Halticus, HAHN, 1832

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 : 604-607

HALTICUS HAHN ( FIGS 3 , 30–32 )

Astemma Latreille, 1829: 199 (nom. nud.); junior homonym of Astemma Lepeletier & Serville, 1825 [ Heteroptera : Pyrrhocoridae ]; type species: Acanthia pallicornis Fabricius, 1794 = Cicada aptera Linnaeus, 1758 by monotypy; Westwood, 1840: 121 (description); Amyot & Serville, 1843: 284 (description)

Halticus Hahn, 1833: 113 (gen. nov.; type species: Acanthia pallicornis Fabricius, 1794 = Cicada aptera Linnaeus, 1758 by monotypy); Burmeister, 1835: 277 (descr.; Spinola, 1850: 43 (key); Fieber, 1858: 312 (key); Fieber, 1861: 69, 281 (key; descr.); Reuter, 1875a: 23 (key); Reuter, 1875b: 86, 89 (key; descr.); Saunders, 1875: 287 (key); Atkinson, 1890: 117 (cat.); Reuter, 1891: 17, 157 (descr.; key); Saunders, 1892: 267 (key; descr.); Distant, 1904: 479 (descr.); Kirkaldy, 1906: 130 (cat.); Hueber, 1906: 2, 42 (key; descr.); Oshanin, 1910: 802 (cat.); Reuter, 1910: 147 (cat.); Poppius, 1914: 84 (key; descr.); Van Duzee, 1916: 211, 374 (cat.); Knight, 1923: 498 (key); Blatchley, 1926: 800 (eastern US species key); Stichel, 1933: 235 (Palaearctic key); Hedicke, 1935 (Palaearctic key); Knight, 1941: 17, 75, 77 (Illinois key; descr.); Hsiao, 1942: 253 (Chinese key); China, 1943: 268 (nomenclature); Zimmerman, 1948: 199 (Hawaiian key); Slater, 1950: 45 (female genitalia); Kiritshenko, 1951: 127 (USSR key); Wagner, 1952: 95, 96 (Palaearctic key; descr.); Carvalho & Leston, 1952: 245 (British key); Carvalho, 1952: 74 (list); Carvalho, 1955: 65 (key); Carvalho, 1956: 51 (key Micronesian spp.); Carvalho, 1958: 12 (cat.); Southwood & Leston, 1959: 244 (British key); Kelton, 1959: 33, Fig. 81 (male genitalia); Wagner, 1961: 48 (diag.; Palaearctic key); Kerzhner, 1964a: 968 (diag.; USSR key); Maldonado, 1969: 57 (diag.; US key); Alayo, 1974: 15 (diag.); Schuh, 1974: 29 (discussion); Wagner, 1973: 6 (descr.; Palaearctic key); Kelton, 1980: 194 (diag.); Henry, 1983: 610 (Western Hemisphere key); Linnavuori, 1994: 5 (African key); Cassis & Gross, 1995: 186 (synonymy; Australian cat.); Schuh, 1995: 53 (synonymy; world cat.).

Eurycephala Laporte, 1832: 36 (gen. nov.; Lygaeus luteicollis Panzer, 1805 by subsequent designation Kirkaldy, 1906: 170); Brullé, 1836: 140 (descr.); Spinola, 1840: 191 (descr.); Blanchard, 1840: 140 (descr.); Reuter, 1891: 17 (synonymy); Carvalho, 1958: 12 (cat.)

Halticocoris Douglas & Scott, 1865: 478 (unnecessary nom. nov. for Halticus Hahn nec Haltica Illiger [ Coleoptera ]); Carvalho, 1958: 13 (cat.);

Halticiellus Villiers, 1952: 186 (gen. nov.; type species: Halticus tibialis Reuter, 1891 by monotypy); Carvalho, 1958: 12 (synonymy)

Diagnosis: Halticus is characterized by a combination of the following: length of first labial segment less than gena height; elongate, flattened right paramere; left paramere with twisted apophysis; scoop-shaped secondary gonopore and vestigial endosoma lacking spicules; large, thin sclerotized rings.

Redescription: Usually small and robust, macropterous or coleopteroid. Coloration ( Fig. 3 ): almost all species black, sometimes with paler yellow, orange, red, or brown colouring on head, antennae, and legs, rarely uniformly brown. Surface and vestiture ( Figs 3 , 30A–E, G, H ): variable, generally glossy, sometimes wholly impunctate, in others pronotum is shallowly punctate; frons and vertex with faint creases radiating from centreline. Body generally with two forms of setae – thin and simple and white, flat, and scale-like; antennae and legs with short semi-erect spines and spine-like setae. Structure: head ( Figs 3 , 30A, B, D ): transverse and short, ranging from broader than tall to taller than broad; vertex with transverse sulcus, posterior margin carinate, concave, generally broader than anterior of pronotum; head height variable, usually> two times eye height; eyes sometimes wrapping around pronotum, sometimes touching; eyes tall and short; clypeus narrow, dorsal margin weakly sulcate; mandibular plate depressed; maxillary plate raised and carinate. Labium ( Fig. 30D ): length variable, LI greatly swollen. Antennae ( Figs 3 , 30A–D ): insertion typically in line with or slightly below lower margin of eye; long, thin, and whip-like; AI slightly thicker than other segments, about as long as eye height. Thorax ( Figs 3 , 30A, D–F ): pronotum trapezoidal (macropterous) to subrectangular (coleopterous), weakly declivent, laterally sloping, collar thin and flat to absent; callosite region ill-defined to obsolete, posterior margin thinly carinate, straight to weakly medially cleft; metathoracic spiracle prominent, narrowly bordered with evaporative bodies; MTG external efferent system swollen, occupying lower half of metepimeron, ostiole opens laterally, peritreme rounded, orientated posteriorly, surrounded by evaporative bodies. Hemelytra ( Fig. 3 ): macropterous: costal margin weakly curved, broadest above cuneal fracture; clavus longitudinally folded, with lateral side steeply sloping; corium laterally sloping; membrane and cuneus deflected. Coleopterous: broadly rounded, laterally and posteriorly sloping; claval suture and medial fracture faint, cuneal fracture very short. Legs: long, metafemur greatly swollen; pretarsi with fleshy pulvilli. Male genitalia ( Figs 30H , 31A–E ): pygophore conical, posterior margin sinuate, with broad deep concavity below left paramere; phallotheca simple and elongate-oval; left paramere apically twisted with sinuate apophysis; right paramere longer than left, extends out of pygophore, broad, flat, sometimes angled laterally at midpoint, apex rounded with a small apicolateral apophysis; ductus seminis attenuate, with flexible ribbing; secondary gonopore partly sclerotized, in form of elongate scoop, nearly filling phallotheca; endosoma extremely reduced, without spicules. Female genitalia ( Figs 31F–H , 32A– C ): dorsal labiate plate mostly membranous; sclerotized rings large, transverse, medially subcontiguous, sclerotized margins thin, lateral margins weakly upturned; VLP weakly sclerotized on posterior margin, laterally sometimes with paired, scimitarshaped sclerotizations, curving laterally and reaching up to dorsal labiate plate; posterior wall of bursa copulatrix variable, for the most part membranous, often with narrow paired sclerotized bands or rods convergent anteriorly; vestibulum symmetrical, with medial margins of first gonapophyses swollen outwards and sclerotized, vestibular opening rounded and strongly sclerotized.

Diversity and distribution: Halticus has 16 species and a mainly Holarctic distribution, with most species confined to the Palaearctic. A single species, Halticus tibialis , is found in Africa, South-East Asia, and the south-west Pacific.

Included species: Halticus apterus ( Linnaeus, 1758) * Palaearctic

Halticus asperulus Horváth, 1898 Armenia

Halticus beganus Linnavuori, 1984 Iraq

Halticus darbandikhaus Linnavuori, 1984 Iraq

Halticus henschii Reuter, 1888 Italy

Halticus insularis Usinger, 1946 Mariana Is. ; Guam; Korea

Halticus intermedius Uhler, 1904 USA

Halticus luteicollis ( Panzer, 1804) * Europe; Mediterranean

Halticus macrocephalus Fieber, 1858 Mediterranean

Halticus major Wagner, 1951 Europe

Halticus obscurior Kerzhner & Muminov, 1974 Asiatic Russia

Halticus puncticollis Fieber, 1870 Montenegro

Halticus pusillus ( Herrich-Schäeffer, 1835) * Palaearctic

Halticus rugosus Reuter, 1894 Syria

Halticus saltator ( Geoffroy, 1785) Europe

Halticus tibialis Reuter, 1891 * Africa; South-East Asia, South Pacific

Biology and host plant associations: Limited host records exist for Halticus . Halticus apterus and H. beganus have both been collected from the family Fabaceae ( Vicia striata and Phaseolus vulgaris , respectively). Halticus beganus has also been collected from rotten oak ( Fagaceae ) ( Henry & Smith, 1979). Halticus pusillus is said to live on and beneath Galium verum (Rubiaceae) ( Wagner, 1973) ( Table 1). Remarks: Most recent definitions of Halticus are broad, inclusive of great morphological heterogeneity (e.g. Wagner, 1973), and pay little regard to genitalic features. This is the first study to account for the morphological diversity exhibited across species of Halticus , and to examine this in a phylogenetic context. As a consequence we have subdivided Halticus sensu lato into three genera. Based primarily on male and female genitalic characters, and supported by our phylogeny, we remove ten species examined that do not follow the model of the type species, H. apterus (left paramere L-shaped with apically curved apophysis; ductus seminis short; secondary gonopore elongate and scoop-shaped, endosoma obsolete and without spicules; sclerotized rings large and subcontiguous). H. bractatus , H. canus , H. chrysolepis , H. minutus , and H. spegazzini are removed, and along with both species of Cafayatina , form the resurrected genus Microtechnites . Similarly, the Asian species H. bicoloratus , H. comitans , H. maculipes , H. niger , and H. fuscous are all transferred to Ectmetopterus .

We have not seen all species of Halticus , and it is therefore possible that with further examination other species may yet be removed from the genus.

Figures

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

Figure 30. Scanning electron micrograph images of Halticus apterus (male). A, head and pronotum, dorsal view; B, head, anterior view; C, antenna; D, head and thorax, lateral view; E, thorax, lateral view; F, detail of external efferent system of the metathoracic scent gland evaporative bodies; G, tarsus; H, pygophore, ventral view.

Figure 31. Male and female genitalia of Halticus apterus: A, phallotheca; B, aedeagus, phallotheca removed; C, secondary gonopore, ventral view; D, left paramere; E, right paramere; F, bursa copulatrix, dorsal view; G, posterior wall; H, bursa copulatrix, ventral view.

Figure 32. Female genitalia of Halticus luteicollis. A, bursa copulatrix, dorsal view; B, first gonapophyses, ventral view; C, posterior wall.