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Intraspecific Variation and Asymmetry in Wing Shape of Dolichopodid Flies (Diptera, Dolichopodidae)

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Intraspecific Variation and Asymmetry in Wing Shape of Dolichopodid Flies (Diptera, Dolichopodidae) International Journal of Entomology Research ISSN: 2455-4758 Impact Factor: RJIF 5.24 www.entomologyjournals.com Volume 2; Issue 5; September 2017; Page No. 10-20 Intraspecific variation and asymmetry in wing shape of dolichopodid flies (Diptera, Dolichopodidae) Mariya A Chursina Department of Ecology and Zoology of Invertebrates, Voronezh State University, Voronezh, Russia Abstract Intraspecific variation in wing shape of six species of the Dolichopodidae family was analyzed by the method of Procrustes superimposition. Specifically, we examined the patterns of sexual dimorphism, interpopulation variation, variation among individuals, directional asymmetry and fluctuating asymmetry (within-individual variability). The results suggest that sexual dimorphism is the most significant sources of intraspecific variation. Highly significant differences in wing shape were observed between populations of Poecilobothrus regalis, it reflects their adaptations to different environmental conditions. The analysis revealed significant directional asymmetry for wing shape of Poecilobothrus regalis from one of the studied populations; directional asymmetry had a significant allometric component. Four types of pattern of individual variation were discovered. It was shown that individual variation and fluctuating asymmetry have similar patterns. Based on previous studies of wing development, we made conclusions about the developmental processes, which cause different types of intraspecific variation. Keywords: diptera, dolichopodidae, interpopulation variation, sexual dimorphism, fluctuating asymmetry, directional asymmetry 1. Introduction Drosophila melanogaster subgroup [11]. Characters of the wing morphology have significant Fluctuating asymmetry (FA) has been defined as a random, importance in the systematic and taxonomy of the non-directional deviation from the perfect symmetry of the Dolichopodidae family [1, 2]. Together with the structure of bilateral traits [12]. In recent studies it has been shown that male genitalia, wing morphology gives more information on forms of fluctuating asymmetry are not inherited [13, 14]. It is the taxonomic position of dolichopodid flies than the other also suggested that FA is a result of small genetic perturbation parts of the body. The large interspecific variety of wing and environmental stressors during the ontogeny of an morphologies within the family suggests that flight plays an organism [15, 16]. The results of recent studies show FA is important role in the dolichopodid’s survival by providing stress, trait, and population specific [17, 18, 19]: in addition, FA varying levels of maneuverability and velocity. Differences in showed low repeatability. the model of flight can be the force influencing the evolution Many studies have devoted more attention to one or another of other morphological structures. type of morphological variation among individuals (for The study of the variability of wings venation in all taxonomic instance, sexual dimorphism or FA): however, there are a few levels can provide data on the evolutionary path of a tax on [3, studies concerning the comparison of different types of 4, 5]. It is important to examine the range and degree of variation with each other. In our study we focused on intraspecific variations in order to study an adaptive optimum intraspecific variability of wing shape in species of the of a morphological trait [6]. In the first instance, patterns of Dolichopodidae family. sexual dimorphism, interpopulation variability, and various Wings of dolichopodid flies usually oval, about as long as types of asymmetries must be taken into consideration. body, hyaline, but sometimes with darkening spots. The Sexual dimorphism of wing shape is a feature of many species following veins are present (Fig. 1): costa (C), subcosta (Sc), a [7, 8, 9] of Diptera . It is obvious that differences exist because of radial sector with three branches (R1, R2+3, R4+5), a medial the various roles of flight in males and females, so sexual vein (M1+2), which sometimes forked near middle of last part shape dimorphism can be related to selective pressures of the and formed arcuate M1 (in the Sciapodinae subfamily), a habitat and sexual selection. Maneuverability traits may be single cubital vein (CuA1). Anal vein is short, sometimes subject to rapid evolution. There is an alternative explanation reduced or absent. for sexual shape dimorphism, related with the ability of males Most variation of wing structure is concentrated in a several to produce acoustical signals, but currently these facts have morphological traits: posterior region, including medial and not verified statistically. cubital veins and the posterior crossvein (dm-cu), and the Allometry is a subject of many studies as factor playing an anterior margin – costal vein and the point of its intersections [10] important role in sexual shape dimorphism . Allometric with R1 and R2+ 3. The characters can vary within subfamilies, relationships are central to the evolution of traits that are used genera and species. in courtships or in aggressive interactions with sexual rivals. Costal vein forms the anterior margin of the wing and it For example, allometric relationships have been discussed as experiencing the greatest air resistance during the flight. The an important component of wing sexual dimorphism in the costa reveals a trend toward thickening in the region of its 10 International Journal of Entomology Research meeting of vein R1. The second apparent form of variation is order to construct a robust phylogenetic hypothesis. To the variation of R2+3 length. Then the ratio of costal section implement this goal in this study we focused on following between the base of wing and R1 to the wing length is used as questions: generic character in the subfamilies Achalcinae and 1. Different types of intraspecific variability in wing shape of Dipahorinae. CuA1 and dm-cu are present stably, although the Dolichopodidae species were examined and the significant variations in its length and position of its meeting presence of a taxonomic significant of these traits was point are present. An interesting area for investigation is M2 tested. vein, which is present at Sciapodinae species and present as 2. We considered the questions of whether there are special rudimentary stub at some Dolichopodinae species. patterns in different types of intraspecific variability or if In the family Dolichopodidae, there are two most probable the same processes control their development. factors influencing the wing shape. First of all, dolichopodid 3. The role of allometry in each case was studied. flies are predators and widespread in various habitats, so flight performance is an important feature, which is responsible for 2. Materials and methods survivability. Type of habitat (open area of a shore or dense 2.1 Flies growth of damp area in a forest) acts on both flight A study of 6 species belonging to 4 genera and 3 subfamilies performance and wing shape. Secondly, for several species of of the Dolichopodidae family was conducted to examine the the Dolichopodidae family the courtship ceremonies are intraspecific variability in wing shape: Poecilobothrus regalis described. A male of Poecilobothrus nobilitatus L. stand in (Meigen, 1824), Poecilobothrus varicoloris (Becker, 1917), front or to on side of a female and its wings vibrate three or Dolichopus latilimbatus Macquart, 1827, Dolichopus more times. The milk-white spots of the apex of wings made ungulatus (Linnaeus, 1758) of the Dolichopodinae subfamily, specific visual image [20, 21, 22, 23]. Campsicnemus lumbatus Loew, 1857 of the Sympycninae Since the functional consequences of wing form determine the subfamily and Sciapus wiedemanni (Fallen, 1823) of the locomotors performance of flight then different performance Sciapodinae subfamily (Table 1). The specimens were requirements lead to different combinations of these traits. A collected from the natural environments in 2015-2016. clear understanding of the directions of changes is required in Table 1: Material examined. Species Collection sites and year for the species and number of specimens Population 1 Population 2 Campsicnemus lumbatus 20 ♀, 36 ♂, 2016 12 ♀, 32 ♂, 2016 55° 87´ N, 48° 67´ E 51° 81´ N, 39° 39´ E 58 ♀, 40 ♂, 2016 Dolichopus latilimbatus 55° 87´ N, 48° 67´ E 20 ♀, 36 ♂, 2016 Dolichopus ungulatus 55° 87´ N, 48° 67´ E Population 1 Population 2 Poecilobothrus regalis 40 ♀, 84 ♂, 2015 18 ♀, 58 ♂, 2015 49° 97´ N, 39° 68´ E 51° 85´ N, 40° 78´ E 34 ♀, 36 ♂, 2015 Poecilobothrus varicoloris 44° 19´ N, 39° 96´ E 43 ♀, 39 ♂, 2016 Sciapus wiedemanni 51° 81´ N, 39̊° 39´ E 2.2 Measurements and statistical analysis [24]. Wing shape was described by 9 landmarks located at vein Wing were placed on a glass slide and covered with a cover intersections with each other and wing margin (Fig. 1). Two- glass. All slides were photographed by using a Levenhuk C dimensional Casterian coordinates of the landmarks were NG microscopic camera and digitizes. Our analysis of digitized from the photo using tps Dig-2.32 software [25]. intraspecific variability was based on the Procrustes technique Fig 1: Landmarks used in the study (Dolichopus ungulatus, male). 11 International Journal of Entomology Research By definition FA is a very small deviation of organism sides dimorphism in two populations of Poecilobothrus regalis). If from the perfect symmetry, and measurements of FA are any two factors affect the same developmental processes, the highly sensitive to
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