Research Article |
Corresponding author: Dominic Wanke ( dominic.wanke@smns-bw.de ) Academic editor: Théo Léger
© 2023 Dominic Wanke, Axel Hausmann, David C. Lees, Kyung Min Lee, Geoff Martin, Pasi Sihvonen, Hermann Staude, Hossein Rajaei.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Wanke D, Hausmann A, Lees DC, Lee KM, Martin G, Sihvonen P, Staude H, Rajaei H (2023) The African endemic species “ Nychiodes” tyttha Prout, 1915 (Lepidoptera, Geometridae, Ennominae) belongs to the genus Aphilopota Warren, 1899. Nota Lepidopterologica 46: 1-17. https://doi.org/10.3897/nl.46.94940
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An extensive examination of the external and internal morphological characters of the genus Nychiodes shows that “Nychiodes” tyttha Prout, 1915 is incorrectly placed in this genus. The systematic position of this species was investigated by using a multigene analysis, including one mitochondrial and up to nine protein-coding nuclear gene regions, and morphological characters. These results support a re-classification of this species as Aphilopota tyttha, comb. nov. A re-description supported by illustrations of morphological characters for A. tyttha is provided.
Recently, the genus Nychiodes has undergone intensive integrative taxonomic revisions (
Specimens used in this study are deposited the following collections (acronyms after Evenhuis 2007):
HSS Private Collection of Hermann Staude, South Africa;
For the documentation of external characters, a Visionary Digital photography system (LK Imaging System, Dun. Inc., equipped with a Canon EOS 5DSR camera), an Olympus E3 digital camera, as well as a Leica digital microscope (Z16 APO) were used. Standard techniques were followed for the preparation of genitalia (e.g.
Extraction of DNA and amplification of the “DNA barcode” fragment (658 base-pairs of the 5’ terminus) of the mitochondrial Cytochrome-C Oxidase I of the holotype of Nychiodes tyttha, was carried out at the Canadian Centre for DNA barcoding (CCDB, Guelph), in the framework of the Lepidoptera Campaign of the international Barcode of Life program (iBOL; www.lepbarcoding.org), using a protocol for old museum specimens based on Next-Generation-Sequencing (
Multiple sequences were aligned using Muscle algorithms as implemented in MEGA11 (
Phylogenetic position of Aphilopota tyttha, comb. nov. (marked with a star) within the tribe Boarmiini, supporting the tentative combination in genus Aphilopota. The numbers above the branches are the bootstrap values of the maximum likelihood IQ-TREE analysis. The complete tree is shown in Appendix
Three different analyses where performed. First, COI fragments of “Nychiodes” tyttha (sequences of holotype and two non-type specimens) were compared to available sequences in the Barcode of Life Datasystems (BOLD) identification engine to search for the genetically nearest neighbor. Second, a neighbor-joining tree (K2P on BOLD) was constructed with the sequence of the holotype of “Nychiodes” tyttha and 99 samples suggested by BOLD as related taxa to find the genetically nearest neighbor. Finally, the minimum p-distance of “Nychiodes” tyttha from Nychiodes dalmatina was calculated, to calculate the distance to the genus Nychiodes.
The molecular data set partitioned by gene and codon position was analysed using maximum likelihood as implemented in IQ-TREE 2.1.3 (
The comparison of the COI fragments only of “Nychiodes” tyttha (holotype and two non-type specimens) with data from the BOLD database, suggested that the genetically nearest neighbors are in the genera Jankowskia Oberthür, 1884, Tephronia Hübner, 1825 and Peribatodes Wehrli, 1943 (genetic distances of 6.4–7.9%). When a neighbor-joining tree (K2P on BOLD) was constructed using the holotype DNA barcode sequence with the 99 nearest samples provided by BOLD, an Australian species, Aeolochroma sp. ANIC1 (BOLD:AAV4042), which is 8.33% divergent by p-distance, separated “N.” tyttha from the above and other genera. Sequences from the other two specimens of “N.” tyttha (BC
Additionally, five out of the eleven target genes of a single non-type specimen of “Nychiodes” tyttha were successfully amplified and sequenced (COI-1,COI-2, wgl, Ca-ATPase, Nex9). In the multi-gene phylogenetic analysis “Nychiodes” tyttha clustered as sister to other species of Aphilopota Warren, 1899 (Fig.
The results of our multi-gene molecular phylogenetic analysis show that “Nychiodes” tyttha groups as sister to Aphilopota (UFB = 97%). The phylogenetic analysis would allow us either to classify “N.” tyttha in a monotypic genus as sister to Aphilopota, or to combine it with other Aphilopota. The classification as sister to Aphilopota may not hold when more species of this genus are added to the dataset. Currently the genus Aphilopota consists of 44 species, distributed exclusively in Africa and Madagascar (
In the following diagnosis (see taxonomy part), the morphological characters of Aphilopota tyttha comb. nov. are compared to the type species of the genera Aphilopota (A. interpellans (Butler, 1875)) and Nychiodes (N. obscuraria (Villers, 1789)), which support the new combination.
Nychiodes tyttha
Prout, 1915. Novitates zoologicae: a journal of zoology in connection with the Tring Museum, 22, 363. Holotype ♂ (Eritrea, Caraiai). Deposited in
Holotype, ♂, Eritrea, Caraiai, 21.xi.1905, N. Beccari, Geometridae genitalia slide No. 4976, Rothschild Bequest B.M. 1939-1, NHMUK010920109, DNA barcode sample ID BC
1♀, Eritrea, Caraiai, 21.xi.1905, N. Beccari, Geometridae genitalia slide No. 4977, NHMUK014173598; 1 ♂, Kenya, Kitale, 14.9.[19]25, leg. G.W. Jeffery, NHMUK010920119; 1 ♂, [Namibia], Sissekab, N.W. of Otavi, 1300 m, leg. K. Jordan, 11.xi.1933, NHMUK010920120; all in
The genus Aphilopota needs taxonomic revision, based on a broad integrative taxonomic approach. Therefore, a comparison with other species of this genus, except of the type species A. interpellans, is not possible and also not necessary here.
In A. tyttha labial palpi thin, about two thirds of the diameter of the eye (labial palpi thick, about one diameter of the eye in A. interpellans and N. obscuraria) (Fig.
According to the molecular phylogenetic analysis (Fig.
Wingspan ♂ 21–25 mm, ♀ 28 mm, average length of forewing 11.2 mm (n = 7). Antennae bipectinate in both sexes. Frons weakly convex, just reaching over the eyes, densely scaled. Labial palpi thin, about two third of the diameter of the eye. Proboscis reduced, represented by barely visible rudimentary slats (Fig.
Ground colour of wings beige brown, transverse lines present in dark brown to black. Terminal line continuous, concolorous with transverse lines. In forewing antemedial line curved towards termen. Postmedial line curved between R5/M1 and M2. Medial area with more darker scales intermixed. In hindwing antemedial line curved towards termen on M1. Discal spots only present on underside (Figs
In forewing, vein R1 arising from the cell, not reaching costa, R2 merged with R1, R3–5 with a common stalk arising from the cell. In hindwing Sc+R1 strongly curved in basal area, approximating to the cell, M2 absent, A3 and A1+2 originating separately (Fig.
Wing venation drawings of male specimens of A. Aphilopota tyttha (Prout) comb. nov.; B. Aphilopota interpellans (Butler), and C. Nychiodes obscuraria (Villers). In the forewing of A. tyttha and A. interpellans vein R2 is fused with R1 (rectangle A1), vein R1 and R2 share a common stalk in N. obscuraria (rectangle C1). Remark: as in the genus Nychiodes, the veins R1 and R2 are on a common stalk. This suggests that the veins R1 + R2 are fused to one vein in the genus Aphilopota; therefore, here we name this vein R1+R2.
Wing pattern of Aphilopota tyttha (Prout), comb. nov., Aphilopota interpellans (Butler) and Nychiodes obscuraria (Villers). 5–11. A. tyttha (5. Holotype, Eritrea, Caraiai, g.prep. 4976, NHMUK010920109; 6. Ethiopia, Awassa, Awassa Lake; 7. Kenya, Kitale, NHMUK010920119; 8. [Namibia], Sissekab, N.W. of Otavi, NHMUK010920120; 9. Ethiopia, Bahar Dar, g.prep. 0732/2020 D. Wanke; 10. South Africa, Mkuze Chaos; 11. Eritrea, Caraiai, NHMUK014173598); 12–14. A. interpellans (12. South Africa, KwaZuluNatal, Vryheid, g.prep. 1314/2022 D. Wanke; 13. [South Africa], Transvaal, Bords du Limpopo, g.prep. 1316/2022 D. Wanke; 14. South Africa, KwaZuluNatal, Vryheid, g.prep. 1315/2022 D. Wanke); 15–16. N. obscuraria (15. Spain, Albarracin, g.prep. 2096/2017 H. Rajaei; 16. Spain, Albarracin, g.prep. 2097/2017 H. Rajaei); a = upperside; b = underside. Scale bar: 1 cm.
In male genitalia uncus strongly sclerotized, short, basally broad and triangular, apically pointed. Gnathos well developed and strongly sclerotized, triangular. Costa of valva sclerotized, valva thin, without any ampulla or harpe. Juxta forked and big, reaching up to gnathos. Saccus tapering. Aedeagus thin tapered, carrying one long and sclerotized cornutus. Cornutus almost same length as aedeagus (Figs
Male genitalia of Aphilopota tyttha (Prout), comb. nov., Aphilopota interpellans (Butler) and Nychiodes obscuraria (Villers). 17–18. A. tyttha (17. Holotype, Eritrea, Caraiai, g.prep. 4976, NHMUK010920109; 18. Ethiopia, Bahar Dar, g.prep. 0732/2020 D. Wanke); 19. A. interpellans (South Africa, KwaZuluNatal, Vryheid, g.prep. 1314/2022 D. Wanke); 20. N. obscuraria (Spain, Albarracin, g.prep. 2096/2017 H. Rajaei). a = genitalia capsule; b = aedeagus. Scale bar: 1 mm.
Female genitalia thin and long, with strongly elongated ovipositor. Apophyses posteriores very long, apophyses anteriores 1/3 length of apophyses posteriors. Antrum sclerotized. Ductus bursae short, bend. Corpus bursae tube-like, elongated. Signum absent (Fig.
Female genitalia of Aphilopota tyttha (Prout), comb. nov., Aphilopota interpellans (Butler) and Nychiodes obscuraria (Villers). 21. A. tyttha (Eritrea, Caraiai, g.prep. 4977, NHMUK014173598); 22–23. A. interpellans (South Africa, KwaZuluNatal, Vryheid, g.prep. 1315/2022 D. Wanke; 22. Ventral view; 23. Lateral view); 24. N. obscuraria (Spain, Albarracin, g.prep. 2097/2017 H. Rajaei). Scale bar: 1 mm.
Adults observed from November to May.
Unknown.
Investigated specimens collected at elevations from 220 to 1300 m in dry savanna ecoregions.
In East Africa (Eritrea, Ethiopia, Kenya), south-western Africa (Namibia) and South Africa (
The copyright of images of the specimens deposited in
Complete phylogenetic analysis from IQ-TREE, showing the phylogenetic position of Aphilopota tyttha comb. nov. (marked with a star) within the tribe Boarmiini. Support values are indicated above the branch. Node confidence values were estimated based on 1000 ultrafast bootstrap replicates.
Taxa used in this study
Data type: table (excel file).
Explanation note: Taxa used in this study, with identification, process code, and GenBank accession numbers for each gene. Data from