Orthocephalus, FIEBER, 1858

Tatarnic, Nikolai J. & Cassis, Gerasimos, 2012, The Halticini of the world (Insecta: Heteroptera: Miridae: Orthotylinae): generic reclassification, phylogeny, and host plant associations, Zoological Journal of the Linnean Society 164 (3), pp. 558-658 : 623-626

ORTHOCEPHALUS FIEBER ( FIGS 4 , 45–46 )

Orthocephalus Fieber, 1858: 316 (gen. nov.; type species: Lygaeus brevis Panzer, 1798 by subsequent designation Reuter, 1888: 76); Fieber, 1858: 316 (gen. nov., key); Fieber, 1860b: 291, 1861 (key, descr.); Douglas & Scott, 1865: 429 (descr.); Thompson, 1871: 432 (key); Reuter, 1875b: 24 (key); Reuter, 1875a 1:86; 2:93 (key, descr.); Saunders, 1875: 289 (key); Provancher, 1887: 136 (descr.); Reuter, 1891: 43, 158 (key, descr.); Saunders, 1892: 269 (key, descr.); Hueber, 1906: 3, 15 (key, descr.); Kirkaldy, 1906: 131 (cat.); Oshanin, 1910: 791 (cat.); Reuter, 1910: 148 (cat.); Van Duzee, 1914: 389 (cat.); Knight, 1923: 498 (key); Blatchley, 1926: 804 (descr.); Stichel, 1933: 235 (key); Hedicke, 1935: 59 (key); Knight, 1941: 75, 81 (key); Hsiao, 1942: 253 (key); China, 1943: 267 (cat.); Slater, 1950: 46 (female genitalia); Kiritshenko, 1951: 127 (key); Wagner, 1952: 96, 103 (key, descr.); Carvalho & Leston, 1952: 245 (key, fig.); Carvalho, 1952: 74 (cat.); Carvalho, 1955: 67 (key); Carvalho, 1958: 22 (cat.); Southwood & Leston, 1959: 247 (key); Wagner, 1961: 50 (diag., key); Kerzhner, 1964a: 966 (diag., key); Wagner & Weber, 1964: 267 (descr., key); Wagner, 1973: 41 (descr., key); Schuh, 1995: 63 (world cat.); Namyatova & Konstantinov, 2009 (generic revision, phylogeny).

Anapomella Putshkov, 1961: 25 (syn. by Namyatova & Konstantinov, 2009)

Oraniella Reuter, 1894: 138 (syn. by Namyatova & Konstantinov, 2009)

Diagnosis: Orthocephalus can be recognized by the following combination of characters: head transverse with frons flat to moderately convex; eyes sessile; AI short with two or three mesial spine-like setae, dorsum impunctate, usually covered with robust, bristle-like setae, apical portion of ductus seminis bowl-shaped, strongly dilated and sclerotized, apically funnel-shaped and dorsoventrally compressed, with distinct scale-like sculpturing.

Redescription: Coloration ( Fig. 4 ): body dark brown to black, sometimes with yellow to reddish-brown colouring, particularly on legs; hemelytra of brachypterous morphs sometimes pale tan and brown. Surface and vestiture ( Figs 4 , 45A–H ): dorsum impunctate, pronotum, scutellum, and hemelytron sometimes rugulose; body clothed with long, black, bristle-like semi-erect setae; AI with two or three spines; femora sometimes with a few spines, tibiae strongly spinose; some species also densely clothed in white, scale-like setae. Structure : degree of wing polymorphism varies across species, both sexes macropterous or brachypterous. Head ( Figs 4 , 45A–C ): transverse, slightly broader than anterior of pronotum, always narrower than posterior of pronotum in macropterous males; posterior margin of vertex rounded to weakly carinate; frons flat to moderately convex, steeply declivent; gena height slightly greater than eye height; eyes round, substylate. Labium ( Fig. 45C ): variable, reaching from before procoxae to slightly beyond metacoxae; LI short and swollen. Antennae ( Figs 4 , 45A–C ): insertion close to and in line with lower margin of eye; thin and cylindrical, shorter than body length; AI sometimes somewhat swollen, short, sometimes slightly longer than eye height; AII sometimes apically clavate; AIII and AIV thin. Thorax ( Figs 4 , 45A–D ): pronotum trapezoidal, in macropterous morphs declivent with thin collar, in brachypterous morphs sometimes more rectangular and flat without collar; callosite region present but sometimes ill-defined, lateral margins rounded in brachypterous morphs, posterior margin straight to weakly concave; mesoscutum sometimes concealed by pronotum in brachypterous morphs; scutellum flat to somewhat swollen; metathoracic spiracle conspicuous and exposed, often large, narrowly surrounded with evaporative bodies that extend up to dorsolateral margin of metathorax; MTG efferent system swollen and triangular, extending as high as base of mesocoxae, ostiole vertical, opens laterally, peritreme oval, extending posterodorsally from ostiole. Hemelytra ( Fig. 4 ): macropterous – long and parallel-sided, all divisions present, cuneus long and narrow, membrane with two cells; brachypterous – membrane absent, all remaining divisions faint but present, cuneal fracture very short, posterior margin rounded. Legs ( Figs 4 , 45E ): metafemora moderately swollen; pretarsi without fleshy pulvilli. Male genitalia ( Figs 45G, H , 46A–D ): pygophore conical, posterior margin weakly asymmetrical, sometimes with thin apical serrate flange (e.g. Orthocephalus modarresi ); parameres of roughly equal length; left paramere L-shaped, sometimes with pronounced sensory lobe, apex of apophysis hooked, sometimes bifid; right paramere with spoon-like apex deflected laterally, with small apical apophysis; ductus seminis attenuate, with flexible ribbing, secondary gonopore moderately sclerotized, basally constricted to form dorsoventral bowl shape, apex dorsoventrally compressed and laterally expanded, with scale-like sculpturing primarily on ventral wall, dorsal wall longer than ventral wall; endosoma usually with two or three spinose spicules, rarely with none. Female genitalia ( Fig. 46E, F ): sclerotized rings widely separated, subtriangular, lateral and medial margins of DLP adjacent to sclerotized rings weakly sclerotized and upturned; VLP divided into anterior band attached to rami and medial plate, both portions weakly sclerotized; posterior wall of bursa copulatrix a simple, undivided, weakly sclerotized plate; vestibulum symmetrical, margins weakly sclerotized and unmodified .

Diversity and distribution: Orthocephalus includes 23 species and has a Holarctic distribution. One species, Orthocephalus coriaceus , is found both in Europe and eastern North America, whereas all other species are restricted to the Palaearctic region.

Included species: Orthocephalus arnoldii ( Putshkov, 1961) * Kazakhstan; Ukraine

Orthocephalus bivittatus Fieber, 1864 Turkey; Turkestan, East Asia (USSR)

Orthocephalus brevis ( Panzer, 1798) * Palaearctic

Orthocephalus championi Saunders, 1894 Corsica

Orthocephalus coriaceus ( Fabricius, 1777) * Europe; North America

Orthocephalus fulvipes Reuter, 1904 * Mediterranean; Asia Minor

Orthocephalus funestus Jakovlev, 1881 China ; Siberia; north-east Asia; Vladivostok

Orthocephalus medvedevi Kiritshenko, 1951 Ukraine

Orthocephalus melas Seidenstücker, 1962 Turkey

Orthocephalus minimus Drapolyuk & Kerzhner, 2000 Kazakhstan

Orthocephalus modarresi Linnavuori, 1997 Iran

Orthocephalus proserpinae ( Mulsant & Rey, 1852) Mediterranean

Orthocephalus putshkovi (Namyatova & Konstantinov) Ukraine; Kazakhstan

Orthocephalus rhyparopus Fieber, 1864 south Russia

Orthocephalus saltator ( Hahn, 1835) Holarctic

Orthocephalus scorzonerae Drapolyuk & Kerzhner, 2000 Uzbekistan; Turkmenistan; Kazakhstan

Orthocephalus sefrensis Reuter, 1895 Algeria

Orthocephalus solidus ( Seidenstücker, 1971 Turkey

Orthocephalus styx Reuter, 1908 Turkey; Russia; Kazakhstan

Orthocephalus tibialis ( Reuter, 1894) Algeria; Morocco; Tunisia

Orthocephalus tristis ( Reuter, 1894) Algeria

Orthocephalus turkmenicus Namyatova & Konstantinov, 2009 Turkmenistan; Iran

Orthocephalus vittipennis ( Herrich-Schäeffer, 1835) Palaearctic

Biology and host plant associations: Host records exist for 13 species, all of which have been collected mainly on asterids in the family Asteraceae , with a single species ( O. brevis ) known only from the Campanulaceae ( Hoberlandt, 1963; Göllner-Scheiding, 1972; Wagner, 1973). In addition to feeding on asterids, three species have also been collected off rosids, with O. coriaceus found on oak ( Fagaceae ) ( Ehanno, 1960, 1965), O. vittipenis on Ononis spinosa (Fabaceae) ( Göllner-Scheiding, 1972), and O. saltator on Spiraea sp. ( Namyatova & Konstantinov, 2009). Orthocephalus saltator has also been collected on unidentified grass ( Poaceae ) ( Ehanno, 1960; Gravestein, 1978), several species of Fabaceae ( Reuter, 1891; Saunders, 1892; Tamanini, 1981; Seidenstücker, 1959), as well as Thymus serpyllum (Lamiaceae) ( Reuter, 1891) and Salix sp. (Salicaceae) ( Namyatova & Konstantinov, 2009) ( Table 1).

Remarks: Orthocephalus is superficially similar to Dasyscytus , Pachytomella , and Anapus , but can be readily distinguished by the unique structure of the secondary gonopore. In a recent revision of Orthocephalus , Anapomella Putshkov and Oraniella Reuter were designated junior synonyms of Orthocephalus , based primarily on the secondary gonopore structure ( Namyatova & Konstantinov, 2009). This was supported through a phylogenetic analysis, which also proposed a sister relationship between Orthocephalus and Pachytomella . Our phylogeny supports this sister taxon relationship as well, based on two synapomorphies (69-1: opening of secondary gonopore slit-like; 83-1: DLP with pair of sickle-shaped sclerites adjacent to sclerotized rings). Interestingly, although we incorporated the coding of Namyatova & Konstantinov (2009) into our analysis, only the slit-shaped secondary gonopore opening was found to be synapomorphic in both analyses. This appears to be the result of differences in taxon sampling, as Namyatova & Konstantinov (2009) sampled all Orthocephalus , not all of which possess paired sickle-shaped sclerites on the DLP.

Figures

Figure 4. Photographs of Halticini genera: Labops–Strongylocoris. Abbreviations: F, female; M, male.

Figure 45. Scanning electron micrograph images of Orthocephalus brevis (male). A, head and thorax, dorsal view; B, head, anterior view; C, head and prothorax, lateral view; D, meso- and metathorax, lateral view; E, tarsus; F, pygophore, dorsal view; G, pygophore, posterior view; H, pygophore, ventral view.

Figure 46. Male and female genitalia of Orthocephalus brevis. A, aedeagus; B, secondary gonopore; C, left paramere; D, right paramere; E, posterior wall; F, bursa copulatrix, dorsal view.