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Downloaded from Brill.Com09/27/2021 08:29:41AM Via Free Access T  E,  141, 1998 SEI-WOONG CHOI Finnish Museum of Natural History, Helsinki SYSTEMATICS OF THE GENUS HETEROTHERA INOUE (LEPIDOPTERA, GEOMETRIDAE: LARENTIINAE) S.-W. Choi, 1998, Systematics of the genus Heterothera Inoue (Lepidoptera, Geometridae: Lar- entiinae). – Tijdschrift voor Entomologie 141: 19-47, figs. 1-57. [ 0040-7496]. Published 30 November 1998. The genus Heterothera Inoue, occurring widely in the Palaearctic and in the high mountains of the Oriental region, is revised and twenty-three species are recognized. Nine species are de- scribed as new: Heterothera hoenei Choi sp. n., H. yunnanensis Choi sp. n., H. triangulata Choi sp. n., H. eclinosis Choi sp. n., H. stamineata Choi sp. n., H. obscurata Choi sp. n., H. distinc- tata Choi sp. n., H. mussooriensis Choi sp. n. and H. kurenzovi Choi, Viidalepp & Vasjurin sp. n., and five new combinations are suggested: Heterothera tephroptilus (Fletcher) comb. n., H. serrataria (Prout) comb. n., H. etes (Prout) comb. n., H. comitabilis (Prout) comb. n. and H. undulata (Warren) comb. n. A key and illustrations of adults and genitalia are given. The monophyly of Heterothera and the species relationships are discussed. In addition, four poorly known taxa of Thera sensu Prout, T. cyphoschema Prout (= atrinotata Joannis syn. n.), T. exan- gulata Warren, and Pennithera distractata Sterneck comb. n., are redescribed and the taxonomy of these species is briefly discussed. Correspondence: Sei-Woong Choi, Department of Entomology, American Museum of Natural History, Central Park West at 79th St., New York, NY 10024, USA. E-mail: [email protected] Key words. – Systematics; Geometridae; Larentiinae; Heterothera; Palaearctic; Oriental. The genus Heterothera Inoue consists of medium- ular cornuti, well developed sterigma, and a broad sized geometrid moths in the subfamily Larentiinae, ductus bursae. which are widely spread in the Palaearctic and Orien- However, the genus Heterothera s.s. was found to tal regions. The genus was erected by Inoue (1943) be paraphyletic in relation to Viidaleppia and the lat- based on the following characters: the absence of un- ter was synonymized with Heterothera in a cladistic cus, long anal tube, stout and flat saccus, weakly scle- analysis of the Cidariini sensu Herbulot (Choi 1997; rotized costa and a well developed sacculus. The mo- see ‘Diagnosis and monophyly’ for synapomorphies). nophyly of the genus Heterothera s.s. was first defined by Viidalepp (1980). He listed seven apomorphies for Since the works of Prout (1914, 1938, 1941), the the genus: the bifid saccus, the absence of an uncus, taxonomy of Thera s.l. has been considerably changed, the membranous ductus and corpus bursae without due to the discovery of many new species and a dif- signum, the simple sterigma, the distinct sacculus, the ferent analytical approach (e.g. Viidalepp 1980, Choi presence of cornuti on the vesica, and the filiform 1997). As a result of the cladistic analysis, the genus male antenna. Two synapomorphies out of seven, the Heterothera s.l. is characterized by several derived bifid saccus and the absence of uncus, were unique to characters and, based on these, many undescribed Heterothera s.s. However, the character ‘absence of species of the genus have been recognized, mainly uncus’ was found to be incorrect. from southwestern China and northern India. The Inoue (1982) proposed a new genus Viidaleppia for purpose of the present study is to revise the species of the species of Asaphodes sensu Viidalepp (1980). Later, Heterothera. While the taxonomy of several poorly he (Inoue 1986) described the characters of the genus known species of Thera s.l., such as T. exangulata, T. Viidaleppia as: doubly bipectinated male antenna, cyphoschema, T. atrinotata and T. distractata, is uncer- strongly sclerotized costa and pointed apex of fore- tain, another purpose is to redescribe these species in wing, sclerotized and plate-like sacculus, well devel- order to understand their relationships better. oped apical projection of the sacculus, numerous spin- 19 Downloaded from Brill.com09/27/2021 08:29:41AM via free access T E, 141, 1998 1 2 3 4 56 Figs. 1-6. Adults of Heterothera. – 1, H. postalbida; 2, H. tephroptilus; 3, H. mussooriensis; 4, H. undulata; 5, H. hoenei; 6, H. quadrifulta. M , private collection of M. Fibiger, Copenhagen , Swedish Natural History Museum, The study is based on the material from the follow- Stockholm ing museums and private collections: , Institute of Botany and Zoology, Tartu , American Museum of Natural History, New , Zoologisches Forschungsinstitut und York Museum Alexander Koenig, Bonn , British Museum (Natural History), London , Zoological Museum, Helsinki , Bulgarian Natural History Museum, Sofia , Zoologische Staatssammlung München, , Hungarian Natural History Museum, München. Budapest , private collection of Katsumi Yazaki, Tokyo Examination and dissection of the genitalia, in- 20 Downloaded from Brill.com09/27/2021 08:29:41AM via free access CHOI: Systematics of Heterothera cluding everting the vesica, follow Hardwick (1950), slanted, bent, often toothed, medially bulged; central while the terminology of the morphology including fascia variable in width, form constant throughout fas- the genitalia follows Forbes (1948) and Klots (1970). cia or thinner at middle and dorsum, discoidal dot blackish, often indistinct by uniting with costal part of antemedial line (etes, firmata), dorsum distinct with S blackish scales (incerta, obscurata, dentifasciata, distinc- tata, triangulata, quadrifulta, undulata, stamineata); Heterothera Inoue, 1943 subterminal line sometimes present, blackish, scal- Heterothera Inoue, 1943: 12. Type species: Cidaria postalbi- loped (serraria, serrataria, kurenzovi); termen blackish da Wileman, 1911 (original designation). or dark ochreous. Hindwing ground colour whitish Viidaleppia Inoue, 1982: 283. Type species: Cidaria quadri- fulta Prout, 1938 (original designation). (postalbida, yunnanensis, sororcula, incerta, dentifascia- ta, distinctata, triangulata, consimilis, quadrifulta, ser- raria, serrataria, kurenzovi, hoenei, stamineata, ecli- Diagnosis and monophyly nosis), yellowish white (etes, firmata), greyish or Species of Heterothera are characterized by the scle- blackish (obscurata, tephroptilus, taigana, undulata, co- rotized costa and the triangular sacculus of the male mitabilis); discoidal dot usually small, often large; genitalia, the well developed sterigma and the greatly postmedial line blackish, usually medially bulged; ter- modified ductus bursae of the female genitalia. The men sometimes tinged with blackish (incerta, triangu- species of Heterothera have male antenna and a wing lata, consimilis, quadrifulta, serrataria, kurenzovi, un- pattern similar to the species of Pennithera Viidalepp, dulata) or with waving subterminal line (serraria). Praethera Viiddalepp and Thera Stephens. However, Male genitalia. – Uncus weakly sclerotized, tapered Heterothera can be distinguished from other taxa by at bottom or slender, length varies from moderate the triangular and sclerotized sacculus, several large (postalbida) to long (yunnanensis). Tegumen smaller spinular cornuti, the well developed sterigma and the than the total length of vinculum and saccus, dome- greatly modified ductus bursae. shaped or triangular. Saccus concave or medially in- The monophyly of Heterothera s.l. has been de- vaginated (postalbida, dentifasciata, yunnanensis, ob- fined by Choi (1997), and seven synapomorphies scurata, serraria, serrataria, kurenzovi, undulata, support the clade of Heterothera: (1) small process of eclinosis), broad and flat at bottom (incerta, tephrop- sacculus (or harpe), (2) long hairs on the cucullus, (3) tilus, distinctata, consimilis, taigana) or round (hoenei, medially invaginated saccus, (4) scattered cornuti, (5) firmata). Juxta sclerotized, broad, occasionally juxta semi-circular lamella antevaginalis, (6) relatively thick and transtilla form a long tubular structure (postalbi- ductus bursae, and (7) sclerites on the wall of the duc- da, sororcula); anellus lobe conspicuous, varies in tus bursae. shape, from simple and digitiform (postalbida, soror- cula, obscurata, distinctata, consimilis, firmata, taigana, Description serrataria, kurenzovi, undulata, hoenei) to large and tri- Antenna of male filiform (postalbida, yunnanensis, angular (yunnanensis, eclinosis) or small and nipple- sororcula, obscurata, eclinosis), bipectinate with short shaped (incerta, tephroptilus, dentifasciata, quadrifulta, pectinations (dentifasciata, distinctata, quadrifulta, serraria), usually long hairs present at apex. Valva dis- taigana, undulata) or with long pectinations (incerta, toventrally expanded, with long hairs; costa sclero- tephroptilus, consimilis, mussooriensis, firmata, serraria, tized, varies in width, distally wider (postalbida, soror- serrataria, kurenzovi, hoenei). Frons smooth, covered cula, obscurata, tephroptilus, dentifasciata, distinctata, with blackish or dark ochreous and whitish scales. consimilis, taigana, undulata, eclinosis), medially ex- Labial palp variable in length, often about twice as panded (yunnanensis, incerta, quadrifulta, hoenei, long as eye diameter. Interantennal fillet dark brown- mussooriensis) or medially and distally expanded (fir- ish in colour, often distinct by white scales. Legs mata, serraria, serrataria, kurenzovi); sacculus distinct, blackish or dark ochreous and whitish, with distinct usually sclerotized, with a distal process (yunnanensis, whitish tibial joints. Metathorax mediodorsally white,
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