Anapus, STAL, 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 : 573-578

ANAPUS STÅL ( FIGS 3 , 7–10 )

Anapus Stål, 1858: 188 (gen. nov.; type species: Anapus kirshbaumi Stål, 1858: 189 by monotypy); Walker, 1873: 160 (list); Reuter, 1891: 69, 159 (key; descr.); Kirkaldy, 1906: 131 (list); Hueber, 1906: 5 (key); Oshanin, 1910: 787 (cat.); Reuter, 1910: 147 (cat.); Stichel, 1933: 235 (key); Kiritshenko, 1951: 126 (key); Wagner, 1952: 96, 106 (key; descr.); Carvalho, 1955: 66 (key); Carvalho, 1958: 6 (cat.); Kerzhner, 1964a: 965 (diag., key to spp.); Wagner, 1973: 55 (descr., key to spp.); Schuh, 1995: 45 (world cat.).

Labops (Merotrichaea) Reuter, 1875c: 24 (subgen. nov.; type species: Orthocephalus freyi Fieber, 1864 , by subsequent designation Kirkaldy, 1906); Reuter, 1875b: 87 (key); Reuter, 1891: 79 (synonymy).

Merotrichia Carvalho, 1958: 6 (cat.; generic status).

Platyporus Reuter, 1890: 246 (gen. nov.; type species: Platyporus dorsalis Reuter, 1890 by monotypy); Carvalho, 1958: 30 (cat.); Wagner, 1973: 53 (descr.); Schuh, 1995: 69 (world cat.). New synonymy. Diagnosis: Recognized by the following combination of characters: substylate eyes; antennal insertion below ventral margin of eye; AI swollen and longer than height of eye, with several elongate spines; hemelytra of brachypterous individuals very short; male ductus seminis long, with elongate apical section lacking ribbing; secondary gonopore rounded, opening ventrally, with prised operculum; and, female posterior wall of bursa copulatrix often with bilaterally symmetrical, sclerotized inter-ramal tumescences.

Redescription: Coloration ( Fig. 3 ): Body uniformly black with minor yellow and orange markings; appendages black. Surface and vestiture ( Figs 3 , 7A–E, G ): Body impunctate, sometimes pronotum, scutellum and hemelytra rugulose, often with wrinkles on vertex and frons radiating from midline. Body with erect and semi-erect simple setae, sometimes intermixed with dense decumbent, white, scalelike setae. Head with a few elongate thin erect black setae. Antennae with semi-erect, spine-like setae; AI with a few to numerous elongate spines, especially in males. Legs with elongate sharp spines, particularly on metatibiae. Structure: both sexes usually brachypterous, rarely macropterous. Head ( Figs 3 , 7A–C, E ): transverse, triangular in shape in dorsal view, with posterior margin straight to very weakly concave; head in lateral view angled somewhat caudally below antennae; height of head below eye approximately twice height of eye; clypeus strongly declivent, not projecting forward; posterior margin of vertex not carinate; frons rounded, strongly declivent in lateral view; posterior margin of head wider than anterior of pronotum; eyes substylate; Antennae ( Figs 3 , 7A–C, E ): antennal insertion well below eye; subequal to body length; AI thickened, sometimes incrassate; remaining segments thin. Labium ( Fig. 7E ): length variable, reaching from mesocoxae to basal abdomen sternites; LI incrassate. Thorax ( Figs 3 , 7A–D ): pronotum short, weakly declivent in lateral view, rectangular in brachypterous forms, trapezoidal in male macropterous individuals; collar absent; callosite region sometimes distinct, lateral margins rounded; posterior margin weakly concave; mesoscutum only visible in macropterous individuals; scutellum triangular, flat to tumescent; metathoracic spiracle large and exposed, sometimes with evaporative bodies along posterior margin; MTG external efferent system broad, swollen and triangular, generally reaching lower margin of metathoracic spiracle; peritreme oval, medially on metepisternum, subparallel to posterior margin of pleuron. Hemelytra ( Figs 3 , 7A–B ): usually brachypterous, sometimes reduced to a small pad, in macropterous morphs with single membrane cell, minor cell obsolete. Legs ( Figs 3 , 7A–B, E–F ): metafemora strongly incrassate; foretibiae usually swollen apically and weakly arcuate; pretarsi with elongate fleshy parempodia (except Anapus americanus ). Abdomen ( Figs 3 , 7G ): pear-shaped, elongate oval, or oval. Male genitalia ( Figs 8A–C , 7B, E, G , 9A–C ): pygophore triangular, posterior margin concave at left paramere insertion; right paramere sometimes projecting slightly from pygophore; phallotheca simple, sometimes moderately sclerotized apically; left paramere sensory lobe broad, apophysis thin, hooked; right paramere broad, spoonshaped, with small apical apophysis; phallotheca simple, without processes; ductus seminis elongate with flexible ribbing, sometimes very slender (as in Anapus freyi ), subapically with elongate section, sometimes heavily sclerotized, lacking flexible ribbing; secondary gonopore elongate, U-shaped, with semiclosed operculum; endosoma with one or two simple spicules of varying size and shape, in most species also with paired, elongate, and sclerotized spiculate strips near apex. Female genitalia ( Figs 8D, E , 10A–C, 10A–F ): sclerotized rings moderate to large, elongate, lateral margins weakly upturned along with adjacent portion of DLP, DLP sometimes with heavily sclerotized inter-ramal bridge; margin of VLP adjacent to rami sclerotized, medially with a small flange; posterior wall of bursa copulatrix variable, but always with bilaterally symmetrical, sclerotized structures (inter-ramal tumescences), sometimes toothed or serrate; margins of vestibular opening symmetrical with weak sclerotization.

Diversity and distribution: Anapus has seven Holarctic species, one from the western USA and the remainder Palaearctic.

Included species: Anapus americanus Knight, 1959 * USA

Anapus dorsalis ( Reuter, 1890) * comb. nov. Russia; Turkey

Anapus freyi ( Fieber, 1864) * Russia

= Pachytoma jakovloeffi Reuter, 1879 (syn. Reuter, 1881)

Anapus kirschbaumi Stål, 1858 * Russia

= Orthocephalus opacus Jakovlev, 1875 (syn. Jakovlev, 1877)

= Anapus pectoralis Horváth, 1904 (syn. Kerzhner, 1996)

Anapus longicornis Jakovlev, 1882 * Hungary; Russia

= Labops (Pachytoma) arenarius Horváth, 1884 (syn. Reuter, 1890)

= Anapus flavicornis Reuter, 1904 (syn. Kerzhner, 1970)

Anapus pachymerus ( Reuter, 1881) Spain

Anapus rugicollis ( Jakovlev, 1877) * Russia

= Pachytoma sibirica Sahlberg, 1878 (syn. Kerzhner, 1962)

= Pachytoma nigrita Jakovlev, 1881 (syn. Kerzhner, 1962)

Biology and host plant associations: Host plant records are known for four species of Anapus ( Table 1; Schuh, 1995). Anapus freyi is known from a grass species, and the other three species are known from either euasterid or eurosid angiosperms.

Remarks: Herein we propose Platyporus as a junior synonym of Anapus , resulting in the new combination: Anapus dorsalis ( Reuter, 1890) . This species conforms to the generic diagnosis of Anapus given in this work, and shares significant overlap in salient morphology, and striking similarities in the male aedeagus and female posterior wall of the bursa copulatrix. This synonymy is supported by our phylogenetic analysis.

In all the Old World species of Anapus examined the male genitalia are nearly identical, with the following similarities: the distal portion of the ductus seminis is sclerotized, the endosoma is relatively small, bearing one or two spicules and apically with sclerotized fields of small spines, and the left paramere has a broad sensory lobe. However, in the New World species, An. americanus , the distal portion of the ductus seminis is weakly sclerotized, the endosoma is much larger with the apical fields of small spines only faintly visible, and the left paramere lacks the large sensory lobe present in other species. Additionally the body is much more ant-like in shape, the hemelytra are reduced to very small pads, the pretarsi lack fleshy pulvilli, and the first antennal segment is thinner, more elongate, and has fewer black spines. Despite these differences we consider An. americanus to represent a divergent New World lineage within Anapus rather than a new monotypic genus, based primarily on commonalities in the structure of the female posterior wall of the bursa copulatrix. Again, this is supported by our phylogenetic results.

Anapus shares some notable female genitalic peculiarities with several other taxa: in particular, the sclerotized inter-ramal bridge of the DLP and the inter-ramal lobes of the posterior wall (characters 82 and 88) are both shared with Labops ( Slater, 1950; Slater, 1954), with the latter of these structures also found in Scirtetellus . In these other genera and An. americanus , the inter-ramal lobes are hollow, apically tapered, and medially convergent, and spinose. These appear homologous to the inter-ramal lobes found in the Orthotylini ( Slater, 1950; also known as K-structures), although they differ in that they are generally broader and much less narrowly constricted basally (89-1).

Aside from An. pachymerus , all species of Anapus were examined for this study.

Figures

Figure 3. Photographs of Halticini genera: Acratheus–Halticus. Abbreviations: F, female; M, male.

Figure 7. Scanning electron micrograph images of Anapus longicornis (male). A, dorsum; B, lateral view; C, head and thorax, lateral view; D, external efferent system of the metathoracic scent gland; E, venter; F, tarsus; G, abdomen, lateral view.

Figure 8. Male and female genitalia of Anapus kirshbaumi. A, aedeagus; B, left paramere; C, right paramere; D, bursa copulatrix (dorsal view); E, posterior wall. Abbreviations: EN, endosoma; IRL, inter-ramal lobe; SG, secondary gonopore; SP, spicule.

Figure 9. Male and female genitalia of Anapus dorsalis. A, aedeagus; B, left paramere; C, right paramere; D, bursa copulatrix, dorsal view; E, bursa copulatrix, ventral view; F, posterior wall, ventral view.

Figure 10. Female genitalia of Anapus americanus. A, bursa copulatrix, dorsal view; B, posterior wall; C, first gonapophyses, ventral view. Abbreviations: IRL, inter-ramal lobe; SR, sclerotized ring.