Comparative palynology and seed morphology in annual pansies (Viola sect. Melanium,Violaceae): implications for species delimitation

Abstract

This integrated study provides new insights into pollen and seed morphology and pollen heteromorphism of four closely related annual taxa of Viola sect. Melanium. The plant material, both fresh and dried, was collected in Italy and studied using light and scanning electron microscope. Palynological data for V. hymettia together with a detailed comparative analysis of seed morphology and micromorphology of the four species are reported for the first time. Results of this work highlight some pollen and seed features as useful diagnostic characters. The pollen size proves to be of diagnostic value to easily separate V. kitaibeliana, having the smallest pollen grains, from the others, especially from V. arvensis with the largest ones. Exine ornamentation is microreticulate, showing no relevant differences among species. We can partially confirm the diagnostic value of the prevailing pollen morph as it can be useful only for V. arvensis (five-aperturate) versus V. tricolor and V. hymettia (four-aperturate). The macro- and micromorphology of seeds provide additional useful distinguishing characters. Particularly, seed size was found to be a good delimitating character, especially to distinguish V. kitaibeliana (with the smallest seeds) from V. arvensis, and easy to be measured with no need of particular equipment.     

Ecological genetics of vernalization response in Bromus tectorum L. (Poaceae)

BACKGROUND AND AIMS: Bromus tectorum (cheatgrass or downy brome) is an exotic annual grass that is dominant over large areas of former shrubland in western North America. To flower in time for seed production in early summer, B. tectorum plants generally require vernalization at winter temperatures, either as imbibed seeds or as established seedlings. METHODS: Variation in response to increasing periods of vernalization as seeds or seedlings for progeny of ten full-sib families from each of four B. tectorum populations from contrasting habitats was studied. KEY RESULTS: As vernalization was increased from 0 to 10 weeks, the proportion of plants flowering within 20 weeks increased, weeks to initiation of flowering decreased, and seed yield per plant increased, regardless of whether plants were vernalized as seeds or seedlings. Most of the variation was accounted for by differences among populations. Plants of the warm desert population flowered promptly even without vernalization, while those of the cold desert, foothill and montane populations showed incremental changes in response variables as a function of vernalization period. Populations differed in among-family variance, with the warm desert population generally showing the least variance and the cold desert population the most. Variation among populations and among families within populations decreased as vernalization period increased, whereas the non-genetic component of variance showed no such pattern. CONCLUSIONS: Variation in vernalization response was found to be adaptively significant and apparently represents the result of contrasting selection regimes on a range of founder genotypes.     

Quantitative genetic diversity and conservation strategies for an allogamous annual species,Dasypyrum villosum (L.) Candargy (Poaceae)

Dasypyrum villosum (L.) Candargy is a weedy annual diploid (2n = 14, VV genomes) allogamous grass species (Poaceae, Triticeae). Genetic variation for 12 traits was studied in 43 natural populations (31 from Italy and 12 from Croatia and Montenegro of former Yugoslavia) grown in a common field environment in California. Although 7 of 12 traits followed the theoretical prediction that a larger proportion of genetic variation was distributed within populations than among populations, exceptions were found for spike length, plant height, and days to flag-leaf emergence, heading, and anthesis. Covariate analysis showed that developmentally closely related characters were more likely correlated at both population and family within population levels. Geographically closer populations shared more genetic similarity than distant populations as indicated by mean coefficients of variation and cluster analysis of the Euclidean distances among populations. As few as five populations, each population with five or more half-sib seeds taken randomly from 5 plants, is expected to capture more than 95% of the total genetic variation of this species in the region sampled, but sampling a much larger number of seeds per population (> 1000) for long-term storage would supply research and plant breeding needs for several decades. If seed regeneration is required, populations can be sampled from clusters having similar genetic variation, and grown in reproductive isolation or bulked seed samples from all populations of each cluster group can be grown in isolation. The former is recommended if population integrity is desired while the latter is sufficient to provide genetic resources for plant-breeding purposes.     

Patterns of genetic differentiation in two annual bromegrasses,Bromus lanceolatus andB. hordeaceus (Poaceae)

Infraspecific genetic differentiation was analysed in two tetraploid annual bromegrasses,Bromus lanceolatus (from N Africa) andB. hordeaceus (from N Africa and France). Genetic analysis of populations was based on allozyme polymorphisms at 17 loci. Different fixed heterozygous phenotypes were scored in both species, according to their allopolyploid origin. In N Africa, more variation occurred among populations ofB. lanceolatus than ofB. hordeaceus. The variation was not randomly distributed among populations of both species. InB. lanceolatus, differentiation was correlated with climatic variables rather than with geographic distance between populations. Higher correlation of genetic differentiation with geographic distance occurred inB. hordeaceus, particularly at large geographic scale, between French and N African populations. Within each region, the populations appeared weakly genetically differentiated, even when belonging to different subspecies.     

Evolutionary history of the European whitefish Coregonus lavaretus (L.) species complex as inferred from mtDNA phylogeography and gill-raker numbers

We compared mitochondrial DNA and gill-raker number variation in populations of the European whitefish Coregonus lavaretus (L.) species complex to illuminate their evolutionary history, and discuss mechanisms behind diversification. Using single-strand conformation polymorphism (SSCP) and sequencing 528 bp of combined parts of the cytochrome oxidase b (cyt b) and NADH dehydrogenase subunit 3 (ND3) mithochondrial DNA (mtDNA) regions, we documented phylogeographic relationships among populations and phylogeny of mtDNA haplotypes. Demographic events behind geographical distribution of haplotypes were inferred using nested clade analysis (NCA) and mismatch distribution. Concordance between operational taxonomical groups, based on gill-raker numbers, and mtDNA patterns was tested. Three major mtDNA clades were resolved in Europe: a North European clade from northwest Russia to Denmark, a Siberian clade from the Arctic Sea to southwest Norway, and a South European clade from Denmark to the European Alps, reflecting occupation in different glacial refugia. Demographic events inferred from NCA were isolation by distance, range expansion, and fragmentation. Mismatch analysis suggested that clades which colonized Fennoscandia and the Alps expanded in population size 24 500-5800 years before present, with minute female effective population sizes, implying small founder populations during colonization. Gill-raker counts did not commensurate with hierarchical mtDNA clades, and poorly with haplotypes, suggesting recent origin of gill-raker variation. Whitefish designations based on gill-raker numbers were not associated with ancient clades. Lack of congruence in morphology and evolutionary lineages implies that the taxonomy of this species complex should be reconsidered.     

Morphological and embryological characters of three middle Asian Allium L. species (Alliaceae)

Morphological and embryological characters of the xerophilous species Allium crystallinum Vved., A. filidens Regel, and A. fritschii Khassanov et Yengalycheva (Allium subg. Allium) were compared. The morphology of flowers, the rate of development of male and female embryonic structures and correlations with bud dimensions were studied. The overall similarity of the embryonic processes and structures underlines the close taxonomic relationship of the species investigated as concluded from morphological and anatomical characters. However, several differences in embryological characters between A. filidens and A. fritschii confirm the separation of the latter species at species level.     

Recombination indices in species of Erythrina L. (Leguminosae, Papilionoideae)

Darlington's recombination indices were calculated for eight species of Erythina by adding the mean chiasma frequency and the haploid chromosome number of each species. The recombination indices distinguished subgenus Erythraster (two species with greater mean chiasma frequency) from subgenera Micropteryx and Eythrina. This discrimination corroborates literature data on pollen morphology. It was not possible to separate groups at sectional level using recombination index. The recombination indices were also compared with data on breeding system and pollinators, available in literature, to investigate Grant's hypothesis on the occurrence of compensatory mechanisms in the regulation of recombination in plants. The data on breeding systems in Eythrina are very incomplete and it was not possible to correlate values of recombination indices with autogamy or allogamy. Species visited by hummingbirds showed lower recombination indices than species visited by other kind of pollinators.     

Chloroplast DNA variation in diploid and polyploid species of Bromus (Poaceae) subgenera Festucaria and Ceratochloa

Summary Chloroplast DNA (cpDNA) restriction endonuclease patterns are used to examine phylogenetic relationships between Bromus subgenera Festucaria and Ceratochloa. Festucaria is considered monophyletic based on the L genome, while Ceratochloa encompasses two species complexes: the B. catharticus complex, which evolved by combining three different genomes, and the B. carinatus complex, which is thought to have originated from hybridization between polyploid species of B. catharticus and diploid members of Festucaria. All species of subgenus Ceratochloa (hexaploids and octoploids) were identical in chloroplast DNA sequences. Similarly, polyploid species of subgenus Festucaria, except for B. auleticus, were identical in cpDNA sequences. In contrast, diploid species of subgenus Festucaria showed various degrees of nucleotide sequence divergence. Species of subgenus Ceratochloa appeared monophyletic and phylogenetically closely related to the diploid B. anomalus and B. auleticus of subgenus Festucaria. The remaining diploid and polyploid species of subgenus Festucaria appeared in a distinct grouping. The study suggests that the B. catharticus complex must have been the maternal parent in the proposed hybrid origin of B. carinatus complex. Although there is no direct evidence for the paternal parent of the latter complex, the cpDNA study shows the complex to be phylogenetically very related to the diploid B. anomalus of subgenus Festucaria.     

鼠尾草属不同物种的雄蕊杠杆机制对传粉者空间变异的进化响应

被子植物虫媒传粉植物的物种分化通常被认为是花性状响应传粉环境(传粉者)的空间变异而发生适应性分化的结果。通过对鼠尾草属(Salvia) 3个物种(共4个居群)传粉互作系统的比较, 探索了花性状对不同传粉环境的进化响应。结果表明: 各居群的传粉者组成、主要传粉者类型及其大小各不相同, 杠杆状雄蕊及相关花部性状大小在不同居群间具有显著差异; 各居群均表现出腹部传粉和背部传粉2种传粉模式, 但背部传粉仍然是最有效的传粉方式; 居群间杠杆状雄蕊长度与传粉者体长表现出极显著的正相关, 然而花冠长与传粉者体长表现出负相关; 花冠口高度和柱头高度与传粉者胸厚也表现出一定的协同变异。鼠尾草属植物的杠杆状雄蕊及相关花部性状在传粉系统的进化过程中表现出高度的可塑性, 表明雄蕊杠杆传粉机制对传粉环境的变异非常敏感, 在该属植物的物种分化过程中具有关键作用。     

Fitness-related traits of allozyme genotypes in Bromus hordeaceus L. (Poaceae) associated with field habitat and experimental flooding

Genetic variation at alcohol dehydrogenase and phosphoglucose isomerase loci in Bromus hordeaceus has in an earlier study been found to show substantial microgeographic spatial structuring. The present study reports differences in fitness related characters between the enzyme genotypes, both from a field study and a greenhouse experiment. The field study showed overall differences in seed set between allozyme genotypes and also that Pgi-1b genotypes differed in number of seeds set at different levels of herb cover in their habitat. In the greenhouse, dry, normal or flooded conditions were applied. Seeds from individuals with the Adh-1b-11 genotype matured faster in the dry and slower in the flooded treatments than did seeds from individuals with the Adh-1b-22 genotype. Individuals containing Pgi-1b-1f1f alleles and Adh-1b-11 alleles are more plastic than individuals with other allele combinations, meaning that allozyme variation could partly explain what could be seen as adaptive phenotypic plasticity. Mean seed weight was different between dry and flooding treatments for Pgi-1b genotypes. There were also direct effects of allozyme genotype on the probability of survival, total plant weight, weight of reproductive parts, seed weight, days to seed maturation and the percentage of reproductive parts out of the total plant weight.     

Evolution in Carex L. sect. Spirostachyae (Cyperaceae): A molecular and cytogenetic approach

Carex sect. Spirostachyae comprises 25 species displaying the centre of diversity in Eurasia. Phylogenetic analysis of the ITS nrDNA region of 20 species of sect. Spirostachyae, six species of sect. Ceratocystis, five species of sect. Elatae, and eight outgroup species reveals that neither sect. Spirostachyae nor sect. Elatae is monophyletic. With the exclusion of Carex cretica, the 19 species of sect. Spirostachyae studied form a clade with the five species of tropical-subtropical sect. Elatae. Taxonomy of the core Spirostachyae is not only mostly in agreement with our phylogenetic hypothesis, but also with ecological and new cytogenetic results. Two main groups with different chromosome numbers and edaphic preferences are identified in the core Spirostachyae. One includes primarily acidophilous species with high chromosome numbers (2n=(64)68–84), whereas the other one includes mainly basophilous species with lower chromosome numbers (2n=60–74(75)). Chromosome-number variation is extremely different in the core Spirostachyae, showing great stability in some widespread species (e.g. Carex extensa) but an active chromosome evolution – faster chromosomal rearrangements, fusion and fission events than ITS nucleotide substitutions – in more restricted species (e.g. Carex troodi). Biogeography of the two amphiatlantic pairs of species reveals two independent colonizations of South America from the European continent. The geographical barrier of the Strait of Gibraltar has played different roles in the course of evolution of this section, acting as an effective barrier to gene flow in one case (Carex helodes) but as a limited barrier or recent separation in two others (Carex distans, Carex punctata).     

Meiotic configuration and chromosome number in some Iranian species of Elymus L. and Agropyron Gaertner (Poaceae: Triticeae)

Chromosome numbers and analyses of meiotic metaphase I are reported for the following taxa: Agropyron cristatum subsp. incanum (2n= 42), A. cristatum subsp. pecttnatum (2n=28 – 33), Elymus elongatus subsp. ponticus (2n= 69, 70), E. hispidus var. hispidus (2n= 41 43), var. podperae (2n= 42) and var. villosus (2n= 41, 42), E. libanoticus (2n= 14), E. pertenuis (2n= 28, 28+1B), E. repens (2n= 42), E. transhyrcanus (2n= 40–42), E. hispidus var. villosus x E. cf. repens (2n= 42). Chromosome numbers only are reported for the following taxa: E. gentri (2n= 41, 42), E. nodosus subsp. dorudicus (2n= 28), and E. elongatiformis (2n= 56, 57). The haploid genomic constitution SP is reported for Elymus pertenuis. Variable chromosome numbers (2n= 28–32) were observed in the meiotic metaphase I within single anthers of Agropyron cristatum subsp. pectinatum, and the supernumerary chromosomes in this taxon are assumed to have originated from crosses with hexaploids. Partial elimination of these supernumerary chromosomes probably occurs during archesporial mitotic divisions or at an early stage in the meiotic cycle. A hybrid, morphologically intermediate between E. hispidus and E. repens, was obtained from a seed of E. hispidus collected in the field. The meiotic metaphase I configuration in this E. hispidus hybrid suggests that the pollen parent may itself be a hybrid or hybrid derivative of E. repens x E. hispidus.     

Meiotic configuration and chromosome number in some Iranian species of Ely mus L. and Agropyron Gaertner (Poaceae: Triticeae)

Chromosome numbers and analyses of meiotic metaphase I are reported for the following taxa: Agropyron cristatum subsp. incanum (2 n = 42), A. cristatum subsp. pecttnatum (2 n =28 – 33), Elymus elongatus subsp. ponticus (2 n = 69, 70), E. hispidus var. hispidus (2 n = 41 43), var. podperae (2 n = 42) and var. villosus (2 n = 41, 42), E. libanoticus (2 n = 14), E. pertenuis (2 n = 28, 28+1B), E. repens (2 n = 42), E. transhyrcanus (2 n = 40–42), E. hispidus var. villosus x E. cf. repens (2 n = 42). Chromosome numbers only are reported for the following taxa: E. gentri (2 n = 41, 42), E. nodosus subsp. dorudicus (2 n = 28), and E. elongatiformis (2 n = 56, 57). The haploid genomic constitution SP is reported for Elymus pertenuis. Variable chromosome numbers (2 n = 28–32) were observed in the meiotic metaphase I within single anthers of Agropyron cristatum subsp. pectinatum , and the supernumerary chromosomes in this taxon are assumed to have originated from crosses with hexaploids. Partial elimination of these supernumerary chromosomes probably occurs during archesporial mitotic divisions or at an early stage in the meiotic cycle. A hybrid, morphologically intermediate between E. hispidus and E. repens , was obtained from a seed of E. hispidus collected in the field. The meiotic metaphase I configuration in this E. hispidus hybrid suggests that the pollen parent may itself be a hybrid or hybrid derivative of E. repens x E. hispidus.     

克隆整合对无芒雀麦在异质性盐分环境中存活和生长的影响

无芒雀麦是浑善达克沙地植物群落中占优势的多年生根茎禾草.研究了克隆整合特性对无芒雀麦在异质性盐分环境中存活和生长的影响.结果表明,克隆整合显著提高了无芒雀麦分株在高盐环境中的存活能力,耗-益分析表明无芒雀麦在高盐斑块中分株的生物量、分株数、根茎节数和根茎总长显著受益于克隆整合,而与之相连的非盐分斑块中的分株却没有产生显著的损耗.因而,克隆整合特性是无芒雀麦对异质性环境形成的重要适应对策,它使无芒雀麦能够扩展到不适合植物生长的高盐分斑块中,从而增加了无芒雀麦在浑善达克沙地中的存活和生长,提高了其在半干旱沙化地区的适合度.     

An evaluation of vertebrate seed dispersal syndromes in four species of black nightshade (Solanum sect. Solanum)

We examined the ecological relevance of bird versus mammal dispersal syndromes in four species of Solanum, S. americanum Type A, S. americanum Type B, S. ptychanthum, and S. sarrachoides. These plants were selected because their morphological characteristics, such as fruit color, mass, and persistence, resembled those typically associated with classically-defined bird and mammal dispersal syndromes. We monitored persistence of tagged fruits, compared physical and chemical chaacteristics, performed fruit preference trials with northern bobwhite quail (Colinus virginianus), deer mice (Peromyscus maniculatus), and American robins (Turdus migratorius), and assessed differences in use of olfactory cues by foraging deer mice. We predicted that principal components analysis of physical and chemical characteristics would place fruits of these species along a clear gradient from bird to mammal dispersal syndromes (S. americanum Type A, S. americanum Type B, S. ptychanthum, and S. sarrachoides). However, physical and chemical characteristics did not consistently follow the gradient. Also, contrary to expectations, both birds and mammals demonstrated a preference for S. americanum Types A and B, both bird fruits. Deer mice and bobwhite quail showed much less discrimination among fruit types than did American robins. While the relatively strong odor of the green-fruited S. sarrachoides suggested a mammalian attractant, deer mice discovered the relatively odorless S. americanum Type A significantly more quickly. We conclude that in Solanum, suites of morphological characteristics resembling bird and mammal dispersal syndromes are not good predictors of fruit choice by birds and mammals. We note, however, that this conclusion is based on a sample of three animal species. Alternative explanations for fruit character suites should be considered. For example, the adaptive significance for an association of green coloration of ripe fruit with impersistence (dropping when ripe), as in S. sarrachoides and some S. ptychanthum, may relate more to photosynthesis and carbon balance in detached fruits than to disperser attraction.     

Correlates of geographic range size in New Zealand Chionochloa (Poaceae) species

Aim To test for correlations between plant traits and geographic range size. Location: New Zealand. Methods Trait data were derived from comparative experiments, in which plants were grown in pots or in a common garden, that tested for intrinsic differences among the species in traits relating to growth, reproduction and dispersal. Controlled experiments were used to test for differences in responses to drought and waterlogging stress. Geographic range size was measured as the number of 10 km grid squares in the New Zealand region containing at least one occurrence of the species. Results Growth rate, dispersal capacity and environmental tolerance were all positively related to geographic range size. Geographically restricted species tended to have more variable flowering between years. Flowering intensity, reproductive allocation, seed set, diaspore size, and responses to single environmental factors were not related to geographic range size. Main conclusions The differences between range‐restricted and widespread Chionochloa species appear to represent alternative strategies of coping with environmental change in a dynamic landscape. Range‐restricted species are specialized to temporally persistent habitats that are of limited geographic extent. As a consequence, they have evolved traits that conflict with persistence in widespread habitats. The implication for conservation management is that the conservation of rare plants will frequently depend on protection of their habitats. The widespread Chionochloa species possess traits that enable them to disperse to and occupy a greater range of habitats. These traits have allowed some of these species to expand their ranges following environmental changes that favoured an increase in grassland extent.     

中国野青茅属(禾本科)一些种类的修订

在标本室研究和野外考察的基础上,对中国野青茅属的一些种类进行了修订。首次归并了5个类群,即将Deyeuxia pulchella var． laxa P．C．Kuo et S．L. Lu,D．gyirongensis P．C. Kuo et S．L．Lu,Calamagrostis megalantha Keng ex Keng f.并入 D. pulchella (Griseb.)Hook．f.作为异名;将D．venusta Keng,Calamagrostis longiflora Keng ex Keng f．并入D.flavens Keng作为异名;并确认D. compacta Munro ex Hook．f.与D.holciformis (Jaub．et Spach．)Bor是同一种。     

A reassessment of some Asian species of Heterophyllium (Schimp.) Kindb. (Pylaisiadelphaceae,Bryophyta)

Introduction. Heterophyllium albicans Thér., H. henryi Tixier, H. micro-alare (Broth. & Paris) Broth. and H. tonkinense (Broth. & Paris) Broth. were all either poorly known species or known only from the type collections since their inception.

Methods. Examination of the type material confirmed that the alar cell structure in these species do not conform to the generic concept of Heterophyllium (Schimp.) Kindb. The study also revealed some new features previously unreported in these species, i.e., the filamentous pseudoparaphyllia, one of the characteristic features of the genus Isopterygium Mitt.

Key Results & Conclusions. Three new combinations, Isopterygium albicans (Thér.) Y.Jia & S.He, I. micro-alare (Broth. & Paris) Y.Jia & S.He, and I. tonkinense (Broth. & Paris) Y.Jia & S.He are proposed. Heterophyllium henryi Tixier is treated as a new synonym of Brotherella henonii (Duby) M.Fleisch.