Evolutionary relationships of wing venation and wing size and shape in Aphidiinae (Hymenoptera: Braconidae)

We explored evolutionary changes in wing venation and wing size and shape in Aphidiinae, one of the well-known groups of parasitic wasps from the family Braconidae. Forewings of 53 species from 12 genera were examined, for which a molecular phylogeny was constructed on the basis of the mitochondrial barcoding gene COI. By covering all types of wing venation within the subfamily Aphidiinae and by using landmark-based geometric morphometrics and phylogenetic comparative methods, we tested whether evolutionary changes in wing shape correlate to the changes in wing venation and if both changes relate to wing size. The relationship between wing morphology and host specificity has been also investigated. We found that six types of wing venation, with different degree of vein reduction, could be recognized. Wing venation type is largely genus specific, except in the case of maximal reduction of wing venation which could be found across examined Aphidiinae taxa. The reconstruction of evolutionary changes in wing venation indicates that evolutionary changes in wing shape are related to the changes in wing size, indicating that miniaturization play a role in evolution of wing morphology while host specialization does not affect the wing shape within the subfamily Aphidiinae.     

Change in the system of wing venation of Drosophila melanogaster under heat shock and selection

A change in the system of wing venation of Drosophila melanogaster appeared in response to heat shock and positive selection pressure directed to restoring the normal formation of wing radial vein, L2, that had been violated earlier by the recessive mutation of radius incompletes. Positive selection was effective, L2 having been formed correctly and completely to 35 generation. Besides, (+)-selection was accompanied by appearance of a small fragment of an additional vein at the wing tip. Selection directed to increase of size of this fragment resulted in the sufficient changes in the system of wing venation as a whole. It is suggested that, during evolution, transformation of wing venation of Drosophila was effected by the change of the way of prepattern realization, whereas the cells of wing plate continued to allow the formation of veins practically over a whole area.     

Dpp/BMP transport mechanism is required for wing venation in the sawfly Athalia rosae

The pattern of wing venation varies considerably among different groups of insects and has been used as a means of species-specific identification. However, little is known about how wing venation is established and diversified among insects. The decapentaplegic (Dpp)/bone morphogenetic protein (BMP) signaling pathway plays a critical role in wing vein formation during the pupal stages in Drosophila melanogaster. A key mechanism is BMP transport from the longitudinal veins (LVs) to the posterior crossvein (PCV) by the BMP-binding proteins, short gastrulation (Sog) and twisted gastrulation2/crossveinless (Tsg2/Cv). To investigate whether the BMP transport mechanism is utilized to specify insect wing vein patterns in other than Drosophila, we used the sawfly Athalia rosae as a model, which has distinct venation patterns in the fore- and hindwings. Here, we show that Ar-dpp is ubiquitously expressed in both the fore- and hindwings, but is required for localized BMP signaling that reflects distinct wing vein patterns between the fore- and hindwings. By isolating Ar-tsg/cv in the sawfly, we found that Ar-Tsg/Cv is also required for BMP signaling in wing vein formation and retains the ability to transport Dpp. These data suggest that the BMP transport system is widely used to redistribute Dpp to specify wing venation and may be a basal mechanism underlying diversified wing vein patterns among insects.     

Zoraptera wing structures: evidence for new genera and relationship with the blattoid orders (Insecta: Blattoneoptera)

Abstract. The order Zoraptera has traditionally been thought to contain only one family (Zorotypidae) and one genus ( Zorotypus Silvestri). An analysis of known zorapteran wings shows that the wing venation contains character sets indicative of the existence of seven genera: Zorotypus, Brazilozoros gen.n., Centrozoros gen.n., Floridazoros gen.n., Latinozoros gen.n., Meridozoros gen.n. and Usazoros gen.n. The wing venation of Meridozoros leleupi (Weidner) from the Galapagos Islands, Ecuador and Venezuela is described here for the first time.
The major wing structures show that Zoraptera belong to the blattoid lineage. Head and abdomen characters indicate that Zoraptera probably diverged from the Blattoneoptera stock early, almost certainly before the (Protelytroptera + Dermaptera) line, and much before the (Isoptera + (Blattodea + Mantodea)) line. A homologized wing vein system is proposed for the Isoptera.
The homologized wing vein system is based on the hypothesis that the Pterygota originated with the development of protowings, which then diverged through separate but characteristic adaptations for flapping flight. Therefore the basic wing venation pattern is monophyletic, but the changes in wing musculature, articulation and basic braces between main veins are different in the major (super-ordinal) pterygote lineages (Pleconeoptera, Orthoneoptera, Blattoneoptera, Hemineoptera and Endoneoptera). Thus, these characters provide an extremely useful, almost untapped, source of data for higher-level systematics. Both higher-level and lower-level wing characters have been applied here to the phylogeny of Zoraptera and are discussed.     

Phylogeny of the Sympetrinae (Odonata: Libellulidae): further evidence of the homoplasious nature of wing venation

Abstract.  Sympetrinae is the largest subfamily of the diverse dragonfly family Libellulidae. This subfamily, like most libellulid subfamilies, is defined currently by a few wing venation characters, none of which are synapomorphies for the taxon. In this study, we used DNA sequence data from the nuclear locus elongation factor-1α and the mitochondrial loci 16S and 12S rRNA, together with 38 wing venation characters, to test the monophyly of the Sympetrinae and several other libellulid subfamilies. No analysis recovered Sympetrinae as monophyletic, partly because of the position of Leucorrhinia (of the subfamily Leucorrhininae) as a strongly supported sister to Sympetrum (of Sympetrinae) in all analyses. The subfamilies Brachydiplactinae, Leucorrhininae, Trameinae and Trithemistinae were also found not to be monophyletic. Libellulinae was the only subfamily supported strongly as monophyletic. Consistency indices and retention indices of wing venation characters used to define various subfamilies were closer to zero than unity, showing that many of these characters were homoplasious, and therefore not useful for a classification scheme within Libellulidae.     

中生代线蛉科化石(昆虫纲,脉翅目)

线蛉科是脉翅目中的一个灭绝科,仅发现于中国中侏罗统和蒙古晚侏罗统地层中。线蛉科昆虫脉序独特,为其分类地位的确定以及整个脉翅目翅脉演化的研究提供了重要的线索;此外,线蛉科昆虫前后翅上均具有条带状的斑纹,初步推测这种斑纹在其生存繁衍过程中发挥了积极作用,也可能是导致其灭绝的最直接因素之一。目前线蛉科已发表3属9种,本文对其进行了简要的回顾,并编制了线蛉科化石种检索表,对当时的古环境特征进行了简单推测,初步探讨了翅斑在其生存和灭绝中的作用。     

Wing and haltere venation in larvae of the Austrosimulium (Austrosimulium) australense group from New Zealand (Diptera: Simuliidae)

The venation pattern found in the developing wing and haltere buds of larvae is compared with the adult venation in the Austrosimulium (Austro-simulium) australense group of Simuliidae.     

Integration of wings and their eyespots in the speckled wood butterfly Pararge aegeria

We investigated both the phenotypic and developmental integration of eyespots on the fore- and hindwings of speckled wood butterflies Pararge aegeria. Eyespots develop within a framework of wing veins, which may not only separate eyespots developmentally, but may at the same time also integrate them by virtue of being both signalling sources and barriers during eyespot development. We therefore specifically investigated the interaction between wing venation patterns and eyespot integration. Phenotypic covariation among eyespots was very high, but only eyespots in neighbouring wing cells and in homologous wing cells on different wing surfaces were developmentally integrated. This can be explained by the fact that the wing cells of these eyespots share one or more wing veins. The wing venation patterns of fore- and hindwings were highly integrated, both phenotypically and developmentally. This did not affect overall developmental integration of the eyespots. The adaptive significance of integration patterns is discussed and more specifically we stress the need to conduct studies on phenotypic plasticity of integration.     

Evocation of venation pattern in the wing of a moth,Manduca sexta

The insect wing is formed from an epithelial sheet that folds during development to establish a saclike tissue with an upper and a lower epithelial monolayer. The adult cuticle formed by the upper and lower monolayers has a distinctive pattern of thickened regions called veins. The venation pattern on the lower surface matches that on the upper surface. As demonstrated by transposition of grafts from the upper monolayer, determination of venation pattern occurs prior to pupation in both wing monolayers. However, the pattern is not expressed until later in adult development. Expression of this determined pattern occurs autonomously in most circumstances. One circumstance in which the pattern fails to be expressed is in pieces of the upper monolayer that are isolated from the lower monolayer before adult cuticle deposition and expression of venation pattern. The only evident interaction between the two monolayers of the wing occurs during a 3-day period, 6–8 days after pupation. During this time, the basal laminae segregating upper monolayer from lower monolayer disappear, and the basal ends of cells form desmosomal junctions at the interface between upper and lower monolayer. Transposition as well as isolation of tissue fragments from the upper monolayer suggest that this interaction between the basal surfaces of the two monolayers is a prerequisite for evocation of venation pattern.     

THE VENATION OF THE MATURE NYMPH WING PAD OF EPHEMEROPSIS TRISETALIS EICHWALD (EPHEMEROPTERA: HEXAGENITIDAE)

Abstract  The complete mature nymph wing pad with distinct venation has been collected from Jiuquan Basin, Northwest China, which is almost without precedent in the fossil history of insect. The venation with large radial and costal fields is considered as archaic feature which very probably represents the primitive stage in the evolution of Ephemeroptera. Furthermore, as compared with the modern adult wing, the stronger articular sclerites in the fossil nymphal wing pad of Ephemeropsis trisetalis probably shows that the reduction of the articulation in modern mayfly wing is a secondarily acquired character.     

三尾类蜉蝣成熟稚虫的翅芽脉相(蜉蝣目)(英语)

本文记述了三尾类蜉蝣Ephemeropsis trisetalis稚虫的脉相和翅关节区特征为化石昆虫保存史上罕见材料,脉相中显出颇大的径脉区和前缘区十分相似于某些原始的古生代蜉蝣,或许可以提示发达的径脉区和前缘脉区是蜉蝣翅进化史中的初期特征.再者,现代蜉蝣的翅关节区退化,一般由一组不发达,分化很弱的骨板和肩板组成;这个化石种的翅关节区具发达的骨板和肩板似指示现化蜉蝣翅关节区退化是后生获得的性状.     

现存蜉蝣目昆虫的原始特征和独特性状

现存蜉蝣具有一系列引人注目的特征 ,它们为重建原始昆虫模式、探讨翅的起源、脉相的演化、附肢的演变等起到重要作用。该文系统总结了现生蜉蝣具有的原始特征和独特性状 (如原变态、稚虫水生、交尾行为、脉相、翅面皱褶、翅位、较多的附肢和蜕皮次数以及口器、生殖系统等等 ) ,并讨论比较了有关这些特征起源和演化的主要观点。     

The present state of nomenclature of wing venation in the Braconidae (Hymenoptera); its origins and comparison with related groups

The problem of nomenclature of wing venation in the Hymenoptera is discussed, and the main systems of terminology and their variations are reviewed.     

Fluctuating asymmetry in the honey bee,Apis mellifera: effects of ploidy and hybridization

We present a new measure of morphological asymmetry that avoids most of the statistical problems inherent in character-by-character analysis of size or shape. The method is an application of Procrustes analysis, which computes best-fitting super-positions of configurations of landmarks to the left and right sides of a single specimen. The Procrustes method combines subtle deviations in all aspects of the landmark configuration into one net asymmetry score. Directional asymmetry is separated from fluctuating asymmetry in a simple partition of a net sum-of-squares, and geometrical details of either component can be inspected by traditional methods of multivariate statistical analysis of landmarks. We demonstrate this method in a comparison of wing venation asymmetry in male (haploid) and female (diploid) honey bees (Apis mellifera). In addition we investigate the effects of ploidy and inter-subspecies hybridization on asymmetry and wing venation abnormalities, using the subspecies A. m. mellifera, A. m. carnica, and the hybrid strain “Nigra”. Results suggest that while the haploid males showed a higher frequency of wing venation abnormalities and greater total asymmetry than the diploid females, most of the asymmetry difference between males and females was in the form of directional, not fluctuating, asymmetry. Hybrid females had a higher frequency of wing venation abnormalities than females of either subspecies, but there were no significant differences in the mean level of asymmetry among females of A. m. mellifera, A. m. carnica and hybrid Nigra. Hybrid males had higher absolute frequency of wing venation abnormalities and asymmetry than males of either subspecies. However the mean frequency of venation abnormalities did not differ significantly between Nigra and A. m. carnica males, and mean asymmetries were not significantly different between Nigra and A. m. mellifera males. We discuss the relationship which is assumed to exist between developmental stability and fluctuating asymmetry in light of our result.     

The clavus and jugum venation in the wings of beetles (Coleoptera) and its genesis

The homology and nomenclature, as well as hypothesized pathways of the historical development of the clavus and jugal lobe of the beetle hind wings are discussed. The reconstructed plan of the clavus venation is largely similar to the venation patterns observed in some representatives of Corydalidae (Megaloptera). Its main apotypic characters are the following: the first anal cell was reduced at the wing base and the second anal or cuneiform cell appeared. This venation pattern is supposed to result from consolidation of the area around the claval furrow base at the earlier stages of the beetle wing evolution. In particular, longitudinal compressing the bases of RA, M, CuP, and 1A resulted in the development of a complex sclerite composed of the distal median plate and the bases of CuP and 1A. After this new reinforced connection between the remigium and the clavus appeared, the proximal parts of CuP and 1A were partly or completely reduced since they were no longer needed to maintain the structural integrity of the beetle wing.     

Discovery of the genus Iasvia Zalessky, 1934 in the Upper Permian of France (Lodève basin) (Orthoptera, Ensifera, Oedischiidae)

New material attributed to the species Iasvia reticulata ZALESSKY, 1934 and to a new species in this genus is described from the Salagou Formation (Saxonian Group, Lodève basin). Preamble to the taxonomic section, the wing venation pattern of Orthoptera is discussed. The numerous described specimens yield decisive information about variability of wing venation within the genus, previously based on a single specimen from the Russian Permian. I. reticulata is the first species from the Lodève basin that is already known from another site. The biostratigraphic implications are discussed.     

中国锯天牛族后翅翅脉研究(鞘翅目,天牛科,锯天牛亚科,锯天牛族)(英文)

对中国锯天牛族的后翅基部关节和后翅翅脉特征进行了研究,发现利用Kukalová-Peck和Lawrence (2004)的后翅命名系统能够很好地对中国锯天牛族后翅翅脉进行命名。但是在中国锯天牛族中,后中脉( MP)和前肘脉(CuA)在后缘并不合并;当前臀脉( AA3)和前肘脉(CuA3 +4)与后肘脉(CuP)相遇时,前臀脉(AA3)消失,前肘脉(CuA3 +4)和后肘脉(CuP)合并,因此楔室(W)仅由肘脉(Cu)的分支脉围成。尽管基部翅关节在研究的各属和各种之间没有表现出差异,但是后翅翅脉在土天牛属Dorysthenes和锯天牛属Prionus不同种类之间差异明显,这些特征包括径室的长宽比例和各边的长度关系、r3存在与否及其长度、后径脉的长度、楔室的长宽比例、以及后中脉(MP3 +4)和前肘脉(CuA3 +4)端部是否分叉等。因此,后翅翅脉特征在土天牛属Dorysthenes和锯天牛属Prionus分种时可能具有分类学意义。     

Evolution of wing shape in hornets: why is the wing venation efficient for species identification?

Wing venation has long been used for insect identification. Lately, the characterization of venation shape using geometric morphometrics has further improved the potential of using the wing for insect identification. However, external factors inducing variation in wing shape could obscure specific differences, preventing accurate discrimination of species in heterogeneous samples. Here, we show that interspecific difference is the main source of wing shape variation within social wasps. We found that a naive clustering of wing shape data from taxonomically and geographically heterogeneous samples of workers returned groups congruent with species. We also confirmed that individuals can be reliably attributed to their genus, species and populations on the basis of their wing shape. Our results suggested that the shape variation reflects the evolutionary history with a potential influence of other factors such as body shape, climate and mimicry selective pressures. However, the high dimensionality of wing shape variation may have prevented absolute convergences between the different species. Wing venation shape is thus a taxonomically relevant marker combining the accuracy of quantitative characters with the specificity required for identification criteria. This marker may also highlight adaptive processes that could help understand the wing's influence on insect flight.