Patterns of variation in wing morphology in the cactophilic Drosophila buzzatii and its sibling D. koepferae

Drosophila buzzatii and D. koepferae are two sibling species that breed on the necrotic tissues of several cactus species and show a certain degree of niche overlap. Also, they show differences in several life history traits, such as body size and developmental time, which probably evolved as a consequence of adaptation to different host plants. In this work we investigate the ecological and genetic factors affecting wing morphology variation both within and between species. Three wing traits were scored, distal and proximal wing length and width in isofemale lines reared in two of the most important host cacti: Opuntia sulphurea and Trichocereus terschekii. Our results revealed that differences between species and sexes in wing size and shape were significant, whereas the cactus factor was only significant for wing size. Intraspecific analyses showed that differences among isofemale lines were highly significant for both size and shape in both species, suggesting that an important fraction of variation in wing morphology has a genetic basis. Moreover, the line by cactus interaction, which can be interpreted as a genotype by environment interaction, also accounted for a significant proportion of variation. In summary, our study shows that wing size is phenotypically plastic and that populations of D. buzzatii and D. koepferae harbour substantial amounts of genetic variation for wing size and shape. Interspecific differences in wing size and shape are interpreted in terms of spatial predictability of the different host plants in nature.     

库蠓亚属6种库蠓翅形变化分析及亲缘关系探讨

本研究基于几何形态学测量分析方法对库蠓亚属6种库蠓(渐灰库蠓Culicoides grisescens、光胸库蠓Culicoides impunctatus、日本库蠓Culicoides nipponensis、灰黑库蠓Culicoides pulicaris、刺螫库蠓Culicoides punctatus、天祥库蠓Culicoides tienhsiangensis)的翅脉形态进行定量分析,以探讨6种库蠓种间的亲缘关系。首先对库蠓亚属6种、24头标本翅脉使用几何形态学标点法标记12个地标点,并通过普氏叠加、典型变量分析法、聚类分析法等对6种库蠓翅进行比较分析以及种间亲缘关系探讨。结果显示,翅形变化与亲缘关系结果一致,即6种库蠓的翅形均有显著差异(P<0.05),其中光胸库蠓和渐灰库蠓的翅形变化最小,亲缘关系最近,而日本库蠓与其他5种库蠓的翅形变化最大,亲缘关系最远。初步证实了几何形态学能够做为一种辅助工具对蠓科昆虫进行种间关系及分类研究。     

Selection on Wing Allometry in Drosophila Melanogaster

Five bivariate distributions of wing dimensions of Drosophila melanogaster were measured, in flies 1) subjected to four defined environmental regimes during development, 2) taken directly from nature in seven U.S. states, 3) selected in ten populations for change in wing form, and 4) sampled from 21 long inbred wild-type lines. Environmental stresses during development altered both wing size and the ratios of wing dimensions, but regardless of treatment all wing dimensions fell near a common allometric baseline in each bivariate distribution. The wings of wild-caught flies from seven widely separated localities, and of their laboratory-reared offspring, also fell along the same baselines. However, when flies were selected divergently for lateral offset from these developmental baselines, response to selection was rapid in every case. The mean divergence in offset between oppositely selected lines was 14.68 SD of the base population offset, after only 15 generations of selection at 20%. Measurements of 21 isofemale lines, founded from wild-caught flies and maintained in small populations for at least 22 years, showed large reductions in phenotypic variance of offsets within lines, but a large increase in the variance among lines. The variance of means of isofemale lines within collection localities was ten times the variance of means among localities of newly established wild lines. These observations show that much additive genetic variance exists for individual dimensions within the wing, such that bivariate developmental patterns can be changed in any direction by selection or by drift. The relative invariance of the allometric baselines of wing morphology in nature is most easily explained as the result of continuous natural selection around a local optimum of functional design.     

Quantitative variation of four morphological traits in Drosophila melanogaster under larval crowding

Effects of three different larval densities (low, intermediate and high) on phenotypic and genetic variation of four morphological traits (thorax and wing length, sternopleural and abdominal bristle number) were studied in Drosophila melanogaster using the isofemale line technique. Phenotypic variation was found to increase at high larval density in all traits examined. Environmental variance for three traits (exception was sternopleural bristle number) and fluctuating asymmetry for both bilateral traits were also increased under high density conditions. For estimates of genetic variability (among isofemale lines variance, heritability and evolvability), no statistically significant differences among density regimes were detected. However, the trends in changes of these estimates across densities indicated a possibility for enhanced genetic variation under larval crowding for all traits except abdominal bristle number. For the latter trait, genetic variation seemed not to be dependent on density regime. Generally, two metric traits (thorax and wing length) were more affected by larval crowding than two meristic ones (sternopleural and abdominal bristle number). The Results are in complete agreement with those previously obtained for D. melanogaster using extreme temperatures as stress-factors.     

Geometric morphometric analysis of Eysarcoris guttiger,E.annamita and E.ventralis (Hemiptera: Pentatomidae)

The genus Eysarcoris can be easily distinguished from other genera through the two spots in the basal angle of the scutellum.Nevertheless,Eysarcoris species show complex variances.Geometric morphometric methods have been increasingly applied to distinguish species and to define the boundary of genera among insects.In the present study,geometric morphometric approach was firstly employed to evaluate the shape variation of three characters (fore wing,hind wing and pygophore) of E.guttiger,E.annamita and E.ventralis using E.aeneus as outgroup to ascertain whether this approach is a reliable method for the taxonomy of Eysarcoris.Analysis was conducted on the landmarks of the three characters of these species.Multivariate regression of procrustes coordinates against centroid size was conducted to test the presence of allometry.Principal component analysis (PCA),canonical variate analysis (CVA)and cluster analysis were utilized to describe variations in shapes among the studied species.For all of the three characters,though PCA analysis showed some overlap among species,p-values for procrustes distance and mahalanobis distance were all less than 0.0001.The distribution of the three studied species corresponds with their species status.This study demonstrates that the geometric morphometrics of both the fore wing and the hind wing might represent a possible tool for the identification of species within this genus.     

A Geometric Morphometric Approach to the Analysis of Lip Shape during Speech: Development of a Clinical Outcome Measure

Objective assessments of lip movement can be beneficial in many disciplines including visual speech recognition, for surgical outcome assessment in patients with cleft lip and for the rehabilitation of patients with facial nerve impairments. The aim of this study was to develop an outcome measure for lip shape during speech using statistical shape analysis techniques. Lip movements during speech were captured from a sample of adult subjects considered as average using a three-dimensional motion capture system. Geometric Morphometrics was employed to extract three-dimensional coordinate data for lip shape during four spoken words decomposed into seven visemes (which included the resting lip shape). Canonical variate analysis was carried out in an attempt to statistically discriminate the seven visemes. The results showed that the second canonical variate discriminated the resting lip shape from articulation of the utterances and accounted for 17.2% of the total variance of the model. The first canonical variate was significant in discriminating between the utterances and accounted for 72.8% of the total variance of the model. The outcome measure was created using the 95% confidence intervals of the canonical variate scores for each subject plotted as ellipses for each viseme. The method and outcome model is proposed as reference to compare lip movement during speech in similar population groups.     

Latitudinal shifts in body size of Enallagma cyathigerum (Odonata)

Aim Survey of the latitudinal body size pattern for populations of Enallagma cyathigerum (Odonata) across a south‐north transect. Location A transect covering the whole distribution range from south to north across Europe was sampled. Methods Newly emerged adults were collected from five major sites across Europe and one to four localities were sampled within each site. In total 253 adults were collected from fourteen localities. Body size was measured using thorax length, length of right front wing and length of right hind tibia. These body size estimates were thereafter related to latitude and mean temperature in January and July. Results Body size showed a U‐shaped pattern with latitude, being large at low and high latitudes and small at intermediate latitudes. The same U‐shaped pattern was found for mean January and July temperature, with large animals at low and high temperatures. Conclusion The U‐shaped relationship between body size and latitude is suggested to be a combination of two effects: (1) the length of the season favourable for growth and development, and (2) variation in life cycle length with latitude.     

Morphometric variability on a microgeographical scale in two inshore seabirds

The range of morphometric variation among two inshore seabirds, the Kerguelen cormorant Phalacrocorax atriceps verrucosus and the gentoo penguin Pygoscelis papua was investigated on the Kerguelen and Crozet archipelagos. Four measurements (weight, culmen, flipper and body length for gentoos; weight, culmen, wing and tarsus length for cormorants) were compared in the same seven localities on Kerguelen. In addition, culmen and wing length were compared in four localities on Crozet for gentoo penguins. The two species exhibited similar trends in variation within Kerguelen although the range of variation was greater in cormorants. Compared to the largest cormorants and gentoo of Kerguelen, the smallest stocks were 60 and 36% of the weights, respectively. There is little evidence elsewhere of similar broad morphometric variability in seabirds on a microgeographical scale. The variation observed did not follow a regular cline and appeared to be less related to physical factors (sea temperatures) than to the local biological factors during the breeding season.     

Morphological and genetic variability of Snow Petrels Pagodroma nivea

Snow Petrels vary greatly in size, the largest birds being over twice as heavy as the smallest. Two morphological types have been distinguished, but the Adelieland population alone shows continuous variation spanning the complete range of the species. There is no evidence of size-related assortative mating. Size variation cannot be attributed to nest site location nor to individual age, nor is the large sexual dimorphism sufficient to account for the situation. The great individual variation in size is accompanied by an important genetic variability as revealed by a starch gel electrophoretic study.
Comparisons of wing length data from many localities around the Antarctic Continent suggest a cline, with a peak at the Balleny Islands. Taking into account the extent of size variation at breeding places only, it appears that morphological variability is low at Balleny Islands, where only large form birds are known, and at Antarctic localities inhabited exclusively by small form birds. In contrast, several localities on the continent in the vicinity of Balleny Islands, and especially Adelieland, exhibit a high degree of variability.
We propose that during the quaternary glaciations, the Antarctic continent provided no refugia for this species, but, when glaciation ended, Snow Petrels re-established colonies, coming from two refugia, a low latitude one inhabited by small birds Pagodroma nivea nivea , and one in higher latitudes characterized by large birds P. n. major. Reproductive isolation broke down, and a hybridization zone was created. The high proportion of polymorphic loci exhibited by the Adelieland population is then attributable to the mixing of two previously isolated genetic pools.     

Multidimensional analysis of Drosophila wing variation in Evolution Canyon

Environmental stress has been suggested to be a major evolutionary force, both through inducing strong selection and because of its direct impact on developmental buffering processes that alter the evolvability of organisms. In particular, temperature has attracted much attention because of its importance as an ecological feature and the relative ease with which it can be experimentally manipulated in the lab. Evolution Canyon, Lower Nahal Oren, Israel, is a well studied natural site where ecological parameters are suspected to drive evolutionary differentiation. In this study, using Drosophila melanogaster isofemale lines derived from wild flies collected on both slopes of the canyon, we investigated the effect of developmental temperature upon the different components of phenotypic variation of a complex trait: the wing. Combining geometric and traditional morphometrics, we find only limited evidence for a differentiation among slopes. Investigating simultaneously phenotypic plasticity, genetic variation among isofemale lines, variation among individuals and fluctuating asymmetry, we could not identify a consistent effect of the stressful conditions encountered on the south facing slope. The prevailing structuring effect is that of the experimentally manipulated temperature which clearly influences wing mean size and shape. Variability, in contrast, is not consistently affected by temperature. Finally, we investigated the specific relationship between individual variation and fluctuating asymmetry. Using metric multi-dimensional scaling we show that the related patterns of wing shape variation are not identical, supporting the view that the underlying developmental processes are to a certain extent different.     

Variation in leaves and petals of Ranunculus subgenus Batrachium on the Aegean Islands, analysed by multivariate analysis

Variation in leaves and petals was studied using canonical variate and cluster analyseS. In total of 33 populations from eight islands leaf variation in laminar, floating leaves was studied in Ranunculus peltatus subsp. peltatus, subsp. baudotii and subsp. saniculifolius, and in R. Tripartitus, all of which are heterophyllous in the Aegean area. The petal data set also included the homophyllous R. Trichophyllus. The results of the analyses show that the populations are morphologically well differentiated and that petal shape has a tendency to discriminate between the populations somewhat better than leaf shape does. There was no basis for subdividing the populations into taxa on these characterS. Analyses showed that up to 70% of the variation was distributed between populations, the remainder (5–18%) being within populations.     

Levels of variation in stress resistance in drosophila among strains, local populations, and geographic regions: patterns for desiccation, starvation, cold resistance, and associated traits

Stress resistance traits in Drosophila often show clinal variation. Although these patterns suggest selection, there is generally no attempt to test how large differences at the geographical level are relative to levels of variation within and between local populations. Here we compare these levels in D. melanogaster from temperate Tasmania versus tropical northern Queensland by focusing on adult resistance to desiccation, cold and starvation stress, as well as associated traits (size, lipid content). For starvation and desiccation resistance, levels of variation were highest among strains from the same population. whereas there was little differentiation among local populations and a low level of differentiation at the geographic level. For adult cold resistance, there was local differentiation and strain variation but no geographic variation. For size (thorax length), geographic differentiation was higher despite some overlap among strains from the tropical and temperate locations. Finally, for lipid levels there was only evidence for variation among strains. The low level of differentiation among geographic locations for stress resistance was further verified with the characterization of isofemale strains from 18 locations along a coastal transect extending from Tasmania to northern Queensland. Crosses among some of the isofemale strains showed that results were not confounded by inbreeding effects. Strains derived from a cross between a tropical and temperate strain differed for all traits, and variation among strains for body size was higher than strain variation within the geographic regions. Unlike in previous studies, lipid content and starvation resistance were not correlated in any set of strains, but there was a correlation between cold resistance and lipid content. There was also a correlation between desiccation resistance and size but only in the geographic cross strains. These findings suggest a large amount of variation in stress resistance at the population level and inconsistent correlation patterns across experimental approaches.     

Wing shape as a potential discriminator of morphologically similar pest taxa within the Bactrocera dorsalis species complex (Diptera: Tephritidae)

Four morphologically cryptic species of the Bactrocera dorsalis fruit fly complex (B. dorsalis s.s., B. papayae, B. carambolae and B. philippinensis) are serious agricultural pests. As they are difficult to diagnose using traditional taxonomic techniques, we examined the potential for geometric morphometric analysis of wing size and shape to discriminate between them. Fifteen wing landmarks generated size and shape data for 245 specimens for subsequent comparisons among three geographically distinct samples of each species. Intraspecific wing size was significantly different within samples of B. carambolae and B. dorsalis s.s. but not within samples of B. papayae or B. philippinensis. Although B. papayae had the smallest wings (average centroid size=6.002 mm±0.061 SE) and B. dorsalis s.s. the largest (6.349 mm±0.066 SE), interspecific wing size comparisons were generally non-informative and incapable of discriminating species. Contrary to the wing size data, canonical variate analysis based on wing shape data discriminated all species with a relatively high degree of accuracy; individuals were correctly reassigned to their respective species on average 93.27% of the time. A single sample group of B. carambolae from locality 'TN Malaysia' was the only sample to be considerably different from its conspecific groups with regards to both wing size and wing shape. This sample was subsequently deemed to have been originally misidentified and likely represents an undescribed species. We demonstrate that geometric morphometric techniques analysing wing shape represent a promising approach for discriminating between morphologically cryptic taxa of the B. dorsalis species complex.     

A multivariate analysis of geographic variation in morphology inPersoonia mollis (Proteaceae)

This study highlights the utility and limitations of multivariate procedures for the analysis and taxonomic interpretation of geographic variation within a complex species. Morphological data from herbarium specimens ofPersoonia mollis (Proteceae) were analyzed by two ordination procedures, multidimensional scaling (MDS) and canonical variate analysis (CVA). Support for the earlier recognition of nine infraspecific taxa was found. However, plots of canonical variate scores against latitute or longitude across parapatric subspecies boundaries revealed unexpected phenetic homogeneity within subspecies with sharp transitions between subspecies. The geographic variation in morphology withinP. mollis can be best described as a mosaic of nine recognisably distinct and internally homogeneous allopatric or parapatric taxa, where neighbouring taxa share surprisingly narrow zones of morphological transition at ecotones. The utility of these procedures was highlighted by the recovery of unexpected patterns of variation. However, questions of rank remain unresolved by phenetic analysis alone.     

GENETIC VARIATION IN CACTOPHILIC DROSOPHILA FOR OVIPOSITION ON NATURAL YEAST SUBSTRATES

Theory predicts that environmental heterogeneity in space or in time can maintain genetic polymorphism. Stable polymorphisms are expected to be more readily maintained if there are genotype specific habitat preferences. Genotype specific preferences for oviposition sites in Drosophila could be a major factor promoting habitat selection, and thus the maintenance of genetic variation. This hypothesis is being tested using the cactophilic species, D. buzzatii and D. aldrichi, where available evidence indicates a potential for such habitat selection, the habitats (oviposition sites) being yeast species found in the natural environment of these flies (cactus rots). Genetic variation for oviposition preferences was tested using isofemale lines—for D. buzzatii, a total of 60 lines from seven localities widely distributed through the species range in Australia, and for D. aldrichi, 21 lines from three of these localities. Females were given a choice of five yeast species as oviposition sites. Genetic variation for oviposition preferences on these natural substrates was demonstrated. There was significant variation among isofemale lines within populations in their patterns of preferences for oviposition on the five yeast species. However, analyses of preferences for each yeast species separately showed that the genetic variation for preferences relates to only three of the five species. Heritabilities of individual female preferences for these three species were low, ranging up to 9%. Little geographic differentiation was apparent among populations, most likely due to similar selection regimes within each population. Within populations, this kind of habitat selection could act to maintain polymorphisms, both at loci determining the habitat preferences and at other loci in linkage disequilibrium with them.     

Morphological variation in diadromous and landlocked populations of the spotted galaxias,Galaxias truttaceus,in Tasmania,south-eastern Australia

Synopsis A comparison of a suite of morphometric measurements and meristic counts of individuals of two landlocked lacustrine and two diadromous riverine populations of Galaxias truttaceus was carried out utilising both univariate and canonical variate analyses. Lacustrine fish had fewer dorsal and anal fin rays than did riverine fish. Differences were not as clear for gill rakers and vertebrae. Comparisons of serial counts were made with two derived lacustrine species, G. auratus and G. tanycephalus, also from Tasmania. Lacustrine G. truttaceus varied in the same direction as the derived species, relative to riverine G. truttaceus. From an analysis of 12 body measurements, the first canonical variate clearly separated lacustrine fish from riverine fish largely based on measurements associated with fins (pre-anal fin length, length of anal base, pre-dorsal fin length, maximum length of dorsal fin and inter-orbital width). An overall value for the correct classification of fish into groups based on locality was 84%. The percentage of fish classified into the wrong habitat (lake or stream) was much less than the percentage classified between localities within habitats. Overall morphological variation was greater between than within habitats. It is suggested that the differences in water movement and food type may in part account for the differences shown and that selective pressures peculiar to the lacustrine environment may be causing the lake populations to diverge from the riverine populations.     

Phenotypic stability of the P-M system in wild populations of Drosophila melanogaster

The P element appears to be one of the most recently invaded transposons of D. melanogaster. To study the dynamics and long-term fate of P elements in natural populations of D. melanogaster, 472 isofemale lines newly collected from 27 localities of Japan were examined for the P element-associated characteristics (abilities to induce and repress of P element transposition) and genomic P element composition (size classes and their numbers). There was variation in the P element-related phenotypes among local populations, but genomic P composition did not correlate strongly with the phenotype of each line: full-size P and KP elements predominated in their genomes (FP+ KP predominance). Comparison with previous results suggests a stability in the P-M system in local populations over about 15 years. In some populations, phenotypic stability for particularly long times was found: for 30 years or more Q strains predominated in Hikone and Tanushimaru, P or Q strains around Inakadate, and M' or Q strains around Tozukawa. There was no clear evidence of structural destruction underlying functional variation of P elements during this period. These results suggest that the current evolutionary status of P elements in the gene pool of D. melanogaster is not intermediary stage predicted by the original recent invasion hypothesis, and that several other factors such as the position effect play important roles.     

Discrimination of Eubazus (Hymenoptera, Braconidae) sibling species using geometric morphometrics analysis of wing venation

Abstract.  Complexes of sibling and cryptic species are encountered frequently in parasitic Hymenoptera. Geometric morphometrics is a useful tool to detect minimal morphological variations, which often are undetectable by traditional morphological studies and even by classical morphometric approaches. We applied geometric morphometrics to wing venation to assess a complex case of sibling species in the genus Eubazus (Hymenoptera, Braconidae), parasitoids of conifer bark weevils of the genus Pissodes (Coleoptera, Curculionidae). The results and methods were compared with previous taxonomic studies on the same species, involving classical multivariate morphometrics, isoenzyme analyses, cross-mating experiments and biological observations. Geometric morphometrics confirmed the previous division into four distinct species. However, this approach enabled the four species to be separated simultaneously, with a reliability of 98.6% for well-classified females and 93.1% for males. A similar result in previous studies was obtained only by combining isoenzyme analyses and several canonical variate analyses, including many morphometric characters. Furthermore, measurements of wing venation were less time-consuming, more reliable and required less prior knowledge of braconid taxonomy than the measurements needed for the classical morphometrics methods. Geometric morphometrics was used also to test the effect of host species on wing shape. Several female populations of Eubazus semirugosus originating from three different Pissodes spp. were compared. Significant differences were found in wing shape between conspecific Eubazus from different host species. The results are discussed in relation to reproductive isolation and genetic flow between the four species.     

Population differentiation in scalation of the Iberian rock lizard (Lacerta monticola)

The rock lizard Lacerta monticola is a climatic relict confined to three mountain ranges and one coastal region of the Iberian peninsula. Scalation was studied in specimens from localities encompassing all major parts of its range. Population differentiation was analysed using analyses of variance, z-transformations of within-locality character-correlations, canonical variate analyses, principal components analysis, and matrix association tests. Substantial differentiation was found between Pyrenees specimens and those from other localities, supporting a recent study which indicated that the Pyrenees populations could represent a different species. Considerable geographic variation is also found among populations from the remaining parts of its range, the pattern of which is not compatible with existing racial categories designated for this species.     

Causes of variation in wing loading among Drosophila species

We examined influences on wing and body size in 11 species (12 strains) of Drosophila. Six measures of wing length and width were closely correlated with wing area and suggested little variation in wing shape among the species. Among ten species wing loading, an important factor in flight costs and manoeuvrability, increased as body mass increased at a rate consistent with expectations from allometric scaling of wing area and body mass to body length. Intraspecific variation in wing loading showed similar relationships to body mass. Density and temperature during larval development influenced wing loading through general allometric relations of body size and wing area. Temperature during the pupal stage, but not during wing hardening after eclosion, influenced wing area independently of body size. Wing area increased as growth temperature decreased. Individuals reared at cooler temperatures thus compensated for a potential allometric increase in wing loading by differentially enlarging the wing area during pupal development.