Most Paleozoic Protorthoptera are ancestral hemipteroids: major wing braces as clues to a new phylogeny of Neoptera (Insecta)

1992 ◽  
Vol 70 (12) ◽  
pp. 2452-2473 ◽  
Author(s):  
Jarmila Kukalová-Peck ◽  
Carsten Brauckmann
Keyword(s):  
Sister Group ◽  
New Method ◽  
Cantabrian Mountains ◽  
Wing Vein ◽  
Ground Plan ◽  
Northwestern Spain ◽  
The Media ◽  
Thoracic Legs ◽  
Phylogenetic Scheme ◽  
Cross Vein

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)))).

EVOLUTION OF THE HIND WING IN COLEOPTERA

1993 ◽  
Vol 125 (2) ◽  
pp. 181-258 ◽  
Author(s):  
Jarmila Kukalová-Peck ◽  
John F. Lawrence
Keyword(s):  
Recent Literature ◽  
Hind Wing ◽  
Sister Group ◽  
Sister Group Relationship ◽  
Wing Vein ◽  
Folding Mechanism ◽  
Evolutionary Radiation ◽  
Wing Veins ◽  
Phylogenetic Scheme ◽  
Wing Folding

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.

The male genitalia of Agathiphaga (Lepidoptera: Agathiphagidae) and the lepidopteran ground plan

1984 ◽  
Vol 15 (2) ◽  
pp. 151-178 ◽  
Author(s):  
Niels P. Kristensen
Keyword(s):  
Male Genitalia ◽  
Ejaculatory Duct ◽  
Posterior Part ◽  
Sister Group ◽  
Sister Group Relationship ◽  
Ground Plan ◽  
Accessory Glands ◽  
Internal Genitalia ◽  
Genital Structure ◽  
Segment Viii

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.

The "Uniramia" do not exist: the ground plan of the Pterygota as revealed by Permian Diaphanopterodea from Russia (Insecta: Paleodictyopteroidea)

1992 ◽  
Vol 70 (2) ◽  
pp. 236-255 ◽  
Author(s):  
Jarmila Kukalová-Peck
Keyword(s):  
Plant Community ◽  
Flapping Flight ◽  
Lower Permian ◽  
Selective Agent ◽  
Ural Mountains ◽  
Ground Plan ◽  
Selection For ◽  
Thoracic Legs ◽  
Maxillary Palps ◽  
Present Plant

The paleomorphology of extinct Paleozoic Diaphanopterodea (Paleodictyopteroidea: Paleoptera) in exceptionally well-preserved Lower Permian Uralia (Parelmoidea) and Paruralia (Paruraliidae) from Tshekarda in the Ural Mountains, Russia, was used to identify the basic morphological components of the pterygote head, body, and genitalia, as follows: a 6-segmented head with sutures between the somites and between the cephalic terga and epicoxae; antennae 2 with four enlarged basal elements and a flagellum; mandibles (shown as fully homologous to the gnathobase of other arthropods) with a sliding anterior articulation; leglike maxillary palps, labial palps, abdominal leglets, and gonostyli, all starting with prefemur, bearing a fully articulated patella (as in spiders) and ending in double claws; polyramous thoracic legs (i.e., with several outer rami or exites) and a prefemur not associated, as it usually is, with the trochanter; a thoracic pleuron (wall support) formed from a flattened subcoxa; an abdominal pleuron clearly composed of three flattened leg segments: the subcoxa, coxa, and trochanter; and genitalia revealing their derivation from legs bearing coxal and trochanteral endites. It is suggested that the homoeotic mutant ophthalmoptera is a morphologically acceptable indication that the cephalic epicoxa surrounds the eye. The presence of several rami (exites) on the legs of primitive Paleozoic and modern Crustacea, Chelicerata, and Insecta shows that they share a polyramous arthropod leg; "biramy," as well as "uniramy," is always secondary and does not define higher arthropod taxa. The Atelocerata Heymons, 1901 (= Myriapoda + Hexapoda) is a natural taxon. "Uniramia" was a taxon based mainly on an erroneous "ground plan" of the arthropod leg and mandible, and should be completely dropped from use. In marine arthropods, one of the upper exites often convergently develops into a long swimming oar ramus, and the legs become functionally "biramous." A complex aquatic epicoxal oar ramus of the ancestral Atelocerata may be the only appendage suitably preadapted to become a protowing, modifiable by selection for flapping flight. Paleodictyopteroids with sucking rostra (about 50% of Paleozoic entomofauna) are probably the most important selective agent that initiated the change from the Paleozoic plant community into the present plant community.

Lower Permian "mecopteroid-like" insects from central Europe (Insecta, Endopterygota)

1990 ◽  
Vol 27 (3) ◽  
pp. 459-468 ◽  
Author(s):  
Jarmila Kukalova-Peck ◽  
Rainer Willmann
Keyword(s):  
New Species ◽  
Kuznetsk Basin ◽  
Phylogenetic Position ◽  
Lower Permian ◽  
Wing Venation ◽  
Medial Branch ◽  
New Genera ◽  
Ground Plan ◽  
The Family ◽  
The Media

Five new genera and eight new species of endopterygote insects are described from Lower Permian (Asselian and Artinskian) strata from Moravia, Czechoslovakia. Three of the genera belong to the family Protomeropidae: Pseudomerope n.gen. (including Pseudomerope mareki n.sp., Pseudomerope havlati n.sp., Pseudomerope oborana n.sp., Pseudomerope gallei n.sp.); Pseudomeropella n.gen. (including Pseudomeropella nekvasilovae n.sp.); and Stenomerope n.gen. (including Stenomerope spinari n.sp.). The fourth genus, Moravochorista n.gen. (including Moravochorista Carolina n.sp.), is similar to Pinnachorista and Kaltanochorista from the Lower Permian of the Kuznetsk Basin, USSR, but has not been assigned to a higher taxon. The phylogenetic position of both the Protomeropidae and Moravochorista and of their allies, within the endopterygotes is unclear. The fifth genus, Microptysmella n.gen. (including Microptysmella moravica n.sp.), may be the earliest known member of Amphiesmenoptera, since it exhibits almost the same wing venation as the amphiesmenopteran Microptysma sibiricum Martynova from the Lower Permian of the USSR.Wing-venation symbols homologous within the pterygote ground plan have been used in the descriptions. The vein "M5" of earlier authors is regarded as homologous to a convex cross-vein (strut) between the media posterior and the cubitus anterior, which is shared primitively by all endopterygotes, and is not a "fifth medial branch."

The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera)

Zootaxa ◽  
2006 ◽  
Vol 1180 (1) ◽  
pp. 1 ◽  
Author(s):  
BRADLEY J. SINCLAIR ◽  
JEFFREY M. CUMMING
Keyword(s):  
Feeding Habits ◽  
Cladistic Analysis ◽  
Sister Group ◽  
Morphological Characters ◽  
Male And Female ◽  
Ground Plan ◽  
Incertae Sedis ◽  
Higher Taxa ◽  
The Family

A cladistic analysis of the Empidoidea and basal lineages of the Cyclorrhapha, based on morphological characters, confirms the monophyly of both groups as well as that of the                    Eremoneura. The resulting final trees are used to revise the classification of the Empidoidea to include the following five families: Empididae, Hybotidae, Atelestidae (including Nemedininae n. subfam.), Brachystomatidae rev. stat. (comprising the subfamilies Brachystomatinae, Ceratomerinae and Trichopezinae), and Dolichopodidae s.lat. The family Microphoridae is not recognized, and the Microphorinae and Parathalassiinae are assigned to the Dolichopodidae s.lat. The Dolichopodidae s.str. includes 15 subfamilies that were previously recognized within the family. Within the Empidoidea we found support for Atelestidae as the sister group to the Hybotidae and for the monophyly of Parathalassiinae + Dolichopodidae s.str. The Empididae remains poorly defined and the genera Homalocnemis Philippi, Iteaphila Zetterstedt, Anthepiscopus Becker, and Oreogeton Schiner are classified as incertae sedis within the                   Empidoidea. In addition, the following higher taxa are proposed: Symballophthalmini n. tribe, Bicellariini n. tribe, Oedaleinae rev. stat., and Trichininae rev. stat., which are all assigned to the Hybotidae. The genus Sematopoda Collin is tentatively assigned to Trichopezinae, and Xanthodromia Saigusa is transferred from Hemerodromiinae to Brachystomatinae.        All morphological characters are extensively discussed and illustrated, including details of the antennae, mouthparts, internal thoracic structures, wings, and male and female terminalia. In addition, a key to families and unplaced genus groups of the Empidoidea is provided. Feeding habits are also discussed in terms of the empidoid ground plan condition.

A new method to denude the endothelium without damage to media: structural, functional, and biomechanical validation

2004 ◽  
Vol 286 (5) ◽  
pp. H1889-H1894 ◽  
Author(s):  
Xiao Lu ◽  
Xiaomei Guo ◽  
Carlos Linares ◽  
Ghassan S. Kassab
Keyword(s):  
Stress State ◽  
Femoral Artery ◽  
Contractile Function ◽  
Common Femoral Artery ◽  
Biomechanical Properties ◽  
New Method ◽  
Balloon Injury ◽  
Stress Strain Relation ◽  
The Media ◽  
The Right

The intimial thickening that occurs in human and animal atherogenesis can be induced by mechanical injury to the endothelium. The objective of the present study was to develop a new method to induce arterial endothelial injury without damage to the media for future investigations of mechanisms of intimal thickening and atherogenesis. A specifically designed catheter was inserted into the common femoral artery of Wistar rats ( n = 9) through an arteriotomic mouth. After application of Tyrode solution containing 0.14 M KCl on the surface of the vessel, the vessel contracted onto the catheter. The catheter was then moved back and forth to scrape away the endothelium. The left common femoral artery of the same rat was subjected to the standard balloon injury model. The two models were evaluated structurally, functionally, and biomechanically. Structurally, we verified that both techniques remove the endothelium, but the balloon method damages the media. Functionally, we examined the contractile response of the artery to [K+] and norepinephrine 2 days after the denudation. We found that the right femoral artery underwent contraction in response to [K+], whereas the left artery did not. Furthermore, neither artery responded to norepinephrine. Biomechanically, we measured the pressure-diameter relationship and the zero-stress state of the vessel and computed the stress-strain relation. The circumferential stretch ratios at 120 mmHg were 1.38 ± 0.08 for the control, 1.41 ± 0.08 ( P > 0.05) for the new method, and 1.56 ± 0.09 for the balloon injury ( P < 0.05). The opening angles at the zero-stress state were 113 ± 21° for the control, 102 ± 18° for the new method ( P > 0.05), and 8 ± 13° for the balloon injury ( P < 0.001). In conclusion, the new method removes the endothelium while maintaining the structure, contractile function, and biomechanical properties of the vessel.

scholarly journals NOTE ON THE DIURNALS

1898 ◽  
Vol 30 (8) ◽  
pp. 201-203
Author(s):  
A. Radcliffe Grote
Keyword(s):  
Common Species ◽  
Hawk Moth ◽  
The Media ◽  
The Cross ◽  
Parallel Case ◽  
The Individual ◽  
Cross Vein

Mancipium brassicœ.—Dr. Chapman writes me that certain specimens of this common species examined by him showed the very short veinlet III. 3+4. This veinlef constantly diminishes in size, progressing towards the tip of the wing to finally vanish, through many forms fo the Pieridœ. I had indeed expected it to be occasionally prersistent in brassicœ, although my preparations did not show it. It has disappeared in Pontia daplidice, in Nathalis iole, and strange to say in that curious and now isolated Pierid, Gonophlebia paradoxa. This variability, in one and the same species, is interesting because it follows the general evolutionary direction of the changes in the venation. Always the radial branches in the Pierids and other groups tend to diminish in number. Always the disintegration of the Media advances, until it finally disappears, as a system, from the surface of the wing (Rothschildia, Samia, Potamis, etc.). A parallel case to that of brassicœ is offered by Copismerinthus ocellata. In some specimens of this Hawk Moth, vein IV, is still thrown off from the cross vein of the hind wings, instead fo the Radius, which it has usually ascended to beyond the cell. We must regard these as instances of generalization in the individual, of a reversion to what was formerly the rule and is now becoming, by slow degrees, the exception.

scholarly journals Patterns of movement of released female brown bears in the Cantabrian Mountains, northwestern Spain

Ursus ◽  
2017 ◽  
Vol 28 (2) ◽  
pp. 165-170 ◽  
Author(s):  
Vincenzo Penteriani ◽  
María del Mar Delgado ◽  
José Vicente López-Bao ◽  
Pablo Vázquez García ◽  
Juan S. Monrós ◽  
...  
Keyword(s):  
Cantabrian Mountains ◽  
Brown Bears ◽  
Northwestern Spain
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