A 100 million-year-old snake-fly larva with an unusually large antenna


Authors: Haug JT, Müller P, Haug C

Published in: Bulletin of Geosciences, volume 95, issue 2; pages: 167 - 177; Received 4 April 2019; Accepted in revised form 19 March 2020; Online 16 May 2020

Keywords: Neuropterida, Raphidioptera, Cretaceous, Burmese amber, morphological diversity,

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Snakeflies is the vernacular name of the representatives of the group Raphidioptera, an ingroup of Holometabola. The group Raphidioptera is rather species-poor, with only slightly more than 200 species. Snake-fly larvae are rather uniform in appearance and possess an elongate body; the trunk has similar appearing segments and lacks specialisations such as protruding structures. Such larvae are also known from the fossil record, namely from fossils preserved in amber that largely resemble extant forms. Here we report a new fossil snake-fly larva from Burmese amber. The specimen is very unusual in the morphology of the antennae. In extant snake-fly larvae, the antenna is quite short and thin, often inconspicuous at first sight on normal habitus photographs. In the new fossil, the antenna is much longer than the head and also appears very robust, comparable to the thoracic appendages. Together with another snake-fly larva previously described from Burmese amber, this find demonstrates that there were larval morphologies of snakefly larvae back in the Cretaceous that are absent in the modern fauna.


Aristov, D.S., Novokshonov, V.G. & Pan’kov, N.N. 2006. Taxonomy of the fossil grylloblattid nymphs (Insecta: Grylloblattida). Paleontological Journal 40(1), 79-89.View article

Aspöck, H. 2002. The biology of Raphidioptera: A review of present knowledge. Acta Zoologica Academiae Scientiarum Hungaricae 48(2), 35-50.

Aspöck, H. & Aspöck, U. 2009. Raphidioptera-Kamelhalsfliegen. Ein Überblick zum Einstieg. Entomologica Austriaca 16, 53-72.View article

Aspöck, U. & Aspöck, H. 1999. Kamelhälse, Schlammfliegen, Ameisenlöwen. Wer sind sie? (Insecta: Neuropterida: Raphidioptera, Megaloptera, Neuroptera). Stapfia 60(138), 1-34.

Aspöck, U. & Aspöck, H. 2007. Verbliebene Vielfalt vergangener Blüte. Zur Evolution, Phylogenie und Biodiversität der Neuropterida:(Insecta: Endopterygota). Denisia 20, zugleich Kataloge der obersterreichischen Landesmuseen Neue Serie 66(2007), 451-516.

Aspöck, U., Haring, E. & Aspöck, H. 2012a. The phylogeny of the Neuropterida: long lasting and current controversies and challenges (Insecta: Endopterygota). Arthropod Systematics & Phylogeny 70(2), 119-129.

Aspöck, H., Liu, X.Y. & Aspöck, U. 2012b. The family Inocelliidae (Neuropterida: Raphidioptera): A review of present knowledge. Mitteilungen der Deutschen Gesellschaft für allgemeine und angewandte Entomologie 18, 565-573.

Aspöck, U., Plant, J.D. & Nemeschkal, H.L. 2001. Cladistic analysis of Neuroptera and their systematic position within Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera). Systematic Entomology 26(1), 73-86.View article

Badano, D., Engel, M.S., Basso, A., Wang, B. & Cerretti, P. 2018. Diverse Cretaceous larvae reveal the evolutionary and behavioural history of antlions and lacewings. Nature Communications 9, 3257.View article

Baranov, V., Hoffeins, C., Hoffeins, H.-W. & Haug, J.T. 2019. More than dead males: Reconstructing the ontogenetic series of terrestrial non-biting midges from the Eocene amber forest. Bulletin of Geosciences 94(2), 187-199.View article

Beutel, R.G. & Friedrich, F. 2008. Comparative study of larval head structures of Megaloptera (Hexapoda). European Journal of Entomology 105, 917-938.View article

Beutel, R.G. & Ge, S.Q. 2008. The larval head of Raphidia (Raphidioptera, Insecta) and its phylogenetic significance. Zoology 111(2), 89-113.View article

Beutel, R.G., Friedrich, F. & Aspöck, U. 2010. The larval head of Nevrorthidae and the phylogeny of Neuroptera (Insecta). Zoological Journal of the Linnean Society 158, 533-562.View article

Clavier, S., Contreras-Ramos, A. & Guillemet, L. 2010. First report of Protosialis Weele, 1909 in French Guiana (Megaloptera, Sialidae). Ephemera 11(2), 135-140.

Contreras-Ramos, A. & Harris, S.C. 1998. The immature stages of Platyneuromus (Corydalidae), with a key to the genera of larval Megaloptera of Mexico. Journal of the North American Benthological Society 17, 489-517.View article

Cruickshank, R.D. & Ko, K. 2003. Geology of an amber locality in the Hukawng Valley, northern Myanmar. Journal of Asian Earth Sciences 21, 441-455.View article

Delclos, X., Penalver, E., Arillo, A., Engel, M.S., Nel, A., Azar, D. & Ross, A. 2016. New mantises (Insecta: Mantodea) in Cretaceous ambers from Lebanon, Spain, and Myanmar. Cretaceous Research 60, 91-108.View article

Engel, M.S. 2002. The smallest snakefly (Raphidioptera: Mesoraphidiidae): a new species in Cretaceous amber from Myanmar, with a catalog of fossil snakeflies. American Museum Novitates 3363, 1-22.View article

Gepp, J. 1984. Erforschungsstand der Neuropteren. Larven der Erde (mit einem Schlüssel zur Larvaldiagnose der Familien, einer Übersicht von 340 beschriebenen Larven und 600 Literaturzitaten), 183-239. In Progress in Worlds Neuropterology. Proceedings of the First International Symposium on Neuropterology, Graz.

Grebennikov, V.V. 2004. Grub-like larvae of Neuroptera (Insecta): a morphological review of the families Ithonidae and Polystoechotidae and a description of Oliarces clara. European Journal of Entomology 101, 409-418.View article

Grimaldi, D.A. 1997. A fossil mantis (Insecta, Mantodea) in Cretaceous amber of New Jersey: with comments on the early history of the Dictyoptera. American Museum Novitates, 3204, 1-11.

Grimaldi, D. 2003. A revision of Cretaceous mantises and their relationships, including new taxa (Insecta: Dictyoptera: Mantodea). American Museum Novitates 3412, 1-47.View article

Grimaldi, D. & Nascimbene, P.C. 2010. Raritan (New Jersey) Amber, 167-191. In Penney, D., Green, D. & Mellish, C. (eds) Biodiversity of Fossils in Amber from the Major World Deposits. Siri Scientific Press, Manchester, UK.

Grimaldi, D.A. & Ross, A.J. 2004. Raphidiomimula, an enigmatic new cockroach in Cretaceous amber from Myanmar (Burma)(Insecta: Blattodea: Raphidiomimidae). Journal of Systematic Palaeontology 2, 101-104.View article

Gurney, A.B. 1947. Notes on Dilaridae and Berothidae, with special reference to the immature stages of the Nearctic genera (Neuroptera). Psyche: A Journal of Entomology 54(3), 145-169.View article

Hagen, H. 1854. Über die Neuropteren der Bernsteinfauna. Verhandlungen des zoologisch-botanischen Vereins in Wien 4, 221-232.

Harmer, S.F. & Shipley, A.E. 1895. The Cambridge natural history. Macmillan and Company, London, New York.View article

Haug, C., Herrera-Flórez, A.F., Müller, P. & Haug, J.T. 2019a. Cretaceous chimera - an unusual 100-million-year old neuropteran larva from the “experimental phase” of insect evolution. Palaeodiversity 12, 1-11.View article

Haug, C., Shannon, K.R., Nyborg, T. & Vega, F.J. 2013b. Isolated mantis shrimp dactyli from the Pliocene of North Carolina and their bearing on the history of Stomatopoda. Bolétin de la Sociedad Geológica Mexicana 65, 273-284.View article

Haug, J.T. 2020. Why the term “larva” is ambiguous, or what makes a larva? Acta Zoologica 101, 167-188.View article

Haug, J.T., Haug, C. & Ehrlich, M. 2008. First fossil stomatopod larva (Arthropoda: Crustacea) and a new way of documenting Solnhofen fossils (Upper Jurassic, Southern Germany). Palaeodiversity 1, 103-109.

Haug, J.T., Briggs, D.E.G. & Haug, C. 2012. Morphology and function in the Cambrian Burgess Shale megacheiran arthropod Leanchoilia superlata and the application of a descriptive matrix. BMC Evolutionary Biology 12(1), 162.View article

Haug, J.T., Müller, P. & Haug, C. 2018. The ride of the parasite: a 100-million-year old mantis lacewing larva captured while mounting its spider host. Zoological Letters 4(1), 31.View article

Haug, J.T., Müller, P. & Haug, C. 2019b. A 100-million-year old predator: a fossil neuropteran larva with unusually elongated mouthparts. Zoological Letters 5, 29.View article

Haug, J.T., Müller, P. & Haug, C. 2019c. A 100-million-year old slim insectan predator with massive venom-injecting stylets - a new type of neuropteran larva from Burmese amber. Bulletin of Geosciences 94(4), 431-440.View article

Haug, J.T., Müller, C.H.G. & Sombke, A. 2013a. A centipede nymph in Baltic amber and a new approach to document amber fossils. Organisms Diversity & Evolution 13, 425-432.View article

Haug, J.T., Haug, C., Kutschera, V., Mayer, G., Maas, A., Liebau, S., Castellani, C., Wolfram, U., Clarkson, E.N.K. & Waloszek, D. 2011. Autofluorescence imaging, an excellent tool for comparative morphology. Journal of Microscopy 244, 259-272.View article

Henry, C.S. 1976. Some aspects of the external morphology of larval owlflies (Neuroptera: Ascalaphidae), with particular reference to Ululodes and Ascaloptynx. Psyche: A Journal of Entomology 83(1), 1-31.View article

Hoffman, K.M. & Brushwein, J.R. 1992. Descriptions of the larvae and pupae of some North American Mantispinae (Neuroptera: Mantispidae) and development of a system of larval chaetotaxy for Neuroptera. Transactions of the American Entomological Society 118, 159-196.

Hörnig, M.K., Haug, C., Schneider, J.W. & Haug, J.T. 2018. Evolution of reproductive strategies in dictyopteran insects-clues from ovipositor morphology of extinct roachoids. Acta Palaeontologica Polonica 63, 1-24.View article

Kerp, H. & Bomfleur, B. 2011. Photography of plant fossils - new techniques, old tricks. Review of Palaeobotany and Palynology 166, 117-151.View article

Liu, X.Y., Lu, X.M. & Zhang, W.W. 2016. New genera and species of the minute snakeflies (Raphidioptera: Mesoraphidiidae: Nanoraphidiini) from the mid Cretaceous of Myanmar. Zootaxa 4103(4), 301-324.View article

Liu, X., Shi, G., Xia, F., Lu, X., Wang, B. & Engel, M.S. 2018. Liverwort mimesis in a Cretaceous lacewing larva. Current Biology 28, 1475-1481.View article

MacLeod, E.G. 1970. The Neuroptera of the Baltic Amber. I. Ascalaphidae, Nymphidae, and Psychopsidae. Psyche: A Journal of Entomology 77(2), 147-180.View article

Mansell, M.W. 1983. New Crocinae (Neuroptera: Nemopteridae) from South America, with descriptions of larvae. Journal of the Entomological Society of Southern Africa 46(1), 115-130.

Monserrat, V.J. 2014. Los megalópteros de la Península Ibérica (Insecta, Neuropterida, Megaloptera, Sialidae). Graellsia 70(2), e009.View article

Monserrat, V.J. & Papenberg, D. 2015. Los rafidiópteros de la Península Ibérica (Insecta, Neuropterida: Raphidioptera). Graellsia 71(1), e024.View article

New, T.R. 1989 Planipennia, Lacewings. Handbuch der Zoologie, Vol. 4. Arthropoda: Insecta, Part 30. 132 pp. Walter de Gruyter, Berlin.

New, T.R. 1992. The lacewings (Insecta, Neuroptera) of Tasmania. Papers and Proceedings of the Royal Society of Tasmania 126, 29-45.View article

New, T.R. 2004. Insecta: Neuropteroidea, 491-500. In Yule, C.M. & Yong, H.S. (eds) Freshwater Invertebrates of the Malaysian Region. Academy of Sciences Malaysia, Kuala Lumpur.

Nicoli Aldini, R., Letardi, A. & Pantaleoni, R.A. 2012. State of the art on Neuropterida of Sicily and Malta. Biodiversity Journal 3, 445-458.

Pérez-de la Fuente, R., Delclos, X., Penalver, E., Speranza, M., Wierzchos, J., Ascaso, C. & Engel, M.S. 2012b. Early evolution and ecology of camouflage in insects. Proceedings of the National Academy of Sciences 109(52), 21414-21419.View article

Pérez-de la Fuente, R., Delclos, X., Penalver, E. & Engel, M.S. 2016. A defensive behavior and plant-insect interaction in Early Cretaceous amber-the case of the immature lacewing Hallucinochrysa diogenesi. Arthropod Structure & Development 45(2), 133-139.View article

Pérez-de la Fuente, R., Engel, M.S., Azar, D. & Penalver, E. 2019. The hatching mechanism of 130-million-year-old insects: an association of neonates, egg shells and egg bursters in Lebanese amber. Palaeontology 62(4), 547-559.View article

Pérez-de la Fuente, R., Penalver, E., Azar, D. & Engel, M.S. 2018. A soil-carrying lacewing larva in Early Cretaceous Lebanese amber. Scientific Reports 8(1), 16663.View article

Pérez-de la Fuente, R., Penalver, E., Delclos, X. & Engel, M.S. 2012a. Snakefly diversity in early Cretaceous amber from Spain (Neuropterida, Raphidioptera). ZooKeys 204, 1-40.View article

Perrichot, V., & Engel, M.S. 2007. Early Cretaceous snakefly larvae in amber from Lebanon, Myanmar, and France (Raphidioptera). American Museum Novitates 3598, 1-11.View article

Riek, E.F. 1970. Neuroptera, 472-494. In Evans, H.E. (ed.) The Insects of Australia. Melbourne University Press, Canberra.

Stürzer, C. & Gepp, J. 2004. Beiträge zur Larvalbiologie und -morphologie von Conwentzia pineticola Enderlein, 1905 und C. psociformis (Curtis, 1834) (Neuroptera: Coniopterygidae). Entomologica Austrica 11, 7-10.

Tauber, C.A., Tauber, M.J. & Albuquerque, G.S. 2003. Neuroptera (lacewings, antlions), 785-798. In Resh, V.H. & Cardé, R.T. (eds) Encyclopedia of insects. Academic Press, Amsterdam.

Vršanský, P. 2009. Albian cockroaches (Insecta, Blattida) from French amber of Archingeay. Geodiversitas 31(1), 73-99.View article

Wachmann, E. & Saure, C. 1997. Netzflügler, Schlamm- und Kamelhalsfliegen: Beobachtung, Lebensweise. Naturbuch-Verlag, Augsburg.

Weidner, H. 1958. Einige interessante Insektenlarven aus der Bernsteininklusen-Sammlung des Geologischen Staatsinstituts Hamburg (Odonata, Coleoptera, Megaloptera, Planipennia). Mitteilungen aus dem Geologisch-Paläontologischen Institut Hamburg 27, 50-68.

Weitschat, W. & Wichard, W. 1998. Atlas der Pflanzen und Tiere im baltischen Bernstein. Verlag Dr. Friedrich Pfeil, München.

Weitschat, W. & Wichard, W. 2002. Atlas of Plants and Animals in Baltic Amber. Verlag Dr. Friedrich Pfeil, München.

Wichard, W., Gröhn, C. & Seredszus, F. 2009. Aquatic insects in Baltic amber. Kessel, Remagen.

Woglum, R.S. & McGregor, E.A. 1958. Observations on the life history and morphology of Agulla bractea Carpenter (Neuroptera: Raphidiodea: Raphidiidae). Annals of the Entomological Society of America 51(2), 129-141.View article