ZOOSYSTEMATICA ROSSICA ISSN 2410-0226 Zoological Institute, Russian Academy of Sciences, St Petersburg ▪ https://www.zin.ru/journals/zsr/ [ onl ine] 0320-9180 Vol. 28(2): 228–237 ▪ Published online 2 September 2019 ▪ DOI 10.31610/zsr/2019.28.2.228 [ print] RESEARCH ARTICLE On the status of the genera complex Acalypta, Dictyonota, Kalama and Derephysia (Heteroptera: Tingidae: Tinginae) having common morphological and karyological features О статусе комплекса родов Acalypta, Dictyonota, Kalama и Derephysia (Heteroptera: Tingidae: Tinginae), имеющих общие морфологические и кариологические особенности V.B. Golub & N.V. Golub В.Б. Голуб, Н.В. Голуб Victor B. Golub, Voronezh State University, 1 Universitetskaya Sq., Voronezh 394018, Russia. E-mail: [email protected] Nataliya V. Golub, Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034, Russia. E-mails: [email protected]; [email protected] Abstract. A complex of morphological features of the genera Dictyonota Curtis, 1827, Kalama Puton, 1876 and Derephysia Spinola, 1837 (Heteroptera: Tingidae: Tinginae), inconsistent with the characters of the tribe Ypsotingini Drake et Ruhoff, 1965, is discussed. The common morphological features of these genera together with those of Acalypta Westwood, 1840, which do not correspond to the characters of Tingis Fabricius, 1803, as the type genus of the tribe Tingini Laporte, 1832, are presented. The data on the sex chromosome system X0/XX of these four genera, radically different from that of all the other 17 so far karyotyped Tinginae genera (XY/XX), are summarised. The closely related genera Acalypta, Dictyonota, Kalama and Derephysia are united in a tribe under the resurrected valid name Acalyptini [Acalyptaini; ICZN Case 3813, in press] Blatchley, 1926 according to the principle of priority (ICZN, Article 23). A diagnosis of the resurrected tribe Acalyptini based on morphological and karyological characters and an improved key to the genera Acalypta, Dictyonota, Kalama and Derephysia are given. Резюме. Обсуждён комплекс морфологических особенностей родов Dictyonota Curtis, 1827, Kalama Puton, 1876 и Derephysia Spinola, 1837 (Heteroptera: Tingidae: Tinginae), не соответству- ющих признакам трибы Ypsotingini Drake et Ruhoff, 1965. Представлены общие морфологиче- ские особенности этих родов и Acalypta Westwood, 1840, не соответствующие признакам Tingis Fabricius, 1803, типового рода трибы Tingini Laporte, 1832. Обобщены данные о половой хромосо- мной системе этих четырех родов (X0/XX), кардинально отличающейся от таковой всех других 17 изученных к настоящему времени родов Tinginae (XY/XX). Таксономически близкие роды Acalypta, Dictyonota, Kalama и Derephysia объединены в трибу с восстановленным валидным на- званием Acalyptini [Acalyptaini; ICZN Case 3813, in press] Blatchley, 1926 в соответствии с принци- пом приоритета (МКЗН, статья 23). Даны диагноз восстановленной трибы Acalyptini на основе морфологических и кариологических признаков и усовершенствованный ключ для определе- ния родов Acalypta, Dictyonota, Kalama и Derephysia. Key words: morphological and karyological features, restoration of tribe, diagnosis, key, lace bugs, Aca- lyptini, Acalypta, Dictyonota, Kalama, Derephysia Ключевые слова: морфологические и кариологические особенности, восстановление трибы, диагноз, ключ, клопы-кружевницы, Acalyptini, Acalypta, Dictyonota, Kalama, Derephysia ZooBank Article LSID: urn:lsid:zoobank.org:pub:5E52C342-D961-477D-B821-8BE35995F573 © 2019 Zoological Institute RAS and the Author(s) V.B. Golub & N.V. Golub. On the status of the genera complex Acalypta, Dictyonota, Kalama and Derephysia Introduction evolutionary and phylogenetic significance of the morphological features of imagines (Guilbert, There is no consensus on the classification of 2001) and larvae (Guilbert, 2004). Guilbert et al. the subfamily Tinginae (Heteroptera: Tingidae) at (2014), based on the results of cladistic analysis the suprageneric level. Various numbers of tribes using a complex of morphological and molecular and other family-group taxa were recognised in data, confirmed the opinion of Guilbert (2004) this subfamily by different authors. The lists of all and defined the tribes Ypsotingini and Litadeini the previously proposed tribes and genera groups as synonyms of Tingini. These authors consid- of different ranks were provided by Drake & Ru- ered only two tribes within Tinginae: Tingini and hoff (1965) and Péricart & Golub (1996). Phatnomini. Lis (1999) had previously excluded The tribe Acalyptini Blatchley, 1926 was cre- Phatnomini from Cantacaderinae, including them ated for two genera, Acalypta Westwood, 1840 in the Tingidae as Phatnominae and raised Can- and Drakella Bergroth, 1922. Drake (1928) re- tacaderini to the family rank. The systematic po- duced Drakella to a synonym of Acalypta. Later, sition of Phatnomini is not discussed here, since Drake & Ruhoff (1965) included Acalypta in the it has been reviewed by Golub (2001), Guilbert tribe Tingini Laporte, 1832. (2004), Schuh et al. (2006), Guilbert et al. (2014), The tribe Ypsotingini Drake & Ruhoff, 1965 and Golub & Popov (2016). originally included two Oriental (Ypsotingis Below we present the results of comparative Drake, 1947 and Dictyotingis Drake, 1942), two analysis of the common morphological and karyo- Australian (Chorotingis Drake, 1961 and Eua- logical features of the genera Acalypta Westwood, ulana Drake, 1945) and three Holarctic genera 1840, Dictyonota Curtis, 1827, Kalama Puton, (Biskria Puton, 1874, Dictyonota Curtis, 1994 1876, and Derephysia Spinola, 1837, and also the and Derephysia Spinola, 1837). Later the ge- characters of the tribes Ypsotingini and Tingini in nus Biskria was synonymised with Dictyonota which they were placed by the previous authors, in (Golub, 1975), and the subgenus Kalama Puton, an attempt to clarify their position in the suprage- 1876 of the genus Dictyonota was erected as the neric classification of Tinginae. valid genus Kalama (Péricart, 1982). Froeschner (2001) retained in the tribe Ypsotingini the indi- Material and methods cated Oriental and Australian genera as well as three widespread Holarctic genera: Dictyonota, Our present contribution is based on the study Kalama and Derephysia. of the extensive material on the species in question Thus, the authors who developed the modern from different regions of the Palearctic, as well as classification of Tinginae tribes, Drake & Ruhoff single specimens of some species from the Nearc- (1965) and Froeschner (2001), recognised three tic [Acalypta parvula (Fallén, 1807), Kalama tri- tribes within Tinginae: Tingini Laporte, 1832, cornis (Schrank, 1801), and Derephysia foliacea Litadeini Drake et Ruhoff, 1965, and Ypsotingini foliacea (Fallén, 1807)] and Oriental [Dictyo- Drake et Ruhoff, 1965, the former tribe compris- tingis monticula Drake, 1956] Regions, kept in the ing the vast majority of recent genera in the world Zoological Institute of the Russian Academy of Tinginae fauna. The Litadeini, which was initially Sciences, St Petersburg, Russia (ZISP) and in the described as a monotypic tribe, currently includes private collection of the first author. Altogether, 13 genera whose species are distributed in the 1.5 thousand specimens of Tingidae were studied. Neotropical, Oriental and Ethiopian Regions, in Photos of some specimens were taken using an Madagascar and Oceania (Froeschner, 2001). MBS-10 stereoscopic microscope, and the digital However, a number of subsequent authors who images were processed using the freeware stack- used the system of the whole subfamily Tinginae ing program CombineZP and Adobe Photoshop did not recognise any tribes within the subfamily CS5. The material used for chromosome analysis Tinginae (Péricart, 1983; Péricart & Golub, 1996, is listed in Table 1. and others). For karyological analysis the male specimens Guilbert included both the Litadeini and Ypso- were fixed in a field in 3:1 Karnoy solution (96% tingini in the Tingini based on his analysis of the ethanol: glacial acetic acid) and stored at 4 °C. In ( Zoosystematica Rossica, Vol. 28, No. 2, pp. 228–237 229 V.B. Golub & N.V. Golub. On the status of the genera complex Acalypta, Dictyonota, Kalama and Derephysia Table 1. Material used for chromosome analysis, with publications in which the chromosome number was report- ed for the first time Number of males/ Locality and dates of collecting Species chromosomal References (all in Russia) plates Acalypta carinata Grozeva, Nokkala, Voronezh Prov., 30.IV.2017 1/35 (Panzer, 1806) 2001 Derephysia (Paraderephysia) Voronezh Prov., 7.VI.2017 22/76 Golub et al., 2018 longispina Golub, 1974 Dictyonota strichnocera Voronezh Prov., 20.VI–1.VII.2017 3/62 Golub et al., 2018 Fieber, 1844 Tingis (Tingis) cardui Env. of Inzer Settlm., South Ural Nature 2/19 Golub et al., 2015 (Linnaeus, 1758) Reserve, Bashkortostan, 4.VIII.2014 Tingis (Tropidocheila) reticulata Herrich- Voronezh Prov., 20.VI–4.VII.2017 20/65 Golub et al., 2018 Schaeffer, 1835 Tingis (Neolasiotropis) Voronezh Prov., 8.VI–25.VI.2017 10/22 Golub et al., 2018 pilosa Hummel, 1825 a laboratory, the testes were dissected in a drop of very long, greatly extended in front of the eyes, 45% acetic acid and squashed on the slide. Covers- usually surpassing apex of first antennal segment, lips were removed using the dry ice technique. The sometimes even that of the second; all tarsi slender preparations were stained by the Feulgen-Giemsa as in Tingini” (Drake & Ruhoff, 1965). Froesch- method according to Grozeva