EVOLUTION OF THE HIND WING IN COLEOPTERA

1993 ◽  
Vol 125 (2) ◽  
pp. 181-258 ◽  
Author(s):  
Jarmila Kukalová-Peck ◽  
John F. Lawrence

AbstractA survey is made of the major features of the venation, articulation, and folding in the hind wings of Coleoptera. The documentation is based upon examination of 108 Coleoptera families and 200 specimens, and shown in 101 published figures. Wing veins and articular sclerites are homologized with elements of the neopteran wing groundplan, resulting in wing vein terminology that differs substantially from that generally used by coleopterists. We tabulate the differences between currently used venational nomenclature and the all-pterygote homologous symbols. The use of the neopteran groundplan, combined with the knowledge of the way in which veins evolved, provides many strong characters linked to the early evolutionary radiation of Coleoptera. The order originated with the development of the apical folding of the hind wings under the elytra executed by the radial and medial loop. The loops, which are very complex venational structures, further diversified in four distinctly different ways which mark the highest (suborder) taxa. The remaining venation and the wing articulation have changed with the loops, which formed additional synapomorphies and autapomorphies at the suborder, superfamily, and sometimes even family and tribe levels. Relationships among the four currently recognized suborders of Coleoptera are reexamined using hind wing characters. The number of wing-related apomorphies are 16 in Coleoptera, seven in Archostemata + Adephaga–Myxophaga, four in Adephaga–Myxophaga, seven in Myxophaga, nine in Archostemata, and five in Polyphaga. The following phylogenetic scheme is suggested: Polyphaga [Archostemata (Adephaga + Myxophaga)]. Venational evidence is given to define two major lineages (the hydrophiloid and the eucinetoid) within the suborder Polyphaga. The unique apical wing folding mechanism of beetles is described. Derived types of wing folding are discussed, based mainly on a survey of recent literature. A sister group relationship between Coleoptera and Strepsiptera is supported by hind wing evidence.

1992 ◽  
Vol 70 (12) ◽  
pp. 2452-2473 ◽  
Author(s):  
Jarmila Kukalová-Peck ◽  
Carsten Brauckmann

The enigmatic and artificial Protorthoptera is the largest Paleozoic order of Neoptera. Typical hemipteroid head characters (inflated clypeus, styletal mouthparts) are here reported for "Protorthoptera" fossils and linked with the basic venational braces of hemipteroid wings. The "order" is recognized as being mostly composed of extremely primitive hemipteroids. Two new Late Carboniferous gerarid wings are described: Osnogerarus trecwithiensis n.gen., n.sp. from the Westphalian D of Osnabrück, Germany, and Cantabrala gandli n.gen., n.sp. from the Lower Stephanian (Cantabrian) of the Cantabrian Mountains, northwestern Spain. Several ground-plan characters of hemipteroids are described: head with visible segmentation; polyramous thoracic legs bearing exites and with trochanter not fused to prefemur; ovipositor with cutting ridges; the medial wing vein is shown as not the complete media (= M), but only the media posterior (= MP); the arculus is diagnosed as a cross-vein turned into an important brace; two radial sectors (RA and RP) originate separately from the radial basivenale; and the anal brace is formed by AA1. The hemipteroid stem-assemblage comprised (i) the gerarid line (extinct) with a long MP–CuA fusion replacing the arculus (derived) and with a repeatedly forked CuP (primitive), and (ii) the paoliid line with an arculus (a convex mp–cua cross-vein) (primitive) and a simple CuP (derived). The paoliids are the probable ancestors of modern hemipteroids. The Hemiptera (Sternorrhycha + Auchenorrhyncha + Coleorrhyncha + Heteroptera) lost the ScA + vein to a V-shaped notch or a flexion line (synapomorphy). Geraridae, Paleozoic and Recent Insecta, and all Arthropoda have demonstrably polyramous legs in the ground plan. Use of the taxon "Uniramia" is erroneous in every aspect. A new method using major venational braces is offered to define the wings of all higher neopteran taxa and to resolve the uncertain relationships between some modern orders. Polyneoptera (plecopteroids + orthopteroids + blattoids) is a polyphyletic taxon, and should be abandoned. Blattoids are not directly related to orthopteroids, but are the sister-group of hemipteroids + endopterygotes. An updated basic phylogenetic scheme of Neoptera proposed here comprises the following clades: (Pleconeoptera (Orthoneoptera (Blattoneoptera (Hemineoptera + Endoneoptera)))).


1985 ◽  
Vol 224 (1236) ◽  
pp. 287-313 ◽  

Described are all the sensilla and non-innervated hairs observable by scanning electron microscopy on the thorax and wings of the minute (0 .4- 0.8 mm) parasitoid wasp, Trichogramma minutum . Most of the sensilla appear mechanosensory, and occur on the wing veins, around the wing bases and near the spiracles. The number, position and morphological type of the sensilla were constant between individuals. Dorsally, the thorax bears 18 hair sensilla, between 5 and 50 μm long. Around each forewing base are nine hair sensilla, and three unique sensilla less than 1 μm long; two of the latter form bifurcated ‘pronged’ structures. Of the nine, four form the anterior wing base hairplate, and two the basalare hairplate; these probably contact the wing at extreme anterodorsal and anteroventral positions. Each hindwing base is equipped with three hairplates, of either two or three 4-6 μm hairs. Two of these are located appropriately to contact the wing during flight, the third only when the wings are folded. Posterior to the hindwing base are two socketed hairs and an unusual grooved hair 10—12 μm long, all located near the opening of the third thoracic spiracle. Each membranous forewing is surrounded by 110-120 evenly spaced, unsocketed fringe hairs. A parallel secondary row of similar hairs occurs on both dorsal and ventral surfaces. The forewing surfaces also bear numerous hairs, unsocketed marginally, but with increasingly distinct sockets centrally. Dorsally, these hairs mainly occur in rows radiating from the wing vein. Ventrally the surface hairs are only partly arranged in rows, and vary more in form. Transmission electron microscopy shows these hairs to be non-innervated. In contrast, the wing veins bear innervated structures, comprising 19 hair-like, and 22-23 campaniform, sensilla. The rod-like hindwings also bear a fringe of hairs similar to those of the forewing, although longer and confined to the posterior margin. The sensilla of the hind wing comprise four ventral and one dorsal campaniform sensilla, and six hairs. The function of the described structures is discussed in relation to the flight mechanism and its control in these very small insects.


Zootaxa ◽  
2019 ◽  
Vol 4674 (4) ◽  
pp. 482-490 ◽  
Author(s):  
HONGYU LI ◽  
BO WANG ◽  
XINGYUE LIU

The male of Cretaconiopteryx grandis Liu & Lu, 2017, which is the only representative species of the extinct dustywing subfamily Cretaconiopteryginae, is described for the first time from the Upper Cretaceous Burmese amber. The male genitalia, well preserved in the examined specimen, show a number of plesiomorphic characters, which support the sister group relationship between Coniopterygidae and the rest of extant lacewing families. 


2003 ◽  
Vol 40 (4) ◽  
pp. 527-556 ◽  
Author(s):  
Michael deBraga

A morphological study of the postcranial skeleton of Procolophon trigoniceps from the Lower Triassic of South Africa and Antarctica is undertaken. Procolophon shares a sister-group relationship with the procolophonid Tichvinskia from the Lower Triassic of Russia and is a basal member of Procolophonidae. This clade also includes the enigmatic taxon Sclerosaurus, believed most recently to be a pareiasaur relative. Owenettids form a separate lineage from Procolophonidae and are predominantly restricted to the Permian of both South Africa and Madagascar. A phylogenetically based assessment is considered, in which specialized modern taxa (sand lizards) are compared to their nonfossorial sister clade, allowing for "key innovations" to be identified. A similar comparison between owenettids and procolophonids reveals a number of apparent "key innovations" within procolophonids that are suggestive of a burrowing lifestyle for Procolophon.


1993 ◽  
Vol 30 (10) ◽  
pp. 2128-2138 ◽  
Author(s):  
Donald B. Brinkman ◽  
Jiang-Hua Peng

Ordosemys leios, n.gen., n.sp., from the Early Cretaceous Luohandong Formation, Zhidan Group, Ordos Basin, Inner Mongolia, is a primitive aquatic turtle with a reduced, fenestrated plastron. It shares with the members of the Centrocryptodira the presence of well-formed articular surfaces on the cervical and caudal vertebrae. Within the Centrocryptodira, characters of the cervical vertebrae suggest it is more closely related to the Polycryptodira than is the Meiolaniidae. Ordosemys shares with the Chelydridae the presence of two procoelous anterior caudals, but this character may be primitive for the Polycryptodira. Characters of the basicranial region of the braincase shared by Ordosemys and the Chelonioidea support a sister-group relationship between these two taxa, but a sister-group relationship between Ordosemys and the Polycryptodira is more strongly supported by characters shared by the Chelonioidea and other members of the Polycryptodira.


1984 ◽  
Vol 15 (2) ◽  
pp. 151-178 ◽  
Author(s):  
Niels P. Kristensen

AbstractThe genital segments and internal genitalia of Agathiphaga vitiensis are described. Sternum VIII is anteriorly produced into blunt paired apophyses and posteriorly into a tongue-shaped lobe. Segment IX is a complete ring, very short in the dorsal and ventral midlines; its anterolateral lobes are largely apodemal. The long and curved gonopod ("valva") consists of a single piece. There is no median sclerite between the gonopod bases, but an open, softwalled "subgenital crypt" below the entrance of the phallocrypt may be homologous with the "median plate" in other primitive homoneurous moths. Tergum X bears a pair of broad "superior lobes" and the postgenital complex terminates in a medially intended, sclerotized "terminal lobe" above the eversible perianal area. The roof of the posterior part of the genital chamber bears a median aggregation of cuticular spines (the "spiny plate"), and a pair of smooth lateral sclerotizations ("presocii") tentatively attributed to venter X: a pair of setose sclerites (socii) are tentatively attributed to the paraprocts. The area bearing the spiny plate and presocii may in repose be folded down behind the phallus, thereby closing the phallocrypt. The phallus comprises a tubular phallotheca and an eversible aedeagus; the thick basal margen of the phallotheca is posteriorly expanded and forms the floor of the greater part of the phallocrypt; there is no ventral aedeagal branch. The musculature comprises two IX/X muscles, a segment IX muscle inserting on the subgenital crypt, phallic pro- and retractors (the former originating in the gonopod), intrinsic phallic muscles, a single segment IX muscle (adductor) to the gonopod and five intrinsic muscles of the postgenital complex. Each testis comprises four large, separate follicles. The spermatozoa do not remain grouped in discrete bundles in the vas deferens. Seminal vesicles are located on the vasa deferentia close to the testis and are doubtfully homologous with the vesicles in other Lepidoptera. The unpaired ejaculatory duct is very short. The evidence bearing on a reconstruction of the ground plan of the lepidopteran male genitalia is reviewed. Segment VIII was similar to the preceding segments. It is tentatively suggested that tergum and sternum IX were fused, that the gonopod was undivided and that a tubular, partly sclerotized aedeagus was present; deviations from these states within the order are therefore considered to be autapomorphic. The base of the aedeagus was probably surrounded by a short, collarlike phallotheca. It is suggested that there was a median sclerite between the gonopod bases, but the presence of discrete, paired and muscular "valvellae" in the lepidopteran ground plan is considered doubtful. It is further suggested that dorsum X bore a pair of lobes and that there were paired sclerotizations in venter X. The X/XI boundary is very difficult to trace. Seventeen muscle sets are ascribed to the lepidopteran ground plan; it is considered an autapomorphy of this ground plan that the phallic protractor originates within the gonopod. The testes presumably had large, separate follicles and there may have been two pairs of tubular accessory glands. The testes and the double set of accessory glands of Agathiphaga could be cited in support of a sistergroup relationship to all other Lepidoptera whereas the phallic structure (and possibly the "spiny plate") might support a sister group relationship to the Heterobathmiina. There is no support in male genital structure for a sistergroup relationship to the Heterobathmiina + Glossata; the latter phylogenetic hypothesis may be preferable on other grounds, however.


Insects ◽  
2021 ◽  
Vol 12 (9) ◽  
pp. 779 ◽  
Author(s):  
Ke-Ke Xu ◽  
Qing-Ping Chen ◽  
Sam Pedro Galilee Ayivi ◽  
Jia-Yin Guan ◽  
Kenneth B. Storey ◽  
...  

Insects of the order Phasmatodea are mainly distributed in the tropics and subtropics and are best known for their remarkable camouflage as plants. In this study, we sequenced three complete mitochondrial genomes from three different families: Orestes guangxiensis, Peruphasma schultei, and Phryganistria guangxiensis. The lengths of the three mitochondrial genomes were 15,896 bp, 16,869 bp, and 17,005 bp, respectively, and the gene composition and structure of the three stick insects were identical to those of the most recent common ancestor of insects. The phylogenetic relationships among stick insects have been chaotic for a long time. In order to discuss the intra- and inter-ordinal relationship of Phasmatodea, we used the 13 protein-coding genes (PCGs) of 85 species for maximum likelihood (ML) and Bayesian inference (BI) analyses. Results showed that the internal topological structure of Phasmatodea had a few differences in both ML and BI trees and long-branch attraction (LBA) appeared between Embioptera and Zoraptera, which led to a non-monophyletic Phasmatodea. Consequently, after removal of the Embioptera and Zoraptera species, we re-performed ML and BI analyses with the remaining 81 species, which showed identical topology except for the position of Tectarchus ovobessus (Phasmatodea). We recovered the monophyly of Phasmatodea and the sister-group relationship between Phasmatodea and Mantophasmatodea. Our analyses also recovered the monophyly of Heteropterygidae and the paraphyly of Diapheromeridae, Phasmatidae, Lonchodidae, Lonchodinae, and Clitumninae. In this study, Peruphasma schultei (Pseudophasmatidae), Phraortes sp. YW-2014 (Lonchodidae), and species of Diapheromeridae clustered into the clade of Phasmatidae. Within Heteropterygidae, O. guangxiensis was the sister clade to O. mouhotii belonging to Dataminae, and the relationship of (Heteropteryginae + (Dataminae + Obriminae)) was recovered.


Fossil Record ◽  
2019 ◽  
Vol 22 (1) ◽  
pp. 1-23 ◽  
Author(s):  
Gloria Arratia ◽  
Hans-Peter Schultze ◽  
Helmut Tischlinger

Abstract. A complete morphological description, as preservation permits, is provided for a new Late Jurassic fish species (Tharsis elleri) together with a revision and comparison of some morphological features of Tharsis dubius, one of the most common species from the Solnhofen limestone, southern Germany. An emended diagnosis of the genus Tharsis – now including two species – is presented. The new species is characterized by a combination of morphological characters, such as the presence of a complete sclerotic ring formed by two bones placed anterior and posterior to the eye, a moderately short lower jaw with quadrate-mandibular articulation below the anterior half of the orbit, caudal vertebrae with neural and haemal arches fused to their respective vertebral centrum, and parapophyses fused to their respective centrum. A phylogenetic analysis based on 198 characters and 43 taxa is performed. Following the phylogenetic hypothesis, the sister-group relationship Ascalaboidae plus more advanced teleosts stands above the node of Leptolepis coryphaenoides. Both nodes have strong support among teleosts. The results confirm the inclusion of Ascalabos, Ebertichthys and Tharsis as members of this extinct family. Tharsis elleri n. sp. (LSID urn:lsid:zoobank.org:act:6434E6F5-2DDD-48CF-A2B1-827495FE46E6, date: 13 December 2018) is so far restricted to one Upper Jurassic German locality – Wegscheid Quarry near Schernfeld, Eichstätt – whereas Tharsis dubius is known not only from Wegscheid Quarry, but also from different localities in the Upper Jurassic of Bavaria, Germany, and Cerin in France.


2017 ◽  
Author(s):  
Christopher E. Laumer ◽  
Harald Gruber-Vodicka ◽  
Michael G. Hadfield ◽  
Vicki B. Pearse ◽  
Ana Riesgo ◽  
...  

AbstractThe phylogenetic placement of the morphologically simple placozoans is crucial to understanding the evolution of complex animal traits. Here, we examine the influence of adding new genomes from placozoans to a large dataset designed to study the deepest splits in the animal phylogeny. Using site-heterogeneous substitution models, we show that it is possible to obtain strong support, in both amino acid and reduced-alphabet matrices, for either a sister-group relationship between Cnidaria and Placozoa, or for Cnidaria and Bilateria (=Planulozoa), also seen in most published work to date, depending on the orthologues selected to construct the matrix. We demonstrate that a majority of genes show evidence of compositional heterogeneity, and that the support for Planulozoa can be assigned to this source of systematic error. In interpreting this placozoan-cnidarian clade, we caution against a peremptory reading of placozoans as secondarily reduced forms of little relevance to broader discussions of early animal evolution.


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