Palaeoentomology 003 (2): 188–195 ISSN 2624-2826 (print edition) https://www.mapress.com/j/pe/ PALAEOENTOMOLOGY Copyright © 2020 Magnolia Press Article ISSN 2624-2834 (online edition) PE https://doi.org/10.11646/palaeoentomology.3.2.8 http://zoobank.org/urn:lsid:zoobank.org:pub:20A34D9A-DC69-453E-9662-0A8FAFA25677

A new fishfly species (Megaloptera: Corydalidae: Chauliodinae) from Eocene Baltic amber

XINGYUE LIU1, * & JÖRG ANSORGE2 1College of Life Science and Technology, Hubei Engineering University, Xiaogan 432000, China �[email protected]; https://orcid.org/0000-0002-9168-0659 2Institute of Geography and Geology, University of Greifswald, Friedrich-Ludwig-Jahnstraße 17a, D-17487 Greifswald, Germany �[email protected]; https://orcid.org/0000-0002-1284-6893 *Corresponding author. �[email protected]

Abstract and Sialidae (alderflies). Species of Megaloptera have worldwide distribution, but most of them occur mainly in The fossil record of Megaloptera (Insecta: Holometabola: subtropical and warm temperate regions, e.g., the Oriental, Neuropterida) is very limited. Both megalopteran families, i.e., Corydalidae and Sialidae, have been found in the Eocene Neotropical, and Australian Regions (Yang & Liu, 2010; Baltic amber, comprising two named species in one genus Liu et al., 2012, 2015a). The phylogeny and biogeography of Corydalidae (Chauliodinae) and four named species in of extant Megaloptera have been intensively studied in two genera of Sialidae. Here we report a new species of Liu et al. (2012, 2015a, b, 2016) and Contreras-Ramos Chauliodinae from the Baltic amber, namely Nigronia (2011). prussia sp. nov.. The new species possesses a spotted hind Compared with the other two orders of Neuropterida wing with broad band-like marking, a well-developed stem (Raphidioptera and Neuroptera), the fossil record of of hind wing MA subdistally with a short crossvein to MP, a Megaloptera is considerably scarce. Thus, the early single straight RP branch separated between 1ra-rp and 2ra- rp in hind wing, and the hind wing A3 with anterior branch evolution of this order is little known. Jepson & Heads proximally touching A2. A tentative placement of the new (2016) presented a comprehensive overview on the fossil species in Nigronia Banks, 1908, which is an extant genus Megaloptera of the world. However, the extinct families endemic to eastern North America, is discussed in detail. Our listed as Megaloptera in Jepson & Heads (2016), i.e., finding provides new evidence indicating that the fishflies Parasialidae and Corydasialidae, have been removed related to the extant species from eastern North America had out of Megaloptera in recent studies (Liu et al., 2017; occurred in Europe during the Early Tertiary. Engel et al., 2018). Parasialidae is considered to be stem group Raphidioptera (Wang et al., 2017; Engel et Keywords: Neuropterida, Nigronia, fossil, Europe, Tertiary al., 2018), while Corydasialidae belongs to Neuroptera, being close to Chrysopoidea (Liu et al., 2017). Thus, the oldest fossil of definite Megaloptera known so far Introduction is Dobbertinia reticulata Handlirsch, 1920 (Sialidae) from the Lower Jurassic of Germany (Ansorge, 2001), The order Megaloptera (dobsonflies, fishflies, and while the oldest fossil of Corydalidae is recorded from alderflies) belongs to the superorder Neuropterida and the Middle Jurassic of China, including two genera and is the second smallest order of Holometabola. Within two species of Chauliodinae (Wang & Zhang, 2010; Liu Neuropterida, adult megalopterans are characterized et al., 2012). Totally, there have been 21 named fossil by the prognathous head and the broad anal area of species of Megaloptera, including nine species in nine hind wing, while their larvae are exclusively aquatic, genera from the Mesozoic and 12 species in six genera campodeiform, with seven or eight pairs of lateral from the Cenozoic (Jepson & Head, 2016; Huang et al., tracheal filaments on elongate abdomen (New & 2016). Among them, eight species in seven genera belong Theischinger, 1993). Currently, there are nearly 380 to Corydalidae, while the remaining 13 species in eight described species of Megaloptera worldwide (Oswald genera belong to Sialidae. & Machado, 2018). Extant Megaloptera consist of only The Eocene Baltic amber is a major source yielding two families, i.e., Corydalidae, with two subfamilies megalopteran fossils. Currently, there are six named Corydalinae (dobsonflies) and Chauliodinae (fishflies), species and two undetermined species respectively of 188 Submitted: 15 Feb. 2020; accepted by D.-Y. Huang: 13 Mar. 2020; published: 30 Apr. 2020 Indosialis Lestage, 1927 and Chauliodes Latreille, 1796 Genus Nigronia Banks, 1908 (Hagen, 1856; Wichard, 1997, 2002, 2003; Wichard & Nigronia Banks, 1908: 30. Engel, 2006; Wichard et al., 2009). Chauliodes is the only known corydalid genus from Baltic amber, with two Type species. Chauliodes serricornis Say, 1824: 304, by named species, i.e., Chauliodes priscus Pictet in Hagen, subsequent designation (van der Weele, 1910: 70). 1856 and Chauliodes carsteni Wichard, 2003. Included species. Two extant species: Nigronia In this paper, we describe a new species of serricornis (Say in Keating, 1824) and Nigronia fasciata Chauliodinae (Corydalidae) from the Eocene Baltic (Walker, 1853) from eastern U.S.A. and eastern Canada. amber, which was first mentioned by Ansorge (2007). One extinct species: Nigronia prussia sp. nov. from The new species is tentatively placed in the extant fishfly Eocene Baltic amber. genus Nigronia Banks, 1908 that is endemic to eastern Diagnostic characters. Adults small to medium- North America, namely Nigronia prussia sp. nov.. Our sized (forewing length 19–28 mm). Body generally finding provides a new case that the ancestor of North blackish brown. Antennae pectinate or subserrate in male, American corydalids had occurred in Europe during the serrate or subserrate in female. Wings blackish brown, Early Tertiary. with some whitish markings, which usually form a whitish band-like marking at middle on forewing in extant species; transparent with dark markings at least in hind wing in Material and methods extinct species, i.e., Nigronia prussia sp. nov.. Forewing: RP with four simple branches, posterior branch proximally The Baltic amber forest grew sometime in the Eocene straight, only one RP branch separated between 1ra-rp in southern Fennoscandia. During the Lutetian (Middle and 2ra-rp crossveins; MA simple; MP with two long Eocene, ca. 45 Ma) the amber was re-deposited into the simple branches; A2 with both branches strongly sinuate, marine silt and clay of the Blue Earth Formation, which and with anterior branch not fused with A1. Hind wing: is mined in the Jantarnyi (Palmnicken) quarry in the Venation mostly same to that of forewing; stem of MA Kaliningrad region of the Russian Federation. During present as a long, slightly sinuate veinlet, subdistally with Neogene and Pleistocene Baltic amber was redeposited a short 1ma-mp crossvein (but sometimes absent); cua- all over the North European plain. Since Baltic amber is cup crossvein nearly vertical to CuA; A3 with anterior also a trading good, the whereabouts of a certain piece are branch proximally connecting A2 by a short crossvein often unknown, as in our case. For a general overview on or a short fusion. Male genitalia: Tergum 9 longer than Baltic amber see Weitschat & Wichard (2002). wide, anteriorly concaved, distinctly produced ventrad, Images of the wing were taken with a Nikon Coolpix posterodorsal portion in lateral view angularly convex; 4500 camera with reflected and translucent light. The sternum 9 posteriorly with a membranous lobe; ectoproct drawing is based on a camera lucida sketch and the short, subquadrate in lateral view, with a feebly prominent photographs. callus cercus; gonocoxites 10 fused into a single sclerite, The examined specimens of extant fishflies with a pair of short lateral arms; gonocoxites 11 fused are deposited in the Entomological Museum, China into an arcuate sclerite beneath anus. Female genitalia: Agricultural University (CAU), Beijing, China, and the Gonocoxites 8 fused into a large plate; gonapophyses Fumio Hayashi collection, Tokyo Metropolitan University, absent; gonocoxite 9 distally with a very tiny gonostylus; Tokyo, Japan. ectoproct subtriangular, not bilobed, with a feebly Abbreviations used for wing veins are as following prominent callus cercus. based on the terminology in Kukalová-Peck & Lawrence (2004): A, anal vein; Cu, cubitus; CuA, cubitus anterior; Nigronia prussia sp. nov. CuP, cubitus posterior; MA, media anterior; MP, media (Figs 1–3) posterior; RA, radius anterior; RP, radius posterior; ScP, subcosta posterior; crossveins in small characters. Type material. Holotype RD 300, German Amber Museum, Ribnitz-Damgarten: Amber piece with a nearly complete hind wing of Nigronia prussia sp. nov., eight sciarids (Diptera: Sciaridae), and five mymarids Systematic palaeontology (Hymenoptera: Mymaridae) as syninclusions. Etymology. The name derives from Latin Prussia, Class Insecta Linnaeus, 1758 the former duchy of Prussia (Herzogtum Preußen) a Order Megaloptera Latreille, 1802 region in the East Baltic, now Kaliningrad (Königsberg) Family Corydalidae Leach, 1815 region of Russian federation, where most of the Baltic Subfamily Chauliodinae van der Weele, 1909

A new fishfly species Palaeoentomology 003 (2) © 2020 Magnolia Press • 189 wide at middle. Largely transparent, but with distinct dark markings; costal space with a small spot present between proximal 7th and 8th costal crossveins, and with two stripes respectively proximad and distad pterostigmal region; proximal half with a broad oblique band, which extends from RA to A1 and is distinctly widened posteriad; distal half with several markings, including a subquadrate marking anteriorly connected with costal stripe proximad pterostigmal region, a short stripe anteriorly connected with costal stripe distad pterostigmal region, a large round spot, a small round spot, and some incompletely preserved markings near wing apex. Venation: Two nygmata present between MA and MP, and an additional nygma present between RP and MA near wing apex; costal space feebly narrowed on pterostigmal region, with 22 crossveins preserved; ScP and RA distally fused with each other; preserved part of RP with two branches and with stem distinctly bended at 1ra-rp; three ra-rp crossveins present; one crossvein (and at least one more crossvein by estimation) present between RP branches; MA stem present as a long, slightly sinuate veinlet between RP and MP, distal part of MA probably simple; three rp-ma crossveins present; MP initially branched slightly proximad midpoint, with two main branches, both probably simple distad; four ma-mp crossveins present, 1ma-mp crossvein anteriorly connected with subdistal point of MA stem; two crossveins present branches of MP; CuA initially branched slightly distad midpoint, leaving two branches, both probably simple distad; CuP simple; three mp-cua crossveins present; one cua-cup crossvein present, nearly vertical to CuA; A1 branched at proximal 1/3, with two long, slightly sinuate, simple branches; A2 simple, slightly sinuate; A3 initially branched near wing base, with two long simple branches, and anterior branch proximally touching A2 at a point and abruptly bended FIGURE 1. Nigronia prussia sp. nov., holotype RD 300. A, posteriad; A4 short, simple, arcuately curved distad; one Photograph of whole amber piece including the new species. B, cup-a1 crossvein present; one a1-a2 crossvein present, Photograph of hind wing under top light. C, Photograph of hind anteriorly close to branching point of A1. wing under bottom light. Remarks. The new species differs from the extant Nigronia species by the mostly transparent hind wing with scattered dark markings and the hind wing a1-a2 amber originates from. The name is treated as a noun in approximating the branching point of A1. In extant species apposition. of Nigronia both fore- and hind wings are largely blackish, Diagnosis. Hind wing with a band-like dark marking with narrow and scattered whitish patches, and the hind on proximal half and several dark markings on distal half, wing a1-a2 is distantly proximad the branching point of including a large round marking at distal 1/3. Hind wing A1. It is also notable that there is an additional nygma costal space feebly narrowed on pterostigmal region; MA between RP and MA near the apex of hind wing in the stem long and slightly sinuate, subdistally with a short new species. This feature is very rare in Chauliodinae and may be just aberrant in certain individuals. We have found 1ma-mp crossvein; a1-a2 crossvein slightly proximad but only one specimen of extant fishflies [Neochauliodes close to branching point of A1. sundaicus (van der Weele, 1906); see Liu et al., 2010: Description. Holotype. Hind wing: Wing apex not fig. 11] that has such similar additional nygma in the hind preserved, preserved part of wing 17 mm long and 7 mm wing.

190 • Palaeoentomology 003 (2) © 2020 Magnolia Press liu & ANSORGE FIGURE 2. Nigronia prussia sp. nov., holotype RD 300, drawing of hind wing. Red arrow indicates single and proximally straight RP branch separated between 1ra-rp and 2ra-rp; blue arrow indicates anterior branch of A3 touching A2.

Discussion and Orohermes Evans, 1984) by the absence of five hind wing nygmata and inclined hind wing cua-cup crossvein. Although Nigronia prussia sp. nov. is known as only a The new species also does not belong the Protochauliodes fragment of a hind wing, its megalopteran affinity can clade because it lacks the inclined hind wing cua-cup be confirmed without doubt based on the combination crossvein and has the anterior branch of hind wing A3 of the following characters: 1) broad membranous hind proximally touching A2, while in all extant genera of wing with well developed anal region, 2) presence of the Protochauliodes clade the hind wing cua-cup is nygmata, 3) distal fusion of ScP and RA, and 4) simple slightly inclined and the anterior branch of hind wing A3 CuP. The presence of nygmata, the well-developed MP, is connected with A2 by a distinct crossvein. The new and the presence of long and deeply forked hind wing species can be distinguished from the two extinct genera, A3 indicate that the new species belongs to Corydalidae Cretochaulus Ponomarenko, 1976 and Eochauliodes Liu but not Sialidae, in which nygmata are absent, the stem et al., 2012, in the Protochauliodes clade as the latter two and proximal half of MP in both fore- and hind wings genera have all hind wing RP and MA branches distally are weakened, and the hind wing A3 is simple and short. forked [probably simple in N. prussia sp. nov.]. The new Furthermore, the chauliodine affinity of the new species species shares the hind wing cua-cup straight and nearly is supported by the hind wing cua-cup crossvein that is vertical to CuA with all genera of the Archichauliodes straight and nearly vertical to CuA. In all known genera clade, so it should be placed in this clade. of Corydalinae the hind wing cua-cup is inclined to CuA Considering the hind wing venation, two characters and slightly sinuate in most species. In addition, the new species also differs from most dobsonfly genera except in the new species are remarkable, i.e. the MA stem Chloroniella Esben-Petersen, 1924 by the bifurcated hind subdistally with a short ma-mp crossvein [alternatively wing CuA [hind wing CuA with three or more branches described as basal r-m or MA with a short branch in most dobsonfly genera except Chloroniella, which has connecting to MP in Liu & Yang (2006) and Liu et al. only two branches]. (2012)], and the A3 with anterior branch proximally The subfamily Chauliodinae is separated into touching A2 by short fusion or very short crossvein. four clades in the phylogenetic analysis based on Both characters are probably apomorphic and can be morphological data (Liu et al., 2012). The extinct genus found only in five fishfly genera, all of which belong Jurochauliodes Wang & Zhang, 2010 from the Middle to the Archichauliodes clade, i.e. Ctenochauliodes Jurassic of China represents the basalmost clade. All van der Weele, 1909, Neochauliodes van der Weele, remaining genera, including all extant fishfly genera, 1909, Nigronia, Parachauliodes van der Weele, 1909, belong to the other three clades, i.e., the Dysmicohermes and Sinochauliodes Liu & Yang, 2006 (Fig. 4). The clade, the Protochauliodes clade, and the Archichauliodes latter four genera form a monophyletic group based on clade. The new species greatly differs from the genera of previous phylogenetic analysis, while Ctenochauliodes the Dysmicohermes clade (Dysmicohermes Munroe, 1953 is distantly related to this group (Liu & Yang, 2006; Liu

A new fishfly species Palaeoentomology 003 (2) © 2020 Magnolia Press • 191 species the a1-a2 crossvein is distantly proximad the branching point of A1 in hind wing, being similar to that in Neochauliodes, and there is only one crossvein (i.e., mp1- mp2) between branches of hind wing MP (present close to wing margin in most individuals, but a second mp1-mp2 crossvein also present in a few individuals we examined). In Parachauliodes and Sinochauliodes (the latter genus will be treated as a junior synonym of the former genus based on unpublished molecular data of Xingyue Liu), the a1-a2 crossvein is close to the branching point of A1 in hind wing, which should be an apomorphic state and shared by the new species. However, the new species also has two significantly different hind wing characters from all species of Parachauliodes and Sinochauliodes, i.e. the feebly narrowed pterostigmal region [strongly narrowed, nearly half width of 3rd radial cell (hypostigmal cell) in the latter two genera] and the presence of broad band-like dark marking [hind wing slightly smoky brown, sometimes with a few scattered distal dark spots, or entirely blackish brown in the latter two genera]. The feebly narrowed pterostigmal region is also present in Neochauliodes and Nigronia, but it seems to be plesiomorphic and is not very useful to infer the taxonomic position of the new species. But, the hind wing with broad band-like dark marking FIGURE 3. Nigronia prussia sp. nov., holotype RD 300. A, is probably apomorphic, and the new species shares this Photograph of proximal-anterior part of hind wing, showing the character state with two species of Ctenochauliodes, stem of MA. B, Photograph of proximal-posterior part of hind many species of Neochauliodes, and all extant species of wing, showing the position of a1-a2. Scale bars: 1.0 mm. Nigronia although the detail marking pattern in the new species is unique. According to the above comparison, the new et al., 2012; Liu, 2019). The new species differs from all species should belong to the monophyletic group species of Ctenochauliodes by the posterior branch of RP including Neochauliodes, Nigronia, Parachauliodes, and proximally straight [distinctly bended anteriad proximal Sinochauliodes. But it could not be clearly assigned to any to 1rp-ma crossvein in Ctenochauliodes] and the a1- of these genera solely based on the hind wing characters. a2 crossvein close to branching point of A1 [distantly Furthermore, there is no convincing argument to describe proximad branching point of A1 in Ctenochauliodes] in a new genus based on this new species as we could not hind wing. The new species can be distinguished from find any autapomorphic generic character. The present all species of Neochauliodes by the presence of only one placement of the new species in Nigronia is tentative and RP branch separated between 1ra-rp and 2ra-rp [two or relies on the presence of broad band-like marking, the more RP branches separated between 1ra-rp and 2ra-rp feebly narrowed pterostigmal region, and the presence in most Neochauliodes species except some species of of single RP branch between 1ra-rp and 2ra-rp in hind the Neochauliodes sundaicus group, which, however, wing. have four radial crossveins (see Liu et al., 2010: fig. 1)] The new Baltic amber fishfly species could provide, and the a1-a2 crossvein close to branching point of A1 in if genus placement is confirmed, important data for hind wing [distantly proximad branching point of A1 in understanding the historical biogeography of Chauliodinae Neochauliodes]. from the Northern Hemisphere. Currently, the palaeofauna The new species mostly resembles Nigronia, of Corydalidae from the Eocene of Europe comprises two Parachauliodes, and Sinochauliodes by having the genera of Chauliodinae (i.e., Chauliodes and Nigronia), presence of only one proximally straight hind wing RP both of which are still living but confined to eastern branch separated between 1ra-rp and 2ra-rp, which, North America. These two genera have been repeatedly however, is probably a plesiomorphic state as it is also regarded to be closely related to Asian endemic fishfly present in many genera of the Dysmicohermes clade and the genera in previous phylogenetic analyses (Liu & Yang, Protochauliodes clade. Moreover, in the extant Nigronia 2006; Liu et al., 2012, 2016). Liu et al. (2012) predicted

192 • Palaeoentomology 003 (2) © 2020 Magnolia Press liu & ANSORGE FIGURE 4. Habitus photographs of extant fishflies. A, Ctenochauliodes yangi Liu & Yang, 2006. B, Neochauliodes umbratus Kimmins, 1954. C, Nigronia fasciata (Walker, 1853). D, Nigronia serricornis (Say in Keating, 1824). E, Parachauliodes japonicus (McLachlan, 1867). F, Sinochauliodes maculosus Liu & Yang, 2006. Species illustrated represent the genera that have the stem of hind wing MA subdistally with a short crossvein. Arrow in A indicates anteriorly bended RP; arrows in B indicate two RP branches separated between 1ra-rp and 2ra-rp; arrow in D indicates a1-a2 distantly apart from branching point of A1; arrows in E and F respectively indicate distinctly narrowed pterostigmal region and a1-a2 close to branching point of A1. Scale bars: 5.0 mm. that the ancestral species of the clade including the which is considered to be an important route for the trans- aforementioned genera have managed the dispersal from Atlantic exchange of temperate biota during the Eocene Europe to eastern North America via the Thulean Bridge, (McKenna, 1983; Sanmartín et al., 2001). Besides, they

A new fishfly species Palaeoentomology 003 (2) © 2020 Magnolia Press • 193 assumed that there might have been no dispersal of the (Megaloptera: Corydalidae). Zootaxa, 2862, 1–38. ancestral species of extant Asian fishflies from Asia to https://doi.org/10.11646/zootaxa.2862.1.1 western North America via the Beringian Bridge or the Engel, M.S., Winterton, S.L. & Breitkreuz, L.C.V. (2018) Phylogeny mid-continental area of North America during the Early and evolution of Neuropterida: where have wings of lace Tertiary (Liu et al., 2012). This hypothesis explains why taken us? Annual Review of Entomology, 63, 531–551. the fishfly genera closely related to the Asian fishflies https://doi.org/10.1146/annurev-ento-020117-043127 are confined to eastern North America. Thus, the Baltic Hagen, H.A. (1856) Die im Bernstein befindlichen Neuropteren amber fishflies stand as crucial evidence to support this der Vorwelt bearbeitet von F. J. Pictet-Baraban, und Dr. H. hypothesis. The present discovery of Nigronia from Hagen. In: Berendt, G.C. (Ed.), Die im Bernstein befindlichen Baltic amber is a new example reflecting the consistent organischen Reste der Vorwelt gesammelt, in Verbindung mit biogeographical scenario shown above. Nevertheless, mehreren bearbeitet und herausgegeben von Dr. Georg Carl concerning the past occurrence of extant Oriental endemic Berendt. Bd. 2. Nicolai, Berlin, pp. 41–125. alderflies (i.e., Indosialis; see Wichard et al., 2009) from Huang, D.Y., Azar, D., Engel, M.S., Cai, C.Y., Garrouste, R. & Nel, the Baltic amber, we could not exclude the possibility that A. (2016) A new genus of alderflies (Megaloptera: Sialidae) the extant Oriental fishfly genera had been also present in Upper Cretaceous Burmese amber. Cretaceous Research, in the Eocene of Europe. Actually, despite the generally 64, 7–11. weak dispersal capacity, a few species of Neochauliodes https://doi.org/10.1016/j.cretres.2016.03.012 can be widely distributed as the Nigronia and Chauliodes Jepson, J.E. & Heads, S.W. (2016) Fossil Megaloptera (Insecta: species (Liu et al., 2007). By sharing some similar Neuropterida) from the Lower Cretaceous Crato Formation characters with Neochauliodes, the new Baltic amber of Brazil. Zootaxa, 4098 (1), 134–144. fishfly might alternatively be a stem-group species of https://doi.org/10.11646/zootaxa.4098.1.5 Kukalová-Peck, J. & Lawrence, J.F. (2004) Relationships among Neochauliodes. Hence, additional and better preserved coleopteran suborders and major endoneopteran lineages: material of N. prussia sp. nov. as well as other fishfly taxa evidence from hind wing characters. European Journal of from the Eocene Baltic amber are expected to be found Entomology, 101, 95–144. in the future to further figure out the palaeodiversity, https://doi.org/10.14411/eje.2004.018 phylogeny, and biogeography of Chauliodinae. Liu, X.Y. (2019) Phylogeny of Megaloptera: A review of present knowledge. In: Weihrauch, F., Frank, O., Gruppe, A., Jepson, J.E., Kirschey, L. & Ohl, M. (Eds), Proceedings of the XIII Acknowledgements International Symposium of Neuropterology, 17–22 June 2018, Laufen, Germany. Osmylus Scientific Publishers, We thank Axel Attula, German Amber Museum, Ribnitz- Wolnzach, pp. 39–54. Damgarten for access to the specimen. The first author is https://doi.org/10.5281/zenodo.3572321 Liu, X.Y., Hayashi, F. & Yang, D. (2007) Revision of the grateful to the Chutian Scholars Program for the support Neochauliodes sinensis species-group (Megaloptera: of this study. Corydalidae: Chauliodinae). 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