毛茛叶报春组一存疑种花粉形态的多样性研究

对分布于浙江西北部的毛茛叶报春组(sect. Ranunculoides)一存疑种5个居群共50个植株的花粉形态进行了研究观察。结果表明该存疑种的花粉形态特征在种下具高度多样化,根据萌发孔类型、数量及组成的不同,其花粉可划分为多沟型、沟多孔少型、孔多沟少型、散孔型和环沟型5种类型;而且居群间花粉的大小及花粉类型的组成也存在较大的差异;此外,同一个植株或同一个花粉囊的花粉也可能具有多种类型的花粉。本文进一步证明了花粉形态特征在种内可能存在多样化的现象。花粉形态特征支持该存疑种为一个独立的新物种,它很可能是由具多沟型和散孔型花粉的亲本杂交进化而来。     

半日花花粉形态种内变异及其花粉地理研究

用扫描电镜在个体水平上研究了产自两个地区的半日花花粉,并首次发现花粉种内变异与其地理分布之间的显著相关,半日花花粉形态在种内个体之间存在差异,以餐壁纹饰明显分为穿孔型和条纹型花粉;按形状和大小又可分为近球形较小和长球形较大花粉。外壁纹饰与形状、大小所得类型基本吻合。本种花粉,因所处地理环境的不同而外壁纹饰也有明显差异,条纹型花粉分布于新疆天山地区,而穿孔型花粉出现在内蒙古伊盟桌子山,并认为花粉外壁     

花色多样性与变异的研究进展

花的颜色不仅在不同物种之间有着丰富的多样性,同一物种的不同居群或个体之间也有着花色的多态性,同一花中的不同器官甚至同一类型的器官也有颜色差异。了解花色多样性的形成和维持机制,有助于揭示花的演化。经典的观点认为,花色是植物提供给传粉者的视觉信号,能促进传粉和提高觅食效率。在分析花色多样性的基础上,本文介绍了4种不同的研究方法,并论述了当前解释花色多样性的3个主要假说。提出今后的研究有必要结合系统发育的分析方法,综合考虑传粉者、植食动物、物理环境等多个因子的选择作用,才能深入理解花色的多样性与演化。     

Does intraspecific behavioural variation of pollinator species influence pollination? A quantitative study with hummingbirds and a Neotropical shrub

• Floral visitors differ in their efficacy as pollinators, and the impact of different pollinator species on pollen flow and plant reproduction has been frequently evaluated. In contrast, the impact of intraspecific behavioural changes on their efficacy as pollinators has seldom been quantified.
• We studied a self‐incompatible shrub Palicourea rigida (Rubiaceae) and its hummingbird pollinators, which adjust their behaviour according to floral resource availability. Fluorescence microscopy was used to access pollen tube growth and incompatibility reaction in pistils after a single visit of territorial or intruder hummingbirds in two populations. To characterise the plant populations and possible differences in resource availability between areas we used a three‐term quadrat variance method to detect clusters of floral resources.
• Within‐species variation in foraging behaviour, but not species identity, affected pollinator efficacy. Effectively, hummingbirds intruding into territories deposited more compatible pollen grains on P. rigida stigmas than territory holders in both study areas. Additionally, territory holders deposited more incompatible than compatible pollen grains.
• Our results imply that intraspecific foraging behaviour variation has consequences for pollination success. Quantifying such variation and addressing the implications of intraspecific variability contribute to a better understanding of the dynamics and consequences of plant–pollinator interactions.

     

Evolution of Euphilotes (Lepidoptera: Lycaenidae) by seasonal and host shifts

The lycaenid butterfly genus Euphilotes , consisting of five species and several dozen subspecies, is confined to western North America. Each subspecies utilizes one or a few species of Eriogonum (Polygonaceae) as larval hosts; larvae feed entirely on pollen and developing seeds. Euphilotes populations are (with few exceptions) univoltine and fly in synchrony with the initial (c. 2 weeks) annual flowering periods of their hosts. Euphilotes evolution coincides with shifts to new hosts, frequently with different bloom periods. Often both inter- and intraspecific populations occur in sympatry. Although interspecific populations may fly in synchrony and utilize the same hosts, sympatric intraspecific populations use different hosts and generally fly allochronically. Analyses of Euphilotes populations using biochemical, morphological, and life history characters, do not support either coevolution or sequential evolution with Eriogonum but are more consistent with opportunistic adaptation to new hosts having different bloom periods.     

Variation in Body Shape across Species and Populations in a Radiation of Diaptomid Copepods

Inter and intra-population variation in morphological traits, such as body size and shape, provides important insights into the ecological importance of individual natural populations. The radiation of Diaptomid species (~400 species) has apparently produced little morphological differentiation other than those in secondary sexual characteristics, suggesting sexual, rather than ecological, selection has driven speciation. This evolutionary history suggests that species, and conspecific populations, would be ecologically redundant but recent work found contrasting ecosystem effects among both species and populations. This study provides the first quantification of shape variation among species, populations, and/or sexes (beyond taxonomic illustrations and body size measurements) to gain insight into the ecological differentiation of Diaptomids. Here we quantify the shape of five Diaptomid species (family Diaptomidae) from four populations each, using morphometric landmarks on the prosome, urosome, and antennae. We partition morphological variation among species, populations, and sexes, and test for phenotype-by-environment correlations to reveal possible functional consequences of shape variation. We found that intraspecific variation was 18-35% as large as interspecific variation across all measured traits. Interspecific variation in body size and relative antennae length, the two traits showing significant sexual dimorphism, were correlated with lake size and geographic location suggesting some niche differentiation between species. Observed relationships between intraspecific morphological variation and the environment suggest that divergent selection in contrasting lakes might contribute to shape differences among local populations, but confirming this requires further analyses. Our results show that although Diaptomid species differ in their reproductive traits, they also differ in other morphological traits that might indicate ecological differences among species and populations.     

Discovery of androecium color polymorphism in Epimedium pubescens with habitat preference of anther/pollen color in the genus

Aims Interspecific and intraspecific variation in flower color in natural populations provides an opportunity for us to understand the evolution and maintenance of diversity of floral traits. Compared to corolla color, little is known about the color polymorphism of sexual organs in flowering plants. To explore evolutionary transitions of androecium color and polymorphism within species, interspecific and intraspecific variation in androecium (anther and pollen) color in the genus Epimedium (Berberidaceae) was investigated.Methods To explore the geographical patterns of anther/pollen color variation in Epimedium species, data of 45 species were collected and their phylogeny was constructed based on available DNA sequences. To investigate whether intraspecific variation in androecium color relates to habitat preference, three environmental factors were measured in the field population of Epimedium pubescens in northeastern Sichuan, China, which plants had green or yellow androecia. Vegetative and reproductive traits of this species were compared between the two color morphs.Important findings Androecium (anther and pollen) color polymorphism in field populations of Epimedium pubescens is reported here where nine populations are monomorphic with a green androecium but three populations are dimorphic with individuals having either a green or a yellow androecium. Inflorescence stalk height, stalk diameter, leaf number, flower number and spur length (as well as spur and nectar volume) were not significantly different between two morphs. Compared to the yellow morph, the green morph had relatively larger leaves and anthers, but smaller sepals. The green morph produced more pollen and larger seeds, but the same number of ovules. Seed set was not significantly different between green and yellow morph. Investigations of environmental factors in the color dimorphic populations of E. pubescens indicated that the green morph was more likely to occur in habitats with relatively lower light intensity. The distribution survey of 45 Epimedium species showed that species with a green androecium tended to appear at lower elevations. Comparative phylogenetic analysis showed that transitions from yellow to a green androecium or to androecial color dimorphism occurred at least seven times. This genus, characterized by anther color diversity and containing some species with anther color polymorphism, provides a model system in which to study the evolution and maintenance of colorful sexual organs in flowering plants.     

Mating system variation in Veronica (Plantaginaceae): inferences from pollen/ovule ratios and other reproductive traits

The pollen–ovule ratio (P/O) is commonly used to estimate the mode of sexual reproduction in flowering plants. In previous studies, a clear correspondence has been detected between this character and the degree of autogamy. We here investigate variation in this character and its expected correlates in the genus Veronica (Plantaginaceae). Pollen–ovule ratios of 45 species representing eleven percent of all the species in the genus were investigated and compared with results from crossing experiments from previous studies. In addition, multiple populations of 17 of the 45 studied species were sampled and a controlled‐environment experiment was conducted to evaluate the extent of intraspecific variation. Moreover, relationships between P/O and other primary and secondary reproductive characters of the Veronica flower were investigated in relation to a phylogenetic hypothesis in order to determine the phylogenetic constraints on reproductive characters. The differences in P/O among species correspond well to the diversity of mating systems in Veronica and correlate well with other floral characters such as corolla size. These characters together seem to allow a powerful and fast tool to infer mating systems. However, causes for intraspecific variation of P/O, such as different cytotypes, ecotypes or different growth conditions, need to be considered.     

葫芦科8属11种植物花粉形态的扫描电镜观察

利用扫描电子显微镜,对葫芦科8属10种1变种植物的花粉形态进行观察研究。结果显示:冬瓜属、葫芦属和栝楼属的花粉粒为近球形,具3孔沟;苦瓜属、西瓜属和丝瓜属的花粉粒为长球形,具3沟;黄瓜属的花粉粒为近球形,具3孔;南瓜属的花粉粒为球形,具散孔。花粉粒大小、形状和外壁雕纹属、种间差异显著。     

Shape analysis of different populations of clams in relation to their geographical structure

Bivalves are excellent candidates for geographically based studies of the morphological variation in individuals of different populations based on the analysis of their shape profiles. In this study, we quantified the overall shell shape differences in individuals of different populations of Ruditapes decussatus and Ruditapes philippinarum in relation to their geographical and genetic distances. A total of 395 and 124 individuals of R. decussatus (nine populations) and R. philippinarum (four populations), respectively, were sampled in different Mediterranean and Atlantic coastal locations. Pictures of the left valve were taken from all individuals. Their profiles were analysed using elliptic Fourier analysis. Mean outlines were computed. In order to classify different individuals for species, the coefficients of harmonic equations were analysed by partial least square discriminant analysis and soft independent modelling of class analogy. The results showed a high percentage of correct classification (99%) between the two species in the independent test. We found that the morphological distance between R. philippinarum and R. decussatus is higher than the morphological distance among populations of the same species. The absence of correspondence between the geographical location and the pattern of morphological and genetic variation indicates the occurrence of a reaction norm in the morphological adaptation of shell shapes to different local environmental conditions.     

Pollen size in Carex: The effect of different chemical treatments and mounting media

Pollen size and pollen aperture size for ten species of the genus Carex L., native to Estonia, have been measured using light microscopy. The species selected represent different sections of the genus, a range of habitats and different chromosome numbers. The effects of two basic chemical treatments, two mounting media and the effect of chemically induced dehydration with tertiary butyl alcohol (TBA) on the size of pollen grains were then recorded.

In general pollen size and pollen aperture size of the species examined is highly variable at both intraspecific and interspecific levels. Carex hirta has notably larger pollen grains than any of the other species investigated and, although correlations between size and chromosome number in the species examined are limited, it also has the highest chromosome number. Statistically significant size differences resulted from variations in chemical treatment, mounting media and tertiary butyl alcohol (TBA) induced dehydration. Acetolysed pollen grains are larger than potassium hydroxide (KOH) treated pollen grains. Pollen grains dehydrated after chemical treatment with TBA are larger than pollen grains not dehydrated. Pollen grains mounted in silicon oil are smaller than grains mounted in glycerine. But considering the great size variation of Carex pollen grains, the size changes caused by preparation procedures fall within the size variation range of the species examined.

All the samples contained a high number of deformed pollen grains and pollen grains with hardly distinguishable or no lateral apertures.     

Pollen morphology and localization of Ni in some Ni-hyperaccumulator taxa of Alyssum L. (Brassicaceae)

Pollen morphology of seven Alyssum L. taxa growing on serpentine soils in different places in the European Mediterranean macrobioclimate territory were studied, described and compared. Cluster analysis was performed to show similarity between species and their populations. The shape of the pollen grains varies among the species and among the grains within the same anther. The pollen grains are 3-colpate, prolate, with long and narrow colpi reaching the poles. The ornamentation of the exine varies from micro-reticulate to reticulate between the species. Pollen sterility/fertility was also calculated. The highest percentage of sterile pollen (73.76%) was calculated for Alyssum murale subsp. murale and the lowest (9.54%) for A. bertolonii subsp. bertolonii. All species are representatives of sect. Odontarrhena (C.A. Meyer) Koch well known as Ni-hyperaccumulators. Nickel and other elements present in pollen and stamen were studied by inductively coupled plasma-mass spectrometry. The stamen parts of all species were micromorphologically analyzed by scanning electron microscopy coupled to an energy-dispersive X-ray probe. Accumulation of Ni was detected in the stamens of all studied species and rarely in the pollen grains. The distribution patterns of Ni were similar among the species examined.     

Concordant clines and significant correlation between floral and pollen characters in Asarum heterotropoides var. heterotropoides (Aristolochiaceae)

To examine geographic differentiation in Asarum heterotropoides var. heterotropoides in Hokkaido Is. in the northern Japan, in which two putative cryptic species have been suspected to exist, extensive and detailed morphological research on 794 individuals from 44 populations throughout Hokkaido Is. was performed. Among the characters examined, the angle between and tip shape of the calyx lobes and the supratecta of the pollen grains were significantly correlated and were found in similar geographic clines. Among them, the pollen showed two discrete states in almost distinct distribution. Multidimensional scaling analysis showed that individuals within each of the two pollen types had different trends in flower characters. Consequently, we assumed a cline from south to north on Hokkaido Is. For the causes of the cline, the two hypotheses were proposed, primary geographic differentiation or extensive introgressive hybridization between two distinct geographical species existing in the past.     

Studies in Montia L. and Claytonia L. and Allied Genera

The pollen morphology of several genera in Portulacaceae is described. Particular attention has been paid to the genera of the subfamily Montioideae, as a stage of continued monographical studies. Among genera especially dealt with are Claytonia, Montia, Crunocallis, Naiocrene, Neopaxia, Mona, Maxia, Limnalsine, and Montiastrum. In the taxonomical treatment of these genera the pollen morphology has proved to afford many important additional characters.

The pollen grains of Claytonia are distinguished from those of the remainder in being 3-colpate. The grains of the Claytonia-type have many similarities with those of Lewisia, a genus of the subfamily Portulacoideae. The other genera of Montioideae have pantocolpate pollen grains. Among these genera several different pollen types are distinguished, chiefly with regard to the sexine structures and the aperture membranes. The Montiastrum-type is especially interesting, with tholate grains, a particular pollen type not met with in any other genus in the family. The pollen morphology of some genera in the Portulacoideae is also treated. In some species in Calandrina and Talinum pantotreme pollen grains are observed with apertures transitional between pori and colpi. The apertures of the pantotreme grains are arranged in characteristic patterns.

Particular attention has been given to the variation of the pollen morphological characters. This variation has been examined with regard to the differences between different populations of the same species as well as between different species. The greatest variation has been observed in the shape and size of the grains. The structure and sculpture and thickness of the sexine and the aperture membranes are less variable. Some polyploid taxa are connected with the occurrence of pollen grains with divergent and varying aperture numbers.

In a survey of the genera the taxonomical results of the investigation are presented with particular regard to the pollen morphology. The new genus, Maxia Ö. Nilss., is described. One new species, Montia clara Ö. Nilss., is described and some new combinations are made.

Pollen morphological diagnoses are given for 46 different taxa. The aperture conditions for 96 different species are presented.     

Patterns of seed dispersal and pollen flow in Quercus garryana (Fagaceae) following post-glacial climatic changes

Aim We examined the genetic structure of Quercus garryana to infer post‐glacial patterns of seed dispersal and pollen flow to test the hypotheses that (1) peripheral populations are genetically distinct from core populations and from one another; (2) genetic diversity declines towards the poleward edge of the species’ range; and (3) genetic diversity in the chloroplast genome, a direct measure of seed dispersal patterns, declines more sharply with increasing latitude than diversity in the nuclear genome. We address our findings in the context of known historical oak distribution from pollen core data derived from previously published research. Location Oak–savanna ecosystems from southern Oregon, USA (core populations/non‐glaciated range) northward to Vancouver Island, British Columbia, Canada (peripheral populations/glaciated range). Methods We genotyped 378 trees from 22 sites with five chloroplast and seven nuclear microsatellite loci. For both sets of markers, we estimated genetic diversity and differentiation using an analysis of molecular variance and generated Mantel correlograms to detect genetic and geographical distance correlations. For the nuclear markers, we also used a Bayesian approach to infer population substructure. Results There was a large degree of population differentiation revealed by six chloroplast haplotypes, with little (≤ 3) or no haplotype diversity within sites. Peripheral island locations shared the same, maternally inherited chloroplast haplotype, whereas locations in mainland Washington had greater haplotype diversity. In contrast, genetic diversity of the nuclear markers was high at all locations sampled. Populations clustered into two groups and were significantly positively correlated over large spatial scales (≤ 200 km), although allele richness decreased significantly with latitude. Population substructure was observed between core and peripheral populations because rare alleles were absent in peripheral localities and common allele frequencies differed. Main conclusions The observed pattern of chloroplast haplotype loss at the northern periphery suggests restricted seed dispersal events from mainland sites to peripheral islands. This pattern was unexpected, however, as refugial oak populations remained near the current post‐glacial range even during the Last Glacial Maximum. Using nuclear markers, we found high within‐population diversity and population differentiation only over large spatial scales, suggesting that pollen flow is relatively high among populations.     

Life cycles shape parasite evolution: comparative population genetics of salmon trematodes

Little is known about what controls effective sizes and migration rates among parasite populations. Such data are important given the medical, veterinary, and economic (e.g., fisheries) impacts of many parasites. The autogenic-allogenic hypothesis, which describes ecological patterns of parasite distribution, provided the foundation on which we studied the effects of life cycles on the distribution of genetic variation within and among parasite populations. The hypothesis states that parasites cycling only in freshwater hosts (autogenic life cycle) will be more limited in their dispersal ability among aquatic habitats than parasites cycling through freshwater and terrestrial hosts (allogenic life cycle). By extending this hypothesis to the level of intraspecific genetic variation, we examined the effects of host dispersal on parasite gene flow. Our a priori prediction was that for a given geographic range, autogenic parasites would have lower gene flow among subpopulations. We compared intraspecific mitochondrial DNA variation for three described species of trematodes that infect salmonid fishes. As predicted, autogenic species had much more highly structured populations and much lower gene flow among subpopulations than an allogenic species sampled from the same locations. In addition, a cryptic species was identified for one of the autogenic trematodes. These results show how variation in life cycles can shape parasite evolution by predisposing them to vastly different genetic structures. Thus, we propose that knowledge of parasite life cycles will help predict important evolutionary processes such as speciation, coevolution, and the spread of drug resistance.     

Sex‐specific responses of phenotypic diversity to environmental variation

Identifying the factors generating ecomorphological diversity within species can provide a window into the nascent stages of ecological radiation. Sexual dimorphism is an obvious axis of intraspecific morphological diversity that could affect how environmental variation leads to ecological divergence among populations. In this paper we test for sex‐specific responses in how environmental variation generates phenotypic diversity within species, using the generalist lizard Gallotia galloti on Tenerife (Canary Islands). We evaluate two hypotheses: the first proposes that different environments have different phenotypic optima, leading to shifts in the positions of populations in morphospace between environments; the second posits that the strength of trait‐filtering differs between environments, predicting changes in the volume of morphospace occupied by populations in different environments. We found that intraspecific morphological diversity, provided it is adaptive, arises from both shifts in populations’ position in morphospace and differences in the strength of environmental filtering among environments, especially at high elevations. However, effects were found only in males; morphological diversity of females responded little to environmental variation. These results within G. galloti suggest natural selection is not the sole source of phenotypic diversity across environments, but rather that variation in the strength of, or response to, sexual selection may play an important role in generating morphological diversity in environmentally diverse settings. More generally, disparities in trait–environment relationships among males and females also suggest that ignoring sex differences in studies of trait dispersion and clustering may produce misleading inferences.     

How oilseed rape (Brassica napus) genotype influences pollen beetle (Meligethes aeneus) oviposition

Oviposition of phytophagous insects is determined either by adaptive behaviours allowing evaluation and response to host plant quality and/or by nutritional constraints occurring during oogenesis. Besides differences found among host plant species, plant intraspecific diversity can also affect insect oviposition. However, to date few studies have extensively investigated the factors accounting for the effect of this intraspecific variation. We addressed this question using oilseed rape (Brassica napus) and the pollen beetle (Meligethes aeneus), a phytophagous insect that uses the same plants and plant organs both for feeding and laying eggs. Our objectives were to test for a genotypic effect of oilseed rape on pollen beetle oviposition and identify the origin of the possible intergenotypic differences. We tested three hypotheses: oviposition is directly linked to (1) the amount of food eaten; (2) the nutritional quality of the food eaten; (3) a preference of females for certain plant genotypes. Results showed intergenotypic differences in both the number and the size of eggs laid. The factor that best accounted for most of these differences was the amount of food eaten. Nutritional quality of the pollen was of minor importance and females exhibited no preference among genotypes. These results reveal the importance of adult feeding on subsequent oviposition in phytophagous insects, an often neglected factor which partly determines the amount of energy available for oogenesis. Taking into account this factor may be of crucial importance in studies conducted on synovogenic insect species feeding on the same plant on which they lay eggs.     

Random amplified polymorphic DNA (RAPD) and quantitative trait analyses across a major phylogeographical break in the Mediterranean ragwort Senecio gallicus Vill. (Asteraceae)

Random amplified polymorphic DNA (RAPD) and quantitative trait variation of the widespread and ephemeral Senecio gallicus were surveyed in 11 populations sampled from the Iberian Peninsula and southern France. The aim of the study was to compare population relationships and levels of geographical differentiation with chloroplast (cp) DNA and allozyme variation assessed previously in the same populations. Employing multivariate statistics, a moderate level of intraspecific differentiation was observed among populations from Iberian coastal and inland regions for both RAPDs and quantitative traits. However, RAPDs provided greater resolution in identifying additional population structure within the hypothesized, Pleistocene refugial source area of the species in coastal Iberia. A major part of the geographical subdivision in RAPD and quantitative traits was concordant with the coastal vs. inland divergence as previously inferred from cpDNA haplotype frequencies, but strongly contrasted with the geographical uniformity of the species for allozymes. This concordance across various nuclear and cytoplasmic markers (RAPDs/quantitative traits, cpDNA) suggests that geographical uniformity for allozymes is more attributable to low rates of evolution and/or small genome sampling rather than high rates of pollen dispersal, slow rates of nuclear lineage sorting, or indirect balancing selection. The present study underscores the value of using additional classes of nuclear markers for narrowing the numbers of competing causal hypotheses about intraspecific cpDNA-allozyme discrepancies and their underlying evolutionary processes.     

Species as Explanatory Hypotheses: Refinements and Implications

The formal definition of species as explanatory hypotheses presented by Fitzhugh (Marine Biol 26:155–165, 2005a, b) is emended. A species is an explanatory account of the occurrences of the same character(s) among gonochoristic or cross-fertilizing hermaphroditic individuals by way of character origin and subsequent fixation during tokogeny. In addition to species, biological systematics also employs hypotheses that are ontogenetic, tokogenetic, intraspecific, and phylogenetic, each of which provides explanatory hypotheses for distinctly different classes of causal questions. It is suggested that species hypotheses can not be applied to organisms with obligate asexual, parthenogenetic, and self-fertilizing modes of reproduction. Hypotheses explaining shared characters among such organisms are, instead, strictly phylogenetic. Several implications of this emended definition are examined, especially the relations between species, intraspecific, and phylogenetic hypotheses, as well as the limitations of species names to be applied to temporally different characters within populations.