Drepanogynis insciata (Felder & Rogenhofer, 1875), a South African geometrid moth lost to science rediscovered after more than 140 years (Lepidoptera, Geometridae, Ennominae)

261 Drepanogynis insciata (Felder & Rogenhofer, 1875), a South African geometrid moth lost to science rediscovered after more than 140 years (Lepidoptera, Geometridae, Ennominae) Pasi Sihvonen1, Kyung Min Lee1, Max Söderholm1, Hossein Rajaei2, Axel Hausmann3, Hermann S. Staude4 1 Finnish Museum of Natural History, P. O. Box 17 (Pohjoinen Rautatiekatu 13), FI-00014, University of Helsinki, Helsinki, Finland 2 State Museum of Natural History, Stuttgart, Germany 3 Bavarian State Collection of Zoology (SNSB-ZSM), Munich, Germany 4 Duif str. 2, Hoekwil, South Africa Corresponding author: Pasi Sihvonen ([email protected]) Copyright: © Pasi Sihvonen et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract The geometrid moth Drepanogynis insciata (Felder & Rogenhofer, 1875) has been known only from two specimens in the Natural History Museum, London, UK: the holotype male collected before 1875 from South Africa, and a second male specimen without date and collecting locality data but probably collected before 1879. Unexpectedly, 13 specimens of this species—long considered lost to science—were observed in four locations between 2020 and 2023 in the Western Cape, near the type locality of the species. These observations were based on photographs available on iNaturalist, apart from a single male attracted to light, which was collected and vouchered for study from the Western Cape on 28 January 2022. We illustrate D. insciata and the newly collected specimen using both classical dissection and non-destructive micro-CT imaging. Phylogenetic analysis based on the multi-gene maximum-likelihood method places the species within Ennominae, tribe Drepanogynini. DNA barcodes reveal its nearest genetic neighbor to be Drepanogynis smaragdaria Krüger, 2002, with 5.9% sequence divergence. We discuss the conservation implications of this rediscovery. Key words: DNA barcoding, molecular phylogeny, morphology, rediscovery Introduction Occasionally, rare or presumed extinct species are rediscovered, and such cases often make headlines. Recent examples include the Portuguese spider (Nemesia berlandii Frade & Bacelar, 1931; Arachnida, Nemesiidae), not discovered since 1931 and found again in 2023; the earless Australian dragon (Tympanocryptis pinguicolla Mitchell, 1948; Sauropsida, Agamidae), last seen in 1969 and rediscovered in 2023; and Wallace’s giant bee from Indonesia (Megachile pluto (Smith, 1860); Hymenoptera, Megachilidae), which had vanished since 1981 and was rediscovered in 2019 (Rewild 2025). The Critically Endangered South African geometrid moth Callioratis millari (Hampson, Academic editor: Gunnar Brehm Received: 13 September 2025 Accepted: 12 November 2025 Published: 2 December 2025 ZooBank: https://zoobank.org/ C053757E-B091-4EA6-91766104483B49D6 Citation: Sihvonen P, Lee KM, Söderholm M, Rajaei H, Hausmann A, Staude HS (2025) Drepanogynis insciata (Felder & Rogenhofer, 1875), a South African geometrid moth lost to science rediscovered after more than 140 years (Lepidoptera, Geometridae, Ennominae). ZooKeys 1261: 261–276. https://doi.org/10.3897/ zookeys.1261.171904 ZooKeys 1261: 261–276 (2025) DOI: 10.3897/zookeys.1261.171904 262 ZooKeys 1261: 261–276 (2025), DOI: 10.3897/zookeys.1261.171904 Pasi Sihvonen et al.: Drepanogynis insciata rediscovered 1905) disappeared when its known habitat was destroyed in 1928 and was not found, despite extensive searching, until 1996 when it was rediscovered over 100 km north of its type locality (Staude 2001). Here, we report a similar case from Lepidoptera, family Geometridae. Drepanogynis insciata (Felder & Rogenhofer, 1875) is a distinctive species with greenish-whitish wings, wine-red margins, and a relatively large wingspan of 27 mm (Figs 1, 2). This species was originally described from South Africa in 1875 based on a single male specimen. Although the holotype lacks a collection date (Fig. 1A), there is good reason to believe it was collected in 1857 during the Austrian frigate’s (Fregatte Novara) global expedition. The explorers made their long voyage during 1857–1859 and visited South Africa from 2 to 26 October 1857 (Wüllerstorf-Urbair 1861: 174–226). The holotype was collected from Swellendam, the third oldest town in South Africa. At the time of the Austrian expedition, Swellendam was a well-known settlement frequently visited by many travelers and naturalists (Wikipedia 2025a). A second historical male specimen exists in the Natural History Museum, London, although it lacks collection data (Fig. 1B). This specimen was originally part of the collection of the French entomologist Jean-Baptiste Boisduval (1799–1879), later acquired by French Charles Oberthür (1845–1924), and ultimately donated to the Natural History Museum, London in 1927 (Wikipedia 2024, 2025b). Therefore, based on indirect evidence, the second specimen must have been collected before Boisduval’s death in 1879. Against expectations, 12 individuals of D. insciata—thought to be lost to science—were observed and photographed between 2020 and 2023 on the southern coast of Western Cape province, within about 150 km radius of the type locality. These records are available on iNaturalist, but the specimens were not physically vouchered (Fig. 2). A single male, which was attracted to an unspecified outdoor light in Gondwana Private Nature Reserve, near Mossel Bay on 28 January 2022, was vouchered for study (Fig. 1C). Based on this evidence, the species has remained undocumented in collections for at least 141 years from 1879 to 2020. Taxonomic history of D. insciata has been complex. It was originally combined in Axia(?) (Felder & Rogenhofer 1875), it was later transferred to Axiodes Warren, 1894, as its type species (Warren 1894), then to Drepanogynis by Janse (1932), then again returned to Axiodes by Prout (1929), followed by Scoble (1999), and finally classified as Drepanogynis insciata in a revision of Drepanogynis by Krüger (2002). In that revisionary paper, this species was classified in the D. insciata group containing six species. The group is characterized by greenish wings (except D. trachyacta (Prout, 1922)) and partly scobinate, flattened processes of juxta in the male genitalia. Krüger (2002) also discussed the difficulties of Drepanogynis classification, proposing a broadened genus concept of Drepanogynis that synonymized five genera, including Axiodes, under Drepanogynis. This concept was approximating the earlier view of Janse (1932), but which was not adopted by Prout (1938). Currently, the taxon insciata is classified as Drepanogynis insciata (Krüger 2020; Rajaei et al. 2022), but the broad genus classification hypothesis of Drepanogynis remains to be tested, particularly with extensive molecular and life history data Murillo-Ramos et al. (2019). 263 ZooKeys 1261: 261–276 (2025), DOI: 10.3897/zookeys.1261.171904 Pasi Sihvonen et al.: Drepanogynis insciata rediscovered Until now, Drepanogynis insciata has only been illustrated by a hand-drawn picture (Felder et al. 1875) and a greyscale photograph of the adult and a schematic drawing of the male genitalia (Krüger 2002). No molecular data of this taxon have been available to date. We use this opportunity to raise awareness on this unique case and on South African biodiversity in general. We aim to clarify its identity and provide high-resolution images of the type specimen, the newly collected specimen and its diagnostic characters, describe the collecting site—located in a highly transformed region on South Africa’s southern coast—and determine its phylogenetic position based on molecular data. We also identify its closest known relative based on DNA barcoding. Material and methods Examined material Holotype male: [SOUTH AFRICA, Western Cape]: Swellendam (L. Taats); Novara CXXIX/ f. 10. Axia?/ insciata/ Flr. m. ♂ m; Rothschild/ Bequest/ B.M.1939-1.; Axia/ insciata/ type Felder; Geometridae/ gen. prep./ no. 21066; BMNH(E) # 275235/ Digitally scanned; NHMUK 010891612 (coll. Natural History Museum, London, UK). 2 males: [no locality data, no date data] Ex. Oberthür Coll./ Brit. Mus. 1927– 3.; Ex MUSAEO/ Dris BOISDUVAL; NHMUK 010891613 (coll. Natural History Museum, London, UK). SOUTH AFRICA, Western Cape,/ Mossel Bay, Gondwana Private Nature Reserve,/ main Lodge, −34.0548 21.8997/ 330 m, 28-01-2022/ leg. C. Scott; Axiodes insciata/ (Felder & Rogenhofer, 1875); Pasi Sihvonen/ DNA sample 1561; Pasi Sihvonen/ prep. number 2914; micro-CT scan #164/ Max Söderholm (coll. Hermann Staude, South Africa). Examined photographs: 12 males from South Africa: Western Cape, photos available on iNaturalist as detailed in Table 1. Table 1. Observation details of 12 Drepanogynis insciata males as available on iNaturalist, arranged by observation site. None of these were physically vouchered for study, but their external characters agree with typical D. insciata. Search term for the species on iNaturalist is “Axiodes insciata”. Date Exact site Longitude, Latitude Altitude (m) Observer / iNaturalist username 11.11.2023 Haarwegskloof, Overberg district municipality -34.3383, 20.3260 200 Petra Broddle 29.10.2021 Haarwegskloof, Overberg district municipality -34.3383, 20.3260 200 Grant Forbes 29.10.2021 Haarwegskloof, Overberg district municipality -34.3383, 20.3260 200 Odette Curtis 10.10.2023 Mossel Bay, Hartebeestkuil Game Reserve -34.0714, 22.0098 130 Eon Grobler 29.12.2021 Mossel Bay, Gondwana Private Nature Reserve -34.0548, 21.8997 330 Cameron Scott 11.1.2022 Mossel Bay, Gondwana Private Nature Reserve -34.0548, 21.8997 330 Cameron Scott 7.9.2023 Mossel Bay, Gondwana Private Nature Reserve -34.0548, 21.8997 330 PG Coetsee 29.12.2021 Mossel Bay, Gondwana Private Nature Reserve -34.0548, 21.8997 330 Kevin Koen 7.9.2023 Mossel Bay, Gondwana Private Nature Reserve -34.0548, 21.8997 330 Kevin Koen 7.9.2023 Mossel Bay, Gondwana Private Nature Reserve -34.0548, 21.8997 330 Jolandie3 25.9.2020 Mossel Bay, Gondwana Private Nature Reserve -34.0548, 21.8997 330 Cameron Scott 10/2022 Overberg Renosterveld, Swellendam -34.2372, 20.3312 170 Dirrtyharry 264 ZooKeys 1261: 261–276 (2025), DOI: 10.3897/zookeys.1261.171904 Pasi Sihvonen et al.: Drepanogynis insciata rediscovered Morphological analyses Adult specimens, genitalia, and abdomens were prepared and photographed following methods summarized in Sihvonen et al. (2020). Uneverted vesica and tympanal organs were photographed in situ during dissection to allow optimal angle for observation and illustration. Wing venation was studied using non-destructive micro-CT scanning, as described by Souza Moraes et al. (2023). Photographs were edited in Adobe Photoshop v. CS6, and figures were compiled in CorelDRAW v. 24. The map of sampling sites and photographic records were created using QGIS v. 3.6. Molecular analyses DNA was extracted from a leg of the specimen collected in 2022. The procedure for DNA extraction, purification, amplification, cleaning, and sequencing of both mitochondrial (COI) and protein-coding nuclear gene regions (Wingless, RpS5, Ca-ATPase, Nex9, and EF-1alpha) (Table 2) followed the protocols described by Sihvonen et al. (2020) and Lee et al. (2024). All molecular work was conducted at the DNA laboratory of the Finnish Museum of Natural History. PCR products were enzymatically cleaned and sequenced in the Institute for Molecular Medicine Finland (FIMM; Helsinki, Finland). Sequence alignment, cleaning, model selection, tree search strategies using maximum-likelihood (ML), node support estimation, and tree visualization also followed the aforementioned protocols. Molecular data for Drepanogynis insciata were analysed using a maximum-likelihood approach implemented in IQ-TREE (Trifinopoulos et al. 2016), using the 1206-taxon dataset compiled by Murillo-Ramos et al. (2019). Best-fitting substitution models were selected by ModelFinder (Kalyaanamoorthy et al. 2017), with each partition assigned its own evolutionary rate. The resulting phylogenetic tree was rooted using representative species of the families Sematuridae, Epicopeiidae, Pseudobistonidae and Uraniidae. The tree was visualized and rooted in FigTree v. 1.4.3 (Rambaut 2015) and edited for presentation in Adobe Illustrator v. CS6 and CorelDRAW v. 24. DNA barcoding A 658 bp region near the 5′ terminus of the COI mitochondrial gene (the standard animal DNA barcode) was analysed for the single specimen collected in 2022. Analyses were conducted using the BOLD analytical tools, including BIN (Barcode Index Number) and barcode gap analysis (Ratnasingham and Hebert 2007; Ratnasingham and Hebert 2013). Genetic divergences were calculated as the number of base differences between sequences and reported as a percentage, following the implementation in MEGA 11 (Tamura et al. 2021). Table 2. GenBank accession numbers for the new sequences used in this study. Voucher COI RpS5 Wgl EF-1alpha Ca-ATPase Nex9 PMS1561 PV937082 PV932705 PV932701 PV932703 PV932702 PV932704 265 ZooKeys 1261: 261–276 (2025), DOI: 10.3897/zookeys.1261.171904 Pasi Sihvonen et al.: Drepanogynis insciata rediscovered Results Diagnosis. Drepanogynis insciata belongs to the D. insciata group (sensu Krüger 2002), containing six greenish (except blackish-brown D. trachyacta (Prout, 1922)), noctuid-like stout species characterized by distinct, undulating medial and postmedial lines on forewings, delimiting wide medial area. In the male genitalia the processes of juxta flattened, with partly scobinate surfaces. Key to species based on adults 1 Forewings blackish-brown ....................... D. trachyacta (see Krüger 2002) – Forewings greenish-white .............................................................................2 2 Forewing postmedial line pointing outwards on termen .............D. insciata – Forewing postmedial line pointing down or inwards on termen ................3 3 Hindwing with distinct, white-bordered postmedian line .............................. ................................................................D. kalahariensis (see Krüger 2002) – Hindwing without distinct postmedian line .................................................4 4 Forewing medial line evenly curved ........ D. rhodampyx (see Krüger 2002) – Forewing medial line distinctly angled in middle ........................................5 5 Cilia concolorous ....................................D. smaragdaria (see Krüger 2002) – Cilia chequered purple-and-green ........D. smaragdarioides (see Krüger 2002) Drepanogynis insciata differs from all other species of D. insciata group (sensu Krüger 2002) by the distinctive shape of the forewing antemedial and postmedial lines (other species illustrated by Krüger 2002): antemedial line is concave near costa (line almost straight or angled inwards in other species), postmedial line features a pointed extrusion about halfway across the wing (extrusion round or angled in other species) and the line is angled outwards near the termen (line angled inwards or rectangular near termen in other species) (Fig. 1). Male genitalia possess unique diagnostic characters: the juxta processes are densely dentate, with a broad lateral plate and a narrow, upward-pointing extrusion (upward-pointing extrusion absent in other species). The aedeagus apex is armed with numerous minute teeth (minute teeth not illustrated for other species by Krüger 2002) (Fig. 3). Redescription. External characters and abdomen: wingspan 25–28 mm (n = 3). Adults rest with wings folded over body, in tent like position (Fig. 2A, B). Forewings olive green, medial area darker with red-wine-color distally, margins white. Antemedial line weakly S-shaped, with one extrusion. Postmedial line with two prominent extrusions. Hindwings basally cream, distally grey. Fringes red wine-colored, weakly chequered in both wings. Discal spots absent. Forewing costa concave. Frons, collar, and thorax concolorous with wings, abdomen pinkish dorsally. Wings below pale green, fringes and legs partly red-wine-colored. Antennae bipectinate. Hindtibia with 2 + 2 spurs. Tympanal organs medium-sized, ansa wider near base, apex with curved extension. Abdominal sternites and tergites undifferentiated. The holotype is faded in color, forewings with flesh tone. Wing venation: homology interpretation of forewing R veins is difficult. Only four veins are present as tubular, instead of common state of five. We hypothesize that one of the R1–R3 veins is fused, so that only two of these are present as tubular, and R4 + R5 are present. Forewing with four R veins: R1/R2 + R2/R3 266 ZooKeys 1261: 261–276 (2025), DOI: 10.3897/zookeys.1261.171904 Pasi Sihvonen et al.: Drepanogynis insciata rediscovered stalked, R4 + R5 stalked, these veins running parallel, forming areole. Hindwing M2 absent, but weak fold present. Wing margins strongly crenulate (Fig. 4). Male genitalia: uncus long, curved ventrally, weakly setose. Socii small, setose. Gnathos arms narrow, fused medially, forming sclerotized, sharply upcurved element. Juxta processes large, with narrow upwards pointing extrusion, lateral parts wider, upcurved, densely dentate. Valva simple, extensively setose, dorsal margin weakly sclerotized, ventral margin weakly concave. SacFigure 1. External characters and selected labels of all known Drepanogynis insciata specimens. A. Holotype male, without date, but specimen collected before 1875; B. An old specimen in The Natural History Museum, London, UK. The specimen is without date and location data, but based on indirect evidence, it is probably collected before 1879; C. Specimen collected on 28 January 2022 from Western Cape: Gondwana Private Game Reserve, after 147 years when the species was described. The old specimens have lost their green color. 267 ZooKeys 1261: 261–276 (2025), DOI: 10.3897/zookeys.1261.171904 Pasi Sihvonen et al.: Drepanogynis insciata rediscovered cus narrow, concave medially. Aedeagus weakly curved, apex covered with numerous minute teeth. Vesica opens at approximately 135° angle, basal part weakly enlarged, with row of microcornuti. Vesica apex evenly tapering (Fig. 3). Distribution and abundance. Known from only five localities in South Africa within 150 km radius of each other: Swellendam (the type locality), Overberg Renosterveld in Swellendam, Haarwegskloof, Hartebeesstkuil Game Reserve and Gondwana Private Game Reserve, all in the Western Cape (Fig. 5). Only three male specimens are preserved in the collections: the holotype (collected before 1875) (Fig. 1A), a male collected 28.1.2022 (Fig. 1C), and a male without date, but probably collected before 1879 (Fig. 1B). Twelve male specimens were observed between 2020 and 2022, but these were not vouchered (Table 1). Phenology. The species has been observed between September and January, potentially being bivoltine. Adults appear to be strictly nocturnal, and all individuals were attracted to unspecified outdoor lights. Biology. Unknown. Habitat. Recorded from fynbos habitats at lower elevations (thus far below 330 m) on the southern slopes of the Langeberge and in the coastal plain towards the Indian Ocean from, near Swellendam and Herbertsdale. The collection locality of the specimen from the Gondwana Private Game Reserve is shown in Fig. 6. Phylogeny and genetic data. Based on multi-gene molecular phylogenetic analysis and morphological data, D. insciata is classified within Ennominae, in the tribe Drepanogynini (Fig. 7). The assignment to Drepanogynini was strongly supported in the maximum-likelihood analysis (100%), although internal relationships within the tribe were mostly poorly supported. Comparisons of the DNA barcode against data on BOLD (http://www.barcodinglife.org/) and GenBank (http://www.ncbi.nlm.nih.gov/Genbank) show the nearest genetic match to be D. smaragdaria Krüger, 2002 (voucher BC_ZSM_Lep_118901) with a divergence of 5.9%, and D. bifasciata (Dewitz, 1881) (voucher HSS-18006) Figure 2. A selection of Drepanogynis insciata male specimens available on iNaturalist. The physical specimens were not vouchered for study, but the identity of these, judged by the external characters, agrees with typical D. insciata. All specimens are from South Africa: Western Cape district. A. Mossel Bay district, Hartebeestkuil Game Reserve, 10 November 2023, dorsal view (photo by Eon Grobler); B. Mossel Bay district, Hartebeestkuil Game Reserve, 10 November 2023, lateral view (photo by Eon Grobler); C. Mossel Bay, Gondwana Private Game Reserve, 7 September 2023 (photo by Jolandie3). Photos are reproduced under CC BY-NC 4.0 license, https://creativecommons.org/licenses/by-nc/4.0/ 268 ZooKeys 1261: 261–276 (2025), DOI: 10.3897/zookeys.1261.171904 Pasi Sihvonen et al.: Drepanogynis insciata rediscovered Figure 3. Male genitalia of Drepanogynis insciata. A–C. Holotype (coll. NHMUK); A. Genitalia; B. Aedeagus; C. Abdomen (dissection Geometridae gen. prep. no. 21066 by an unknown person); D–I. Specimen collected 28 January 2022 (coll. H. Staude); D. Genitalia; E. Aedeagus; F. Aedeagus with everted vesica; G. Magnification of the aedeagus apex with dentate sclerotisations and base of the vesica with microcornuti; H. Tympanal organs; I. Ansa (dissection Pasi Sihvonen prep. no. 2914, coll. Hermann S. Staude). 269 ZooKeys 1261: 261–276 (2025), DOI: 10.3897/zookeys.1261.171904 Pasi Sihvonen et al.: Drepanogynis insciata rediscovered Figure 4. Wing venation of Drepanogynis insciata, specimen collected 28 January 2022 (micro-CT scan no.164 Max Söderholm, coll. Hermann S. Staude). Figure 5. The known records of Drepanogynis insciata in South Africa. The species is known from five places in Western Cape only, which are within 150 km from each other. The holotype was collected from Swellendam pre-1875, likely in 1857, while all other records are between 2020 and 2023. 276 ZooKeys 1261: 261–276 (2025), DOI: 10.3897/zookeys.1261.171904 Pasi Sihvonen et al.: Drepanogynis insciata rediscovered Supplementary material 1 Phylogenetic tree Authors: Pasi Sihvonen, Kyung Min Lee, Max Söderholm, Hossein Rajaei, Axel Hausmann, Hermann S. Staude Data type: pdf Explanation note: IQ-TREE showing the full 1206 taxa dataset of Murillo-Ramos et al. (2019), based on the maximum-likelihood approach of up to 11 mitochondrial and nuclear genes, supplemented with voucher number PMS1561_Axiodes insciata (marked with pale orange). Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zookeys.1261.171904.suppl1