Shared traits make flies and bees effective pollinators of oilseed rape (Brassica napus L.)

One of the major aims of pollination ecology has been to understand the role and relative importance of different pollinator species in both natural and agricultural systems. This study explores how the quantity of pollen delivered in a single visit to a stigma (SVD) differs across insect species from a wide range of taxa (Orders: Hymenoptera, Diptera and Coleoptera) and assesses the morphological and behavioural traits that lead to differences in pollen delivery. We used oilseed rape (OSR) (Brassica napus L.), an economically important crop with open self-fertile flowers, as a test system. Behavioural traits and SVD were measured in the field for individuals of all frequent flower-visiting species of OSR. Individuals were collected and morphological traits were measured in the laboratory. The quantities of free pollen on different body parts of flower visitors, or ‘pollen load’, were also measured. Behavioural and morphological traits were then assessed as predictors of SVD. The individuals that delivered most pollen were (in order of greatest median number of pollen grains delivered): bumblebees (Bombus spp.), mining bees (Andrenidae) and honeybees (Apis mellifera). However, all but one family of flies tested delivered a significantly greater median number of pollen grains than the control, and most delivered more than sweat bees (Halictidae). Behavioural and morphological traits were important in determining pollen delivery, with greater body length, hairiness and visit duration all resulting in greater SVD. Pollen load was measured for a proportion of individuals and we found that for momentary flower visits, a greater quantity of pollen on the head of the flower visitor was linked to a greater SVD. This study demonstrates that, while bees are important pollinators in this system, many fly taxa are also effective at delivering pollen and this is linked to morphological and behavioural traits found in both groups.     

Congruence between comparative morphology and molecular phylogenies: Evolution of the male genital skeletal/muscular system in the subtribe Polyommatina (Lepidoptera,Lycaenidae)

The skeleton and musculature of male genitalia were studied in species of a model butterfly group (subtribe Polyommatina, Lycaenidae). In total, we analyzed 45 species of the tribe Polyommatini most of which were previously used in the molecular phylogenetic study (Talavera et al., 2013). The studied morphological characters were mapped on the molecular trees, which allowed us to reveal trends of morphological changes and to estimate the age of their origin. As a result, chronology of evolution of skeleton and musculature traits was established. It was shown that periods of slow morphological evolution alternated in the subtribe Polyommatina with those of a high rate of origin of new traits. For example, topography of the intravalvar muscles has not changed for 26 MY preserving their initial fan-shaped attachment. The evolution of intravalvar muscles started 10 MYA, proceeded slowly during the first 5 MY, and then accelerated during the last 5 MY resulting in the extensive splitting of the musculature in most monophyletic lineages. Mapping the morphological characters on the phylogeny demonstrated that the rates of skeleton and muscle evolution within the skeleton/musculature apparatus were different. In most cases the intravalvar musculature evolved much faster than the skeleton. The cladistic interpretation of states of morphological traits was found to be consistent with phylogenetic reconstructions based on analysis of multiple molecular markers. Moreover, morphological synapomorphies were found for the lineages Alpherakya + Glabroculus and Aricia + (Alpherakya + Glabroculus), which had low statistical support in molecular phylogenetic analysis. Additionally, in some cases molecular studies helped to reveal trends in the evolution of morphological traits. For example, the unpaired uncus and the compact juxta are not plesiomorphic for Cupidina as previously thought; instead, they were shown to have evolved secondarily within this subtribe.     

基于形态及分子标记的濒危植物夏蜡梅自然居群的遗传变异研究

比较了濒危植物夏蜡梅(Sinocalycanthus chinensis)大明山、大雷山、龙须山3个居群的果实与种子的形态变异,采用ISSR分子标记技术分析DNA序列的变异,综合评价夏蜡梅3个居群的遗传变异。结果如下:果实性状中,果柄长、果实长、果实重、每果种子数等指标均以大明山居群最大、大雷山居群次之、龙须山居群最小,它们之间差异显著(P<0.05)。种子重、种子长、种子宽、种子厚等指标均以大雷山居群最大,与大明山居群和龙须山居群差异显著(P<0.05)。基于果实形态特征的表型分化系数(VST)在0.5518—0.9750之间,平均为0.8930,所有果实形态指标的变异大部分存在于居群间。基于种子形态特征的VST在0.1669—0.8678之间,平均为0.6240,除种子长外,其他种子形态指标的变异也大部分存在于居群间。ISSR分析表明,3个居群的多态位点百分率、Shannon信息指数、Nei基因多样性均是大明山居群最高、龙须山居群次之、大雷山居群最低。居群间的遗传分化系数(GST)为0.6050。基于ISSR分子标记数据的聚类结果显示大明山居群先与龙须山居群聚在一起,再与大雷山居群相聚,这与基于种子性状特征的聚类结果相似,而与基于果实性状特征的聚类结果不同。3个居群的种子形态特征变异系数与遗传多样性水平具有显著的正相关,且基于果实形态特征估算的VST要高于GST,而基于种子形态特征估算的VST仅略高于GST,表明夏蜡梅不同居群间的种子形态变异主要由遗传变异所造成的,而环境因子在果实形态变异中起了重要作用。Mantel检验显示3个居群基于分子标记数据的遗传距离矩阵和基于果实及种子形态特征的欧氏距离矩阵之间的相关性不显著,表明果实与种子在居群间出现的表型分化除了受遗传因素影响外,还受到其它环境因子的强烈影响。     

Genetic diversity of Persian shallot (Allium hirtifolium Boiss.) populations based on morphological traits and RAPD markers

Mooseer or Persian shallot (Allium hirtifolium Boiss.) is a bulbous perennial herb belonging to Alliaceae family. Genetic diversity of 16 mooseer populations collected from different parts of Iran were evaluated using 14 morphological traits as well as antioxidant activities and RAPD markers. Statistical analysis showed significant differences among populations for all morphological traits. In a principal component analysis, four components showed 77.5 % of the total morphological variation. Based on the morphological traits, populations were divided into five groups by clustering. Antioxidant capacities differed among all populations, with the Sahneh and Nahavand populations having the highest and the lowest activities, respectively. Genetic diversity of populations was also investigated through RAPD technique using 14 random primers. Cluster analysis based on RAPD markers plus 47 % coefficient of similarity divided the genotypes into six distinct groups. The highest and lowest similarities between populations were detected to be 0.63 and 0.21, respectively. These results indicated that the assessment of genetic diversity in Iranian mooseer populations could be appropriately carried out using RAPD technique which correlates with morphological traits.     

A naturally occurring variant of Hsp90 that is associated with decanalization

The heat shock protein Hsp90 has been the focus of many studies since it was suggested that it acts to mediate the buffering of phenotypic variation. Hsp90-mediated buffering may result in the accumulation of cryptic genetic variation that, when released either as a consequence of environmental or genetic stress, increases the evolvability of a population. Recent studies using laboratory-induced mutations of Hsp90 and/or chemical inhibition to disrupt Hsp90 function confirm that Hsp90 can buffer cryptic genetic variation. We have previously identified a naturally occurring variant in the charged linker region of the Hsp90 gene, and now examine whether this variant is associated with altered levels of trait variability. The variant is associated with the release of cryptic genetic variation for canalized morphological (bristle) traits, but not for uncanalized morphological (wing and bristle) traits, and the effect on canalized traits depends on culture temperature. This suggests that natural genetic variation in Hsp90 may mediate the evolution of canalized morphological traits even if it does not influence the expression of variation for uncanalized traits.     

Morphometric and genetic differentiation of diploid and hexaploid populations of Aster amellus agg. in a contact zone

The aim of this study is to determine the origin, genetic relationships and morphological differences between diploid and hexaploid cytotypes of Aster amellus aggregate (Asteraceae) at their contact zone in the Czech Republic. We collected data on morphological and isozyme variation in a range of populations of the two cytotypes. We also studied the plasticity of the morphological traits in a common garden. The results suggest that hexaploid individuals of Aster amellus aggregate are of autopolyploid origin. The isozyme data indicate that diploids and hexaploids have different evolutionary histories. This, together with previous detailed cytological and ecological analyses, suggests that there is a secondary contact zone between the two cytotypes in the Czech Republic. The results of multivariate morphometric analyses and data on plasticity of the morphological traits indicate that it is not possible to distinguish the two cytotypes morphologically. The previously published morphological distinction between the cytotypes is thus not supported.     

The relevance of morphology for habitat use and locomotion in two species of wall lizards

Understanding if morphological differences between organisms that occupy different environments are associated to differences in functional performance can suggest a functional link between environmental and morphological variation. In this study we examined three components of the ecomorphological paradigm – morphology, locomotor performance and habitat use – using two syntopic wall lizards endemic to the Iberian Peninsula as a case study to establish whether morphological variation is associated with habitat use and determine the potential relevance of locomotor performance for such an association. Differences in habitat use between both lizards matched patterns of morphological variation. Indeed, individuals of Podarcis guadarramae lusitanicus, which are more flattened, used more rocky environments, whereas Podarcis bocagei, which have higher heads, used more vegetation than rocks. These patterns translated into a significant association between morphology and habitat use. Nevertheless, the two species were only differentiated in some of the functional traits quantified, and locomotor performance did not exhibit an association with morphological traits. Our results suggest that the link between morphology and habitat use is mediated by refuge use, rather than locomotor performance, in this system, and advise caution when extrapolating morphology-performance-environment associations across organisms.     

A morphometrical study of oribatid mites (Acari: Oribatida) of the genus Carabodes C.L. Koch, 1835 (Carabodidae) using a confocal laser scanning microscope: an alternative approach to quantitative analysis of various features

In the past, micro-arthropod taxonomists have mainly relied on qualitative characters to identify species, and relatively little attention has been paid to quantitative analysis. However, several taxonomic groups have high morphological variability, which creates problems in identification. Quantitative analysis of morphometrical data of specific characters could aid in understanding differences in the morphology of oribatid species. In this respect, confocal laser scanning microscopy and quantitative analysis were used to describe morphological traits of four oribatid mite species belonging to the genus Carabodes Koch, 1835—C. femoralis (Nicolet, 1855), C. labyrinthicus (Michael, 1879), C. ornatus Storkan, 1925 and C. subarcticus Trägårdh, 1902. All specimens were collected in Latvia. Thirteen morphological traits were measured for 838 adult individuals. The same traits were also measured with transmitted light microscopy, and results were compared. The impact of a number of environmental factors on traits was also assessed. High interspecific variability in size of most morphological features was observed. In many cases, sizes of the measured features correlated between different species. The ranges in length of notogastral setae p3 and h3 of C. femoralis did not overlap with those of C. labyrinthicus or C. ornatus. Significant correlations between body length and size of setal structures, as well as between setal structures, were detected. Size of morphological features was directly affected by the cover of lichens, a main food resource for Carabodes, and indirectly by the effect of soil moisture on lichen communities.     

Ontogenetic variations in leaf morphology of the tropical rain forest species Dipterocarpus alatus Roxb. ex G. Don

Under natural conditions ontogenetic development often coincides with changes in environmental factors. When explaining variations in leaf parameters, analyses based solely on environmental factors will lead to significant errors if the plant shows substantial ontogenetic variations in leaf properties. We evaluated intraspecific variations in eight morphological leaf traits of Dipterocarpus alatus over six architectural development stages under two different light conditions. An architectural analysis was conducted to distinguish precisely and objectively developmental stages of D. alatus. Leaves were collected on the most recent complete growth unit on the trunks of trees growing under two different light conditions. Eight leaf morphological traits were measured and calculated using ImageJ on greyscale images of leaf tracings. One-way ANOVA and Tukey tests were used to determine differences in leaf traits during ontogeny. The correlation coefficients were compared to determine whether leaf traits correlated more strongly with ontogenetic stage than with light intensity. D. alatus develops through a progressive transformation of its structure and architecture that adds one new axis category stage after stage. Specific leaf area, blade shape index and leaf dissection index decreased whereas blade area, perimeter, length, width and blade dry weight increased. Leaf traits correlated more strongly with ontogenetic stage than with light intensity. Our results demonstrated that studies on the responses of leaf traits to the environment may need to be corrected for an ontogeny effect. To strengthen this conclusion, future work should evaluate leaf variations during the ontogeny of different axis orders and/or axis categories.     

Genetic diversity in traditional Ethiopian highland maize accessions assessed by AFLP markers and morphological traits

In the highland regions of Ethiopia the heterogeneity of the land, the climate, and soil favors the presence of a large number of landraces. We analyzed a representative sample of 62 traditional Ethiopian highland maize accessions, using amplified fragment length polymorphism (AFLP^®) markers and morphological traits with the aim to group the accessions based on the their genetic profiles and morphological traits, to study agroecological variation and to assess the level of correlation between phenotypic and genetic distances. Eight EcoRI/MseI primer combinations and 15 morphological traits were used. The accessions varied significantly for all of the measured morphological traits. Of a total of 650 AFLP markers that were scored, 89.5% were polymorphic. Pair-wise genetic distance estimates based on AFLP data revealed dissimilarity coefficients ranging from 0.32 to 0.69 (mean of 0.57). Cluster analysis of the AFLP data grouped most accessions collected from the Northern highlands into one major cluster. It, however, failed to separate the Western and Southern accessions into different clusters. Regardless of the large variation in environmental conditions between agroecologies, only 9% of the total genetic variation was found between agroecologies, whereas 91% was found within agroecologies in Ethiopia. This finding may be explained by long distance seed exchange, continuous seed introduction and gene flow between agroecologies. The relationship between morphological and AFLP-based distances was significant and positive. Based on this study, three groups of highland accessions, with distinctive genetic profiles and morphological traits were identified. This information will be useful for further collections and conservation of the unique diversity included in the highland maize landraces of Ethiopia.     

Morphological variability of Undaria pinnatifida (Harvey) Suringar, 1873 (Phaeophyceae, Laminariales) in Peter the Great Bay, Sea of Japan

The morphological variability of the brown alga Undaria pinnatifida was studied in Peter the Great Bay (Sea of Japan). Comparison of the morphology of algae growing at eight sites of the bay that have different water movement revealed three phenotypically different population groups. A discriminant analysis of a set of morphological traits demonstrated that the width of the undivided part of the lamina and the stipe length make the most significant contribution to the discrimination of the detected groups. Analysis of the environmental dependence of morphological traits showed that lamina width and width of undivided part of lamina depend generally on water movement intensity, whereas the stipe length and thallus length are influenced by light intensity and surf degree.     

QTL Analysis for Flag Leaf Characteristics and Their Relationships with Yield and Yield Traits in Rice

Photosynthesis of carbohydrate is the primary source of grain yield in rice (Oryza sativa L.). It is important to genetically analyze the morphological and the physiological characteristics of functional leaves, especially flag leaf, in rice improvement. In this study, a recombinant inbred population derived from a cross between an indica (O. sativa L. ssp. indica) cultivar and a japonica (O. sativa L. ssp. japonica) cultivar was employed to map quantitative traits loci (QTLs) for the morphological (i.e., leaf length, width, and area) and physiological (i.e., leaf color rating and stay-green) characteristics of flag leaf and their relationships with yield and yield traits in 2003 and 2004. A total of 17 QTLs for morphological traits (flag leaf length, width, and area), 6 QTLs for degree of greenness and 14 QTLs for stay-green-related traits (retention-degrees of greenness, relative retention of greenness, and retention of the green area) were resolved, and 10 QTLs were commonly detected in both the years. Correlation analysis revealed that flag leaf area increased grain yield by increasing spikelet number per panicle. However, the physiological traits including degree of greenness and stay-green traits were not or negatively correlated to grain yield and yield traits, which may arise from the negative relation between degree of greenness and flag leaf size and the partial sterility occurred in a fraction of the lines in this population. The region RM255-RM349 on chromosome 4 controlled the three leaf morphological traits simultaneously and explained a large part of variation, which was very useful for genetic improvement of grain yield. The region RM422-RM565 on chromosome 3 was associated with the three stay-green traits simultaneously, and the use of this region in genetic improvement of grain yield needs to be assessed by constructing near-isogenic lines.     

Contrasting Effects of Long-Term Grazing and Clipping on Plant Morphological Plasticity: Evidence from a Rhizomatous Grass

Understanding the mechanism of plant morphological plasticity in response to grazing and clipping of semiarid grassland can provide insight into the process of disturbance-induced decline in grassland productivity. In recent studies there has been controversy regarding two hypotheses: 1) grazing avoidance; and 2) growth limiting mechanisms of morphological plasticity in response to defoliation. However, the experimental evidence presented for the memory response to grazing and clipping of plants has been poorly reported. This paper reports on two experiments that tested these hypotheses in field and in a controlled environment, respectively. We examined the effects of long-term clipping and grazing on the functional traits and their plasticity for Leymus chinensis (Trin.) Tzvelev (the dominate species) in the typical-steppe grassland of Inner Mongolia, China. There were four main findings from these experiments. (i) The majority of phenotypic traits of L. chinensis tended to significantly miniaturize in response to long-term field clipping and grazing. (ii) The significant response of morphological plasticity with and without grazing was maintained in a hydroponic experiment designed to remove environmental variability, but there was no significant difference in L. chinensis individual size traits for the clipping comparison. (iii) Plasticity indexes of L. chinensis traits in a controlled environment were significantly lower than under field conditions indicating that plants had partial and slight memory effect to long-term grazing. (iv) The allometry of various phenotypic traits, indicated significant trade-offs between leaf and stem allocation with variations in plant size induced by defoliation, which were maintained only under grazing in the hydroponic controlled environment experiment. Taken together, our findings suggest that the morphological plasticity of L. chinensis induced by artificial clipping was different with that by livestock grazing. The miniaturization of plant size in long-term grazed grassland may reflect retained characteristics of dwarf memory for adaptation to long-term grazing by large herbivores.     

Morphological and molecular genetic variations of oat genotypes grown in Kermanshah,Iran

Morphological traits and molecular markers are two common methods for genetic variation studies. Molecular markers, morphological traits methods and relationship between the two were used to study genetic variation among 43 oat genotypes and varieties. For this purpose, an augmented design was conducted in three replicates at 2008–2009 cropping season in the experimental field of Campus of Agriculture and Natural Resources of Razi University, Kermanshah, Iran. Four wild oat accessions (Avena sterilis) were added to evaluated genotypes in molecular experiment. Results showed a significant variation among genotypes for all morphological traits and they were classified based on this variation in four groups by WARD cluster analysis. In molecular experiment, 28 inter simple sequence repeat (ISSR) primers amplified 206 polymorph bands. Based on Jaccard similarity matrix, similarity among genotypes was varied from 0.23 to 0.66 and cluster analysis classified genotypes in seven groups by complete linkage method. The correlation between ISSR marker and morphological traits classifications was not significant. ISSR showed to be a helpful marker for genotype identity and separation as it put wild accessions in a group.     

内蒙古中东部草原不同生境克氏针茅（Stipa krylovii Roshev.）种群的形态差异分析

对内蒙古中东部草原分布的克氏针茅进行了种群内和种群间的形态差异分析。结果表明：（1）种群内不同的形态性状存在不同程度的差异;（2）种群间生殖枝中的一部分形态性状以及营养枝高度和营养枝干重均存在较大差异;（3）每穗小花数、每穗籽粒数、每穗小穗数、生殖枝高、穗干重、生殖枝干重、每穗种子重、种子重/生殖枝重比等性状的变化趋势相同,均表现为随生境条件变差而增加,即在生殖上投入增加以增大适合度;（4）主成分分析表明穗长、种子重/生殖枝重比、每穗小花数、每穗籽粒数、生殖枝高、基盘长、第二芒柱长、营养枝高、千粒重、芒针长是不同种群差异的主要指标。这些形态性状的差异可以看作克氏针茅对不同生境的适应性表现。     

Trait‐based adaptability of Phragmites australis to the effects of soil water and salinity in the Yellow River Delta

Phragmites australis is the dominant species in the Yellow River Delta and plays an important role in wetland ecosystems. In order to evaluate the relationship between phenotypic variation and environmental factors, explore how functional traits respond to changes in electrical conductivity and soil water content, and reveal the ecological strategies of P. australis, we investigated the ecological responses of P. australis to soil properties based on 96 plots along the coastal–inland regions in the Yellow River Delta of China. Within the range of soil water content (SWC, 9.39%–36.92%) and electrical conductivity (EC, 0.14–13.29 ms/cm), the results showed that (a) the effects of salinity were more important than the soil water content for the characterization of the morphological traits and that plant functional traits including leaf traits and stem traits responded more strongly to soil salinity than soil water content; (b) compared with morphological traits such as average height and internode number, physiological traits such as SPAD value, as well as morphological traits closely related to physiological traits such as specific leaf area and leaf thickness, showed stronger stability in response to soil water and salinity; and (c) under the condition of high electrical conductivity, P. australis improved its water acquisition ability by increasing indicators such as leaf water content and leaf thickness. In addition, with the increase in plant tolerance to stress, more resources were used to resist external stress, and the survival strategy was inclined toward the stress tolerator (S) strategy. Under low EC conditions, P. australis increased specific leaf area and leaf area for its growth in order to obtain resources rapidly, while its survival strategy gradually moved toward the competitor (C) strategy.     

Morphology, sociality, and ecology: can morphology predict pairing behavior in coral reef fishes?

Morphology can contain valuable information about the ecological performance of reef fishes, but it has rarely been used in combination with social traits. Social behavior is known to influence the ecological role of fishes; however, the ecological basis for pairing in reef fishes is not well understood. Field observations of 2,753 individuals, in 47 species in six families of biting reef fishes (Acanthuridae, Chaetodontidae, Kyphosidae, Labridae, Pomacanthidae, Siganidae), were used in combination with six morphological measurements, to examine the morphology of fishes in different social systems. A principal components analysis of morphological traits segregated species with high proportions of pairing individuals from non-pairing species along principal component 1, explaining 40.8 % of the variation. Pairing species were characterized by large eyes, concave foreheads, pointed snouts, deep bodies, and small maximum sizes. There was a significant positive relationship between these morphological traits (i.e., scores on PC1) and the prevalence of pairing within the Chaetodontidae (r ² = 0.59; P = 0.026), Siganidae (r ² = 0.72; P = 0.004), and Acanthuridae (r ² = 0.82; P < 0.001). This was consistent when traits were corrected for phylogenetic effects. No pattern was evident in the scarine Labridae (r ² = 0.15; P = 0.17). The morphological characteristics found among pairing species suggest that pairing species share common ecological traits, including foraging for small prey items in micro-topographically complex environments such as reef crevices. These ecological traits may have played a role in the evolution of pairing behavior and subsequently led to the development of reproductive patterns based on monogamy.     

Mapping of chromosome loci determined manifestation of morphological and biochemical traits of quality in <Emphasis Type="Italic">Brassica rapa</Emphasis> L. crops

Mapping chromosomal loci responsible for seven morphological and five biochemical traits of quality in populations of doubled haploid lines of leafy, rooted, and oilseed crops of Brassica rapa L. species was performed for the first time. In total, 140 quantitative trait loci (QTLs) were mapped determining manifestation of the studied morphological and biochemical economically important traits of quality under field and greenhouse conditions; the control of the same traits under different growth conditions or simultaneous control of several traits by one locus were noted. Molecular markers genetically linked with the selected QTLs were detected. The block genomic structure of the genetic components (chromosomal loci and linkage groups) determining manifestation of morphological and biochemical traits of quality that make B. rapa plants nutritionally valuable is discussed for the first time. The conclusion is drawn that the revealed QTLs and identified molecular markers may be of interest for the forthcoming study of genetic control by them of economically important characters and for the marker assisted selection in B. rapa.