Venation Characters Research Articles

An updated phylogeny of Anisoptera including formal convergence analysis of morphological characters

AbstractFamily interrelationships among Anisoptera (dragonflies) are unresolved. Molecular markers applied thus far have not been particularly useful for resolving relationships at the family level. Previous morphological studies have depended heavily on characters of wing venation and articulation which are believed to display considerable degrees of homoplasy due to adaptations to different flight modes. Here, we present a comprehensive anatomical dataset of the head morphology of Anisoptera focusing on muscle organization and endoskeletal features covering nearly all families. The characters are illustrated in detail and incorporated into an updated morphological character matrix covering all parts of the dragonfly body. Phylogenetic analysis recovers all families as monophyletic clades except Corduliidae, Gomphidae as sister group to all remaining Anisoptera, and Austropetaliidae as sister group to Aeshnidae (=Aeshnoidea). The position of Petaluridae and Aeshnoidea to each other could not be resolved. Libelluloidea is monophyletic with Neopetalia and Cordulegastridae as first splits. Chlorogomphidae is sister to monophyletic [Synthemistidae + (‘Corduliidae’ + Libellulidae)]. In addition, we applied a recently published formal approach to detect concerted convergence in morphological data matrices and uncover possible homoplasies. Analyses show that especially head and thorax characters may harbour homoplasies. After exclusion of possible homoplastic characters, Gomphidae is corroborated as sister group to all remaining Anisoptera.

Venation patterns of neotropical blueberries (Vaccinieae: Ericaceae) and their phylogenetic utility

The description of venation patterns in neotropical Vaccinieae has been problematic because of the lack of an operational definition. Moreover, the underlying homologies remain unknown across and within lineages, thus precluding the use of venation characters in phylogenetic analyses. Venation patterns are often obscured in herbarium and living specimens due to the characteristic coriaceous and thick nature of Ericaceae leaves. Therefore, to obtain an unobstructed view of the veins, leaf clearings of 16 putative generic lineages of Vaccinieae were prepared and described. The main venation patterns were then defined and traditional definitions of venation were discussed in the light of the observed patterns. Selected characters were scored and the evolution of venation characters was discussed using a phylogenetic framework derived from molecular data analyzed under Maximum Likelihood. Venation descriptions and photographs are provided for 33 species.

The most mysterious beetles: Jurassic Jurodidae (Insecta: Coleoptera) from China

Jurodidae are thought to be some of the world's most mysterious beetles and display an intriguing mixture of characters occurring in three different suborders. Hitherto, all known fossil and extant Jurodidae were extremely rare and restricted to Russian Siberia and the Far East. Here we describe two new species, Jurodes daohugouensis sp. nov. and Jurodes pygmaeus sp. nov. from the Middle Jurassic of Daohugou, China. A key to all species of Jurodidae and their research history are given, and the detailed morphology of fossil Jurodidae is also presented. Our results confirm the presence of three ocelli in fossil Jurodidae, and reveal that hind wings of J. pygmaeus sp. nov. possess well developed radial and oblong cells, and are very similar to those of extant Jurodidae (Sikhotealinia zhiltzovae). Male genitalia of fossil Jurodidae were described herein for the first time, showing a basal trilobate structure. The presence of exposed propleuron, three pairs of external trochantins, metanepisterna with ridges, primitive characters of wing venation further support the probable placement of Jurodidae in the suborder Archostemata. J. daohugouensis and J. pygmaeus are very similar to S. zhiltzovae in having the big protruding eyes, elevated median portion of frons bearing three ocelli, pubescent body, contiguous procoxae, exposed trachantion of all coxae, elytra with rows of punctures, and wing venation. Our results show that these key characters of extant Jurodidae can be traced back to the Middle Jurassic, revealing that the family has been in a period of evolutionary stasis for at least 160million years. Furthermore, our discoveries widen the paleogeographic distribution of fossil Jurodidae from Russian Siberia to northern China.

Discovery ofNyctiophylax(Trichoptera:Polycentropodidae) in Europe, with the description of a new species

A new species of the genus Nyctiophylax Brauer 1865, Nyctiophylax (Paranyctiophylax) gaditana sp. n., endemic to the southern Iberian Peninsula, is described and illustrated. To date, Nyctiophylax is known in the Western Palearctic region only through fossil representatives, some of which are preserved in Baltic amber. This finding is the first record of a living species of the genus Nyctiophylax for the European fauna. A key is provided to distinguish adult Nyctiophylax from other European genera of Polycentropodidae based on wing venation characters. The new species appears to be related to 2 Eastern Palaearctic species: Nyctiophylax (Paranyctiophylax) hjangsanchonus Botosaneanu 1970 from Korea and Nyctiophylax (Paranyctiophylax) adaequatus Wang and Yang 1997 from China, but it also could be related to the African species. Last, the ecology of the new species is described.

Phylogeny of Ptychopteroidea (Insecta: Diptera)

A phylogenetic tree is proposed for the superfamily Ptychopteroidea, reconstructed taking into account both extinct and extant taxa and based mainly on characters of wing venation.

Venation-Like Rib Layout Design in Plate under Bending Loads

This paper concerns a venation-like design method of rib layout and its application on some plates subjected to bending loads. Main topological characters of dicotyledonous venation and two determinative mechanical parameters in leaf venation morphogenesis theories are extracted, on the basis of those elicitations, a venation growing algorithm is proposed as an attempt at the proper layout of ribs in plate. Energy criterion and shear stress is specified as the factor orienting mainvein and subveins respectively. Vectorial equation equilibrium is used in reorganization to calculate widths and adjust vein cells slightly. Finite element method functions as background technique, under which several simple-shaped plates under bending loads are designed by venation growing algorithm. The resultant venation-like ribs offer multi-balanced improvements.

Species limits of the endemic genus Stemonoporus Thw. : leaf architecture answers

Stemonoporus Thw. is the most species-rich (27 species) endemic dipterocarp genus in Sri Lanka. All its members are categorized as highly threatened or threatened in the IUCN red data book. A recent study carried out in order to evaluate the species limits ofStemonoporus, further strengthened its position as the most species-rich endemic dipterocarp genus in Sri Lanka by further adding a new species and totaling the number to 27. A detailed study on leaf architecture was carried out in order to identify characters for field identification of these 27 species. Leaves were cleared and scored for characters. Twenty leaf architectural characters were identified, including leaf venation type, primary, secondary and tertiary veins, areoles and veinlets. A comprehensive description of leaf architecture for all the species was complied. A combined phylogenetic analysis with other morphological characters was performed. The simple, entire margined, spirally arranged leaves indicated a shared ancestry.Within the genus Stemonoporus, all 27 species are monophyletic and are supported by leaf architectural apomorphies.The leaf venation characters used during the previous works are indistinctly defined and during the present study, novel, strong and stable leaf architectural characters were identified for each of the species. These characters could play a crucial role in field identification of these highly threatened taxa.

Classification of Camellia (Theaceae) Species Using Leaf Architecture Variations and Pattern Recognition Techniques

Leaf characters have been successfully utilized to classify Camellia (Theaceae) species; however, leaf characters combined with supervised pattern recognition techniques have not been previously explored. We present results of using leaf morphological and venation characters of 93 species from five sections of genus Camellia to assess the effectiveness of several supervised pattern recognition techniques for classifications and compare their accuracy. Clustering approach, Learning Vector Quantization neural network (LVQ-ANN), Dynamic Architecture for Artificial Neural Networks (DAN2), and C-support vector machines (SVM) are used to discriminate 93 species from five sections of genus Camellia (11 in sect. Furfuracea, 16 in sect. Paracamellia, 12 in sect. Tuberculata, 34 in sect. Camellia, and 20 in sect. Theopsis). DAN2 and SVM show excellent classification results for genus Camellia with DAN2's accuracy of 97.92% and 91.11% for training and testing data sets respectively. The RBF-SVM results of 97.92% and 97.78% for training and testing offer the best classification accuracy. A hierarchical dendrogram based on leaf architecture data has confirmed the morphological classification of the five sections as previously proposed. The overall results suggest that leaf architecture-based data analysis using supervised pattern recognition techniques, especially DAN2 and SVM discrimination methods, is excellent for identification of Camellia species.

Bechala sommeri Ilger & Brauckmann, 2012 enlightens the Namurian griffenfly diversity (Insecta: Odonatoptera: Bechalidae)

Bechala sommeriIlger & Brauckmann, 2012, the type species of the type genus of the early Late Carboniferous (Namurian) family Bechalidae Ilger & Brauckmann, 2012, is redescribed. It does not belong to the order Megasecoptera as previously proposed. The taxon is clearly attributable to Odonatoptera for the typical venation characters as CuA separating from MP obliquely, a true arculus with concave RP and convex MA emerging from a composite vein R+MA, short ScP, and presence of convex intercalaries IR2 and IR1 between the main branches of RP3/4, RP2 and RP1. We transfer this taxon with the monospecific family Bechalidae to Odonatoptera. A new diagnosis is given for Bechalidae and its type genus Bechala. Furthermore, the presence of an oblique subnodal crossvein very far from the ending of ScP and close to the base of RP2 confirms the hypothesis that the subnodus is a structure originally independent of the nodus with a different function in relation to wing tracheation. The Bechalidae are included in a clade (Meganeuridae–Sinierasipteridae–Bechalidae–Lapeyridae–Nodialata), in contrast to a sister group relationships between the two clades Meganisoptera (=Namurotypidae–Paralogidae–Kargalotypidae–Kohlwaldiidae–Meganeuridae) and Odonatoclada (=Lapeyridae–Nodialata), while the potential relationships between the Campylopteridae and the Lapeyridae and Nodialata are rejected. Bechala represents a ‘damselfly-like’ ecological niche in the Namurian, showing the high diversity of the earliest known Odonatoptera, strongly suggesting an Early Carboniferous, if not Late Devonian age for this pterygote clade.

Minyscapheus dominicanus n. gen., n. sp. (Hemiptera: Cicadidae), a fossil cicada in Dominican amber

A new fossil cicada, Minyscapheus dominicanus n. gen., n. sp. Poinar, Kritsky and Brown (Hemiptera: Cicadidae) is described from Dominican amber. The small size distinguishes the new genus from all extant West Indian cicadas. M. dominicanus aligns with the Hispaniolan Uhleroides clade; however, it differs from the two endemic Uhleroides species and other West Indian members of the genus. Diagnostic characters include a narrow pronotal collar with pointed lateral angles, a narrow scutellum, tegmina with a narrow, elongated costal area (R 2 cell), wing with length of A3 vein greater than half length of vannus and additional venation features. Two erythraeid mites are attached to the mesonotum of the fossil cicada. This is the first description of an adult cicada in amber.

On the edge of parasitoidism: a new Lower Cretaceous woodwasp forming the putative sister group of Xiphydriidae + Euhymenoptera

The fossil woodwasp Cratoenigma articulatagen. et sp.n. (Insecta: Hymenoptera) is described from the Lower Cretaceous Crato formation of Brazil. This fossil cannot be placed in any existing superfamily, but its putative phylogenetic position within Hymenoptera is discussed in detail on the basis of relevant thoracic, abdominal and wing venation characters. These characters are critically evaluated and compared with those of extant and fossil Hymenoptera. The phylogenetic position of C. articulatasp.n. is investigated relative to extant Xyelidae, Tenthredinoidea s.l., Pamphilioidea, Cephidae, Siricoidea, Xiphydriidae, Orussidae and Apocrita, and also to Mesozoic Gigasiricidae, Myrmiciidae, Daohugoidae, Sepulcidae, Anaxyelidae, Paroryssidae and Ephialtitidae. Based on the presence of a synapomorphic transscutal articulation, a plesiomorphic unconcealed mesopostnotum and autapomorphic hindwing venation (cu‐a distinctly basal to fork between M and Cu), C. articulatasp.n. most likely forms the sister group of Xiphydriidae + Euhymenoptera. This would place it well within Unicalcarida, i.e. the clade in which the transition from endophytic to parasitoid lifestyle evolved.

Ordralfabetix sirophatanis gen. et sp. n.—the first Lophopidae from the Lowermost Eocene Oise amber, Paris Basin, France (Hemiptera: Fulgoromorpha)

A new genus and species, Ordralfabetix sirophatanis, is described on the basis of a specimen from the Lowermost Oise amber, Paris Basin, France. It could be ascribed to the Sarebasa + group of Lophopidae in respect to tegmen venation characters, but differs from the genera placed there by significant tegmen features. The evolution and distribution pattern of fossil and recent Lophopidae are also reviewed.

A new Chinese Mesozoic dragonfly clarifies the relationships between Rudiaeschnidae and Cymatophlebiidae (Odonata: Aeshnoptera)

A very well preserved fossil specimen of Rudiaeschna limnobia Ren and Guo, 1996, is described and diagnosis for Rudiaeschna is emended. Its clear morphological structures, including body characters, clarify and confirm the affinities of Rudiaeschna and the Rudiaeschnidae with Cymatophlebia Deichmüller, 1886, and the Cymatophlebiidae, previously inferred only through venational characters.

Abraracourcix curvivenatus n. gen. n. sp. from the Lowermost Eocene Oise amber (Hemiptera: Fulgoromorpha: Ricaniidae)

An extinct genus Abraracourcix n. gen. with A. curvivenatus n. sp. is described based on the specimen from the Lowermost Eocene Oise amber. The venation characters and possible hostplant relationships, as well as taxonomic position and biogeographical pattern of recent Pochazoides generic group are discussed.

Trends in evolution of ant wing venation (Hymenoptera, Formicidae)

Some characters of wings are applied for description of ants and their phylogeny. Seventeen recent subfamilies of Formicidae are analyzed. Sixteen types of forewing venation are distinguished. The main ant subfamilies are compared with respect to their forewing venation and other wing characteristics. The general relations between wing venation and size of ants are described. The specific features of wing venation in representatives of various subfamilies are characterized, as well as differences in the evolutionary trends. The general trends in wing venation are discussed.

Weiwoboidae fam. nov. of ‘Higher’ Fulgoroidea (Hemiptera: Fulgoromorpha) from the Eocene Deposits of Yunnan, China

Abstract:A new family of so‐called ‘higher’ planthoppers, Weiwoboidae fam. nov., from the Lower Eocene of Yunnan is described. A new monotypic genus, Weiwoboa gen. nov., with Weiwoboa meridiana sp. nov. is also described and illustrated. The characters of tegmen venation of the new family, its evolution, and supposed relationships are briefly discussed.

Early Mecopterida and the systematic position of the Microptysmatidae (Insecta: Endopterygota)

In recent times many authors have regarded the Protomeropidae and Microptysmatidae - two essentially Permian groups - as either early trichopteran lineages or members of the stem-group of the Amphiesmenoptera (basically: Trichoptera+Lepidoptera). Actually none of these families possesses, in its ground plan, the most significant derived trait of the amphiesmenopteran forewing, namely a true ‘double-Y loop’ arrangement of the anal veins. Since ‘Carpenter’s organs’, small rounded structures in the costal area of the hindwing, are only known to occur in certain members of the Permochoristidae, Kaltanidae and Protomeropidae, these three families should belong to a fossil clade, which we ascribe to the Mecoptera, suborder Pistillifera sensu lato, mainly on account of a few venational features. Although we maintain the Microptysmatidae in the Mecopterida (=Panorpida, i.e. Amphiesmenoptera, Mecoptera, Diptera, and relatives), we propose to place this family in a separate order: the Permotrichoptera, n. status. Indeed, apparently, Microptysmatidae can be ascribed neither to the Amphiesmenoptera nor to the Antliophora (=Mecoptera-Diptera complex).

Atypical Wing Venation inDialictusandHemihalictusand Its Implications for Subgeneric Classification ofLasioglossum

The subgeneric classification of hundreds of species inLasioglossumCurtissensu latois currently unstable due to differing opinions on the suitability of wing venation characters for differentiating subgenera. The subgeneraDialictusRobertson andHemihalictuswere both originally defined primarily by the forewing having two submarginal cells. I present examples of variation in submarginal cell number in the type species of these two subgenera:L. (Dialictus)anomalum(Robertson) andL. (Hemihalictus)lustrans(Cockerell). These results suggest that submarginal cell number is insufficient for recognizing subgenera inLasioglossum. The variability of this character is used to refute the classification proposed by some authors thatChloralictusRobertson, but notDialictus, be synonymised withEvylaeusRobertson.

Technical Note: Extraction of Wing Venation Characteristics by Image Analysis

Wing venation characteristics, such as the wing diagram and coordinates of vein junctions, are species-specific and are usually used in studies of insect identification. Image analysis methods are quick and precise means to measure these characteristics. An image analysis method is proposed that enables extraction of the wing diagram and automatic identification of vein junctions. A color image of an insect wing is converted to a binary image of wing venation based on thresholding. A distance-to-boundary map of the complement of the binary image is computed using a distance transform function. The vein skeleton is obtained by a watershed transform. The lookup table method is applied to find coordinates of vein junctions. The proposed method is capable of producing a smooth vein skeleton diagram and can extract the coordinates of vein junctions. This approach is not limited to high-quality images and is simple because the characteristics are obtained almost simultaneously. The experimental results showed that the proposed method is repeatable and useful for wing analysis.

Tertiary leaf fossils of Mangifera (Anacardiaceae) from Li Basin, Thailand as examples of the utility of leaf marginal venation characters

The mango (Mangifera) is the most important commercial fruit in Asia and is popular worldwide. Because of uncertainty in its origin and biogeography, the study of Mangifera could be of value in elucidating its current genetic diversity and geographical distribution. We report here two new species of compressed leaves of Mangifera (Anacardiaceae) from upper Oligocene-lower Miocene lacustrine sediments in the Li Basin, northern Thailand. Leaf form and venation characters of fossil and extant leaves are presented with detailed marginal venation patterns that were found to be important in assigning the fossil leaves to Mangifera paleoindica and M. paleocaloneura. These two fossil species of Mangifera and the present occurrence of basal species of Mangifera in Thailand suggest that northern Thailand could be an area with a long history of evolution and diversification of Mangifera, particularly M. indica.