Lepidoptera, Yponomeutoidea, Praydidae) from China with Descriptions of Two New Species

A taxonomic review of Prays Hübner, 1825 (Lepidoptera, Yponomeutoidea, Praydidae) from China with descriptions of two new species

Jae-Cheon Sohn & Chun-Sheng Wu

Our review of the Chinese species of Prays Hübner, 1825 proposes two new species: Prays kalligraphos sp. n., from Sichuan, and P. tineiformis sp. n., from Hainan, and reports one new record of Prays delta Moriuti, 1977. External morphology and genitalia are described and illustrated. Keys to all seven Chinese Prays are provided for the adults and genitalia of each sex where available. Jae-Cheon Sohn*, Department of Entomology, University of Maryland, 4112 Plant Sciences Building, College Park, MD 20742, USA. [email protected] Chun-Sheng Wu, Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Science, Beijing 100101, China. [email protected]

Introduction forewing chorda, antennal scape without pecten, Praydides are a group of Yponomeutoidea whose one-segmented maxillary palpi, and the abdominal systematic position remains uncertain. In the terga without spines (Dugdale et al. 1998). last century, two opinions of their placement in Yponomeutoidea were prevalent: i.e. as a sub- Prays Hübner, 1825 is the largest, if not only, genus family of Yponomeutidae (Kyrki 1990, Dugdale et of Praydidae with globally 40 described species al. 1998) or as a subfamily of Plutellidae (Heppner (Lewis & Sohn, unpublished). The genus may not be 1998). This is apparently due to their mosaic char- familiar even to lepidopterists, but the leaf-mining acteristics which fail to show complete affinity with or bud-boring larvae occasionally cause substantial either family (Pierce & Metcalfe 1935, Friese 1960, economic losses in agriculture (Carter 1984), as seen Moriuti 1977). Recently a status as separate family in two European pest species, the Olive moth, Prays has been proposed based on a rigorous molecular- oleae (Bernard, 1788), and the Citrus blossom moth, phylogenetic study (Mutanen et al. 2010). This new Prays citri (Millière, 1873). Adults of the genus are classification has been accepted by the latest pub- not readily conspicuous and thus they are often lications (Karsholt & Van Nieukerken 2011, Van neglected in field collecting or during specimen cura- Nieukerken et al. 2011 in press). We follow this tion. In the last few decades, a very limited number opinion which assigns the Prays-group to its own of descriptive works for the group have been pub- family Praydidae. lished (e.g. Landry & Landry 1998; Agassiz 2007). Consequently the species diversity remains greatly The family is easily recognized by its unique modi- under-estimated. Given the ill-defined generic fication of the eighth abdominal segment and by boundary of Prays, a critical revision for the species their genitalia. The broadly enlarged male sternum should improve our understanding of the group VIII and the greatly reduced female apophyses ante- The genus Prays is represented in almost all zoogeo- riores are amongst these characteristics (Kyrki 1984). graphic regions but the species are regionally concen- Other characters that help to distinguish them trated. East Asia is one region where high diversity from other yponomeutoid families include loss of of the group is observed and still largely unknown.

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Genus Prays Hübner, 1825 China is especially promising for finding new species, because its fauna remains little explored. A total This genus is closest to Atemelia but differs from of four species of Prays are known from China (Qian the latter by the lack of a pecten on the antennal 1980, Yu & Li 2004), which is much less than the scape, the ascending labial palpus, the absence of 12 species recorded from Japan (Moriuti 1977, Oku a medial process on the eighth tergite of the male 2003). (misinterpreted as “uncus” by Moriuti 1977) and The aims of this paper are to describe two new spe- the presence of a saccular process in the male geni- cies of Prays from China, to update the Chinese talia. Also Moriuti (1977) mentioned these distinc- fauna of the genus with the new record of P. delta tive characteristics between the two genera. In fact, Moriuti, 1977 and to provide keys for the Chinese defining Prays in comparison with other genera has species. been hampered by the heterogeneous character states in the genus. To deal with this heterogeneity, Moriuti (1977) suggested separating them into two groups, Material and methods based on the width of the valvula in male genitalia Dried specimens were obtained from the Institute and the presence/absence of a digitate process on the of Zoology, Chinese Academy of Sciences (IZCAS), lamellae postvaginales. This distinction turned out Beijing, China. The specimens were compared to to be no more than a classification for convenience, species in the United States National Museum of considering its mosaic occurrence; for example, Prays Natural History, Washington DC, USA (USNM) cingulata Yu & Li, 2004 which falls into “Group A” and the Natural History Museum, London, UK according to the female characters and in “Group B” (BMNH). In those cases where actual specimens sensu Moriuti (1977) according to the male genital are not available, descriptions in the literature were characters. referred to, in order to complete our review and keys. The selected specimens were dissected for examina- A checklist of Chinese Prays tion of genitalia and abdominal structures, following Prays alpha Moriuti, 1977 Clarke (1941) except that chlorozal black was used Prays cingulata Yu & Li, 2004 as stain. Genitalia slides were prepared by mounting Prays delta Moriuti, 1977 with Euparal Medium and Essence (BioQuip), and Prays inconspicua Yu & Li, 2004 examined under a microscope (Leica MZ APO and Prays kalligraphos Sohn & Wu, sp. n. Leica LETTZ-DMRX). Terminology follows Klots Prays lobata Yu & Li, 2004 (1970) for genitalia and Forbes (1948) for wing Prays tineiformis Sohn & Wu, sp. n. venation. Key to the Chinese species of Prays based on external appearance Systematic accounts 1. Wingspan not exceeding 10 mm . . . tineiformis – Wingspan over 10 mm ...... 2 Family Praydidae Moriuti, 1977 2. Forewing unicolorous ...... 3 – Forewing mottled ...... 4 Because the world fauna of Yponomeutoidea has 3. Forewing termen oblique ...... inconspicua never been thoroughly studied, the taxonomic – Forewing termen nearly perpendicular to boundary of Praydidae remains unstable. Moriuti apex ...... delta (1977) originally defined the family, based on only 4. Forewing intermixed with white ...... 5 two genera, Prays and Atemelia Herrich-Schäffer, – Forewing without whitish area ...... 6 1853. Distagmos Herrich-Schäffer, 1853 has been 5. Forewing with white predominant; labial added to the group by Karsholt & Razowski (1996) palpus speckled with white on all three seg- in their catalogue of the European Lepidoptera. ments ...... cingulata Kyrki (1990) first reviewed most, if not all, of the – Forewing with gray predominant; labial pal- world genera and synonymized Orinympha Meyrick, pus tinged with white only around apex . . . . 1927 with Atemelia and also included the Nearctic ...... kalligraphos Eucatagma Busck, 1900 in the family. However, his 6. Forewing with well-defined, oblique, fuscous suggestions have not been universally accepted. A median line ...... alpha critical review of all praydid genera to confirm their – Forewing with diffused fuscous median line . relationship is being conducted by the first author ...... lobata (JCS).

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Key to the Chinese species of Prays based on Distribution male genitalia China (Heilongjiang) and Japan. Prays kalligraphos is excluded because the male is unknown. Prays cingulata Yu & Li

1. Valva elongate, slender ...... 2 Prays cingulata Yu & Li, 2004: 15 (type locality: Xishui, – Valva lobate ...... 5 Guizhou, China) 2. Valva divided at apex ...... 3 – Valva undivided at apex ...... 4 Diagnosis 3. A long, strong, spine like cornutus ...... delta This species is superficially similar to P. iota from – Cornuti as a cluster of spinules ...... lobata Japan by its pale grayish forewing with mottled pat- 4. Socius with several denticles ...... alpha tern, but differs from the latter by the paler forewing – Socius without denticles ...... inconspicua ground color and presence of a broad semicircular 5. Socius with two spines ...... tineiformis fuscous area on the middle of costa. The genital fea- – Socius without spine ...... cingulata tures indicate that it belongs to “Group B” sensu Moriuti (1977). Its male and female genitalia are Key to the Chinese species of Prays based on close to P. omicron Moriuti, 1977 but differ from female genitalia the latter by the slender socii, distally broadened sac- Pray inconspicua and P. lobata are excluded, because cus, smaller membraneous area around ostium, and the females are unknown. larger cervix area of corpus bursae.

1. Lamella postvaginalis with digitate process . . . 2 Material examined. 1?, Wunquan, Yunnan Prov., – Lamella postvaginalis without digitate process . . 3 21 VIII 1990, genitalia slide no. IOZ-09039, IZCAS. 2. Digitate process on lamella postvaginalis longer than ductus bursae ...... kalligraphos Distribution – Digitate process on lamella postvaginalis China (Guizhou and Yunnan). shorter than ductus bursae ...... cingulata 3. Signum present ...... 4 Prays delta Moriuti – Signum absent ...... delta Figs 2, 5 & 5a 4. Ductus bursae sclerotized ...... alpha – Ductus bursae membranous ...... tineiformis Prays delta Moriuti, 1977: 122–123 (type locality: Mt. Ta- tesinayama, Honshu, Japan)

Prays alpha Moriuti Diagnosis Fig. 1 Forewing length 7.5 mm (n=1). This species is dif- Prays alpha Moriuti, 1977: 119–121 (type locality: Asa- ficult to distinguish from other unicolorous con- hikawa, Hokkaido, Japan); Qian, 1980: 509. geners such as P. gamma, P. epsilon, P. omicron and P. kappa, all of which were described by Moriuti Diagnosis (1977). Although careful comparison of the head Forewing length 7mm (n=2). This species is super- and forewing colors can be used for primary distinc- ficially similar to P. iota Moriuti, 1977 but differs tion (see Moriuti 1977, for details), only examina- from the latter by the broadly fuscous costal area of tion of their genitalia guarantees correct identifica- the forewing (gray-blotched in P. iota). The genital tion. The genital features (Fig. 5) of P. delta resemble features of P. alpha indicate its relationship with the those of P. epsilon but differ from the latter by the species assigned to “Group A” sensu Moriuti (1977) slender saccus, larger cornutus, and smaller medial which are characterized by a simple lamella postvagi- emargination on the posterior margin of the lamella nalis without a digitate process. The species can be postvaginalis. distinguished from four other species belonging to group A by the combination of a short socius, a ter- Material examined. 1?, Yichun, Heilongjiang Prov., minally undivided valva, and signum longer than 10.vi.1963, reared from Syringa persica L., genitalia ductus bursae. slide no. IOZ-09038, IZCAS.

Material examined. 1? 1/, Kedong, Heilongjiang Distribution Prov., 19 VI 1976, genitalia slide no. 09041(?) & China (Heilongjiang), Korea, and Japan. Here 09049(/), IZCAS. recorded for the first time from China.

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1 2

3 4

Figs 1–4. Adults: 1. Prays alpha; 2. Prays delta, with bright patch on terminal ¼ of forewing by artefacts from light reflection (concolorous with the rest of the forewing in the actual specimen); 3. Prays kalligraphos, holotype; 4. Prays tineiformis, holotype.

Prays inconspicua Yu & Li (longer in P. kalligraphos) and sclerotized area of the Prays inconspicua Yu & Li, 2004: 16 (type locality: Shan, ductus bursae (shorter in P. kalligraphos) make it pos- Henan, China) sible to distinguish these two species. The male of P. kalligraphos is still unknown. Diagnosis This unicolorous species was distinguished from Description P. epsilon Moriuti, 1977 by Yu & Li (2004), based Head. Vertex smooth, dark yellowish brown, sparsely on the male genitalia: the differences between two intermixed with pale brown on posterior half, brown- species in the dentate pattern of the socii and ventral ish white on anterior half; frons white, tinged with margin of the valvula. The female of P. inconspicua dark brown laterally. Antenna filiform, ½ as long as is unknown. forewing; scape dark grayish brown with luster dorsally, white ventrally; flagellum dark gray with luster. Distribution Labial palpus porrect, 1.5 longer than maximum China (known only from type locality). diameter of eye, dark grayish brown, whitish brown around obtuse apex; first segment ½ as long as sec-  Prays kalligraphos Sohn & Wu, sp. n. ond; third segment 1.5 longer than second. Pro- boscis well developed, naked. Figs 3 & 7 Thorax and abdomen. Patagium dark grayish brown, Type material. Holotype: 1/, Mt. Qingcheng, tinged with yellowish white along distal margin; Sichuan Province, China, 4 VI 1979, genitalia slide tegula and mesonotum dark grayish brown, suffused no. IOZ-09037, IZCAS. with yellowish white posteriorly. Foreleg with coxa dark grayish brown on anterior side, brownish white Diagnosis on posterior side, tinged with white distally; femur This species is similar to another Chinese species to tarsi dark brown; tibia and first tarsomere with P. cingulata by possessing a white/gray mottled a narrow white band distally. Midleg with coxa to forewing. It can be distinguished from the latter by tarsi dark brown dorsally, white ventrally; tibia with the broader gray mottles, but examination of genital white marking ventrally, extended dorsad at the base features is necessary for a reliable identification. The and the middle; tibia and tarsomeres with a whitish digitate process on the female lamella postvaginalis band distally. Hindleg with coxa and femur mostly

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5a 6

Figs 5–6. Male genitalia (a – aedeagus): 5. Prays delta; 6. Prays tineiformis, paratype (aedeagus attached).

7 8

Figs 7–8. Female genitalia: 7. Prays kalligraphos, holotype; 8. Prays tineiformis, paratype, enlarged signum in box.

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Prays tineiformis Sohn & Wu, sp. n. yellowish white, narrowly dark brown dorsally; tibia Figs 4, 6 & 8 to tarsi grayish brown, with a white band distally. Forewing length 6.7mm (n=1), relatively narrow, Type material. Holotype: 1?, Tianchi, Mt. Jianfen- narrowly rounded apically, with rather strongly gling, Hainan Prov., 14.xii.2007 (FQ Chen), geni. oblique termen, white, intermixed with gray costally slide no. IOZ-09048, IZCAS. Paratypes: 1/, same and dorsally, speckled with dark brown to form nar- as holotype, genitalia slide no. IOZ-09040, IZCAS; row, arched basal line; median line broad, oblique; 2/, Dujiacun, Mt. Wuzhishan, Hainan Prov., tornal patch triangular; apical strigulae present; 7.xii.2007 (FQ Chen), IZCAS. fringes dark brown around apex, remainder of fringe dark gray, intermixed with white on termen. Hind- Diagnosis wing knife-shaped, dark yellowish gray; fringe dark This species possesses numerous unique characteris- yellowish gray. Abdomen dark grayish brown dor- tics. The mottled, pale grayish, narrow forewings are sally, yellowish white ventrally. similar to the pest species P. citri, but the oblique Female genitalia (Fig. 7). Papilla analis triangular, median line on the forewing is discontinuous in setose; ninth abdominal segment elongate, trapezoi- P. tineiformis, compared to continuous in P. citri. dal, 3 longer than papilla analis, broadly sclero- Genital features cannot be confused with other con- tized laterally; apophysis posterioris slender, ⅔ as long geners: i.e. slender socii with two spines, each at end as ninth abdominal segment; apophysis anterioris and the middle; uncal plate inverted-subtriangular; absent. Lamella postvaginalis slender, as long as duc- eighth sternite swollen medially; ductus and corpus tus bursae (partly broken in the present specimen). bursae reduced. Posterior margin of eighth sternite emarginated subrectangularly, with a medial projection. Ostium Description bursae small, near to posterior margin of eighth ster- Head. Vertex smooth, yellowish gray with luster; nite. Ductus bursae relatively broad, as long as eighth frons pale gray. Antennae filiform, ½ as long as sternite, densely granulated on anterior ½. Corpus forewing; scape yellowish gray; flagellum gray with bursae oval, 1.7 longer than ductus bursae, slightly luster. Labial palpus porrect, yellowish gray dorsally, protruding on cervix; round signum with an ellipti- dark brown ventrally; second segment 2 longer cal and marginally dentate inward projection. than first, with denser scales distally; third segment 1.3 longer than second, obtuse apically. Proboscis Distribution well developed, naked. China (Sichuan). Thorax and abdomen. Patagium dark gray; tegula dark gray, tinged with yellowish gray distally; mesonotum Etymology dark gray with luster. Foreleg and midleg with coxa The species epithet is derived from the Greek and femur yellowish gray; tibia dark brown, with kalligraphos, meaning “beautifully written” and refers three transverse, yellowish gray bands dorsally, yel- to the dark gray- and white-patterned forewing of lowish gray ventrally; tarsomeres dark brown dor- the new species, resembling calligraphy. sally, yellowish gray ventrally. Hindleg with coxa yellowish gray; femur and tibia yellowish gray, narrowly Prays lobata Yu & Li tinged with dark brown dorsally; tarsomeres dark brownish gray, each with a yellowish gray ring dis- Prays lobata Yu & Li, 2004: 16–17 (type locality: Xianfeng, tally. Forewing length 4.3–5.5 mm (n=4), relatively Hubei, China) narrow, of equal width throughout, pale yellowish brown, mottled, apex narrowly rounded, termen Diagnosis oblique; antemedian, median and submarginal lines Yu and Li (2004) stated a resemblance of P. lobata oblique, formed by dark brown mottles; fringe gray. to P. alpha and P. epsilon in the male genitalia. The Hindwing narrow, knife-shaped, gray, semitranspar- former is distinguished from P. alpha by the lack of ent; fringe gray. Abdomen gray with luster dorsally, white scales on the labial palpus and from P. epsi- brownish white ventrally. lon by the mottled forewing. Its male genitalia also Male genitalia (Fig. 6). Uncus rudimentary as a shal- differ from the latter two species by the presence low protuberance on posterior margin of tegumen; of a medial plate on the gnathos, and more distally socius elongate, bent medially, narrower to apex, spaced socii. with two spines, one at apex and other at middle. Tegumen relatively narrow, triangular at distal half; Distribution gnathos band-like, perpendicular at ⅔, fused medi- China (known only from type locality). ally. Valva elongate, relatively narrow, linguiform

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terminally; costa very slender, curved upward, with Special Reference to the British Isles. – Series Entomo- long setae around apex; sacculus extending over basal logica, Vol. 31. Junk, Dordrecht. 431 pp. ½ of valva, setose, with spiniform process apically; Clarke, J.F.C., 1941. The preparation of slides of the geni- membranous zone present beyond saccular process. talia of Lepidoptera. – Bulletin of the Brooklyn Ento- Vinculum V-shaped; saccus with a stout, clubbed mological Society 36: 149–161. process as long as saccular process. Aedeagus broad, Dugdale J.S., N.P. Kristensen, G.S. Robinson & M.J. Sco- straight, gradually narrowed to apex; cornuti consist- ble, 1998. The Yponomeutidae. – In: N.P. Kristensen ing of 5 stout spines, ⅔ as long as aedeagus. (Ed.), Lepidoptera, Moths and Butterflies. Vol. 1: Female genitalia (Fig. 8). Papilla analis long, triangu- Evolution, Systematics, and Biogeography. Handbook lar, setose, sclerotized dorsally. Ninth abdominal seg- of Zoology 4: 119–130. Walter de Gruyter, Berlin & ment as long as papilla analis, sclerotized, protrud- New York. ing ventromedially, with a broad, microscopic-setose Forbes, W.T.M., 1923. The Lepidoptera of New York lobe laterally; apophysis posterioris relatively thick, and neighbouring States. Primitive forms, Micro- lepidoptera, Pyraloids, Bombyces. – Memoirs, Cor- short, ½ as long as papillae analis; apophysis ante- th nell University Agricultural Experiment Station 68: rioris absent. Posterior margin of 8 sternite deeply 729pp. emarginated rectangularly. Ostium bursae as wide as Friese, G., 1960. Revision der paläarktischen Yponomeuti- a diameter of ductus bursae, near to posterior mar- dae unter besonderer Berücksichtigung der Genitalien. th gin of 8 sternite. Ductus bursae moderately broad, – Beiträge zur Entomologie 10 (1/2): 1–131. entirely membranous, short, ¼ as long as corpus bur- Heppner, J.B., 1998. Classification of Lepidoptera, Part 1. sae. Corpus bursae oval, with numerous winkles; sig- Introduction. – Holarctic Lepidoptera 5 (Suppl. 1): num small, round, with an internally dentate plate. 1–148. Herrich-Schäffer, G.A.W., 1853. Systematische Bear- Distribution beitung der Schmetterlinge von Europa, Zugleich China (Hainan). als Text, Revision und Supplement zu Jacob Hübn- ers Sammlung europäischer Schmetterlinge. Vol. 5. Etymology – G.J. Manz, Regensburg. 394 pp. The species epithet is derived from Tinea (a genus of Hübner, J., 1816–[1825]. Verzeichniss bekannter Schmet- tlinge [sic]. – Augsburg. 431pp. Tineidae), and the latin forma, meaning “shape” and Karsholt, O. & J. Razowski, 1996. The Lepidoptera of refers to resemblance of the new species to tineids. Europa, A Distributional Checklist. – Apollo Books, Stenstrup. 380 pp. & CD-ROM. Acknowledgements Karsholt, O. & E.J. van Nieukerken, 2011. Fauna Euro- paea: Lepidoptera, Moths. – Fauna Europaea version We would like to express our cordial thanks to 2.4. – Fauna Europaea. http://www.faunaeur.org/ [vis- Dr. Donald Davis (U.S. National Museum of Natu- ited on: 29-03-2011]. ral History, Washington DC) and Dr. Jon Lewis Klots, A.B., 1970. Lepidoptera. – In: S.L. Tuxen (Ed), Tax- (United States Department of Agriculture) for read- onomist’s Glossary of Genitalia in Insects: 115–130. ing our manuscript and providing relevant advice Munksgaard. and to Dr. Donald Davis (USNM) and Dr. Day- Kyrki, J., 1990. Tentative reclassification of holarctic oung Xue (Chinese Academy of Sciences, Beijing) Yponomeutoidea (Lepidoptera). – Nota lepidoptero- for allowing us to examine their institutional col- logica 13 (1): 28–42. lections. We are also indebted to Dr. Hui-Lin Han Landry B. & J.-F. Landry, 1998. Yponomeutidae of the Galápagos Islands, with description of a new species and Mr. Mujie Qi, both from the Northeast Forestry of Prays (Lepidoptera: Yponomeutoidea). – Tropical University, Harbin, for their assistance during the Lepidoptera 9 (1): 31–40. senior author’s trips and Mrs. Zaile Du (Univerity of Meyrick, E., 1927. Exotic Microlepidoptera. 3 (12): Maryland, College Park) for helping with the trans- 353–384. lation of the Chinese label data. Moriuti, S. 1977. Fauna Japonica, Yponomeutidae s. lat. (Insecta, Lepidoptera). – Tokyo, 327 p. Mutanen, M., N. Wahlberg & L. Kaila, 2010. Compre- References hensive gene and taxon coverage elucidates radiation Agassiz, D., 2007. Prays peregrine sp. n. (Yponomeutidae) patterns in moths and butterflies. – Proceedings of the a presumed adventive species in Greater London. Royal Society B 277: 2839–2848. – Nota lepidopterologica 30 (2): 407–410. Nieukerken, E.J. van, L. Kaila, I.J. Kitching, N. P. Kris- Busck, A., 1900. New American Tineina. – Journal of the tensen, D.C. Lees, J. Minet, C. Mitter, M. Mutanen, New York Entomological Society 8: 234–248. J.C. Regier, T.J. Simonsen, N. Wahlberg, S.-H. Yen, Carter, D.J., 1984. Pest Lepidoptera of Europe with R. Zahiri, D. Adamski, J. Baixeras, D. Bartsch,

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