Journal of South American Earth Sciences 120 (2022) 104055 Contents lists available at ScienceDirect Journal of South American Earth Sciences journal homepage: www.elsevier.com/locate/jsames A new fossil inchworm moth discovered in Miocene Dominican amber (Lepidoptera: Geometridae) Victor Sarto i Monteys a, b, *, Axel Hausmann c, Mónica M. Solórzano-Kraemer d, Jörg U. Hammel e, Joaquín Baixeras f, Xavier Delclòs g, h, Enrique Peñalver i a Institut de Ciència i Tecnologia Ambientals (ICTA), Entomology, Plants and Health, Edifici Z, C/ de les columnes s/n. Campus de Bellaterra, Universitat Autònoma de Barcelona, Bellaterra, Spain b Servei de Sanitat Vegetal, DARP, Generalitat de Catalunya, Barcelona, Spain c SNSB – Zoologische Staatssammlung München, Münchhausenstraße 21, D-81247 Munich, Germany d Senckenberg Forschungsinstitut und Naturmuseum, Paleaontologie und Historische Geologie, Senckenberganlage 25, 60325 Frankfurt am Main, Germany e Institute of Materials Physics, Helmholtz-Zentrum Hereon, Outstation at DESY, Max-Planck-Str. 1, D-21502 Geesthacht, Germany f Institut Cavanilles de Biodiversitat i Biologia Evolutiva, Universitat de València, C/ Catedràtic José Beltran 2, 46980 Paterna (València), Spain g Department Dinàmica de la Terra i de l’Oceà, Facultat de Ciències de La Terra, Universitat de Barcelona (UB), Martí i Franquès s/n, 08028 Barcelona, Spain h Institut de Recerca de la Biodiversitat (IRBio), Universitat de Barcelona (UB), Barcelona, Spain i Instituto Geológico y Minero de España (IGME), CSIC, C/ Cirilo Amorós 42, 46004 València, Spain A R T I C L E I N F O A B S T R A C T Keywords: We report a fossil geometrid moth, a male, virtually complete, preserved in a clear piece of Miocene Dominican Lepidoptera amber dating from 19 to 16 Mya. Fore- and hindwings appear partially overlapped, and all body characters are Desmobathrinae visible externally in dorsal and ventral views, including the outer surface of the valvae of the genitalia. The scale New species pattern on the wing membrane is preserved, whereas the wing color pattern is not. It belongs to the genus Amber Dolichoneura (Geometridae: Desmobathrinae) and is named Dolichoneura jorelisae Sarto i Monteys, Hausmann, Miocene Baixeras and Peñalver sp. n., based on wing features. Because of the poor fossil record of lepidopterans, both in amber and compression rocks, the description of the available well-preserved specimens is of considerable in­ terest for phylogenetic studies. Furthermore, it could also serve for calibrating molecular clocks and for paleo­ biogeographic inferences. 1. Introduction inclusions in ambers, principally because they live and eat far from the trunk or big branches of resiniferous trees, from where the resin would So far, the oldest known fossil lepidopteran, Archaeolepis mane mostly ooze (Solórzano Kraemer et al., 2018). Actually, macro­ Whalley (1985), is from the Lower Jurassic (Sinemurian, ca. 190 Mya) of lepidopteran wings can get caught very easily in resin, so their size is not England (Whalley, 1985). Yet, phylogenomic studies by Kawahara et al. that important. If they get trapped in resin emissions on a tree trunk or (2019) suggest that the most recent common ancestor of crown Lepi­ branch, the ants will have easy access and will eat them before the next doptera appeared perhaps significantly earlier, in the Late Carbonif­ flow arrives covering the carcasses (MMS-K, XD, EP, personal observa­ erous, ca. 300 Mya, although this estimation might be biased (this age is tion, Madagascar). All this contributes to the scarcity of their inclusions very close to the initial appearance of the Holometabola) and should be in ambers. confirmed by further studies. Geometridae, whose age was estimated at about 54 Mya, in the Although Lepidoptera is one of the megadiverse orders of insects and Eocene (credibility interval of 62 to 48 Mya corresponding to Middle one of the most abundant around different terrestrial ecosystems, they Paleocene–Early Eocene) (Yamamoto and Sota, 2007; Wahlberg et al., are, due to taphonomical processes (Martínez-Delclòs and Martinell, 2013), stand as the second most species-rich family of Lepidoptera, 1993), among the rarest findings amid insect orders in deep time, mostly comprising ca. 24,000 described species (van Nieukerken et al., 2011; occurring in ambers. Yet, imagoes of macrolepidopterans are rare as Murillo-Ramos et al., 2019). They belong to the superfamily * Corresponding author. Institut de Ciència i Tecnologia Ambientals (ICTA), Entomology, Plants and Health, Edifici Z, C/ de les columnes s/n. Campus de Bel­ laterra, Universitat Autònoma de Barcelona, Bellaterra, Spain. E-mail address: victor.sarto@uab.cat (V. Sarto i Monteys). https://doi.org/10.1016/j.jsames.2022.104055 Received 8 August 2022; Accepted 27 September 2022 Available online 30 September 2022 0895-9811/© 2022 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by- nc-nd/4.0/). V. Sarto i Monteys et al. Journal of South American Earth Sciences 120 (2022) 104055 Fig. 2. Virtual representation (SRμCT) in different views of the head of †Dolichoneura jorelisae sp. n. (Geometridae: Desmobathrinae), holotype. Both antennae are distally incomplete due to the limits of the scan; in the amber piece one distal end was kept within it (see Fig. 3B). are from amber collected in this area: (1) an adult assigned to Enno­ Fig. 1. Habitus of †Dolichoneura jorelisae sp. n. (Geometridae: Desmoba­ thrinae), holotype. (A) Dominican amber piece with the geometrid specimen, minae, plus (2) a caterpillar and (3) another adult, both unassigned, ventral view. (B) Virtual representation (SRμCT) of the geometrid specimen in figured but undescribed (Grimaldi and Engel, 2005), and (4) an adult ventral view, with the hind wing venation enhanced. assigned to Ennominae, described as Miogeometrida chunjenshihi Zhang et al., 2020 (Zhang et al., 2020). Here we report a fifth geometrid moth Geometroidea, which traces back to 83 Mya (Wahlberg et al., 2013). included in Dominican amber. Geometrids are characterized by the presence of paired tympanal organs placed at the base of the abdomen; they occur throughout the family, in 2. Materials and methods both sexes, although they are secondarily reduced or lost in a very few flightless females (Cook and Scoble, 1992). The prolegs of their larvae The type specimen in amber described herein is housed at the Museo are usually reduced to two pairs causing the larvae to move by ‘looping’ Nacional de Historia Natural “Prof. Eugenio de Jesús Marcano”, Santo (Minet and Scoble, 1999), that is where the names “inchworm”, Domingo, Dominican Republic, with the accession number MNHNSD “loopers” and “Geometridae”, as for ‘earth-measuring’, come from. A FOS 16.01. The piece that includes the moth originates from the eastern comprehensive molecular study by Murillo-Ramos et al. (2019, 2021) amber mines of Hato Mayor-El Valle Region, Dominican Republic proposed recognizing nine mostly monophyletic geometrid subfamilies: (Eastern district in Iturralde-Vinent and MacPhee, 2019); it is oval, Sterrhinae, Larentiinae, Archiearinae, Desmobathrinae, Orthostixinae weighs 3.58 g, with dimensions 31 mm by 20 mm, and a height of 10 (validity controversial), Epidesmiinae, Oenochrominae, Geometrinae, mm–2 mm since it is flattening from one side to the other, no internal and Ennominae. cracks are visible and no syninclusions are observed. The moth is situ­ Despite of being one of the most specious family groups of Lepi­ ated centrally within it, with its head contacting one outer border doptera, the fossil record of Geometridae is comparatively very scarce, (Fig. 1). The piece was delivered to us by Mr. Jorge Caridad, from his with only 19 findings formally reported worldwide, most of them of private collection in the Amber World Museum. Mr. Jorge Caridad is difficult systematic assignment. The affiliation of two of them is disputed known to most authors, and is an established and trusted source. by Sohn et al. (2012). They were all listed and reviewed by Zhang et al. (2020), who noted that Eogeometer vadens Fischer, Michalski and 2.1. Spectroscopy Hausmann, 2020 (Bartonian–Priabonian, Baltic amber), Geometridites larentiiformis Jarzembowski, 1980 (Upper Priabonian, Bembridge Marls, The amber piece was kept at constant temperature in plastic clip bags of the Isle of Wight), and Hydriomena protrita Cockerell, 1922 (Priabo­ within metal boxes during the study to exclude oxygen and light. To nian, Florissant Fossil Beds), all from the Late Eocene, were believed to determine its origin and establish a molecular fingerprint, we carried out be the earliest representatives of Geometridae. a Fourier Transform Infrared Spectroscopy (FTIR) (see results below). To To date, 31 fossil specimens within seven superfamilies of ditrysian ensure the piece containing the moth was amber, and not copal or Lepidoptera had been reported from amber collected in the Dominican Defaunation resin (sensu Solórzano-Kraemer et al., 2020), its FTIR Republic (Grimaldi and Engel, 2005; Sohn et al., 2012; Zhang et al., spectrum was compared to that of two pieces of amber found by us in the 2020), with an estimated age of ca. 19–16 Mya (Iturralde-Vinent and field, one in the Northern district (Palo Quemado) and another in the MacPhee, 2019). Among the known fossil geometrids worldwide, four Eastern district (El Valle - Mina Siete Cañadas), the latter belonging to 2 V. Sarto i Monteys et al. Journal of South American Earth Sciences 120 (2022) 104055 used cool white LED illuminators. Images were prepared for illustration using Adobe Photoshop CS2, and some of them were z-stacked. The body length was measured from the frons to the end of the abdomen. 2.3. Synchrotron radiation micro-computed tomography (SRμ-CT) In order to observe morphological details which could not be prop­ erly seen under a stereomicroscope, the inclusion was additionally studied by synchrotron radiation based x-ray micro tomography (SRμCT). Imaging was performed at the Imaging Beamline – IBL P05 – PETRA III at Deutsches Elektronen Synchrotron (DESY) in Hamburg operated by the Helmholtz-Zentrum Hereon (Greving et al., 2014; Wilde et al., 2016). Due to the size of the specimen imaging was done in an extended field of view (eVoF) mode (~12 mm × 5 mm). 12,602 pro­ jections have been recorded at 6001 angular steps equally spaced be­ tween 0 and π. Projections were recorded at a photon energy of 18 keV and a sample to detector distance of 30 mm using a custum build CMOS detector (Lytaev et al., 2014). Tomographic reconstruction was done with a beamline specific stitching and processing workflow (Moosmann et al., 2014; Beckmann pers. comm., June 2019) implemented in Matlab (Math Works) and IDL (Harris Inc) using the filtered back projection algorithm implemented in the Astra Toolbox (van Aarle et al., 2015, 2016; Palenstijn et al., 2011). Raw projections have been binned two times for processing, resulting in an effective pixel size of 1.28 μm in the reconstructed tomographic volume. 2.4. Systematics Family-level classification follows van Nieukerken et al. (2011). Terminology of veins follows Hausmann (2001). 3. Results 3.1. Systematic paleontology Order Lepidoptera Linnaeus, 1758 Suborder Glossata Fabricius, 1775 Clade Ditrysia Börner, 1925. Superfamily Geometroidea Leach, 1815 Family Geometridae Leach, 1815 Subfamily Desmobathrinae Meyrick, 1886 Fig. 3. Cilia on antennal flagellum of †Dolichoneura jorelisae sp. n. (Geo­ Genus Dolichoneura Warren, 1894 metridae: Desmobathrinae), holotype. (A) Flagellum densely setose ventrally †Dolichoneura jorelisae Sarto i Monteys, Hausmann, Baixeras and (arrowhead), seen on the left antenna, sectioned in the amber surface. (B) Cilia Peñalver, sp. n. shorten progressively as they approach the distal end, where the antennae look Figs. 1–7. serrate (arrowhead). Both images are Z-stacked. LSID:urn:lsid:zoobank.org:act:DC6F5F7F-2CAA-4955-B78E- 26CBEEEB59BB the same location from where the piece including the moth was collected. 3.2. Diagnosis (male) A tiny bit was removed from the amber sample for FTIR analysis at one edge. The latter was conducted using an IR PerkinElmer Frontier The species can be distinguished from other species of the genus by spectrometer that utilizes a diamond ATR system with a temperature the following combination of wing characters: pointed forewing apexes, stabilized DTGS detector and a CsI beam splitter at the Molecular relatively straight forewing termen, a well-developed double areole and Spectrometry Unit of the CCiTUB (Centres Científics i Tecnològics of the absence of anastomosis between Sc + R1 and Rs in the forewing, the size Universitat de Barcelona). To avoid breaking the piece of amber con­ of the forewing discal cell relatively longer than in other species of the taining the moth, the FTIR was performed using a Thermo SCIENTIFIC genus, the point where M2 arises from the forewing discal cell relatively NICOLET iN10 MX, reflexion, detector MCT, overture 100μ, accumula­ closer to M1 than to M3 in comparison to other species of the genus, an tion 64 scans, spectral resolution 4 cm− 1, Kramers-Kronic. evenly rounded hindwing termen, and a rather weak but tubular M2 vein in the hindwing. 2.2. Stereomicroscopy and photography Holotype: Specimen code MNHNSD FOS 16.01, male (Figs. 1–7). Repository: Museo Nacional de Historia Natural “Prof. Eugenio de The specimen was examined and drawn by using an optical micro­ Jesús Marcano”, Santo Domingo, Dominican Republic. scope Olympus BH-2 equipped with an Olympus camera lucida, and Etymology: The specific name is dedicated to Ms. Jorelis Caridad, photomicrographs were taken using a digital camera (Leica M80 Cam­ daugther of Mr. Jorge Caridad, who entrusted us with this piece. She is era) attached to a Leica IC90 E stereomicroscope, at the laboratory of the well-known for her dedication to preserving and promoting the study of Museu Valencià d’Història Natural (Alginet, València). These devices Dominican amber. 3 V. Sarto i Monteys et al. Journal of South American Earth Sciences 120 (2022) 104055 Fig. 4. Tibiae of †Dolichoneura jorelisae sp. n. (Geometridae: Desmobathrinae), holotype. (A) Camera lucida drawings of left tibiae; note that mid and hindtibiae show spurs not present in foretibia. (B) Left foretibia. (C) Left midtibial spurs. (D) Left hindtibia. (E) Left hindtibia: detail of hair-pencil; also two unequal spurs projecting from its central part. (F) Left hindtibia: two unequal spurs projecting from its distal part. Images D and F are Z-stacked. Locus typicus: Amber mine of Hato Mayor-El Valle Region, forewing venation fairly complete with characteristic double areole Dominican Republic. involving distal part of Sc, basal half of R1 and base of Rs, R1 from the Stratum typicum: The piece was found in stratigraphic levels of the areole, R2 and R3+R4 connate in the areole, R4 to costa, R5 from areole to Yanigua Formation, in the boundary between Early–Middle Miocene, termen; discal cell 0.4 the length of the wing, discocellular vein com­ 17–16 Mya (Iturralde-Vinent and MacPhee, 2019). plete, M1, M2 and M3 approximately equidistant at base; CuA1 closer at base to M3 than to CuA2; hindwing venation unmodified, Sc + R1 and Rs 3.3. Description. Male unanastomosed, Sc + R1 rather straight to apex, Rs sinuous on discal cell, M1, M2, M3 rather equidistant on the discal cell, M2 tubular Head: Vertex and frons scaled. Eyes well developed. Antennae although less developed than M1 and M3 (Fig. 6), base of CuA1 closer to dentate, scaled dorsally, densely ciliate-setose ventrally, length of cilia M3 than to CuA2. at basal part of antennae ca. three times the width of flagellum (Fig. 2); Abdomen: Long, slender, covered by appressed scales. Male geni­ cilia shorten progressively as they approach the distal end, where the talia with valvae closed, well delimited externally but structural details antennae look serrate (Fig. 3B). Ocelli not visible. Chaetosemata pre­ poorly visible, uncus apparently robust, prominent, valva elongate with sent, paired. Labial palpi upturned, not extending dorsally beyond frons, rounded edges (Fig. 7). first and second segments about the same length, the second being slender, third segment small, about one fourth the length of the first two 3.4. Infrared absorption spectrum of amber piece studied segments. Maxillary palpi not visible. Haustellum well developed, ca. 4.5 mm in length, coiled, unscaled, with distal part broken and kept next The infrared absorption spectrum of the amber piece that includes to the main part (Fig. 2). the moth is shown in Fig. 8 (in red), where the basic skeleton of amber is Thorax: Scaled. Legs elongate; foretibia without tibial spurs, but an aliphatic structure. It is dominated by an asymmetric telescopic vi­ showing a conspicuous single process arising from its centre; spur for­ bration of ν (CH2) located at 2928 cm− 1 (small C–H stretching band), mula 0-2-4, pairs of spurs unequal; hindtibia with well developed hair- symmetric telescopic vibration of ν (CH2) located at 2865 cm− 1, trans­ pencil, partially retractable in a groove (Fig. 4). formative vibration of δ (CH2, CH3) located at 1465 cm− 1 (intense C–H Wings. Forewing length: 15.7 mm, with pointed apexes; no trace of band), the symmetric transformative vibration of δ (CH2, CH3) located wing color pattern (Fig. 5). Costa slightly convex at base, central portion at 1380 cm− 1, and telescopic vibration of ν (C=O) located at 1725 cm− 1 rather straight, distal part convexly curved from the point where R2 (intense carbonyl band), and telescopic vibration of ν (C–O) located at meets the costa to pointed apex, termen straight, tornus broadly curved; 1245 cm− 1, 1135 cm− 1, and 1030 cm− 1. Hydroxyl bands near 3500 4 V. Sarto i Monteys et al. Journal of South American Earth Sciences 120 (2022) 104055 Fig. 5. Wing venation and preserved scale pattern of †Dolichoneura jorelisae sp. n. (Geometridae: Desmobathrinae), holotype. (A) Camera lucida drawing of fore- and hindwing. Note the absence of anastomosis between Sc + R1 and Rs on the hindwing. (B) Same, showing detail of anterior venation and scale pattern of forewing. (C) Detail of scale pattern preserved on forewing. (D) Costal part of left forewing (underside). (E) Detail of left fore- and hindwing (underside) partially overlapped. (F) Detail of the well-developed double areole on forewing. Images D and F are Z-stacked. 5 V. Sarto i Monteys et al. Journal of South American Earth Sciences 120 (2022) 104055 in the posterodistal margin of the left hindwing, the wings are quite well preserved. We may conjecture that the moth became trapped in resin shortly after emerging and was rapidly and entirely covered by subse­ quent resin flows. Yet, their venation, usually of supraspecific value, is reasonably well marked and allows, in some detail, a taxonomical assignment. The most striking venation characters of this fossil moth are the presence of a well-developed double areole in the forewing, and the absence of anastomosis between Sc + R1 and Rs, as well as a rather weak but tubular M2 vein in the hindwing. A forewing areole is common in some subfamilies of Geometridae, mostly Orthostixinae, Desmoba­ thrinae, Larentiinae and Sterrhinae (Hausmann, 2001). The areole is formed by a variable degree of anastomosis of R1 and Rs veins, occa­ sionally involving Sc. In this fossil, the areole is clearly affecting distally Sc as well as basally R1 and Rs veins. Although Sc and R veins meet the wing edge separately, all of them arise from the areole. In addition, the Fig. 6. Virtual representation (SRμCT) showing hindwing venation of Doli­ areole is subdivided by a cross vein between basal Rs and R1 giving rise choneura jorelisae sp. n. (Geometridae: Desmobathrinae), holotype. Note that M2 to two areoles. The presence of a tubular M2 in the hindwing is a is tubular although less developed than M1 and M3. generalized feature in the above-mentioned subfamilies, but the absence of anastomosis between Sc + R1 and Rs rules out our fossil moth from being a Larentiinae member. Holloway (1996) included representatives of slender-bodied moths with long and narrow appendages in the sub­ family Desmobathrinae. Pointed forewing apex, among other wing shape traits, as present in our fossil, are common among representatives of this subfamily. Examination of this subfamily representatives, and especially of the genus Dolichoneura Warren, 1894 (e.g., D. oxypteraria (Guenée, 1858)), revealed striking similarities in venation and wing shape with our fossil. Indeed, we compared the wing venation of the type species of the genus Dolichoneura, D. oxypteraria, with that of our fossil. Minor differences are found in fore- and hindwing venations. These are (1) the relative size of the forewing discal cell (being longer and narrower in D. oxypteraria vs. shorter and broader in D. jorelisae), (2) the relative size of the hindwing discal cell (being shorter and broader in D. oxypteraria vs. longer and somewhat narrower in D. jorelisae), and (3) the point (in both, fore- and hindwings) where M2 arises from the discal cell (relatively closer to M1 than to M3 in D. oxypteraria vs. equidistant in D. jorelisae) (Fig. 9). There is also a coincidence between D. jorelisae and other Dolichoneura spp. concerning the spur number in legs and antennal features (dentate flagellum, densely ciliate-setose in males, Figs. 2 and 3). Fig. 7. Micrograph showing the male genitalia, delimited by white dots Although the monophyly of Desmobathrinae has been challenged by (ventral view), of †Dolichoneura jorelisae sp. n. (Geometridae: Desmobathrinae), recent multigene analysis (Murillo-Ramos et al., 2019, 2021), the holotype. Z-stacked image. phylogenetic position of Dolichoneura remains unquestionable. The similarity between our fossil and the extant representatives of the genus cm− 1 are also present. The spectrum of the piece that contains the Dolichoneura is so high that the description of a new genus does not seem geometrid is very similar to those from two geological Dominican amber appropriate. Actually, taking into account the relatively little antiquity localities. These are El Valle-Siete Cañadas (2018 own extraction in the of Dominican amber (19–16 Mya), there is no surprise that arthropods Eastern district of the island, in Iturralde-Vinent and MacPhee, 2019, in preserved in it have been assigned to extant genera several times and blue) and Palo Quemado (2019 own extraction in the Northern district, sometimes even to living species (e.g., Arillo and Ortuño, 2005; in green), their main difference being the presence of a small band near Hoernschemeyer et al., 2010). Based on this evidence, the former 1200 cm-1 in the latter. paleoenvironment is considered to be not so different from a typical extant lowland Neotropical rainforest (Grimaldi, 1996). Therefore, we 4. Discussion conclude that, based on wing shape and venation, our amber geometrid is congeneric with Dolichoneura. As usual in the study of fossil Lepidoptera, characters of the genitalia, According to Iturralde-vinent and MacPhee (2019) the main which are of most interest for a suitable identification, are not accessible amberiferous deposits in the Dominican Republic were formed in a despite 3D preservation in amber and having explored the structure single sedimentary basin in the boundary between Early and Middle through SRμ-CT. Miocene, 19–16 Mya, due to accumulation of abundant resin produced The wings show pointed forewing apex, relatively straight forewing by an extinct legume tree, Hymenaea protera Poinar, 1991 (Legumino­ termen and an evenly rounded hindwing termen, rather unusual wing sae). The genus Hymenaea consists today of fourteen extant species, all shape characters that deserve some attention. Except for minor damage but one native to the tropics of the Americas, with one additional species 6 V. Sarto i Monteys et al. Journal of South American Earth Sciences 120 (2022) 104055 Fig. 8. Infrared absorption spectrum of the amber piece containing the moth (red line) from Hato Mayor-El Valle and spectra of two geological samples from El Valle- Siete Cañadas (blue line) and Palo Quemado (green line), for comparison. (Hymenaea verrucosa Gaertner, 1791) on the east coast of Africa and Lepidoptera, the fossil record of Geometridae is comparatively very Madagascar, considered sister to all other Neotropical species (Pinto scarce, with only 19 findings formally reported worldwide, most of them et al., 2018). Although, so far, fossils of Dolichoneura or Desmobathrinae of difficult systematic assignment, so that this fossil is of remarkable have not been reported anywhere from the Miocene, our finding of interest for phylogenetic studies. It belongs to the genus Dolichoneura yDolichoneura jorelisae sp. n. evidences that they were already present at (Geometridae: Desmobathrinae) and is named Dolichoneura jorelisae this time in Central America. Sarto i Monteys, Hausmann, Baixeras and Peñalver sp. n., based on wing The genus Dolichoneura, in which the fossil herein has been classi­ features. Furthermore, it could also serve for calibrating molecular fied, includes eleven extant species distributed from Central to South clocks and for paleobiogeographic inferences. The head and wings of America (between Costa Rica and Argentina) (Scoble and Hausmann, this specimen were analyzed through SRμCT which provided superb 2007; GBIF Secretariat, 2021). The sister genus Pycnoneura Warren, virtual representations. 1894 includes ten species exclusively distributed in South America (Scoble and Hausmann, 2007). Prout (1932) stated that both genera CRediT authorship contribution statement might have to be merged into one. The biology of Dolichoneura is poorly known; all species are nocturnal and have been collected in well sunny Victor Sarto i Monteys: Conceptualization, Investigation, Writing – slope forest and ridge forest in the lowland rainforest of Costa Rica, original draft. Axel Hausmann: Conceptualization, Investigation, where the resiniferous legume trees Hymenaea courbaril Linnaeus, 1753 Writing – review & editing. Mónica M. Solórzano-Kraemer: Data and H. osanigraseminae Aguilar Fernández, Poveda Álvarez and Santa­ curation, Funding acquisition, Investigation, Methodology, Writing – maría-Aguilar, 2018 live today, but not in creek forest (Rabl, 2016). Two review & editing. Jörg U. Hammel: Methodology. Joaquín Baixeras: larvae of an unidentified Dolichoneura species were collected by canopy Investigation, Writing – review & editing. Xavier Delclòs: Funding fogging a Pouteria caimito (Ruiz & Pav.) Radlk, 1882 (Sapotaceae) tree in acquisition, Investigation, Writing – review & editing. Enrique Peñal­ Peru; molecular gut content analysis suggested this tree to be the feeding ver: Conceptualization, Investigation, Writing – review & editing. plant. Declaration of competing interest 5. Conclusions The authors declare that they have no known competing financial We report a fossil geometrid moth, a male, virtually complete, pre­ interests or personal relationships that could have appeared to influence served in a clear piece of Miocene Dominican amber dating from 19 to the work reported in this paper. 16 Mya. Despite of being one of the most specious family groups of 7 V. Sarto i Monteys et al. Journal of South American Earth Sciences 120 (2022) 104055 Fig. 9. Dolichoneura oxypteraria (Guenée, 1858). (A) male, from French Guyana, Cayenne, Rte. Regina, PK 32.5, Piste de Nancibo, 8.II.1986, ex coll. Herbulot; wingspan 29 mm, forewing length 15 mm. (B) Live specimen of Dolichoneura spp. from Canelos, Pastaza, Ecuador, Eerika Schulz, 15.02.2020, Source iNaturalist, http://www.inaturalist.org/pages/privacy. (C) Close-up of head, proboscis comparatively short, labial palpi strongly curved, length equal to diameter of eye. (D) Close-up of male antenna, slightly dentate, ciliate-fasciculate. (E) Forewing venation: typical characters are the double areole with R5 arising from second areole; Hindwing venation: typical characters are the lack of anastomosis between Sc + R1 and Rs and the full number of M-veins (both characters pointing to Desmobathrinae). Data availability Natural “Prof. Eugenio de Jesús Marcano”) for including it in the collection. Robin Kunz (Senckenberg Research Institute, Frankfurt) Data will be made available on request. supported us with the segmentation of the Synchrotron data. Dr. Alberto Martínez Ortí, director of the Museu Valencià d’Història Natural (Algi­ Acknowledgements net, València, Spain), facilitated the use of the optical microscope and stereomicroscope of that scientific institution. We thank the Spanish We thank Mr. Jorge Caridad (World Amber Museum, Santo Domi­ Ministry of Economy and Competitiveness (project “CRE”, Spanish AEI/ ngo, Dominican Republic) and his family for the loan of the piece FEDER, UE CGL 2017-84419) and VolkswagenStiftung (project 90946) MNHNSD FOS 16.01. for studying purposes, and for its donation to the for financial support. Funding sources were not involved in study design, Museo Nacional de Historia Natural “Prof. Eugenio de Jesús Marcano”, analysis and interpretation of data, the writing of the report and in the Santo Domingo. We thank Carlos Suriel (Museo Nacional de Historia decision to submit the article for publication. 8 V. Sarto i Monteys et al. 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