A third fossil argid sawfly (Hymenoptera, Tenthredinoidea) from the Upper Miocene of Ardèche, southern France

Directeur De la publication / Publication director : Gilles Bloch Président du Muséum national d’Histoire naturelle réDactrice en chef / editor-in-chief : Laure Desutter-Grandcolas assistante De réDaction / assistant editor : Anne Mabille ([email protected]) Mise en page / Page layout : Anne Mabille coMité scientifique / scientific board : Nesrine Akkari (Naturhistorisches Museum, Vienne, Autriche) Maria Marta Cigliano (Museo de La Plata, La Plata, Argentine) Serge Gofas (Universidad de Málaga, Málaga, Espagne) Sylvain Hugel (CNRS, Université de Strasbourg, France) Marco Isaia (Università degli Studi di Torino, Turin, Italie) Rafael Marquez (CSIC, Madrid, Espagne) Jose Christopher E. Mendoza (Lee Kong Chian Natural History Museum, Singapour) Annemarie Ohler (MNHN, Paris, France) Jean-Yves Rasplus (INRA, Montferrier-sur-Lez, France) Wanda M. Weiner (Polish Academy of Sciences, Cracovie, Pologne) couverture / cover : Photograph of general habitus of Spinarge miocenicus n. sp., holotype MDA.R2024.8.1. 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Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / https://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (imprimé / print) : 1280-9551/ ISSN (électronique / electronic) : 1638-9387 765 ZOOSYSTEMA • 2025 • 47 (32) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com urn:lsid:zoobank.org:pub:34C90F03-47B9-4C51-809C-79109E5239C3 Ngô Muller V., Riou B. & Nel A. 2025. — A third fossil argid sawfly (Hymenoptera, Tenthredinoidea) from the Upper Miocene of Ardèche, southern France. Zoosystema 47 (32): 765-772. https://doi.org/10.5252/zoosystema2025v47a32. http://zoosystema.com/47/32 ABSTRACT Spinarge miocenicus n.sp., first fossil representative of an extant argine genus, is described from the Upper Miocene lacustrine maar of montagne d’Andance (Coiron Massif, Ardèche, France). The extant species of Spinarge Wei, 1998 are distributed from Japan, China, Korea, and eastern Siberia to Europe and live on broad-leaved trees, which are well represented in this paleoflora. During the Late Miocene the area of the Coiron Massif was probably quite favorable for Argidae Konow, 1890 as two of the three described fossil argids also originate from this area and period. RÉSUMÉ Un troisième fossile de mouche-à-scie argide (Hymenoptera, Tenthredinoidea) du Miocène supérieur d’Ardèche, Sud de la France. Spinarge miocenicus n.sp., premier représentant fossile d’un genre actuel d’Arginae Konow, 1890, est décrit du maar lacustre du Miocène supérieur de la montagne d’Andance (massif du Coiron, Ardèche, France). Les espèces actuelles de Spinarge Wei, 1998 sont réparties du Japon, de la Chine, de la Corée et de la Sibérie orientale jusqu’en Europe et vivent sur des feuillus, qui sont bien représentés dans cette paléoflore. Au cours du Miocène supérieur, la zone du massif du Coiron était probablement assez favorable aux Argidae Konow, 1890, car deux des trois argides fossiles décrits sont également originaires de cette zone et de cette période. Valérie NGÔ MULLER Faculté des Sciences, UFR Sciences du Vivant, Université Paris-Cité, Paris (France); and Institut de Systématique, Évolution, Biodiversité (ISYEB), Muséum national d’Histoire naturelle, CNRS, Sorbonne Université, EPHE-PSL, Université des Antilles, 57 rue Cuvier, CP 50, F-75231 Paris cedex 05 (France) [email protected] Bernard RIOU Muséum de l’Ardèche, 344 Chemin de Cournasou F-07120 Balazuc (France) [email protected] André NEL Institut de Systématique, Évolution, Biodiversité (ISYEB), Muséum national d’Histoire naturelle, CNRS, Sorbonne Université, EPHE-PSL, Université des Antilles, 57 rue Cuvier, CP 50, F-75231 Paris cedex 05 (France) [email protected] Submitted on 22 December 2024 | Accepted on 17 April 2025 | Published on 10 December 2025 A third fossil argid sawfly (Hymenoptera, Tenthredinoidea) from the Upper Miocene of Ardèche, southern France MOTS CLÉS Insecta, ‘Symphyta’, Arginae, Spinarge, Ardèche, Sud de la France, Cénozoïque, espèce nouvelle. KEY WORDS Insecta, ‘Symphyta’, Arginae, Spinarge, Ardèche, southern France, Cenozoic, new species. 766 ZOOSYSTEMA • 2025 • 47 (32) Ngô Muller V. et al. INTRODUCTION With approximately 913 species and 60 genera, Argidae Konow, 1890 are the second-most diverse family of the symphytan grade of Hymenoptera Linnaeus, 1758, lagging behind Tenthredinidae Latreille, 1818 (Taeger etal. 2018). This is paralleled in their known fossil occurrences, with many more fossil tenthredinids known than for Argidae. Only two fossil argid species are currently described, viz. Mioarge azari Nel, 2004 (Arginae Konow, 1890: Upper Miocene of MontCharay, Ardèche, France) and Sterictiphora konowi Rohwer, 1908 (Sterictiphorinae Konow, 1890: Eocene-Oligocene boundary, Florissant Formation, Colorado, USA). Douglas (1991: fig.V-18) and Douglas& Stockey (1996) figured an incomplete forewing they attributed to ‘Argidae/Diprionidae’ from the middle Eocene of the USA. Lastly, Smith& Poinar (1992) recorded an argid larva in Early Miocene amber from the Dominican Republic. Here we describe a third adult of a fossil argid sawfly from the Upper Miocene of montagne d’Andance (Ardèche, France). It is also the first known fossil representative of an extant genus of Arginae. MATERIAL AND METHODS The holotype of Spinarge miocenicus n.sp. originates from the Miocene KonservatLagerstätte of the montagne d’Andance (Saint-Bauzile, Ardèche, France). The montagne d’Andance (altitude 552 m) is located on the site of an ancient maar dating from the Miocene epoch and linked to the Coiron basalt complex. The diatomitic unit from which the specimen originated was deposited in a lake formed by the maar crater (Demarcq etal. 1989), under a warm and humid climate (Brice 1965), with a possibly seasonal character. This lake was probably calm and shallow (Ehrlich 1966), surrounded by a deciduous forest with an undergrowth of ferns (Grangeon 1958; Riou 1988). Over time, this lake was filled with diatomite, forming rock strata up to 30m thick in some localities (Serieyssol& Gasse 1991). This diatomite is a soft siliceous rock in which were preserved the insects that drowned in the lake at the time. These stratigraphic layers of diatomite represent a reducing environment that has allowed a remarkable conservation of mummified organic matter. The age of this deposit corresponds to the late Tortonian, 7.54 ± 0.06 Mya (Pastre etal. 2004). The new fossil was collected by two of us (VNM and BR) during fieldwork in summer 2024, with the authorization of Chemviron France Company, owner of the Quarry of montagne d’Andance. The fossil was fixed using wallpaper glue. Two extant representatives of the genus were observed for comparison with the fossil: – Arge meliosmae Shinohara& Hara, 2011, MNHN.F.EY38826 (Fig. 3); – Spinarge (Arge) metallica Klug, 1834, MNHN.F.EY42476 (Fig. 4). Specimens were observed using a Leica M205C stereomicroscope and photographed using a Sony alpha 7RIV camera. Photographs were exported in NEF format and processed with Nikon NX Studio software. All images are digitally stacked photomicrographic composites of several individual focal planes that were obtained using Helicon Focus 6.7 software. We follow the list of genera of Argidae of Taeger etal. (2010, 2018), and the wing venation nomenclature was adapted from Goulet& Hubert (1993). AbbreviAtions Mya Million years ago; MDA Muséum de l’Ardèche, Balazuc; MNHN Muséum national d’Histoire naturelle, Paris. Morphology A anal vein; C costal vein; Cu cubital vein; cu-a transverse between Cu and A; M medial vein; m-cu transverse between M and Cu; R radial vein; Rs radial sector vein; rs-m transverse between Rs and M; Sc intercostal vein. SYSTEMATICS Class INSECTA Linnaeus, 1758 Order HYMENOPTERA Linnaeus, 1758 Family ArgidAe Konow, 1890 Subfamily ArginAe Konow, 1890 Genus Spinarge Wei, 1998 Spinarge miocenicus n.sp. (Figs1; 2) urn:lsid:zoobank.org:pub:34C90F03-47B9-4C51-809C-79109E5239C3 type mAteriAl.— Holotype. France • body rather well-preserved with thorax, abdomen, one antenna, two complete forewings and partly preserved hind wings; Upper Miocene; montagne d’Andance paleomaar, diatomites, Saint-Bauzile, Ardèche; MDA.R2024.8.1. type strAtigrAphy And locAlity.— Upper Miocene (7.54Mya), montagne d’Andance paleomaar, diatomites, Saint-Bauzile, F-07210 Ardèche, France. diAgnosis.— In addition to the characters of the genus (see Discussion), third antennomere simple, elongate and slightly curved, 6.3× as long as wide; foreand hind wings brown in basal half, more yellow in distal half. description Head hidden in matrix; third antennomere elongate and slightly curved, 1.9mm long, 0.3mm wide. Thorax 3.4mm long, 1.9mm wide; mesoscutellar appendage fused to mesoscutellum. Forewing complete, 8.8mm long, 3.1mm wide; brown in basal half, more yellow in distal half; short anterior 767 A third fossil argid sawfly from the Upper Miocene of Ardèche ZOOSYSTEMA • 2025 • 47 (32) Fig. 1. — Spinarge miocenicus n. sp., holotype MDA.R2024.8.1. Photograph of general habitus. White arrow, median process on the 5th tergum, poorly preserved. Scale bar: 2 mm. Sc branch (intercostal vein) located slightly anteriad Sc+R and 1M junction; pterostigma 1.5mm long, 0.4mm wide, strongly arched along cell 2R1, slightly arched along anterior wing margin; M+Cu slightly curved; 1M slightly longer than 1Cu, curved basally; R clearly deviating between junctions of Sc and 1M; fusion of R with M exceedingly short; R meeting 2M much less than halfway between short anterior branch of Sc and pterostigma; 2M slightly longer than 1rs-m; Rs emerging at pterostigmal base; 2Rs strongly curved, shorter than 3Rs; no r-rs crossvein; cell 2R1 with a distal abscissa ending in wing margin; cell 1R1 trapezoidal, 0.5mm long; cell 1Rs 1.2mm long, enclosed distally by subvertical 2rs-m (the latter slightly longer than 1rs-m); 3M shorter than 4M; cell 2Rs 1.5mm long, distinctly wider along curved 3rs-m; 1Cu and 2Cu nearly straight, 1Cu as long as 2Cu; lm-cu meeting Cu at an angle of ca 90°, 1M and 1m-cu converging; cell 1Cu elongate, longer than cell R; 1cu-a (nervulus) located at middle of cell 1M, distinctly distad point of fusion of R with 1M; cell 2Cu long, trapezoidal; cell 2M fully enclosed distally by long and slightly sinusoidal 2m-cu; 1A long, nearly straight; basal anal cell 1A distally closed; a long fusion of 2A+3A with 1A; cell 2A quite long. Hind wings only partly visible, 6.8 mm long, brown in basal half and more yellow in distal half; cell R1 with a distal abscissa not ending in wing margin; cell Rs distinctly longer than cell 1M, both distally closed. Abdomen 5.7mm long, 2.8mm wide; fifth tergum with a long median process; genitalia not visible. DISCUSSION Following the key to symphytan families of Goulet& Hubert (1993), the current fossil falls within the Argidae sensu lato because of the following characters: mesoscutellar appendage fused to mesoscutellum; forewing without 2r-rs; forewing base of 2A+3A present; antenna with one flagellomere. In addition, the forewing venation is quite similar to those of extant Argidae (Fig.3: Arge meliosmae Shinohara& Hara, 2011). Malagón-Aldana etal. (2021) separated the genus Zenarge Rohwer, 1918 from other Argidae, elevating the Zenarginae Rohwer, 1918 as family Zenargidae Rohwer, 1918. The new fossil differs from Zenarge in a shorter 2A+3A, 768 ZOOSYSTEMA • 2025 • 47 (32) Ngô Muller V. et al. Fig. 2. — Spinarge miocenicus n. sp., holotype MDA.R2024.8.1: A, photograph of third antennomere (note that part of the antennomere is missing) and forewing; B, with nervation indications. Abbreviations: see Material and methods. Scale bar: 2 mm. distal part of Rs as long as Rs+M (vs quite shorter), Rs+M making a strong angle with cell R1 (Malagón-Aldana etal. 2021: fig.5). The Argidae sensu stricto are currently divided into Arginae Konow, 1890, Athermantinae Benson, 1938, Atomocerinae Benson, 1938, Dielocerinae Benson, 1938, Erigleninae Rohwer, 1911, and Sterictiphorinae Rohwer, 1911 (Taeger etal. 2010, 2018). The Erigleninae, Atomacerinae, and Sterictiphorinae have the forewing cell 2R1 and the R1 of hind wing open apically, unlike the new fossil (Benson 1938; Malaise 1937, 1955; Smith 1969). The presence of the intercostal vein also excludes affinities with the Sterictiphorinae (Benson 1938, third antennomere A B pterostigma Rs R C Sc+R M+Cu M R+M Sc M 3rs-m 2rs-m 2m-cu 1A 1m-cu 1cu-a 3Rs 2Rs 1Rs 1R1 1M 1Cu 2Cu 2M 2R1 3M 2 A + 3 A 769 A third fossil argid sawfly from the Upper Miocene of Ardèche ZOOSYSTEMA • 2025 • 47 (32) Fig. 3. — Arge meliosmae Shinohara & Hara, 2011, MNHN.F.EY38826. Photograph of general habitus in dorsal view, together with the cocoon from which the imago emerged. Scale bar: 1 mm. Smith 1969; Togashi 1990, 1997). The Neotropical Dielocerinae also have the hind wing cell R1 opened (Fernández& Sharkey 2006). Benson (1938) first separated the Athermantinae from the Arginae on the basis of the presence vs absence of preapical spurs on the mesoand metatibiae (but see Liston etal., 2017). Both groups share with the new fossil the presence of the intercostal vein and the forewing radial cell closed apically. In the Athermantinae, Pampsilota Konow, 1899has nearly the same forewing venation as the new fossil (Koch 2006; Liston etal. 2017), as do the genera Athermantus Kirby, 1882 and Pseudosinarge Saini, Thind& Kaur, 1998 (Wei 1997; Saini etal. 1998). Pampsilota and Athermantus have a third antennomere much longer compared to its width than in the fossil (Liston etal. 2017; Luo etal. 2019). Athermantus also has the forewing cell 2R1 distinctly broadened basally, not less than half breadth of cell 2R1, unlike in the new fossil. Pseudosinarge differs from the new fossil in the much longer and thin third antennomere, point of fusion of R with 1M situated opposite 1cu-a, and an abscissa of forewing cell 2R1. Cibdela Konow, 1899 also has R+M as long as or longer than the anterior branch of Sc. Sinarge Forsius, 1934 and Tanyphatnidea Rohwer, 1912have R+M punctiform, which is unlike the new fossil. Within the Arginae, the tribes Argini Konow, 1890 and Scobinini Benson 1963 are separated on the basis of characters not visible in this fossil (leg and head structures) (Benson 1963). In the Scobinini, the Australian genus Antargidium Morice, 1919 is characterized by characters not visible in the new fossil (Benson, 1963), and they share an open appendiculate cell in hind wing (Benson 1934). Nevertheless, Antargidium has a third antennomere only twice as long as broad (Benson 1934). Scobina Lepeletier de Saint Fargeau& Audinet-Serville, 1828 differs from the new fossil in the exceedingly weak angle between R1 and R+M (Smith 2008: figs1-2). 770 ZOOSYSTEMA • 2025 • 47 (32) Ngô Muller V. et al. Following the key to Palaearctic and Oriental genera of Argidae (Wei 1997: 296-297), the new fossil would fall in the Argini because of the following characters: short anterior SC branch (intercostal vein) present; forewing and hind wing cells R1 closed with an appendiculate cell. In this group affinities with Alloscenia Enderlein, 1919 are excluded because the anterior branch of Sc is well defined, not faint. Kokujewia Konow, 1902has the forewing 1rs-m absent and the basal anal cell open, as in Asiarge Gussakovskij, 1935. Sjoestedtia Konow, 1907has the forewing basal anal cell absent (Liston etal. 2017). Triarge Forsius, 1931has the forewing 2rs-m absent (Koch, 2006; Liston etal. 2017). Pseudarge Gussakovskij, 1935has the forewing basal anal cell open distally and R+M distinctly longer than in the new fossil (Koch& Goerger 2008). Arge Schrank, 1802 (Calarge Enslin, 1911 is a junior synonym of Arge after Liston etal. (2017)) and Clyparge Pasteels, 1963have very similar wing venation (Pasteels 1963). The new fossil has a forewing venation similar to those of the species in Arge (Choi etal. 2016). Spinarge Wei, 1998 shares with the new fossil R+M shorter than the anterior branch of Sc and the venation of the new fossil is quite similar to those of the species in this genus (Liu etal. 2022). Furthermore, the new fossil has a long median process on the fifth tergum (Fig.1), typical of males in the genus Spinarge (Hara& Shinohara 2006: fig.7). Liu etal. (2022: 231) indicated that ‘[t] he validity of this genus needs further study on the generic phylogeny of Arginae’. Spinarge seems to be close to Arge. The different species of Spinarge are separated on the basis of wing coloration, viz. yellowish orange in S.chrysoptera vs hyaline, dark, yellowish or brownish in the other species (Hara& Shinohara 2006). The new fossil has brown-yellow wings, but their present coloration may be different from that of the living animal during the Miocene. It is thus nearly impossible to specify its exact affinities in the genus. Extant representatives of the genus Spinarge include three chinese species and 11 palearctic species. The only species reported in France (South of France, Ariège) is Spinarge metallica Klug, 1834 (Hara& Shinohara 2006; Noblecourt 2020), a country-wise rare species (Fig.4). Species of Spinarge live on broad-leaved trees such as Betula L., 1753, Prunus L., 1753, Sorbus L., 1753, Tilia L., 1753 (Hara& Shinohara 2006; Savina& Chevin 2012; Liu etal. 2022). The yellow larvae of Spinarge metallica were observed in two localities (Aulus-les-Bains and Bethmale, Ariège piedmont) by Henri Savina on July 2010, and were feeding on Betula (Savina& Chevin, 2012). At least Betula and Tilia have been recorded from the outcrop of Saint-Bauzile (Brice 1965; Riou 1996), representing potential host plants for the fossil species occurring in this paleobiota. Fig. 4. — Spinarge metallica Klug, 1834, MNHN.F.EY42476. Photograph of general habitus of female in dorsal view. Third antennomeres and right forewing missing. Scale bar: 1mm. 771 A third fossil argid sawfly from the Upper Miocene of Ardèche ZOOSYSTEMA • 2025 • 47 (32) CONCLUSION Even though several extant genera in Arginae and Athermantinae have wing venations similar to the new fossil, we could discriminate them and attribute the extinct species to Spinarge on the basis of the structure of the abdominal tergum. Although we lack significant characters to separate the fossil from the extant species in this genus, we propose to name it because it is the first known fossil representative of an extant genus of Arginae. Spinarge miocenicus n.sp. is the earliest occurrence of the genus Spinarge, 7.54 Mya, and may serve as a calibration point for phylogenetic analyses. It is rather remarkable that the new fossil comes from an outcrop quite close to that from where Mioarge azari was recovered. The diatomite outcrop of montagne d’Andance is dated around 7.54±0.06 Mya (Pastre etal. 2004), while the basaltic Coiron Massif (including the interstratified diatomite of Mont-Charay) would be slightly younger, around 6Mya. The sediments in the two outcrops are quite different. In the Mont-Charay outcrop the diatomite is exceptionally hard and filled with a stacking of numerous angiosperm leaves, while at the montagne d’Andance outcrop the matrix is much softer and there is no such stacking of leaves. The localities may correspond to two different paleolakes, though paleoenvironment and paleoclimate were probably similar. It remains puzzling to find so few fossil Argidae in Miocene outcrops. The scarcity of Argidae in the French Miocene outcrops may mirror the scarcity of species diversity among existing argid sawflies in France. Acknowledgements We thank the reviewers for their constructive remarks, which significantly contributed to enhance this work. We also thank M. Jean-Louis Patissier (factory director, Chemviron France Company) and M. Bernard Reboul (quarry manager) for their authorization to collect in the Quarry of montagne d’Andance and for their help with security requirements. Our appreciation also goes to Laure Desutter for her keen advice editing the manuscript and to Antoine Guiguet for providing us access to extant specimens hosted in the MNHN entomological collections. REFERENCES b enson r. b. 1934.— New Australian sawflies. Memoirs of the Queensland Museum 10: 211-229. https://www.biodiversitylibrary.org/page/51236371 benson r. b. 1938. — Sawflies of the subfamily Trichorhachinae (Argidae). Annals and Magazine of Natural History (11) 2: 117-122. benson r. b. 1963.— The affinities of the Australian Argidae (Hymenoptera). Annals and Magazine of Natural History (13)5: 631-635. https://doi.org/10.1080/00222936208651295 b rice d. 1965.— Recherche sur la flore mio-pliocène de la Montagne d’Andance (Coiron, Ardèche). Annales de la Société géologique du Nord 85 (2-3): 189-239. choi J.-K., lee s.-b., Wei m.-c. & lee J.-W. 2016.— Three new species of the genus Arge (Hymenoptera: Symphyta: Argidae) from South Korea with key to species of the subfamily Arginae. Journal of Asia-Pacific Biodiversity 9: 183-193. d emArcq g., m ein p., b Allesio r. & r omAggi J.-p. 1989.— Le gisement d’Andance (Coiron, Ardèche, France) dans le Miocène supérieur de la vallée du Rhône: un essai de corrélation marincontinental. Bulletin de la Société géologique de France (8), 5 (4): 797-806. https://doi.org/10.2113/gssgfbull.V.4.797 douglAs s. d. 1991.— Flower form and pollinator diversity in the Middle Eocene of British Columbia and Washington. M.Sc. thesis, University of Alberta, Edmonton, 342 p. douglAs s. d. & stocKey r. A. 1996.— Insect fossils in middle Eocene deposits from British Columbia and Washington State: faunal diversity and geological range extensions. Canadian Journal of Zoology 74 (6): 1140-1157. https://doi.org/10.1139/z96-126 e nderlein g. 1919.— Symphytologica. i. Zur Kenntnis der Oryssiden und Tenthrediniden. Sitzungsberichte der Gesellschaft Naturforschender zu Berlin 1919: 111-126. https://www.biodiversitylibrary.org/page/10289210 enslin e. 1911.— Über Tenthrediniden aus Africa. Deutsche Entomologische Zeitschrift 55: 657-669. ehrlich A. 1966.— Contribution à l’étude des gisements volcanolacustres à diatomées de la région de Rochessauve et de St Bauzile. Bulletin de la Société Géologique de France (7) 8: 311-321. Fernández F. c. & shArKey m. J. (eds) 2006.— Introducción a los Hymenoptera de la Región Neotropical. Sociedad Colombiana de Entomología y Universidad Nacional de Colombia, Bogotá D.C.: xxx + 894 p. http://hdl.handle.net/20.500.12324/34432 Forsius r. 1931.— Notes on a collection of Ethiopian Oryssoidea and Tenthredinoidea (Hymenoptera). Annals and Magazine of Natural History (10) 8 (43): 1-36. F orsius r. 1934.— New or little known Tenthredinoidea. 1. Notulae Entomologicae 16: 45-52. goulet h. & huber J. t. 1993. — Hymenoptera of the World: an identification guide to families. Research Branch Agriculture Canada Publication, Ottawa, 1894/E: i-vii + 1-668. grAngeon p. 1958.— Contribution à l’étude de la paléontologie végétale du massif du Coiron (Ardèche) (Sud-Est du massif Central Français). Mémoires de la Société d’Histoire naturelle d’Auvergne 6: 1-290. gussAKovsKy v. v. 1935.— Chalastogastra (P. 1), in sernov s. A. & s tAcKelberg A. A. (eds). Insectes, Hyménoptères. Faune de l’URSS, Moscou Leningrad 2 (1): I-XVII + 1-452. hArA h. & shinohArA A. 2006.— The sawfly genus Spinarge (Hymenoptera, Argidae). Bulletin of the National Science Museum of Tokyo (A) 32 (2): 61-94. Kirby W. F. 1882.— List of Hymenoptera with descriptions and figures of the typical specimens in the British Museum. 1. Tenthredinidae and Siricidae Vol. 1. By order of the Trustees, London, 450 p. Koch F. 2006.— A contribution to the sawfly fauna of the winter rainfall area of southern Africa: revision of the genus Triarge Forsius, 1931 with description of a new species of Pampsilota Konow, 1899 (Hymenoptera: Symphyta: Argidae: Arginae, Athermantinae). Mitteilungen aus dem Museum für Naturkunde in Berlin - Zoologische Reihe 82: 223-238. https://doi.org/10.1002/ mmnz.200600008 KonoW F. W. 1890.— Neue paläarctische Blattwespen. Wiener Entomologische Zeitung 9: 8-13. K onoW F. W. 1899.— Einige neue Chalastogastra-Gattungen und Arten. Entomologische Nachrichten 25: 73-79. https://doi. org/10.5281/zenodo.15946810 KonoW F. W. 1902.— Novum genus et novae species Chalastogastrorum Rossiae. Trudy Yaroslavskogo Yestestvenno-Istoricheskogo Obshchestva 1: 1-4. KonoW F. W. 1907.— Neue Argides (Hym.). Zeitschrift für Systematische Hymenopterologie und Dipterologie 7 (4): 306-309. Koch F. & goerge g. 2008.— Annotations to the Palaearctic sawfly genus Pseudarge Gussakovskij, 1935 gen. rev. with a con-