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Premise of research. Many plant species balance resource allocation between sexual and asexual reproduction. Ecological factors that alter this balance can have important effects on plant populations. By eliminating flowers, floral herbivory (florivory) reduces sexual reproduction and consequently shifts reproductive allocation toward asexual reproduction. Over time, the clonal structure of the population may change. To investigate long-term effects of florivory, we characterized clonal structure in patches of the woodland herb Clintonia borealis (Liliaceae), or the bluebead lily. Methodology. A common understory species of North American boreal forests, bluebead lily is a favorite spring food of Odocoileus virginianus, or white-tailed deer. We sampled patches of bluebead lily located on islands in the Great Lakes, which either had or did not have white-tailed deer. Genotyping using newly developed microsatellite markers allowed clones to be identified. Pivotal results. The clonal structure, measured as the genet-to-ramet ratio (G∶N) of a patch, averaged 0.104 genets per ramet on islands with deer, compared with 0.458 genets per ramet on islands without deer. The difference was significant, based on a linear mixed effects model (ΔAICc=3.94, Akaike weight = 0.878), and the effect size, as measured by Cohen’s d was 1.995. We also observed significantly lower clonal diversity and clonal evenness on islands with deer, which in turn could further limit sexual reproduction due to self-sterility or inbreeding. Conclusions. While the effects of white-tailed deer on plant community structure have been well documented, our results suggest that they may also have significant effects on the reproductive dynamics of individual plant species.

Polyploidization plays an important role in plant evolution and biodiversity. However, intraspecific polyploidy compared to interspecific polyploidy received less attention. Clintonia udensis (Liliaceae) possess diploid (2 n  = 2 x  = 14) and autotetraploid (2 n  = 4 x  = 28) cytotypes. In the Hualongshan Mountains, the autotetraploids grew on the northern slope, while the diploids grew on the southern slopes. The clonal growth characteristics and clonal architecture were measured and analyzed by field observations and morphological methods. The diversity level and differentiation patterns for two different cytotypes were investigated using SSR markers. The results showed that the clonal growth parameters, such as the bud numbers of each rhizome node and the ratio of rhizome branches in the autotetraploids were higher than those in the diploids. Both the diploids and autotetraploids appeared phalanx clonal architectures with short internodes between ramets. However, the ramets or genets of the diploids had a relatively scattered distribution, while those of the autotetraploids were relatively clumping. The diploids and autotetraploids all allocated more biomass to their vegetative growth. The diploids had a higher allocation to reproductive organs than that of autotetraploids, which indicated that the tetraploids invested more resources in clonal reproduction than diploids. The clone diversity and genetic diversity of the autotetraploids were higher than that of the diploids. Significant genetic differentiation between two different cytotypes was observed ( P  < 0.01). During establishment and evolution, C. udensis autotetraploids employed more clumping phalanx clonal architecture and exhibited more genetic variation than the diploids.

The whole chloroplast genome of Clintonia udensis, an import Chinese medicinal herb, was determined by Illumina sequencing data in this study. The cp genome is 153,160 bp in length, with a large single-copy region (LSC) of 83,901 bp, a small single-copy region (SSC) of 17,241 bp and a pair of inverted repeat regions (IRs) of 26,009 bp. It contains 132 genes, including 83 protein-coding genes, 36 tRNA genes, 8 rRNA genes, and 5 pseudogenes. The overall GC content was 37.3%, while the corresponding values in the LSC, SSC and IR region are 35.1, 31.0, and 42.8%, respectively. Phylogenetic analysis indicated that Clintonia udensis was related to Lilioideae species in Liliaceae.

Polyploidy is an important factor shaping the geographic range of a species. Clintonia udensis (Clintonia) is a primary perennial herb widely distributed in China with two karyotypic characteristics—diploid and tetraploid and thereby used to understand the ploidy and distribution. This study unraveled the patterns of genetic variation and spatiotemporal history among the cytotypes of C. udensis using simple sequence repeat or microsatellites. The results showed that the diploids and tetraploids showed the medium level of genetic differentiation; tetraploid was slightly lower than diploid in genetic diversity; recurrent polyploidization seems to have opened new possibilities for the local genotype; the spatiotemporal history of C. udensis allows tracing the interplay of polyploidy evolution; isolated and different ecological surroundings could act as evolutionary capacitors, preserve distinct karyological, and genetic diversity. The approaches of integrating genetic differentiation and spatiotemporal history of diploidy and tetraploidy of Clintonia udens would possibly provide a powerful way to understand the ploidy and plant distribution and undertaken in similar studies in other plant species simultaneously contained the diploid and tetraploid. The results showed that the diploids and tetraploids showed the medium level of genetic differentiation; tetraploid were slightly lower than diploid in genetic diversity; recurrent polyploidization was possible to explain the local genotype and was allowed to trace its interplay from the spatiotemporal history. Isolated and different ecological surroundings could be an evolutionary capacitors to preserved distinct karyological and genetic diversity.

The pollen morphology of 54 species and one variety of seven genera in Polygonatae including Clintonia, Disporopsis, Disporum, Maianthemum, Polygonatum, Smilacina and Streptopus was observed and studied in detail; of these, nine species were reported for the first time. Our results showed that the surface ornamentation of pollen grains of the studied materials could be divided into seven types, namely gemmate, granulate-foveolate, perforate, reticulate, rugulate, rugulate-perforate and verrucate. In line with previous studies, we believe that (i) Smilacina ginfushanicum should be classified into the genus Heteropolygonatum rather than the genus Smilacina; (ii) Polygonatum should be divided into section Polygonatum and section Verticillata; (iii) Smilacina and Maianthemum should be combined as one genus, i.e. Maianthemum sensu lato; and (iv) Clintonia, Disporum and Streptopus should be separated from the tribe Polygonatae.

The genetic diversity of 17 populations of Clintonia udensis Trautv. et Mey. was studied by using 15 intersimple sequence repeat primers. The results revealed a relatively high genetic diversity, with percentage of polymorphic \\documentclass{aastex} \\usepackage{amsbsy} \\usepackage{amsfonts} \\usepackage{amssymb} \\usepackage{bm} \\usepackage{mathrsfs} \\usepackage{pifont} \\usepackage{stmaryrd} \\usepackage{textcomp} \\usepackage{portland,xspace} \\usepackage{amsmath,amsxtra} \\usepackage[OT2,OT1]{fontenc} \\newcommand\\cyr{ \\renewcommand\\rmdefault{wncyr} \\renewcommand\\sfdefault{wncyss} \\renewcommand\\encodingdefault{OT2} \\normalfont \\selectfont} \\DeclareTextFontCommand{\\textcyr}{\\cyr} \\pagestyle{empty} \\DeclareMathSizes{10}{9}{7}{6} \\begin{document} \\landscape $$\\mathrm{bands}\\,=98.8\\% $$ \\end{document} , Shannon's information \\documentclass{aastex} \\usepackage{amsbsy} \\usepackage{amsfonts} \\usepackage{amssymb} \\usepackage{bm} \\usepackage{mathrsfs} \\usepackage{pifont} \\usepackage{stmaryrd} \\usepackage{textcomp} \\usepackage{portland,xspace} \\usepackage{amsmath,amsxtra} \\usepackage[OT2,OT1]{fontenc} \\newcommand\\cyr{ \\renewcommand\\rmdefault{wncyr} \\renewcommand\\sfdefault{wncyss} \\renewcommand\\encodingdefault{OT2} \\normalfont \\selectfont} \\DeclareTextFontCommand{\\textcyr}{\\cyr} \\pagestyle{empty} \\DeclareMathSizes{10}{9}{7}{6} \\begin{document} \\landscape $$\\mathrm{index}\\,=0.56$$ \\end{document} , and Nei's gene diversity \\documentclass{aastex} \\usepackage{amsbsy} \\usepackage{amsfonts} \\usepackage{amssymb} \\usepackage{bm} \\usepackage{mathrsfs} \\usepackage{pifont} \\usepackage{stmaryrd} \\usepackage{textcomp} \\usepackage{portland,xspace} \\usepackage{amsmath,amsxtra} \\usepackage[OT2,OT1]{fontenc} \\newcommand\\cyr{ \\renewcommand\\rmdefault{wncyr} \\renewcommand\\sfdefault{wncyss} \\renewcommand\\encodingdefault{OT2} \\normalfont \\selectfont} \\DeclareTextFontCommand{\\textcyr}{\\cyr} \\pagestyle{empty} \\DeclareMathSizes{10}{9}{7}{6} \\begin{document} \\landscape $$\\mathrm{index}\\,=0.38$$ \\end{document} . The genetic diversity at the population level was considerably low, while a high level of genetic differentiation among populations was detected. This was confirmed by analysis of molecular variance (AMOVA). No significant statistical difference was found between geographic regions. Tetraploid populations showed an approximately equal level of genetic diversity compared with that of the diploid populations. No significant differentiation occurred between cytotypes. Shannon diversity index, Nei's gene diversity index, and AMOVA analysis all showed that there was more genetic variation among diploid/tetraploid populations than within diploid/tetraploid populations. The Mantel test revealed no significant correlation between genetic distance and geographic distance at the species level. The biological characteristics of this species, the unique Clintonia-type megasporogenesis and genetic drift, might account for the existence of relatively high interpopulation genetic variation. Cluster analysis gathered the 17 populations into two groups. Group I contained all diploids and made up geographic region I. This might be the center of origin of C. udensis. Group II had two coexisting cytotypes and made up geographic region IIa, IIb-1, and IIb-2, which might represent the secondary diversity centers of C. udensis.

Clonal herbs that attain maximum development in late-seral forest are often assumed to have similar responses to disturbance and to be functionally equivalent. However, little is known about the demographic or physiological responses of these plants to disturbance or to the altered conditions of the post-disturbance environment. Following harvest of a mature coniferous forest, we compared abundance, demographic changes, and physiological acclimation of three clonal herbs (Asarum caudatum, Clintonia uniflora, and Pyrola picta) that differ in belowground morphology and leaf longevity. We measured ramet density, leaf area, and demographic variables (survival, clonal growth, flowering, and seedling establishment) before and for two years after harvest, and in adjacent undisturbed forest. Acclimation to increased solar radiation was assessed two years after harvest, by measuring leaf mass per unit area (LMA) and chlorophyll a:b ratios of leaves produced in the current year. Although initial declines in abundance were similar, demographic responses indicate that patterns of recovery varied greatly among species. Two years after logging, ramet survival and clonal growth (production of new ramets) of Clintonia were greater in the harvest area than in the forest. Asarum had lower survival in the harvest area, but greater clonal growth, and Pyrola showed no difference in either survival or growth between environments. Only Asarum produced seedlings, although their survival was low in the harvest area. All species had higher LMA in the harvest area, but only Clintonia (with annual leaves) had a higher chlorophyll a:b ratio, suggesting the greatest potential for acclimation to increased light. Our results demonstrate that forest herbs with greater rhizome plasticity and shorter leaf duration have greater potential to acclimate after disturbance than those with rigid architectures and persistent leaves. Thus, species with comparable successional roles can vary substantially in their demographic and physiological responses to disturbance, with potential consequences for long-term recovery.

Premise of the study: Over-browsing of the understory vegetation by white-tailed deer has been a cause of decline in many plant populations. Liliaceae are particularly sensitive, yet individual species differ in their tolerance to deer herbivory. In this paper, we examine whether differences in clonal habit, carbon allocation patterns, and phenology influence the capacity of a species to tolerate and recover from repeated herbivory. METHODS: Flowering ramets of Clintonia borealis, Maianthemum canadense, and Trillium erectum were subjected to total defoliation for one or two springs. Key results: Survival was highest in the nonclonal species, T. erectum, most probably due to its very large carbohydrate reserves. Nutrient reserves were less affected than carbohydrate reserves by defoliation, confirming the importance of carbohydrate reserves for survival. However, faster recovery following episodes of defoliation was observed not in the species that sprouted the earliest, T. erectum, but in the clonal species, M. canadense, which had the smallest carbohydrate reserves but also a lower shoot to root ratio than the other clonal species, C. borealis. All plants that were defoliated for 2 years only partially recovered in terms of leaf area, plant biomass, and carbohydrate and nutrient reserves, confirming the overall sensitivity of these species to simulated deer herbivory. CONCLUSIONS: High carbohydrate reserves and consequently low shoot to root ratios appear to increase tolerance to herbivory, whereas clonal species recover faster than nonclonal species. The role played by carbohydrates reserves suggests that these species could benefit from slightly higher light conditions in areas subjected to high deer pressure.

Quametopia gen. n. is established on the basis of recent phylogenetic analysis of taxa formerly assembled under the genus Mumetopia Melander, 1913 to include M. terminalis (Loew, 1863) and two new closely related Nearctic species of Anthomyzidae. The new genus is diagnosed and its phylogenetic relationships discussed. Quametopia terminalis (Loew, 1863) comb. n. is transferred from Mumetopia and redescribed based on revision of the type material (lectotypes of Anthophilina terminalis Loew, 1863 and its synonym Mumetopia nitens Melander, 1913 are designated) and other extensive material. Quametopia clintonia sp. n. and Q. amplistylus sp. n. (both from Canada, USA) are described and relationships of all three Quametopia species discussed. Immature stages of Q. terminalis and Q. clintonia sp. n. obtained by means of adult-to-adult rearing are described (1st- and 2nd-instar larvae for the first time in the family Anthomyzidae) and illustrated. Keys to adults, eggs, larvae and puparia of Quametopia species are presented. Biology (habitat and host-plant associations, life history) of Q. terminalis and Q. clintonia sp. n. are studied in detail and their ecological separation demonstrated. Distribution of all Quametopia species is reviewed. [PUBLICATION ABSTRACT]

The clonal diversity of Clintonia udensis Trautv. et Mey. was detected by ISSR markers among 16 populations, and its correlation with ecological factors was analyzed as well in this work. Results showed that individuals (clonal ramets) per genotype were 1.12 and 1.149 at population and species levels, respectively, and that the 16 populations were all multiclonal. The detected genotypes were localized, without exception, within populations but demonstrated relatively high clonal differentiation among populations. The clonal diversity of the studied populations was high, with the average Simpson's index of 0.975, while the genets showed a clonal architecture of \"guerilla\". The population genetic diversities revealed by genet were consistent with those by ramet, further confirming their genetic differentiation among populations. And its genotype diversity within populations probably resulted largely from the frequent seedling regeneration and self-compatibility. In addition, the correlation analysis further revealed that, among the ecological factors, Simpson's index of C. udensis had a significant positive correlation (P<0.05) with pH values in the soil but not others.