Russian Entomol. J. 13(4): 257266 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2004 A synopsis of the genus Cheiloneurus Westwood, 1833 (Hymenoptera: Chalcidoidea: Encyrtidae) of the New World Êðàòêèé îáçîð ðîäà Cheiloneurus Westwood, 1833 (Hymenoptera: Chalcidoidea: Encyrtidae) Íîâîãî Ñâåòà Vladimir A. Trjapitzin1,2 and Robert L. Zuparko2,3 Âëàäèìèð À. Òðÿïèöûí1,2, Ðîáåðò Ë. Çóïàðêî2,3 1 Zoological Institute of Russian Academy of Sciences, Universitetskaya nab. 1, St. Petersburg 199034, Russia Çîîëîãè÷åñêèé èíñòèòóò Ðîññèéñêîé àêàäåìèè íàóê, Óíèâåðñèòåòñêàÿ íàá. 1, Ñàíêò-Ïåòåðáóðã 199034, Ðîññèÿ 2 Centro de Investigación, Unidad Académica Multidisciplinaria Agronomía y Ciencias, Universidad Autónoma de Tamaulipas, 87149 Ciudad Victoria, México. E-mail: [email protected] or [email protected] 3 Essig Museum of Entomology, University of California, Berkeley, CA 94720, U.S.A. and California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118, USA. E-mail: [email protected] or [email protected] KEYWORDS. Cheiloneurus, Encyrtidae, hyperparasitoids. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ. Cheiloneurus, Encyrtidae, ãèïåðïàðàçèòîèäû. ABSTRACT. The New World species of the genus the type specimens of New World Cheiloneurus. The Cheiloneurus are examined. One new name is pro- results are presented here, which we hope will provide a posed: C. giraulti for C. pulcher Girault, and C. ampli- foundation which other researchers can use to produce a cornis Gahan is determined to be a junior synonym of C. revision of the genus. banksi Howard. A redescription of C. marilandia (Gi- MATERIALS AND METHODS. Type specimens rault) is provided. and other material were examined at the Essig Museum of Entomology, University of California, Berkeley ÐÅÇÞÌÅ. Èçó÷åíû âèäû ðîäà Cheiloneurus Íî- (EMEC), Entomological Research Museum, University âîãî Ñâåòà. Äëÿ C. pulcher Girault ïðåäëîæåíî çàìå- of California, Riverside (UCR), California Academy of ùàþùåå íàçâàíèå C. giraulti, à C. amplicornis Gahan Sciences, San Francisco, California (CAS), the United óñòàíîâëåí â êà÷åñòâå ìëàäøåãî ñèíîíèìà C. banksi States National Museum of Natural History, Washing- Howard. Ïðèâîäèòñÿ ïåðåîïèñàíèå C. marilandia ton, D.C. (USNM) and Universidad Autónoma de (Girault). Tamaulipas, Ciudad Victoria, Tamaulipas, México (UAT). Additional specimens from the University of The cosmopolitan genus Cheiloneurus Westwood Central Florida, Orlando, Florida (UCFC) were viewed (Hymenoptera: Encyrtidae) has never been revised in the by the junior author. Terminology follows Gibson et al. limits of the New World, other than a review of the (1997). The new taxonomic status of C. giraulti is solely Argentinian species [De Santis, 1964]. Nevertheless, attributable to V.A. Trjapitzin. both the Nearctic and Neotropical faunas are likely to be very rich in their number of species and morphological Genus Cheiloneurus Westwood, 1833 diversity. For example, nearly each specimen of Cheilo- neurus collected in Mexico by the senior author and half Type species: Encyrtus elegans Dalman 1820. Mo- of the specimens collected in California by the junior notypic. author, appear to represent several different undescribed Synonyms: Chilonevrus Agassiz, 1846; Chiloneu- species, while Noyes [1980: 185] mentions ten or more rus Förster, 1856; Chrysopophagus Ashmead ,1894; undetermined species from the Neotropics. Blatticida Ashmead, 1904; Saronotum Perkins, 1906; To date, all Cheiloneurus species whose biologies Cristatithorax Girault, 1911; Eusemionella Girault, are known have proven to be hyperparasitoids, and in 1915, Chrysopophagoides Girault, 1915; Paracheilo- several cases, of economically important pests. Al- neurus Girault ,1915; Epicheiloneurus Girault, 1915; though the exact role of hyperparasitism in biological Eusemionopsis Girault, 1918; Procheiloneurus Girault, control programs is still debated, there is a general 1920; Raphaelana Girault, 1926; Lepidoneurus Hoffer, consensus that it is deleterious [Doutt & DeBach, 1964; 1957; Metacheiloneurus Hoffer, 1957. Rosen, 1978; Sullivan, 1987]. The impact of Cheilo- The genus Cheiloneurus belongs to the subfamily neurus species on beneficial species has been discussed Encyrtinae Walker, 1837, and has been placed in the by Compere [1925], Bennett & Hughes [1959], De tribe Cheiloneurini Hoffer, 1955, and the subtribe Cheilo- Santis [1964] and Weseloh [1969]. neurina Hoffer, 1955 [Trjapitzin 1973; Trjapitzin & In 2001, the senior author visited several entomo- Gordh 1978; Trjapitzin 1989], although Anis & Hayat logical collections in the United States, largely to study [2002] recently synonymized Cheiloneurini with Ectro- 258 V. A. Trjapitzin and R. L. Zuparko hosts. Most of these species (41) were reared from Ho- moptera, including 38 from Coccidae (including Euleca- nium, Mesolecanium, Parthenolecanium, Physokermes, Pulvinaria, Saissetia). Other Homoptera recorded as secondary hosts include Aclerdidae (Aclerda, Aclerdox), Asterolecaniidae (Asterodiaspis), Eriococcidae (Erio- coccus), Kermesidae (Nanokermes, Kermes), Margar- odidae (Icerya), Pseudococcidae (Antonina, Phenacoc- cus, Planococcus), Psylloidea, Flatidae (Ormenis) and Delphacidae. Cheiloneurus species have also been reared from larvae of Anthophoridae (Hymenoptera), puparia of Diptera, cocoons of Ceraeochrysa, Chrysopa and Chrysoperla (Neuroptera: Chrysopidae), immature Co- leoptera, and oothecae of Blattodea. Such a broad range of secondary hosts is typical for hyperparasitoids. Secondary parasitism by Cheiloneurus has been2 proved for only 12 species. In most cases the primary hosts are immature stages of Encyrtidae (e.g. Diversin- ervus, Isodromus, Metaphycus, Microterys) and Aphe- Fig. 1. Cheiloneurus compressicornis, female [after Clancy 1946, schematized]. linidae, but Cheiloneurus also attacks Perilampidae Ðèñ. 1. Cheiloneurus compressicornis, ñàìêà [ïî Clancy 1946, (Perilampus), Platygastridae (Platygaster) Ichneu- ñõåìàòèçîâàíî]. monidae (Gelis) and Dryinidae (Haplogonatopus, Neo- dryinus, Pseudogonatopus,). matini. At least 120 species have been described in the One paper [McCoy & Selhime, 1970] reported that genus, distributed among the following regions as fol- upon inspection of Sassetia oleae (Olivier) hosts from lows 30 Australian, 36 Indo-Malayan, 23 Palearctic, which C. iminicus adults emerged, the larval remains of 13 Afrotropical, 4 Oceania, 18 Nearctic, and 13 Neotro- only a single parasitoid per scale could be found, and pical (13 species, C. chrysopae Fullaway, therefore they concluded that it was a primary parasi- C. compressicornis (Ashmead), C. cupreicollis Ash- toid. However, as Kfir & Rosen [1981] documented, a mead, C. elegans (Dalman), C. flaccus (Walker), Cheiloneurus larva can completely consume the prima- C. flavoscutatus (Nikolskaya), C. gonatopodis Perkins, ry parasitoid host, and any remaining fragments could C. inimicus Compere, C. kuisebi Prinsloo, C. noxius be broken up by the emerging adult hyperparasitoid. Compere, C. pulvinariae Dozier, C. quadricolor (Gi- Reports on the biology of Cheiloneurus species rault) and C. yasumatsui Trjapitzin, have been reported include studies of C. bonariensis De Santis [De Santis from two or more regions). & Virla, 1991], C. claviger (Thomson) [Saakian-Bara- Female Cheiloneurus can typically be distinguished nova et al., 1971], C. compressicornis (Clancy 1946), from other genera by the following combination of C. inimicus [Compere, 1925; Maple, 1947], C. noxius characters: scutellum with a apical tuft of setae [the tuft Compere [Le Pelley, 1937; Maple, 1947; Weseloh, is absent in three Palearctic species, C. submuticus 1969, 1971a, 1971b, 1972; Weseloh & Bartlett, 1971] Thomson, C. flavoscutatus (Nikolskaya) and C. redic- and C. paralia [Kfir & Rosen, 1981]. All studies of ulus (Trjapitzin & Khlopunov), and reduced to a row of larvae indicate the first instar larvae are endoparasitic, 5 short hairs in the Nearctic C. marilandia (Girault)]; caudate and apneustic. forewing infuscate; long marginal vein and short stig- Keys to species of Cheiloneurus have been published mal and postmarginal veins; hypopygium extending at for the United States [Gahan, 1914], Argentina [De San- most 0.5x length of metasoma; ovipositor never exsert- tis 1964], the Afrotropical region [Compere, 1938], the ed more than 0.33x length of metasoma (Fig. 1). Palaearctic region [Trjapitzin, 1971, 1989], Britain and Sexual dimorphism is marked for the genus, and the Scandinavia [Claridge ,1958], Japan [Ishii, 1928], India males of many species are either unknown or can be [Hayat et al., 1975; Khan & Agarwal, 1978; Anis & very difficult to distinguish, hence identification keys Hayat, 2002] and Hawaii [Beardsley, 1976]. are useful pro tem only for females. Three Nearctic species, C. annulicornis (Ashmead), C. dubius Howard Synopsis of species and C. reate (Walker), are known only from males. Another species, C. kuisebi Prinsloo, originally de- scribed from Namibia, was reported from the New 1. Cheiloneurus albicornis (Howard, 1881) World by Löhr et al. [1990]. We suspect this was a Fig. 2. misidentification, but in lieu of seeing the specimens, Howard, 1881: 363364 (Chiloneurus); Girault, 1916: 300 include it in our synopsis. 301; Peck, 1951: 498; 1963: 436437; Oatman et al., 1964: 981; It is presumed that all members of the genus are Gordh, 1979: 933934. MATERIAL EXAMINED. 1 $ USA, Iowa, type series hyperparasitoids of various insects. Of the 120 described (USNM); 2 $$ New Hampshire, Mt. Washington, 5500, 20 species, 60 (50%) have been reared from their secondary