Abstract
The non-European taxa of the genus Nychiodes Lederer, 1853 are revised. Type specimens of all described species and a large series of about 800 additional specimens were morphologically examined. More than 400 genitalia preparations were made and analyzed along with distributional and DNA barcode data. As a result of our integrative taxonomic approach, Nychiodes waltheri saerdabica Wehrli, 1938 syn. nov., is synonymized with N. waltheri Wagner, 1919; N. palaestinensis libanotica Zerny, 1933 syn. nov. is synonymized with N. palaestinensis Wagner, 1919 and the synonymy of N. persuavis Wehrli, 1929 syn. rev. with N. palaestinensis is confirmed; N. admirabila safidaria Wiltshire, 1943 syn. nov. is synonymized with N. admirabila Brandt, 1938; N. agatcha Brandt, 1938 syn. nov., N. subvirida disjuncta Wehrli, 1941 syn. nov. and N. subvirida taftana Brandt, 1941 syn. nov. are synonymized with N. subvirida Brandt, 1938. Also, N. variabila variabila Brandt, 1938 syn. nov., N. variabila opulenta Brandt, 1941 syn. nov., N. divergaria elbursica Wehrli, 1937 syn. nov., N. divergaria fallax Wehrli, 1939 syn. nov. and N. divergaria achtyca Wehrli, 1939 syn. nov. are synonymized with N. divergaria Staudinger, 1892. Nychiodes convergata sp. nov. from Israel, N. mirzayansi sp. nov. from the Iran and N. eberti sp. nov. from Turkey are described. Lecto- and paralectotypes are designated for N. palaestinensis, N. antiquaria, N. divergaria. Furthermore, N. antiquaria is reported as a new species for Pakistan, N. rayatica is reported as a new species for Iran and the hypothetical occurrence of N. amygdalaria in Iran is confirmed. Additionally, N. tyttha needs to be excluded from the genus. Wing pattern, male and female genitalia and diagnostic characters of all examined species are illustrated and distribution maps are provided. Illustrated keys based on genitalia, as well as a complete checklist of the genus is given here.
References
Brandt, W. (1938) Beitrag zur Lepidopteren-Fauna von Iran. Neue Gattungen, Arten und Formen (Macrolepidoptera). Entomologische Rundschau, 55, pp. 1–72.
Brandt, W. (1941) Beitrag zur Lepidopteren-Fauna von Iran (4). Einige neue Geometriden. Mitteilungen der Münchner Entomologischen Gesellschaft, 31, 864–886.
Evenhuis, N.L. (2007) The Insect and Spider Collections of the World website of the Bishop Museum, Honolulu, Hawaii. Available from: http://hbs.bishopmuseum.org/codens/codens-r-us.html (accessed 14 May 2018)
Expósito Hermosa, A. (1984) Es Nychiodes andalusiaria Milliere, 1865 bona species?. Shilap revista de lepidopterología, 12 (447), 202–204.
Expósito Hermosa, A. (2005) Una nueva especie del gènero Nychiodes Lederer, 1853 en España (Lepidoptera: Geometridae). Shilap Revista de Lepidopterologia, 33 (132), 461–466.
Fazekas, I. (1996) Revision der Nychiodes waltheri F. Wagner, 1919 Art. 1. Beitrag Die Balkanhalbinsel (Lepidoptera: Geometridae). Entomologica Romanica, 1, 3–16.
Fazekas, I. (1997) Revision der Nychiodes obscuraria Villers, 1789 Artenkomplexes, mit ergänzenden Angaben zu Nychiodes dalmatina F. Wagner, 1909 (Lepidoptera: Geometridae). Entomologica Romanica, 2, 13–44.
Leraut, P. (2009) Moths of Europe. Vol. 2. N.A.P. Editions, Verrières le Buisson, pp. 1-808.
Hausmann, A. (1991) Beitrag zur Geometridenfauna Palästinas: Die Spanner der KLAPPERICH-Ausbeute aus Jordanien (Lepidoptera, Geometridae). Mitteilungen der Münchner Entomologischen Gesellschaft, 81, 111–163.
Hausmann, A. (1994) Beitrag zur Geometridenfauna Zyperns. Zeitschrift der Arbeitsgemeinschaft der österreichischen Entomologen, 46 (3/4), 81–98.
Herrich-Schäffer, G.A.W. (1848) Systematische Bearbeitung der Schmetterlinge von Europa. Dritter Band. Die Spanner. In Commission bei G. J. Manz, Regensburg, pp. 1–184.
Hübner, J. (1799) Sammlung europäischer Schmetterlinge. 5. Geometrae. Jakob Hübner Augsburg, 113 pls.
Ivanova, N.V., deWaard, J.R. & Hebert, P.D.N. (2006) An inexpensive, automation-friendly protocol for recovering high-quality DNA. Molecular Ecology Notes, 6, 998–1002.
https://doi.org/10.1111/j.1471-8286.2006.01428.x
Kimura, M. (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16, 111–120.
https://doi.org/10.1007/BF01731581
Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Molecular Biology and Evolution, 35 (6), 1547–1549.
https://doi.org/10.1093/molbev/msy096
Lederer, J. (1853) Die Spanner (Als Fortsetzung des im vorigen Jahre dieser Zeitschrift enthaltenen Aufsatzes: „Versuch, die europäischen Lepidopteren in möglichst natürliche Reihenfolge zu stellen.“). Verhandlungen des Zoologisch-Botanischen Vereins in Wien, 3, 167–255.
Millière, P. (1865) s.n. In: Iconographie et description de chenilles et lépidoptères inédits. II. F. Dumoulin, Lyon, pp. 69–100.
Millière, P. (1884) Cataloge Raisonné des Lépidoptères des Alpes-Maritimes. Supplement 2. Il naturalista siciliano, giornale di scienze naturali, 3 (7), 195–199.
Murillo-Ramos, L., Brehm, G., Sihvonen, P., Hausmann, A., Holm, S., Reza Ghanavi, H., Õunap, E., Truuverk, A., Staude, H., Friedrich, E., Tammaru, T., & Wahlberg, N. (2019) A comprehensive molecular phylogeny of Geometridae (Lepidoptera) with a focus on enigmatic small subfamilies. PeerJ, 7, e7386.
https://doi.org/10.7717/peerj.7386
Müller, B., Erlacher, S., Hausmann, A., Rajaei, H., Shivonen, P. & Skou, P. (2019) Ennominae II. In: Hausmann, A., Rajaei, H., Shivonen, P. & Skou, P. (Eds.), The Geometrid Moths of Europe, 6, pp. 1–906.
https://doi.org/10.1163/9789004387485_001
Prout, L.B. (1915) New genera and species of African Geometridae. Novitates zoologicae: a journal of zoology in connection with the Tring Museum, 22, 311–385.
https://doi.org/10.5962/bhl.part.1923
Prout, L.B. (1912–1916) Die spannerartigen Nachtfalter. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde. Vol. 4. A. Kernen Verlag, Stuttgart, pp. 1–479.
Rajaei, H., Stüning, D. & Trusch, R. (2012) Taxonomic revision and zoogeographical patterns of the species of Gnopharmia Staudinger, 1892 (Geometridae, Ennominae). Zootaxa, 3360 (1), 1–52.
https://doi.org/10.11646/zootaxa.3360.1.1
Rákosy, L. (1997) Genus Nychiodes Lederer 1853 in den Sammlungen des Biologiezentrums in Linz (Lepidoptera, Geometridae). Entomologica Romanica, 2, 13–44.
Robinson, G.S. (1976) The preparation of slides of Lepidoptera genitalia with special reference to the Microlepidoptera. Entomologist’s Gazette, 27, 127–132.
Redondo, V.M., Gastón, F.J. & Gimeno, R. (2009) Geometridae Ibericae. Apollo Books, Stenstrup, 361 pp.
https://doi.org/10.1163/9789004261013
Schmidt, O. (2017) Comparison of the structure and musculature of male terminalia in the tribes Operophterini, Phileremini and Triphosini (Lepidoptera: Geometridae: Larentiinae). Zootaxa, 4237 (2), 347–358.
https://doi.org/10.11646/zootaxa.4237.2.8
Scoble, M.J. (1999) Geometrid Moths of the World: A Catalogue (Lepidoptera, Geometridae). Vol. 1 & 2. CSIRO Publishing, Collingwood, Victoria and Apollo Books, Stenstrup, 1200 pp.
Scoble, M.J. & Hausmann, A. (2007) Online list of valid and available names of the Geometridae of the World. Updated. Available from:
http://www.herbulot.de/globalspecieslist.htm (accessed 18 November 2018)
Sihvonen, P. (2001) Everted vesicae of the Timandra griseata group: methodology and differential features (Geometridae, Sterrhinae). Nota lepidopterologica, 24 (3), 57–64.
Staudinger, O. (1892) Neue Arten und Varietäten von paläarktischen Geometriden aus meiner Sammlung. Deutsche Entomologische Zeitschrift Iris, 5, 141–260.
Staudinger, O. (1871) Catalog der Lepidopteren des Europaeischen Faunengebiets. I. Macrolepidoptera. Dr. O. Staudinger, Dresden, 426 pp.
https://doi.org/10.5962/bhl.title.9475
Staudinger, O. (1901) Catalog der Lepidopteren des palaearctischen Faunengebietes. 1. Friedländer & Sohn, Berlin, 411 pp.
https://doi.org/10.5962/bhl.title.120482
Saitou, N. & Nei, M. (1987) The neighbor-joining method: A new method for reconstructing phylogenetic trees. Molecular Biology and Evolution, 4, 406–425.
Viidalepp, J. (1996) Checklist of the Geometridae (Lepidoptera) of the former U.S.S.R. Apollo Books, Stenstrup, pp 1–111.
Villers, C. de (1789) s.n. In: Caroli Linnaei entomologia, faunae Suecicae descriptionibus aucta, DD. Scopoli, Geoffroy, de Geer, Fabricii, Schrank etc. speciebus vel in systemate non enumeratis, vel nuperrime detectis, vel speciebus Galliae Australis locupletata, generum specierumque rariorum iconibus ornata. Tomus Primus. Piestre & Delamollière, Lugduni pp. j–xvj + 1–765.
Wagner, F. (1909) Einige neue Lepidopterenformen. Entomologische Zeitschrift, Centralorgan des Entomologischen Internationalen Vereins, 23 (4), 17–19.
Wagner, F. (1919) Ueber Nychiodes obscuraria Vill. (=lividaria Hb.). Deutsche Entomologische Zeitschrift, Iris, 33, 105–116.
Wanke, D., Bigalk, S., Krogmann, L., Wendt, I. & Rajaei, H. (2019) The Fixator–A simple method for mounting of arthropod specimens and photography of complex structures in liquid. Zootaxa, 4657 (2), 385–391.
https://doi.org/10.11646/zootaxa.4657.2.11
Wanke, D., Hausmann, A. & Rajaei, H. (2019) An integrative taxonomic revision of the genus Triphosa Stephens, 1829 (Geometridae: Larentiinae) in Middle East and Central Asia, with description of two new species. Zootaxa, 4603 (1), 039–065.
https://doi.org/10.11646/zootaxa.4603.1.2
Wanke, D. & Rajaei, H. (2018) An effective method for the close up photography of insect genitalia during dissection: a case study on the Lepidoptera. Nota lepidopterologica, 4, 219–223.
https://doi.org/10.3897/nl.41.27831
Warren, W. (1902) Drepanulidae, Thyrididae, Uraniidae, Epiplemidae and Geometridae from the Oriental Region. Novitates zoologicae: a journal of zoology in connection with the Tring Museum, IX, 340–372.
Wehrli, E. (1929a) Ueber die Arten und Formen der Gattung Nychiodes Led. Mitteilungen der Münchner Entomologischen Gesellschaft, 19 (1), 29–36.
Wehrli, E. (1929b) Ueber die Arten und Formen der Gattung Nychiodes Led., Fortsetzung. Mitteilungen der Münchner Entomologischen Gesellschaft, 19 (2–4), 37–56.
Wehrli, E. (1929c) Ueber die Arten und Formen der Gattung Nychiodes Led., Nachtrag. Mitteilungen der Münchner Entomologischen Gesellschaft, 19 (5–9), 171–172.
Wehrli, E. (1929) Zwei neue Nychiodes-Arten (Lepid. Geometr.). Internationale Entomologische Zeitschrift, 22 (42), 385–387.
Wehrli, E. (1938) Neue Gattungen, Untergattungen, Arten und Rassen von Geometriden (Lep.). Entomologische Rundschau, 55, 354–360.
Wehrli, E. (1939) Neue Gattungen, Arten und Rassen aus China und Transkaukasien. Entomologische Rundschau, 56, 363–366.
Wehrli, E. (1939-1954) Die Spanner des Palaearktischen Faunengebietes. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde. Vol. 4. Supplement. A. Kernen, Stuttgart. pp. 254–766.
Wiltshire, E.P. (1943) Early stages of oriental palearctic Lepidoptera. V. Journal of the Bombay Natural History Society, Bombay, 43, 621–634.
Wiltshire, E.P. (1957) The Lepidoptera of Iraq. Ministry of Agriculture, Government of Iraq, Baghdad, 162 pp.
Wiltshire, E.P. (1966) Österreichische entomologische Iran-Afghanistan-Expeditionen, Beiträge zur Lepidopterenfauna, Teil 10, Subfamilien Larentiinae und Ennominae (sensu lato) (Lepidoptera, Geometridae). Middle East Lepidoptera XXIII. Zeitschrift der Wiener Entomologischen Gesellschaft, 51, 138–152.
Zerny, H. (1933) Lepidopteren aus dem nördlichen Libanon. Deutsche entomologische Zeitschrift Iris, 47, 60–109.