, with as few as one-third and as many as one-half of sampled plants producing unreduced grains. Eight individuals were found to produce >1% unreduced pollen, with highest observed frequencies of 7.0, 13.2 and 15.8%. Experimental crosses using high unreduced pollen producers as male parents generated viable seeds. However, the frequency of neohexaploids in the progeny of experimental crosses (0.388%) was similar to that observed in progeny of randomly selected, open-pollinated control parents (0.465%). These results suggest that unreduced eggs are the most likely source of new polyploids. In spite of the inefficiency of unreduced pollen in unilateral sexual polyploidization, the overall rate of neohexaploid formation (one in 233) was several orders of magnitude greater than estimates of genic mutation rates.