Pinthaeus sanguinipes (Fabricius, 1781)

Zhao, Qing, Rédei, Dávid & Bu, Wenjun, 2013, A revision of the genus Pinthaeus (Hemiptera: Heteroptera: Pentatomidae), Zootaxa 3636 (1), pp. 59-84 : 61-70

Pinthaeus sanguinipes (Fabricius, 1781)

Figs. 1–32

Cimex sanguinipes Fabricius, 1781: 344 . Syntype (s): Italy; ZMUC!

Asopus genei Costa, 1842: 299 . Holotype: Italy, Astroni, near Naples; ZIUN. Synonymized by Baerensprung (1860: 5). Pentatoma rufipes (non Linnaeus, 1758): Kato (1933: [legend to plate 5]). Misidentification (Kiritshenko 1961: 444). Pinthaeus humeralis Horváth, 1911: 432 . Holotype (Ƥ): China: Yunnan, “Tali” [= Dali]; HNHM! New synonym. Pinthaeus sangvinipes: Reuter (1909: 94) . Incorrect subsequent spelling.

References. Fabricius 1787: 283 (diagnosis), Villers 1789: 491 (diagnosis, distribution), 692 (listed), Petagna 1792: 631 (diagnosis, distribution), Fabricius 1794: 93 (diagnosis, redescription, distribution), Fabricius 1803: 156 (diagnosis, distribution), Latreille 1804: 188 (diagnosis, distribution), Schilling 1844: 180 (diagnosis), Herrich- Schäffer 1835: 93 ( catalogue ), Herrich-Schäffer 1839: 101 (diagnosis, habitus, record), Costa 1847: 396 ( genei , redescription, record), Gorski 1852: 120 (redescription, record), Dohrn 1859: 7 ( catalogue , distribution), Baerensprung 1860: 5 ( catalogue , synonymy, distribution), Fieber 1861: 348 ( sanguinipes and genei , redescription, distribution), Mulsant & Rey 1866: 336 (redescription, distribution), Walker 1867: 123 (listed, records), Garbiglietti 1869: 48 ( sanguinipes , genei , listed, distribution), Stål 1870: 46 ( catalogue , distribution), Scott 1874: 289 (listed), Puton 1875: 15 ( catalogue , distribution), Puton 1881: 78 (redescription, records, distribution), Horváth 1883: 25 (record), Autran & Reuter 1888: 199 (record), Reuter 1888: 515 ( catalogue ), Reuter 1890: 238 (records), Coubeaux 1891: ccclxxxix (listed), Hüeber 1891: 76 (distribution), Griffini 1894: 87 (redescription, records), Lethierry & Severin 1893: 216 ( catalogue , distribution), Giglio-Tos 1894: 4 (doubtful record), Horváth 1897: 21 (records), Puton 1899: 17 ( catalogue ), Chopard 1905: 130 (record), Oshanin 1906: 152 ( catalogue , distribution), Montandon 1907: 59 (records), Schouteden 1907: 40 ( catalogue , distribution, habitus), Reuter 1909: 94 ( sangvinipes , record, habitat), Kirkaldy 1909: 14 ( catalogue , distribution), Oshanin 1910: 62 ( catalogue , distribution), Schumacher 1910a: 264 (in key), Schmacher 1910b: 380 (habitus, records, habitat, phenology), Kiritshenko 1911: 41 (record, distribution), Oshanin 1912: 18 ( catalogue , distribution), Kiritshenko 1916: 80 (record), Kiritshenko 1918: 55 (records), Gulde 1921: 356 (records, distribution, habitat, phenology, prey, larva), Seabra 1924: 25 (redescription, habitus, record), Stichel 1925: 34 (diagnosis, biology, distribution), Seabra 1926a: 161 (redescription, habitus, food, phenology, record), Seabra 1926b: 59 (record), Seabra 1929: 78 (diagnosis, distribution, prey, phenology), Hoffmann 1932: 7 ( humeralis , listed), Reclaire 1932: 82 (distribution), Rosenkranz 1939: 304 (midgut crypt symbionts), Gulde 1934: 170 (redescription, distribution, habitat, prey), Lindberg 1934: 6 (records), Hoffmann 1935: 107, 174 ( humeralis , catalogue , distribution), Stichel 1935: 362 ( catalogue ), Yang 1935: 106 ( humeralis , reproduction of original description and figure), Bodenheimer 1937: 200 (listed), Ookubo 1937: 115 (records), Stichel 1938: 436 (distribution), Tempel 1939: 51 (photo of larva, records, mass occurrence, prey, distribution), Seabra 1941: 13 (listed), Balthasar 1942: 26 (record), Dupuis 1947: 54 (description and figures of 5th instar larva), Ishihara 1947: 67 (distribution), Gravestein 1949: XXV (record, distribution, diagnostic characters), Dupuis 1949: 238 (in key), 241 (in key to larvae), 245 (figure of head and pronotum, distribution, habitat, prey, phenology), Vidal 1949: 204 (redescription, distribution), Kiritshenko 1951: 344 (distribution), Reclaire 1951: 4 (record), Esaki 1952: 208 (redescription, habitus, distribution), Takeuchi 1955: 45 (photo, record, distribution, habitat), Hoberlandt 1956: 205 (record, distribution), Nagy 1957: 263 (photo, record, prey, biology, behaviour, phenology, distribution), Halászfy 1959: 78 (redescription, figure, biology, distribution), Putshkov 1961: 309 (redescription, larva, distribution, habitat, biology, prey, phenology), Stichel 1961a: 653 (redescription, habitus, habitat, distribution), Stichel 1961b: 767 ( sanguinipes , humeralis , listed, distribution), Wagner 1961: 159 (diagnosis, habitat), Stichel 1962: 248 ( catalogue , distribution), Yang 1962: 52 ( humeralis , in key), 65 ( humeralis , redescription, host plant, distribution), Zebe 1963: 25 (record, habitat, abundance), Kerzhner & Jaczewski 1964: 844 (diagnosis, distribution), Servadei 1964: 76 (spermatheca, figure), Zimsen 1964: 314 (type material), Wagner 1966: 77 (redescription, figure, habitat, prey, phenology, distribution), Servadei 1967: 520 ( catalogue , distribution), Josifov 1969: 58 (records, distribution), Lughofer 1972: 106 (record), Rieger 1972: 161 (record, distribution), Hsiao & Cheng 1977: 80 ( humeralis , sanguinipes , in key, redescription, distribution), Josifov & Kerzhner 1978: 181 (records), Dioli 1979: 86 (records, distribution), Rieger 1979: 267 (conservation), Tomokuni 1979: 104 (distribution), Zhang et al. 1980: 24 ( humeralis , record, phenology), Josifov 1981: 185 (redescription, figure, distribution), Zhang & Lin 1981: 136 ( humeralis , record, distribution), Gidayatov 1982: 117 (distribution, records), Lodos & Önder 1983: 223 (record, distribution, prey), Chen & Yang 1984: 99 ( humeralis , listed, distribution), Kis 1984: 186 (redescription, habitus, figures, genitalia, biology, distribution), Wu 1984: 37 ( humeralis , listed, records), Dioli 1985: 230 (records, diagnosis, figure of head and pronotum, habitus, biology), Jiang 1985: 59 ( humeralis , records), Josifov 1986: 90 (distribution), Zhang 1986: 426 (listed), Chen 1987: 155 ( humeralis , redescription, habitus, phenology, distribution), Dioli 1987: 617 (listed), Ji 1987: 23 ( humeralis , listed, distribution), Zhang et al. 1987: 126 ( humeralis , in key, distribution, figures), Stehlík 1987: 190 (records, habitat, distribution, phenology, biology, prey, distribution, egg, figure), Kanyukova 1988: 929 (diagnosis, habitat, distribution), Zhang & Lin 1988: 76 ( humeralis , records), Zheng et al. 1988: 2 ( humeralis , record), Aukema 1989: 1 (distribution), Chan et al. 1989: 282 ( humeralis , listed, prey), Miyamoto & Yasunaga 1989: 187 (listed, distribution), Günther & Schuster 1990: 391 (listed), Heiss & Josifov 1990: 146 (distribution), Lis 1990: 67 (records, distribution, biology), Ribes 1990: (record, distribution, diagnostic characters), Ren 1992: 34 ( humeralis , egg, larva), Zhang & Lin 1992: 19 ( humeralis , listed), Zhang et al. 1992: 231 ( humeralis , redescription, habitus, record, distribution), Tomokuni 1993: 235, plate 124 (redescription, habitat, prey, distribution, records), Kim & Lee 1994: 220 (spermatheca), Thomas 1994: 193 ( sanguinipes , humeralis , catalogue , distribution), Zhang 1994: 35 (listed), Chen 1995: 31 ( humeralis , redescription, habitus, larva, distribution, prey), Faraci & Rizzotti Vlach 1995: 46 (listed, distribution), Redl & Kallenborn 1995: 21 (etymology), Ribes & Goula 1995: 59 (record), Rabitsch & Waitzbauer 1996: 207 (record), Rieger 1996: 251 (listed), Bu & Zheng 1997: 191 ( humeralis , redescription, figure, records, distribution), Derzhansky 1997: 19 (listed), Hoberlandt 1997: 270, 277 (record, distribution), Lukashuk 1997: 33 (distribution), Shen & Shi 1998: 231 ( humeralis , listed, records), Dolling et al. 1999: 81 ( catalogue ), Kondorosy 1999: 148 (listed), Lin et al. 1999: 63 (redescription, habitus, record, distribution), Hua 2000: 178 ( sanguinipes , humeralis , listed, distribution, prey), Friess 2000: 68 (photo, record, habitat, prey, distribution), Günther & Schuster 2000: 53 (distribution), Lin et al. 2000: 38 ( sanguinipes , humeralis , distribution), Lis 2000: 66 (redescription, distribution, habitat, prey, phenology), Rieger 2000: 203 (rearing, biology, prey, development, egg, larval instars, photos, figures, records), Ding et al. 2001: 98 (zoogeography), Kment & Bryja 2001: 242 (record), Kondorosy 2001: 130 (listed), Kwon et al. 2001: 403 ( catalogue , records, distribution, habitat), Protiċ 2001: 196 (records), Tolsgaard 2001: 24, 38 (photo, record, biology, map), Rider & Zheng 2002: 111 ( humeralis , sanguinipes , distribution), Baugnée et al. 2003: 49 (listed), Guo & Zheng 2003: 90 ( humeralis , in key), Lin 2003: 111 ( sanguinipes , humeralis , distribution), Bartels et al. 2004: 247 (listed, conservation), Negrobova 2005: 316 (records), Rabitsch 2005: 56 (listed), Wang & Liu 2005: 282 (record, distribution), Bryja & Kment 2006: 290 (records), Dusoulier & Lupoli 2006: 34 (listed, distribution, biology), Gapon & Konstantinov 2006: 810 (records, phallus), Li et al. 2006: 310 ( humeralis , sanguinipes , listed, records), Liu & Ju 2006: 327 ( humeralis , listed), Nikolaeva 2006: 116 (records, distribution), Rabitsch 2006: 509 (listed), Ren et al. 2006: 274 ( humeralis , listed), Rider 2006: 244 ( humeralis , sanguinipes , catalogue , distribution), Enju 2007: 95 (record), Guo et al. 2007: 343 ( humeralis , sanguinipes , listed, diet), He et al. 2007: 95 ( sanguinipes , humeralis , listed, records), Li & He 2007: 349 ( sanguinipes , humeralis , listed, record), Rabitsch 2007: 49, 53, 212 (colour photo, distribution, habitat, prey, conservation), Gogala 2008: 108 (records), Harmat 2008: 48 (record), Hanel & Kment 2009: 34 (diagnostic characters, photos, biology, records), Linnavuori 2008: 7 (record, distribution), Mohaghegh 2008: 2 (listed), Protiċ 2008: 198 (record), Ren et al. 2008: 195 ( sanguinipes , humeralis , record, diet), Wachmann et al. 2008: 89 (photos of adult and larvae, distribution, habitat, prey, phenology), Aukema & Hermes 2009: 74 (record, distribution, prey, habitat, phenology, photo of larva), 87 (listed), Aukema et al. 2009: 29 (record, distribution, habitat, prey, phenology), Esser 2009: 127 (records, distribution), Hebda & Przewożny 2009: 65 (record), Kanyukova & Vinokurov 2009: 60 (record), Kment et al. 2009: 43 (photos, records, distribution, biology, long-term population changes), Neimorovets 2010: 96 (records), Péricart 2010: 205 (redescription, figures, photos, egg. larva. habitat, biology, prey, phenology, records, distribution), Ren & Ba 2010: 139 ( humeralis , record, distribution), Vinokurov et al. 2010: 244 ( catalogue , distribution), Wyniger & Kment 2010: 264 (in key, photo), Baužys 2011: 9: (record), Bury 2011: 1 (record, photos, habitat, biology), Kment & Kejval 2011: 45 (record), Protiċ 2011: 123 (record), Tarnawski 2011: 22 (records, distribution), Kment & Baňař 2012: 598 (records, distribution, habitat, prey).

Diagnosis. Being the single included species of Pinthaeus it can be diagnosed by the combination of the characters of the genus.

Redescription. Colour. Dorsum ( Figs. 3–5 ) dark brown or reddish-brown, with black or brown punctures; head black, lateral margin and midline occasionally with yellow to red callosity ( Fig. 7 ) which is sometimes extensive ( Fig. 8 ); antenna black, except basal two-third of segment IV yellow; rostrum yellowish to light brown, segment IV darker; anterior lobe and humeral processes of pronotum greatly black due to confluent black punctures, lateral margin with yellow to red callosity ( Fig. 7 ) which is sometimes extensive, occasionally occupies most of anterior lobe ( Fig. 8 ); posterior lobe, scutellum, clavus and corium of fore wings brown, lateral callosities and apex of scutellum whitish to yellow, membrane smoky brown; legs yellowish brown to brown, frequently reddish, femora with yellow longitudinal patches ventrally, mid and hind tibiae with a distinct yellow annulus occupying middle third; tarsi brown, apical segment black; dorsum of abdomen black, connexivum with middle third of segments III–VI and posterior two-third of segment VII yellow. Venter yellow to reddish with black markings as follows: ventral surface of mandibular plates; a black spots at ventral surface of humeral process, along callose lateral margin of propleuron, and around prothoracic supracoxal lobes due to confluent black punctation; more or less extensive spots on pterothoracic pleura mainly sublaterally, at supracoxal lobes, and at apex of peritreme of metathoracic scent gland ostiole; most of mesosternum except median longitudinal carina; a large median spot at base of abdominal sternites IV–VII each, patches of variable extension formed by black punctures on sternites III–VII arranged longitudinally submedially and sublaterally; a small spot surrounding spiracles II–VII each; a marginal spot occupying each intersegmental borders from segments II to VII.

Integument and vestiture. Head densely punctured and somewhat rugose, lateral margin and midline occasionally with callose areas of variable extension, sometimes occupying majority of dorsal surface of head ( Fig. 8 ); anterior lobe of pronotum densely, posterior lobe distinctly more sparsely punctured; anterior and lateral margins of pronotum callose, callosities sometimes extensive and occupy most of anterior lobe of pronotum except cicatrices ( Fig. 8 ); punctation on humeral process of pronotum dense, confluent; base of scutellum with a pair of sublateral callosities; venter with dense, colourless punctation and with several black spots which form regularly distributed black patches.

Head subequal in length to width across eyes or slightly longer than broad, about 1.65–1.7 times as broad as interocular distance; anteocular region more narrow immediately anteriad to eye than apically; mandibular plates contiguous anteriad of apex of clypeus ( Fig. 7 ) or separated by a narrow gap ( Fig. 8 ). Pronotum 2.1–2.9 times as broad as long, ratio strongly varies because of variable development of humeri ( Figs. 9–14 ).

External male genitalia. Genital capsule (Figs. 15–16) transversely oval in posterior view, ventral rim excised medially, with a small rounded impression below excision; intersegmental membrane IX–X with a pair of dorsal (Fig. 16: dsim9-10) and a smaller pair of ventral sclerites (Fig. 16: vsim9-10). Paramere (Figs. 21–23) with a sharp, subtriangular, flattened blade directed laterad in rest (Fig. 16) and a short, obtuse dorsal projection. Phallus. Articulatory apparatus: basal plates (Figs. 17–18: bp) extending posterolaterally, basolateral portion broadened, with a pair of short, obtuse basal projections directed anteriad; support bridge complex (Figs. 17–18: sbc) completely fused with basal plates, terminating in short dorsal connective (Fig. 17: dc) connected to a small capitate process (Fig. 17: cap); erection fluid pump (Fig. 17: erp) elongate; ductifer (Fig. 18: df) short; support bridge prolongation (Fig. 18: sbp) tubular, surrounding ductus seminis (Fig. 18: ds), attached to base of second conjunctival process. Hinge (Fig. 18: hi) strongly protruding, rounded, with rather strongly sclerotized wall. Phallotheca short, barrel-shaped (Fig. 17: phth) only slightly longer than its diameter, with an elongate phallothecal shield (Fig. 17: phths) separated from the phallotheca itself by a distinct, deep constriction; apex of phallothecal shield broadly, roundely produced ventrolaterally, broadly, somewhat angulately emarginate dorsally. Conjunctiva with a short apical lobe (Fig. 18: alc), second conjunctival process (Figs. 17–18: cp-II) divided into a pair of membranous ventrolateral lobes (Figs. 17–18: vll) and a pair of elongate, strongly sclerotized, laterally flattened, posteriorly broadly rounded medial penial plates (Figs. 17–18: mpp) fused around their middle and forming a small dorsomedian lobe of slightly variable shape (Figs. 19–20: arrow). Ductus seminis opening into endophallic reservoir around middle of its ventral surface, endophallic reservoir (Fig. 17: res) large, occupying most of inner lumen of phallotheca; endophallic duct (Figs. 17–18: end) narrow, single-walled, forming a strong loop dorsally, terminating in a narrow, horizontally directed, tubular aedeagus s. str. (Fig. 17: aed).

Male postgenital abdomen. Proctiger (Figs. 15–16: X+XI): segment X large and broad, completely occupying posterior opening of genital capsule, segment XI greatly reduced.

External female genitalia. Terminalia: laterotergites VIII (Figs. 24–25: lt8) broadly rounded, spiracle VIII exposed; valvifers VIII (Fig. 24: vf8) broad, medial margin straight; laterotergites IX (Figs. 24–25: lt9) broad, apically rounded; fused valvifers IX (Fig. 24: vf9) tapezoidal, broadly exposed between laterotergites IX in rest. Gynatrium (Fig. 25: gy) short, with two arms anterolaterally, provided with an oval sclerite surrounding spermathecal orifice (fig. 26: os) and a horseshoe-shaped thickening anteriad of it (Fig. 26: tgw), ring sclerites could not be traced. Spermatheca (Figs. 27–30): proximal duct (Fig. 27: pd) relatively short, thin, of uniform diameter; dilation (Fig. 27: dil) short, transversely striate cuticle restricted at extreme base only; distal invagination (Fig. 27: div) strongly sclerotized, rather short and thick; distal duct (Fig. 27: dd) very short, approximately as long as intermediate part of spermatheca; intermediate part rather variable (Figs. 28–30), proximal flange usually rather thin and nearly conically surrounds distal duct; apical receptacle (Fig. 27: ar) rather variable, but invariably globose, lacking processes (Figs. 28–30).

Measurements (in mm; N = 5 33 6 ƤƤ). Body length 14.0–16.0 (3) / 15.0–20.0 (Ƥ); length of head 2.5–3.1, width across eyes 2.5–2.95, interocular distance 1.5–1.7; lengths of antennal segments (I) 0.3–0.4: (IIa) 1.5–1.7: (IIb) 1.6–1.8: (III) 1.7–2.0: (IV) 1.8–2.0; median length of pronotum 3.1–4.1, humeral width between apices of humeral projections 6.5–10.2; length of scutellum 4.3–5.8, width of scutellum 4.0–4.7.

Egg. Egg was described and illustrated by Stehlík (1987), Ren (1992, as humeralis ), Rieger (2000), and Péricart (2010). SEM photos of particular structures were presented by Ren (1992, as humeralis ). Egg batch was photographed by Tomokuni (1993).

Larval instars. Larvae can be recognized by the diagnostic characters provided above for the genus. 1st–5th instars were described and photographed by Rieger (2000), 3rd–5th instars were photographed by Tomokuni (1993), 5th instar by Tempel (1939). The 5th instar larva was described by Gulde (1921), Dupuis (1947), Putshkov (1961), Chen (1995, as humeralis ) and Péricart (2010) some of these authors also presented illustrations. The 4th instar was photographed by Aukema & Hermes (2009).

Habitat and biology. It is an arboricolous species primarily occurring in mesophilous deciduous forests and forest edges. It was recorded to occur in fir-beech forests from the sub-formation Dentario glandulosae-Fagenion and Galio odorati-Fagenion, moreover oak-hornbeam (Galio-Carpinetum) and linden-oak-hornbeam (Tilio- Carpinetum) forests (Bury 2011). In montane regions of Bulgaria it occurs in both the Fagus - and the Pinus -zone (Heiss & Josifov 1990).

It was recorded from species of the genera Prunus , Sambucus , Populus , Carpinus , Alnus , Pinus , Picea (earlier literature reviewed by Schumacher 1910b, further records by Dioli 1985, Friess 2000, Rieger 2000, Kment et al. 2009), also from Quercus spp. (Dupuis 1949, Ribes & Goula 1995, Wachmann et al. 2008), Corylus avellana L. (Lodos & Önder 1983), species of Fagus , Salix , and Tilia (Wachmann et al. 2008) and occasionally from shrubs ( Rubus ) and herbaceous plants (Rieger 2000, Kment et al. 2009). Although it apparently does not show an obvious preference to any tree species (Putshkov 1961), Stehlík (1987) found it only on Alnus glutinosa (L.) Gaertn. in the Czech Republic; some of the above authors also emphasized its frequent occurrence on alders.

FIGURES 15–20. Pinthaeus sanguinipes , external male genitalia and postgenital abdomen. 15–16, genital capsule: 15, ventral view; 16, posterior view; 17, phallus ( China: Chongqing, Mt. Jinyun), dorsal view, right capitate process removed; 18, same, lateral view; 19, apical portion of phallus ( Hungary: Kiskunfélegyháza); 20, same, lateral view. Scales in mm. Lettering: aed = aedeagus s. str.; alc = apical lobe of conjunctiva; bp = basal plates; cap = capitate process; cp-II = second conjunctival process; cre = central chamber of endophallic reservoir; dc = dorsal connective; dej = ductus ejaculatorius; df = ductifer; ds = ductus seminis; dsim9-10 = dorsal sclerite of intersegmental membrane IX–X; end = endophallic duct; erp = erection fluid pump; hi = hinge; lp = left paramere; mpp = medial penial plates; res = endophallic reservoir; sbc = support bridge complex; sbp = support bridge prolongation; phth = phallotheca; phths = phallothecal shield; vll = ventrolateral lobe of conjunctiva; vsim9-10 = ventral sclerite of intersegmental membrane IX–X; X+XI = proctiger.

FIGURES 21–30. Pinthaeus sanguinipes , external male (21–23) and female genitalia and postgenital abdomen (24–30). 21–23, right paramere, three different aspects; 24, terminalia, posteroventral view; 25, female genital and postgenital abdomen, dorsal view after removal of tergite VIII; 26, structures of dorsal wall of vagina surrounding spermathecal orifice; 27, spermatheca; 28–30, intermediate part and apical receptacle of spermatheca of different species (28: Slovakia: Humenné; 29: China: Heilongjiang, Ning’an; 30: China: Sichuan, Mt. Emei). Scales in mm. Lettering: ar = apical receptacle; dd = distal duct of spermatheca; dil = dilation of spermatheca; div = distal invagination of spermatheca; gy = gynatrium; lt8, lt9 = laterotergites VIII, IX; os = sclerite surrounding orifice of spermatheca; pd = proximal duct of spermatheca; tgw = thickening of gynatrial wall; vf8, vf9 = valvifers VIII, IX; X = anal tube.

First instar larvae do not accept prey, but they readily feed on leaves and drink from drops of water (Rieger 2000). Older larvae and adults feed on various insects, although they also regularly suck plants; Nagy (1957) reported regular feeding from leaves of Acer negundo L. In the field they primarily attack larvae of beetles and lepidopterans; literature records are available about the following taxa as prey: Coleoptera : larva of Agelastica alni (Linnaeus, 1758) , Melasoma populi (Linnaeus, 1758) , Chrysomela sp. [all Chrysomelidae ]; Lepidoptera : Acronicta rumicis (Linnaeus, 1758) , Subacronicta megacephala (Denis & Schiffermüller, 1775) , Panolis flammea (Denis & Schiffermüller, 1775) [all Noctuidae ], Calliteara pudibunda (Linnaeus, 1758) , Lymantria dispar (Linnaeus, 1758) [ Lymantriidae ], Aglia tau (Linnaeus, 1758) [ Saturniidae ], Hyphantria cunea (Drury, 1773) [ Arctiidae ], pupae of unspecified Lepidoptera and Hymenoptera : Tenthredinidae (Gulde 1921, 1934, Devantoy 1948 cited by Dupuis 1949, Nagy 1957, Putshkov 1961, Zebe 1963, Wagner 1966, Stehlík 1987, Friess 2000, Rieger 2000, Wachmann et al. 2008, Bury 2011). On the contrary, Stehlík (1987) claimed that it did not accept Agelastica alni as prey under laboratory conditions, therefore the precise predator-prey relationship with this chrysomelid needs further investigation. Besides of unspecified beetles and lepidopterans, in East Asia it was reported to feed on the gypsy moth, Lymantria dispar (Chan et al. 1989) . Occasionally it feeds on eggs of lepidopterans, e.g. L. dispar (Nagy 1957) . Tempel (1939) and Stehlík (1987) listed various other insects the larvae of which were accepted under laboratory conditions. Preying efficiency is generally lower than that of Arma custos (Fabricius, 1794) , a common species of Asopinae having more or less similar habits, but the preying behaviour is similar: it paralyses the prey by piercing with the stylets, and consumes usually by hanging them down (Nagy 1957). They also suck out dead insects (Stehlík 1987). Cannibalism was observed among larvae only in lack of prey (Rieger 2000). Although all instars feed on plants too (Stehlík 1987), records of this species as a forestry pest (Yang 1962, Ren et al. 2008) are certainly erroneous.

It seems rather rare all over of its distribution area. In several areas of Europe the population apparently shows oscillation in rather long time scale, and in certain regions it can disappear for several decades; the reasons of this phenomenon are uncertain (Kment et al. 2009). Under exceptional circumstances, e.g. on alder strongly infected by Agelastica alni , larvae of L. dispar or other caterpillars occasionally it can be abundant (Tempel 1939, Zebe 1963). In such circumstances it contributes to the control of the populations of various foliage-feeding forestry pests, but its significance is usually much smaller than that of Arma custos (Nagy 1957) .

This species is univoltine in Europe. Adults overwinter in litter or under loose bark (Putshkov 1961, Stehlík 1987, Wachmann et al. 2008). Individuals leave their refugia in May or June (Schumacher 1910b); sporadic captures of freshly emerged larvae in late autumn might indicate that occasionally larvae can overwinter too (Wachmann et al. 2008). Copulation occurs mostly in May; eggs are laid from beginning of June; last instar larvae and freshly emerged adults were observed around end of July and beginning of August in France (Dupuis 1947, 1949), Hungary (Nagy 1957), Ukraine (Putshkov 1961), Czech Republic (Stehlík 1987) and Germany (Rieger 2000, Wachmann et al. 2008); adults were collected in June and July in China (Zhang 1980, Chen 1987, both as humeralis ) and Japan (Ishihara et al. 1953). Young adults do not copulate but move to overwintering refugia (Rieger 2000).

Rieger (2000) studied the reproduction and postembryonic development under laboratory conditions. Copulation takes several hours (a copula was observed for 31 hours). Eggs are laid on the upper surface of leaves. A single female was observed to lay 10 egg batches each composed of 32– 62 eggs (44 in average); an additional, 11th egg batch containing only 8 eggs was observed too. They hatch 12 days later in average; first instar larvae aggregate usually on lower sides of leaves. Developing from first instar to adult took 23 days in average.

Conservation. The species is included in the Red List of threatened Heteroptera of Baden-Württemberg (Rieger 1979) and Sachsen-Anhalt (Bartels et al. 2004) states of Germany as well as Lower Austria (Rabitsch 2005). Because of its scarce and sporadical records and the unsatisfying knowledge on its biology and the factors controlling its abundance, its conservation status is uncertain (Rieger 1979, Bartels et al. 2004, Rabitsch 2005).

Figures

FIGURES 1 – 6. Habitus photos of various specimens of Pinthaeus sanguinipes. 1 – 2, lectotype of P. sanguinipes (Fabricius): 1, dorsal view; 2, its labels; 3, non-type female (Hungary: Simontornya), dorsal view; 4 – 6, holotype of P. humeralis Horváth: 4, dorsal view; 5, lateral view; 6, its labels. Scale in mm; labels not to scale. Figs. 1 – 2: © ZMUC, photographed by N. Akkari; Figs. 4 – 6: © HNHM.

FIGURES 7 – 14. Intraspecific variability of characters of head and pronotum (7 – 8) and humeral angle (9 – 14) of Pinthaeus sanguinipes. 7, North Korea: Kumgan-san (♂); 8, China: Hebei, Qinglong Forest Farm (Ƥ); 9, China: Heilongjiang, Ning’an (Ƥ); 10, Hungary: Simontornya (Ƥ); 11, China: Hebei, Qinglong Forest Farm (Ƥ); 12, Sichuan: Baoxing (Ƥ); 13, Sichuan: Mt. Emei (Ƥ); 14, Yunnan: Maguan (Ƥ). Figs. 7 – 8: most exposed (anterodorsal) view; Figs. 9 – 14: dorsal view. Scale in mm; Figs. 9 – 14 not to scale.

FIGURES 31 – 32. Distribution of Pinthaeus sanguinipes in the West Palaearctic (Fig. 31) and East Asia (Fig. 32). In Fig. 32 numbers – refer to localities of specimens with differently developed humeral angle as illustrated in the left; photos of the same specimens are shown in Figs. 9 and 11 – 14.