Megymenum brevicorne ( Fabricius, 1787 )

Hemala, Vladimír, Kocorek, Anna & Lis, Jerzy A., 2020, Megymenum tuberculatum, a new species of Megymenini from Java and a review of distribution of M. brevicorne (Hemiptera: Heteroptera: Dinidoridae), Zootaxa 4808 (3), pp. 439-454 : 444-448

Megymenum brevicorne ( Fabricius, 1787)

( Figs 4 , 6 , 14 )

Cimex brevicornis Fabricius, 1787: 294–295 (original description, distribution). Lectotype: ♂, China (ZMUC) (designated by Durai 1987: 257).

Cimex brevicornis: Gmelin (1790) : 2152 (diagnosis); Fabricius (1794): 118 (redescription, distribution).

Edessa brevicornis: Fabricius (1803) : 154 (new combination, diagnosis, distribution). Erroneously designated as paralectotypes (2 ♀♀, locality unknown (ZMUC)) by Durai (1987: 257), but these specimens can not be the paralectotypes because they evidently are not a part of the type series (Durai apparently had examined the specimens and she stated these specimens as ‘ Edessa brevicornis ’ while she parallely stated the lectotype male as ‘ Cimex brevicorne ’, therefore only the lectotype designation by Durai (1987: 257) is correct).

Amaurus brevicornis: Burmeister (1834a) : 294 (new combination, distribution); Burmeister (1835): 350 (diagnosis, distribution).

Pseudaradus brevicornis: Burmeister (1834b) : 26 + pl. 16: fig. 1 (new combination, list of species, figure of antenna).

Platydius brevicornis: Westwood (1835) : 446 (new combination, diagnosis).

Edessa brevicornis: Spinola (1837) : 304 (diagnosis); Herrich-Schaeffer (1840): 62 (synonymization with M. cupreum Guérin- Méneville, 1838); Amyot & Serville (1843): 182 (diagnosis, rejecting synonymy with M. cupreum ).

Megymenum Brevicorne : Westwood (1837): 6 (list of species, distribution).

Amaurus inermis Herrich-Schaeffer, 1840: 62 + pl. 163 [not 164]: figs. G, H (description, diagnosis, distribution, figure of antenna and pronotum). Syntype (s): Bengal [= India or Bangladesh] (lost) (synonymized by Kocorek & Lis 2000: 22 with M. brevicorne ).

Amaurus inermis: Amyot & Serville (1843) : 182 (diagnosis).

Megymenum brevicorne: Dallas (1851) : 364 (catalogue, distribution); Dohrn (1859): 23 (list of species, distribution); Mayr (1868): 70 (list of species, catalogue, distribution); Stål (1868): 38 (catalogue, distribution, diagnosis); Walker (1868): 501 (list of species, distribution); Atkinson (1889): 92 (catalogue, redescription, diagnosis, distribution); Lethierry (1891): CXLIII (list of species, distribution); Lethierry & Severin (1893): 239 (catalogue, distribution); Breddin (1900): 332 (list of species, distribution); Distant (1901): 104 (list of species, distribution); Distant (1902): 286 (redescription, diagnosis, distribution); Kuhlgratz (1902): 1129 (list of Breddin’s (1900) record in literary research); Van Duzee (1905): 211 (only briefly mentioned with some further Megymenum species in note about M. insulare Westwood, 1837 (= M. affine Boisduval, 1835 )); Paiva (1906): 352 (information about presence of species in the Indian Museum in Calcutta collection); Kirkaldy (1910): 109 (list of species, distribution); Oshanin (1911): 332 (catalogue, distribution); Longstaff (1912): 374 (distribution); Distant (1921): 166 (list of species); Miller (1929): 421 –436 (distribution, host plants, control, description of egg, immatures and adult, biology, figures); Corbett (1933): 43 (sucking record on plant, distribution); Schouteden (1933): 52 (list of species, distribution); Miller (1934): 517, 525 (description and figure of egg burster); Evans (1952): 108 (list of species); Miller (1956): 48 (note to the oviposition); Southwood (1956): 184 (note to the oviposition); Banerjee (1958): 11, 16, pl. 2: fig. 2 (list of species, food plants, distribution, spermatogonial chromosomes); Stichel (1962a): 725 (list of species); Stichel (1962b): 205 (list of species); Goodchild (1967): 194 (citation of information about a food strategy from Miller 1929); Takenouchi & Muramoto (1969): 9 (list of species—citation of Banerjee 1958); Nuamah (1982): 16 (list of species); Durai (1987): 256 –257 (catalogue, redescription, diagnosis, distribution); Schaefer & Ahmad (1987): 30 (list of species, host plants); Zhang & Lin (1988): 85 (list of species, distribution); Lis (1990): 137, 143 (catalogue, distribution, list of species); Lis (1992): 41 (distribution); Rolston et al. (1996): 68 –70 (catalogue, distribution); Lis & Kocorek (1996): 251 (distribution); Lis & Kocorek (1997): 570, 578 (description and figures of hind wing venation); Lin et al. (2000): 18 (list of species); Kocorek & Lis (2000): 14, 16, 17, 22–23 (cladistics, diagnosis); Kocorek & Danielczok-Demska (2002): 94, 96 (description and figure of spermatheca); Lis et al. (2002): 167, 180 (material examined, figures of pretarsal structures); Lis (2003): 299, 303 (description and photograph of tympanum on female hind tibia); Kerzhner et al. (2004): 18 (list of species); Tsai et al. (2004): 795 (list of species); Lis (2006): 231 –232 (catalogue, distribution); Tsai et al. (2006): 21 (list of species); Chakraborty & Bal (2007): 262 (list of species, distribution); He et al. (2007): 95 (list of species, distribution); Paini et al. (2010): 18, 68 (lists of species); Lis et al. (2011): 19, 22–23, 25–26 (list of species, distribution, similarity of obtained 12S and 16S mtDNA sequences with Macroscytus subaeneus ( Dallas, 1851) ( Heteroptera : Cydnidae ), electropherograms of selected 12S and 16S mtDNA fragments); Kocorek & Ghate (2012): 33 –34, 38 (diagnosis, figures of head with antenna, pronotum, connexivum, paramere and ejaculatory reservoir); Lis et al. (2012): 62 –63, 65–66 (mitochondrial 12S and 16S rDNA sequences); Rakowiecka & Lis (2012): 60 –61 (list of species, distribution, comparison of obtained DNA sequence with DNA sequence of Saccoglossus kowalevskii (Agassiz, 1873) (Hemichordata: Enteropneusta : Harrimaniidae ) available from GenBank); Zheng & Lin (2013): 141 (morphology, host plant, larvae, distribution); Eschen et al. (2014): 106 (list of species); Leavengood (2015): 107, 115 (list of species, using of the species in phylogenetic analysis and ancestral state reconstruction of Hemiptera ); Liang et al. (2014): 377 (list of species, distribution); Lis et al. (2015): 612 (list of species); Li et al. (2017): 7 (listed in chronogram showing phylogeny of Hemiptera with divergence time estimates); Lis et al. (2017): 484 –485, 490–492 (use of 28S rDNA sequence in Bayesian phylogenetic analysis of Pentatomoidea ); Zhao et al. (2018): 2, 8–9 (use of mitochondrial genes in phylogenetic analysis of Pentatomomorpha); Zhou & Rédei (2020): 2 (list of species in examined taxa).

Megymenum inerme: Dallas (1851) : 364 (catalogue, distribution); Dohrn (1859): 23 (list of species, distribution); Walker (1868): 501 (list of species, distribution); Distant (1879): 45 (list of species, distribution); Atkinson (1882): 170 (list of species, distribution); Atkinson (1889): 93 (catalogue, diagnosis, distribution); Lethierry & Severin (1893): 239 (catalogue, distribution); Distant (1902): 286 (redescription, distribution); Paiva (1906): 352 (information about presence in the Indian Museum in Calcutta collection); Breddin (1909): 282 (redescription, diagnosis, distribution); Kirkaldy (1910): 109 (list of species, distribution); Hoffmann (1948): 24 (catalogue); Stichel (1962a): 725 (list of species); Stichel (1962b): 205 (list of species); Yang (1962): 47, 51 (key, redescription); Sienkiewicz (1964): 113 (list of species, records); Chang [= Zhang] (1974): 357 (list of species, distribution); Hsiao et al. (1977): 71 –72 + pl. 9: fig. 130 (figure of head with pronotum, key, habitus photograph); Ahmad & Khan (1979): 5, 8 + fig. 4 (list of species, habitus figure, distribution); Jiang (1985): 58 (list of species, distribution); Zhang & Lin (1986): 60 (list of species, distribution); Chen (1987): 145, 149 (key, diagnosis, morphology, line drawing of pronotum and head, distribution); Durai (1987): 246, 257–258 (key, redescription, diagnosis, distribution); Satapathy & Patnaik (1988): 50, 54, 55–58 (karyotype, photographs of meiosis); Hua (1989): 44 (list of species, distribution); Lis (1990): 143 (list of species, distribution); Zhang & Lin (1990): 2 (list of species, distribution); Jiang (1993): 11 (list of species, catalogue); Chakraborty et al. (1994): 474 (list of species, distribution); Zhang (1994): 32 (list of species, distribution); Zhang et al. (1994): 63 (catalogue, distribution); Zhang (1995): 24 (catalogue, distribution); Rolston et al. (1996): 72 (catalogue, distribution); Easton & Pun (1997): 576 (distribution); Waterhouse (1998): 332 (list of species, distribution); Lin et al. (1999): 56 (list of species, distribution); Lin et al. (2000): 18 (list of species); Kocorek & Lis (2000): 22 –23 (diagnosis, synonymization); Chen & Gu (2000): 53 (list of species, host plants); Wang et al. (2000): 253 (list of species, distribution); Xie et al. (2000): 78 (list of species, distribution); Yang et al. (2005): 42 (list of species, distribution); Biswas & Bal (2007): 303 (list of species, distribution); Li et al. (2008): 212 (list of species, distribution); Li (2011): 162 (list of species, distribution, color photograph of living adult); Xie et al. (2011): 79 (list of species, distribution).

Megymenum (Pseudaradus) brevicorne: Stål (1870) : 86 (new combination, catalogue, distribution); Walker (1873): 29 (list of species); Schouteden (1913): 15 + pl. 1: fig. 13 (catalogue, distribution, figure of head with antennae).

Megymenum (Pseudaradus) inerme: Stål (1870) : 86 (new combination, catalogue, distribution); Walker (1873): 29 (list of species); Kirkaldy (1909): 259 (catalogue, distribution); Schouteden (1913): 15 (catalogue, distribution); Hoffmann (1935): 121 –122 (catalogue, distribution); Yang (1940): 43 –45 (redescription, diagnosis, distribution, figures of connexivum, pronotal margin, antenna and spermatheca).

Megymenum (Pseudaradus) brevicornis (incorrect gender agreement): Kirkaldy (1909): 259 (catalogue, distribution); Hoffmann (1931): 144 (host plants); Tang (1935): 357 (catalogue, distribution); Yang (1940): 42 –43 (redescription, distribution, figures of connexivum, antenna, head, pronotum and spermatheca).

Megymenum brevicornis (incorrect gender agreement): Hoffmann (1932a): 1019 –1020 (distribution, host plants, biology); Hoffmann (1932c): 140 (list of species, distribution); Wu (1933): 220 (list of species, catalogue, distribution); Cheo (1935): 30 (list of species, host plants, distribution); Hoffmann (1935): 120 –121 (catalogue, distribution); Hoffmann (1948): 24 (cata- logue); Yang (1962): 47, 51–52 (key, redescription, diagnosis); Sen (1965): 488 (list of species, distribution, host plant); Hsiao et al. (1977): 71 + pl. 8: fig. 129 (key, figure of head with pronotum, habitus photograph); Jiang (1985): 58 (list of species, distribution); Zhang (1985): 57 –58 + pl. 12: figs. 28.1–28.5 (redescription, description of egg and immatures, figure of head with pronotum, eggs, immatures and adult); Durai (1986): 4 –6 (list of species, habitus photograph, photograph of adult feeding on host plant); Zhang (1986): 426 (list of species, distribution); Zhang & Lin (1986): 60 (list of species, distribution); Chen (1987): 145 –148 (key, morphology, line drawing of pronotum and head, figure with development stages, distribution, biology); Durai (1987): 245, 256–257 (key, redescription, diagnosis, distribution); Zhang & Lin (1987): 76 (list of species, distribution); Hua (1989): 44 (list of species, distribution); Chen (1990): 116 (list of species, distribution); Hu (1990): 90 (list of species, distribution); Zheng & Jin (1990): 142 (list of species, distribution); Wany et al. (1992): 19 (list of species, distribution); Lin et al. (1992): 170 (list of species, distribution); Zhang & Lin (1992): 19 (list of species, distribution); Durai (1993): 219 –228 (life history, host plants, figure of eggs after eclosion, photographs of eggs and immatures); Jiang (1993): 10 (list of species, catalogue); Zhang (1994): 32 (list of species, distribution); Zhang et al. (1994): 63 (catalogue, distribution); Vir & Verma (1996): 354 (list of species); Lei & Zhou (1998): 36 (list of species, distribution); Lin et al. (1999): 55 –56 (list of species, distribution); Hua (2000): 170 (list of species, distribution, host plants); Wang et al. (2000): 253 (list of species, distribution); Liu (2006): 154 (list of species, distribution); Li et al. (2008): 212 (list of species, distribution); Chen et al. (2009): 1353 (list of species, distribution, ecology, competition with lac insects); He et al. (2011): 58 (list of species, distribution); Wang & Tong (2014): 144 (list of species, distribution).

Megarhamphus (Pseudaradus) brevicornis (incorrect genus agreement): Hoffmann (1932b): 9 (list of species).

Megarhamphus (Pseudaradus) inermis (incorrect genus agreement): Hoffmann (1932b): 9 (list of species).

Megymenus brevicorne (incorrect subsequent spelling): Wu (1932): 82 (list of species, distribution).

Megymenum (Pseudaradus) inerma (incorrect subsequent spelling): Tang (1935): 357 (catalogue, distribution).

Megimenum brevicorne (incorrect subsequent spelling): Bose & Sinha (1963): 269 –270 (hydrogen ion concentration in odoriferous glands of larva and adult).

Material examined. CHINA: unspecified: no exact data, 1 ♀, Nonfried lgt., A. Kocorek det. ( DBOU) . Guangdong: Canton [= Guangzhou], 12.x.1911, 1 ♀, S. V. Mell lgt., J. A. Lis 1988 det. ( DBOU) . Hainan: Qiongshan , vi.1961, 1 ♂ [lost pygophore], from winter melon [= Benincasa hispida ], X. S. Tan lgt., A. Kocorek det. ( DBOU; translated from Chinese by D. Rédei) ; Bawangling Mts., Baotie env., 19°05.3′N 109°07.4′E, 415–800 m a.s.l., 7.– 8.v.2011, 1 ♂, at light, M. Fikáček, V. Kubeček & L. Li lgt., V. Hemala det. ( NMPC) . Yunnan: no exact locality, no date, 1 ♂, no collector, R. Kumar 1973 det. ( DBOU) .— NEPAL: Central Nepal, Chitwan ( Roy. Nat. Park ), Sauraha village , 27.35N 84.30E, 166 m a.s.l., 1 ♀, D. Král lgt., V. Hemala det. ( NMPC) .— THAILAND: Chiang-Mai prov., Chiang-Mai surroundings, 1.vii.1995, 4 ♂♂ 1 ♀ 12 L, W. G. Ullrich lgt., A. Kocorek det. ( WULG) ; North Thailand, Muang Nan, viii.1995, 1 ♂, W. G. Ullrich lgt., A. Kocorek det. ( WULG) ; Chiang Mai-Chiang Dao , 5.–10.vii.1997, 1 ♀, M. Klícha lgt., V. Hemala det. ( NMPC) ; ca. 5 km S of Mae Hong Son, Pai River , high water, flooding detri- tus/vegetation, 19°14′12.2″N 97°53′41.1″E, 23.vii.1997, 1 ♂ 4 ♀♀, W. G. Ullrich lgt., A Kocorek det. ( WULG) ; Chiang Dao , 27.v.–2.vi.2002, 1 ♀, B. Makovský lgt., V. Hemala det. ( NMPC) ; Trat prov., Ko Chang island , 12°05′N 102°21′E, 0–200 m a.s.l., 6.–13.vii.2002, 1 ♂ 1 ♀, R. + H. Fouquè lgt., V. Hemala & A. Kocorek det. ( NMPC) ; Phuket, White Buddha , 7°51′39″N 98°19′30″E, 50 m a.s.l., 21.vii.2010, 1 ♀, L. Juříčková lgt., V. Hemala det. ( NMPC) .— LAOS: no exact data, 1 spec. [lost abdomen], no collector, J. A. Lis 1988 det. ( DBOU) .— VIETNAM: Annam , no date, 2 ♂♂, no collector, R. Kumar 1973 det. ( DBOU) ; Mai lam, NE of Hanoi, 12.–16.iv.1966, 1 ♂, Gy. Topál lgt., A. Kocorek det. ( HNHM) ; Tonkin , no date, 1 ♂, K. U. Tyduna lgt., J. A. Lis 1988 det. ( DBOU) .— MA- LAYSIA: Perak: Talping , x.1977, 1 ♂, P. Pfanner lgt., V. Hemala & A. Kocorek det. ( MHNG) . Federal Territory of Kuala Lumpur: Kuala Lumpur , 16.viii.1983, 1 ♀, S. Zabanski lgt., B. Lis det. ( DBOU) .— INDONESIA: Sumatra: Medan , no date, 1 ♂ 1 ♀, Mjöberg lgt., V. Hemala det. ( NHRS) ; Tjinta Radja , no date, 1 ♀, Mjöberg lgt., V. Hemala det. ( NHRS) .

Redescription of selected structures. Head slightly wider than long; clypeus very small, shorter than mandibular plates, together heart-shaped and strongly concave dorsally ( Fig. 4 ). Anteocular spines present only sporadically (in most specimens absent), instead small rounded anteocular tubercles present ( Fig. 4 ). Base of head wide (widest across eyes); small compound eyes positioned posterolaterally, nearly touching anterior part of pronotum. Antenniferous tubercle very small, short and not visible in dorsal view. Anteocular tubercles visible from dorsal view. Scape (I) very short, shorter than mandibular plate. Pedicel (II) is longest antennal segment, only slightly flattened, more circular in cross-section, about 2.2 times longer than segment I and longer than mandibular plate. Basiflagellum (III) more circular in cross-section, very slightly curved apically and 1.65–1.69 times longer than segment I. Distiflagellum (IV) apically narrowed, relatively short, only 1.46–1.59 times longer than scape. Bucculae round, very short, but longer than labrum. Labial segment I shorter than segment II, clearly reaching posterior margin of head, segment III and IV very short (shortest segments of labium; 2.0–2.26 times shorter than segment I) with nearly of same length or segment IV slightly shorter. Labial segment IV apically narrowed. Pronotum ( Figs 4 , 6 ) trapezoid in dorsal view, with well differentiated collar-like structure due to the distinct concavity of its anterolateral margin. Anterolateral margin apparently or strongly projecting forward to form distinct process, strongly concave anteriad of this process and only slightly concave posteriad of this process. Posterolateral margin slightly rounded, as long as 2/3 of anterolateral margin. Posterior margin along base of scutellum slightly convex. Pronotal surface tuberculate, with distinct medium sized rounded tubercle in middle of its anterior part, narrowly positioned near the anterior margin; this tubercle only 1.12–1.18 times wider than long. Disc with strong depressions at anteriolateral angles. Spermatheca ( Fig. 14 ) with medium sized and spherical apical receptacle (spermathecal bulb), short and distinct intermediate part (pumping region) with distinct and large distal and proximal flanges, large and strongly dilated (sac-like) spermathecal duct with a distinct lateral fold bearing distinct ring sclerite. Spines of spermathecal duct distinct and large, covering only a small longitudinal area posteriorly of the ring sclerite.

Measurements [median (minimum–maximum); in mm]. Females (n = 17; for pedicel: n = 16; for basiflagellum: n = 9; for distiflagellum IV: n = 7; for labial segments: n = 11; for profemora and protibiae: n = 14; for mesofemora and mesotibiae: n = 13; for metafemora and metatibae: n = 12; for tarsi: n = 11). Body length (from apex of clypeus to apex of abdomen) 13.72 (12.79–14.88); head: length (from apex of clypeus to anterior margin of pronotum) 1.86 (1.71–2.17), width (maximum width across eyes) 2.56 (2.40–2.87), height (height across compound eye) 1.55 (1.55–1.86), interocular width (between inner margins of compound eyes) 1.86 (1.71–2.02); lengths of antennal segments: scape (I)—0.85 (0.70–0.93), pedicel (II)—1.86 (1.71–2.09), basiflagellum (III)—1.40 (1.40–1.75), distiflagellum (IV)—1.24 (1.24–1.32); length of labial segments: I—1.40 (1.24–1.55), II—1.55 (1.32–2.16), III—0.62 (0.47–0.78), IV—0.62 (0.47–0.70); pronotum: length (medially in most exposed view) 3.57 (3.41–3.95), anterior width (between anterolateral angles) 4.88 (4.57–5.43), length of anterior tubercle (medially) 1.05 (0.93–1.16), width of anterior tubercle (transversally) 1.24 (1.09–1.50); posterior width (maximum width between humeral angles) 6.36 (6.05–7.29); scutellum: length (medially from base to apex) 4.00 (3.72–4.31), width (maximum width at base) 3.95 (3.64–4.31); corium: length 4.50 (4.19–4.96), width 2.09 (1.86–2.33); abdomen width (maximum width between apexes of lobes on laterotergites IV) 7.36 (6.67–7.91); length of femora: profemur 2.95 (2.71–3.10), mesofemur 3.26 (3.10–3.64), metafemur 4.19 (3.57–4.26); length of tibiae: protibia 2.56 (2.33–2.95), mesotibia 2.95 (2.64–3.57), metatibia 4.19 (3.80–4.65); length of tarsi: protarsus 1.55 (1.55–1.55), mesotarsus 1.71 (1.55–1.71), metatarsus 1.71 (1.55–1.71).

Males (n = 12; for pedicel: n = 11; for basiflagellum: n = 7; for distiflagellum: n = 6; for labial segments, femora, protibia and metatibia: n = 8; for mesotibia and protarsus: n = 7; for mesotarsus: n = 6; for metatarsus: n = 3). Body length (from apex of clypeus to apex of abdomen) 12.12 (11.44–13.02); head: length (from apex of clypeus to anterior margin of pronotum) 1.71 (1.55–1.94), width (maximum width across eyes) 2.48 (2.17–2.64), height (height across compound eye) 1.55 (1.47–1.71), interocular width (between inner margins of compound eyes) 1.71 (1.71–1.86); lengths of antennal segments: scape (I)—0.78 (0.62–0.81), pedicel (II)—1.71 (1.47–1.78), basiflagellum (III)—1.32 (1.24–1.52), distiflagellum (IV)—1.24 (1.16–1.40); length of labial segments: I—1.32 (1.24–1.32), II—1.32 (1.24–1.39), III—0.65 (0.45–0.78), IV—0.55 (0.45–0.62); pronotum: length (medially in most exposed view) 3.30 (3.10–3.64), anterior width (between anterolateral angles) 4.54 (4.19–5.19), length of anterior tubercle (medially) 0.97 (0.85–1.01), width of anterior tubercle (transversally) 1.09 (1.01–1.47); posterior width (maximum width between humeral angles) 6.05 (5.74–6.82); scutellum: length (medially from base to apex) 3.57 (3.49–4.03), width (maximum width at base) 3.57 (3.26–4.03); corium: length 4.23 (3.95–4.65), width 1.86 (1.75–2.09); abdomen width (maximum width between apexes of lobes on laterotergites IV) 6.57 (6.12–7.70); length of femora: profemur 2.64 (2.48–2.95), mesofemur 3.17 (2.87–3.38), metafemur 3.76 (3.64–3.95); length of tibiae: protibia 2.48 (2.33–2.64), mesotibia 2.95 (2.64–3.10), metatibia 3.95 (3.72–4.34); length of tarsi: protarsus 1.55 (1.40–1.55), mesotarsus 1.55 (1.40–1.71), metatarsus 1.55 (1.47–1.71).

Biology. Megymenum brevicorne is trophically bound especially on plants from the families Cucurbitaceae and Passifloraceae sucking on their stems and fruits ( Miller 1929, Hoffmann 1932a, Schaefer & Ahmad 1987). The species was reported most often as a minor pest of Passiflora quadrangularis , Cucurbita maxima and Trichosan- thes cucumerina, on which the larval development was observed ( Miller 1929, Hoffmann 1932a). It was recorded also on other Cucurbitaceae (e.g. Cucumis sativus , Lagenaria siceraria ), on some Fabaceae ( Vigna unguiculata sesquipedalis , Phaseolus lunatus ) and even on some other plant families— Celosia cristata (Amaranthaceae) , Im- patiens balsamina ( Balsaminaceae ), Manihot esculenta (Euphorbiaceae) , Quercus spp. ( Fagaceae ), Cinnamomum camphora (Lauraceae) , Eucalyptus spp. ( Myrtaceae ), on which the development is supposed, but not confirmed yet ( Hoffmann 1931, 1932a; Corbett 1933; Cheo 1935; Chen & Gu 2000; Hua 2000). Sen (1965) stated the species as a

“major pest” of Millettia pinnata (Fabaceae) without giving any further details and therefore this record also needs a confirmation. Feeding of M. brevicorne is probable also on Benincasa hispida (Cucurbitaceae) on which the species was recorded in Qiongshan (Hainan, China) in 1961 (sitting record) (this paper). Megymenum brevicorne was listed also in the list of pests of various leguminous trees ( Fabaceae ) in India without indicating the taxon on which it was observed ( Vir & Verma 1996). Larvae of M. brevicorne were successfully reared on Mormodica charantia ( Cucurbitaceae ), Basella alba (Basellaceae) and Citrus aurantiifolia (Rutaceae) ( Durai 1993). Larval development was not fully described in nature, but only in laboratory conditions ( Miller 1929, Durai 1993). Number of generations per year is not known, but length of life from the egg hatching to adult death was 74–103 days in laboratory conditions ( Durai 1993).

Distribution. CHINA: unspecified ( Fabricius 1787, 1794, 1803; Gmelin 1790, as Sina; Burmeister 1835; Westwood 1837; Dallas 1851; Dohrn 1859; Walker 1868; Stål 1870; Distant 1902; Kirkaldy 1909; Miller 1929; Hoffmann 1935; Durai 1987; Hua 1989; Lis 1990, 1992; Zhang 1995; Waterhouse 1998; Chen & Gu 2000; Lin et al. 2000; Li 2011; this paper), Beijing ( Hua 2000), Chongqing ( Lin et al. 1992), Fujian ( Wu 1932; Hoffmann 1935 [both previous as Fukien]; Tang 1935; Yang 1940 as Fokien; Lin et al. 1999; Hua 2000), Guangdong ( Burmeister 1834a; Kirkaldy 1909; Oshanin 1911; Hoffmann 1931, 1935, 1948; Wu 1933; Cheo 1935; Tang 1935; Yang 1940; Lis 1990; Lis & Kocorek 1996; Hua 2000; this paper), Guangxi ( Zhang et al. 1994; Hua 2000; Xie et al. 2000, 2011; Yang et al. 2005; Liang et al. 2014), Guizhou ( Yang 1940 as Kweichow; Chen 1987; Lin et al. 1992; Hua 2000; Liu 2006), Hainan ( Hoffmann 1932c, 1935; Tang 1935; Yang 1940; Hua 2000; this paper), Hebei ( Hoffmann 1948 as Hopei; Wang et al. 2000), Hong Kong ( Mayr 1868; Hoffmann 1935; Durai 1987), Hubei ( Lei & Zhou 1998; Hua 2000), Hunan ( Lin et al. 1992), Jiangxi (Chang [= Zhang] 1974, as Kiangsi; Zhang & Lin 1986, 1992; Zhang 1994; Hua 2000; Li et al. 2008), Macao ( Kirkaldy 1910; Hoffmann 1935; Easton & Pun 1997), Shaanxi ( He et al. 2007), Sichuan ( Hua 2000), Tibet [= Xizang] (Zhang 1986; Zhang & Lin 1987, 1988; Hu 1990; Zheng & Jin 1990; Wany et al. 1992; Hua 2000), Yunnan ( Hoffmann 1948; Jiang 1985; Zhang & Lin 1990; Hua 2000; Chen et al. 2009; He et al. 2011; this paper; record in Zhang 2011 is misidentification of M. spinosum ), Zhejiang ( Yang 1940 as Chekiang; Chen 1990; Hua 2000; Wang & Tong 2014). TAIWAN ( Lis 1992 as Formosa; Zheng & Lin 2013). INDIA: unspecified ( Lethierry & Severin 1893; Kirkaldy 1909; Hoffmann 1935; Tang 1935; Durai 1987; Lis 1990), “east part” ( Dohrn 1859 as M. inerme ), “North India ” ( Atkinson 1882), Andhra Pradesh ( Biswas & Bal 2007), Assam ( Atkinson 1882, 1889; Distant 1902; Kirkaldy 1909; Schouteden 1913; Hoffmann 1935; Yang 1940; Durai 1987), Jharkhand ( Sen 1965), Meghalaya ( Distant 1902; Miller 1929), Mizoram ( Chakraborty & Bal 2007), Nicobar Islands ( Distant 1902; Kirkaldy 1909; Miller 1929; Hoffmann 1935; Tang 1935; Yang 1940; Durai 1987; Lis 1990), Rajasthan ( Lethierry 1891 from Tetara), Sikkim ( Yang 1940), West Bengal ( Distant 1902; Miller 1929; Banerjee 1958; Chakraborty et al. 1994;? Herrich-Schaeffer 1840;? Hoffmann 1935,? Tang 1935 [previous three ambiguously as Bengal]).? BANGLADESH (? Herrich-Schaeffer 1840;? Hoffmann 1935;? Tang 1935 [all ambiguously as Bengal]; Ahmad & Khan 1979 without any exact locality). NEPAL (new record; the record in Jyoti & Rajbhandari 2015 is misidentification of an unidentified species of the genus Coridius Illiger, 1807 ). BHOUTAN ( Yang 1940). SRI LANKA ( Longstaff 1912; Hoffmann 1935 as Ceylon; Durai 1987; Lis 1990). MYANMAR: unspecified ( Kirkaldy 1909; Hoffmann 1935; Tang 1935; Durai 1987; Lis 1992 [all previous as Burma]; Yang 1940 as Birmaine), Yangon Region ( Distant 1901, 1902; Miller 1929; Tang 1935 [all as Rangoon]). THAILAND ( Hoffmann 1935 as Siam; Durai 1987; Lis 1990; Lis & Kocorek 1996; Lis et al. 2011; Rakowiecka & Lis 2012; this paper). LAOS ( Lis 1990; this paper). CAMBODIA ( Yang 1940, as Cambodge). VIETNAM ( Hoffmann 1948; Sienkiewicz 1964 [both as M. inerme ]; Lis 1992; this paper). MALAYSIA: unspecified (? Corbett 1933 as Malaya [= ‘British Malaya’, the term describing a set of states on the Malay Peninsula and the island of Singapore that were under the British control in 1874–1957 (e.g. Andaya & Andaya 1982)]; Durai 1986, 1987; listed in Lis 1990), Perak (this paper), Federal Territory of Kuala Lumpur (this paper).? SINGAPORE (? Corbett 1933 as Malaya [= ‘British Malaya’, the term describing a set of states on the Malay Peninsula and the island of Singapore that were under the British control in 1874–1957 (e.g. Andaya & Andaya 1982)]). INDONESIA:? Java ( Yang 1940: country is only listed; cited only in Hoffmann 1948), Sumatra ( Breddin 1900; Kirkaldy 1909; Schouteden 1933; Hoffmann 1935; Yang 1940; Durai 1987; Lis 1990, 1992; this paper).

Erroneous records. Japan: Hua’s (2000) reference to the distribution of M. brevicorne in Japan seems to be erroneous, because there is reliable documentation of only one species of the genus Megymenum Guérin-Méneville, 1831 — M. gracilicorne Dallas, 1851 ( Hayashi et al. 2016).

Figures

FIGURES 1–4. Habitus of Megymenum tuberculatum Hemala & Kocorek, sp. nov. and M. brevicorne (Fabricius, 1787). 1–3—female of M. tuberculatum (1—dorsal view, 2—ventral view, 3—lateral view); 4—female of M. brevicorne (dorsal view). Black arrows show the position of anteocular spine/tubercle. (Photo: V. Hemala).

FIGURES 5–12. Visual comparison of pronotum shape and size of dorsal anterior tubercle in eight species of the genus Megy- menum Guérin-Méneville, 1831. 5—Megymenum tuberculatum Hemala & Kocorek, sp. nov.; 6—M. brevicorne (Fabricius, 1787); 7—M. basale Walker, 1868; 8—M. parallelum Vollenhoven, 1868; 9—M. spinosum (Burmeister, 1834); 10—M. mekon- gum Distant, 1921; 11—M. pratti Distant, 1911; 12—M. gracilicorne Dallas, 1851. (Photo: V. Hemala).

FIGURES 13–14. Spermatheca. 13—Megymenum tuberculatum Hemala & Kocorek, sp. nov.; 14—M. brevicorne (Fabricius, 1787). Abbreviations: ar—apical receptacle (spermathecal bulb), dfl—distal flange, ip—intermediate part (pumping region), pfl—proximal flange, s—spines, sd—spermathecal duct, rs—ring sclerite. Scale bars: 0.5 mm. (Orig. A. Kocorek).