Neuroptera; larvae; phylogeny; empodium; Cretaceous; Burmese amber


The larval life phase often represents the major share of the life span of holometabolans, such as beetles or flies. Therefore, knowledge of these larvae is crucial concerning ecological functions, but also concerning the evolutionary history of the different groups. In the holometabolan group Neuroptera (lacewings), larvae have numerous specialisations, among them the venom-injecting mouthparts, the stylets. Another such very characteristic feature of many lacewing larvae is the empodium, an attachment structure at the tip of the leg, which varies in shape in the larvae of the different lacewing groups. One type of empodium is the elongate trumpet-shaped type, which is found in several lacewing groups today as well as in many fossil representatives, especially well visible due to preservation in amber. Based on the pattern of phylogenetic distribution of the elongate trumpet-shaped empodium, different reconstructions of the evolution of this structure have been put forward, some suggesting possibly convergent evolution, others repetitive losses. In this study, new lacewing larvae preserved in 100-million-year-old amber from Myanmar are presented, which expand the number of lineages with an elongate trumpet-shaped empodium. Based on these new data, the implications on the evolutionary history of the different lineages within Neuroptera are discussed. The fossils demand for an earlier evolution in some reconstructions, but especially for more and later losses of this structure.


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

Aspöck U. & Aspöck H. (2008) - Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola). Systematic Entomology, 33(1): 97-127.

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

Avise J.C. & Liu J.X. (2011) - On the temporal inconsistencies of Linnean taxonomic ranks. Biological Journal of the Linnean Society, 102(4): 707-714.

Ax P. (1995) - Das System der Metazoa I - Ein Lehrbuch der phylogenetischen Systematik. Gustav Fischer, Stuttgart, Germany, 226 pp.

Badano D. (2022) - Rediscovery of the myrmecophilous larvae of Italochrysa italica (Insecta: Neuroptera: Chrysopidae). Integrative Systematics: Stuttgart Contributions to Natural History, 5(1): 17-24.

Badano D., Aspöck U., Aspöck H. & Cerretti P. (2017) - Phylogeny of Myrmeleontiformia based on larval morphology (Neuropterida: Neuroptera). Systematic Entomology, 42(1): 94-117.

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(1): 3257.

Badano D., Fratini M., Maugeri L., Palermo F., Pieroni N., Cedola A., Haug J.T., Weiterschan T., Velten J., Mei M., Di Giulio A. & Cerretti P. (2021a) - X-ray microtomography and phylogenomics provide insights into the morphology and evolution of an enigmatic Mesozoic insect larva. Systematic Entomology, 46(3): 672-684.

Badano D., Di Giulio A., Aspöck H., Aspöck U. & Cerretti P. (2021b) - Burrowing specializations in a lacewing larva (Neuroptera: Dilaridae). Zoologischer Anzeiger, 293: 247-256.

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(3): 533-562.

Cai C.Y., Tihelka E., Liu X.Y. & Engel M.S. (2023) - Improved modelling of compositional heterogeneity reconciles phylogenomic conflicts among lacewings. Palaeoentomology, 6(1): 049-057.

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.

Dey S.R. (2014) - Developmental stages of Hemerobius indicus Kimmins (Hemerobiidae : Neuroptera) from western Himalaya, India. The Beats of Natural Sciences, 1(3): 1-8.

Engel M.S., Winterton S.L. & Breitkreuz L.C.V. (2018) - Phylogeny and evolution of Neuropterida: where have wings of lace taken us? Annual Review of Entomology, 63: 531-551.

Grimaldi D. & Engel M.S. (2005) - Evolution of the Insects. Cambridge University Press, Cambridge, England.

Gurney A.B. (1947) - Notes on Dilaridae and Berothidae, with special reference to the immature stages of the Nearctic genera (Neuroptera). Psyche, 54(3): 145-170.

Haring E. & Aspöck U. (2004) - Phylogeny of the Neuropterida: A first molecular approach. Systematic Entomology, 29(3): 415-430.

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

Haug C., Posada Zuluaga V., Zippel A., Braig F., Müller P., Gröhn C., Weiterschan T., Wunderlich J., Haug G.T. & Haug J.T. (2022b) - The morphological diversity of antlion larvae and their closest relatives over 100 million years. Insects, 13: 587.

Haug C., Braig F. & Haug J.T. (2023a) - Quantitative analysis of lacewing larvae over more than 100 million years reveals a complex pattern of loss of morphological diversity. Scientific Reports, 13: 6127.

Haug G.T., Baranov V., Wizen G., Pazinato P.G., Müller P., Haug C. & Haug J.T. (2021b) - The morphological diversity of long-necked lacewing larvae (Neuroptera: Myrmeleontiformia). Bulletin of Geosciences, 96: 431-457.

Haug G.T., Haug C., van der Wal S., Müller P. & Haug J.T. (2022a) - Split-footed lacewings declined over time: indications from the morphological diversity of their antlion-like larvae. PalZ, 96: 29-50.

Haug J.T. & Haug C. (2022) - 100 Million-year-old straight-jawed lacewing larvae with enormously inflated trunks represent the oldest cases of extreme physogastry in insects. Scientific Reports, 12: 12760.

Haug J.T., Baranov V., Müller P. & Haug C. (2021a) - New extreme morphologies as exemplified by 100 million-year-old lacewing larvae. Scientific Reports, 11: 20432.

Haug J.T., Baranov V., Schädel M., Müller P., Gröhn C. & Haug C. (2020) - Challenges for understanding lacewings: how to deal with the incomplete data from extant and fossil larvae of Nevrorthidae? (Neuroptera). Fragmenta Entomologica, 52(2): 137-167.

Haug J.T., Haug G.T., Zippel A., van der Wal S., Müller P., Gröhn C., Wunderlich J., Hoffeins C., Hoffeins H.W. & Haug C. (2021c) - Changes in the morphological diversity of larvae of lance lacewings, mantis lacewings and their closer relatives over 100 million years. Insects, 12(10): 860.

Haug J.T., Haug G.T. & Haug C. (2023b) - Reconstructing the history of lacewing diversification: shape heterochrony and core tree as tools for reconstructing evolutionary processes. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 308/1: 1-21.

Jandausch K., Beutel R.G., Pohl H., Gorb S.N. & Büsse S. (2018a) - The legs of "spider associated" parasitic primary larvae of Mantispa aphavexelte (Mantispidae, Neuroptera) - Attachment devices and phylogenetic implications. Arthropod Structure and Development, 47(5): 449-456.

Jandausch K., Pohl H., Aspöck U., Winterton S.L. & Beutel R.G. (2018b) - Morphology of the primary larva of Mantispa aphavexelte Aspöck & Aspöck, 1994 (Neuroptera: Mantispidae) and phylogenetic implications to the order of Neuroptera. Arthropod Systematics and Phylogeny, 76(3): 529-560.

Jandausch K., Beutel R.G. & Bellstedt R. (2019) - The larval morphology of the spongefly Sisyra nigra (Retzius, 1783) (Neuroptera: Sisyridae). Journal of Morphology, 280(12): 1742-1758.

Killington F.J. (1936) - A monograph of the British Neuroptera. Ray Society, London, 269 pp. + 15 pls.

Lambertz M. & Perry S.F. (2015) - Chordate phylogeny and the meaning of categorial ranks in modern evolutionary biology. Proceedings of the Royal Society B: Biological Sciences, 282(1807): 20142327.

Lambertz M. & Perry S.F. (2016) - Again on the meaning of categorial ranks in modern evolutionary biology? Organisms Diversity & Evolution, 16: 723-725.

Machado R.J.P., Gillung J.P., Winterton S.L., Garzón-Orduña I.J., Lemmon A.R., Lemmon E.M. & Oswald J.D. (2019) - Owlflies are derived antlions: anchored phylogenomics supports a new phylogeny and classification of Myrmeleontidae (Neuroptera). Systematic Entomology, 44(2): 418-450.

MacLeod E.G. (1960) - The immature stages of Boriomyia fidelis (Banks) with taxonomic notes on the affinities of the genus Boriomyia (Neuroptera: Hemerobiidae). Psyche: A Journal of Entomology, 67(1-2): 26-40.

MacLeod E.G. (1964) - A comparative morphological study of the head capsule and cervix of larval Neuroptera (Insecta). Harvard University, Cambridge, Massachusetts, USA.

MacLeod E.G. & Stange L.A. (2005) - Brown Lacewings (of Florida) (Insecta: Neuroptera: Hemerobiidae). UF University of Florida, IFAS Extension, 225: 1-6.

Mansell M.W. (1976) - The larva of Laurhervasia setacea (Klug) (Neuroptera: Nemopteridae: Crocinae) from Southern Africa. Journal of the Entomological Society of Southern Africa, 39(2): 153-158.

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.

Martins C.C., Ardila-Camacho A. & Courtney G.W. (2018) - Neotropical Osmylidae larvae (Insecta, Neuroptera): description of habitats and morphology. Aquatic Insects, 39(2-3): 181-207.

Miller G.L. & Lambdin P.L. (1984) - Redescriptions of the larval stages of Hemerobius stigma Stephens (Neuroptera: Hemerobiidae). Florida Entomologist, 67(3): 377-382.

Minter L.R. (1990) - A comparison of the eggs and first-instar larvae of Mucroberotha vesicaria Tjeder with those of other species in the families Berothidae and Mantispidae (Insecta : Neuroptera). Advances in Neuropterology. Proceedings of the Third International Symposium on Neuropterology: 115-129.

Mitchell R.G. (1962) - Balsam woolly aphid predators native to Oregon and Washington. Technical Bulletin of the Oregon Agricultural Experiment Station, 62: 1-63.

Möller A., Minter L.R. & Olivier P.A.S. (2006) - Larval morphology of Podallea vasseana Navás and Podallea manselli Aspöck & Aspöck from South Africa (Neuroptera: Berothidae). African Entomology, 14(1): 1-12.

Monserrat V.J., Oswald J.D., Tauber C.A. & Díaz-Aranda L.M. (2001) - Recognition of larval Neuroptera. In: McEwen P.K., New T.R. & Whittington A.E. (Eds.) - Lacewings in the Crop Environment: 43-81. Cambridge University Press, Cambridge.

Monserrat V.J. (2008) - Nuevos datos sobre algunas especies de crisópidos (Insecta: Neuroptera: Chrysopidae). Heteropterus Revista de Entomología, 8: 171-196.

New T.R. (1989) - Planipennia, Lacewings. Handbuch der Zoologie, Vol. 4 (Arthropoda: Insecta), Part 30: 1-132. Walter de Gruyter, Berlin.

New T.R. (1991) - Neuroptera (lacewings). In: Naumann I.D. (Ed.) - The Insects of Australia, Vol. I. 2nd edition: 525-542. CSIRO Publishing, Clayton.

New T.R. (1992) - The lacewings (Insecta, Neuroptera) of Tasmania. Papers & Proceedings - Royal Society of Tasmania, 126: 29-45.

Oswald J.D., Contreras-Ramos A. & Penny N.D. (2002) - Neuroptera (Neuropterida). In: Llorente-Bousquets J.J. & Morrone J. (Eds.) - Biodiversidad, Taxonomía y Biogeografía de Artrópodos de México: Hacia una Síntesis de su Conocimiento: 559-581. Universidad Nacional Autónoma de México, Mexico City.

Shi G., Grimaldi D.A., Harlow G.E., Wang J., Wang J., Yang M., Lei W., Li Q. & Li X. (2012) - Age constraint on Burmese amber based on U-Pb dating of zircons. Cretaceous Research, 37, 155-163.

Tauber C.A. (1974) - Systematics of North American chrysopid larvae: Chrysopa carnea group (Neuroptera). The Canadian Entomologist, 106(11): 1133-1153.

Tauber C.A. & Tauber M.J. (1968) - Lomamyia latipennis (Neuroptera: Berothidae) life history and larval descriptions. The Canadian Entomologist, 100(6): 623-629.

Tauber C.A., Tauber M.J. & Albuquerque G.S. (2003) - Neuroptera (Lacewings, Antlions). In: Resh V.H. & Cardé R.T. (Eds.) - Encyclopedia of Insects: 785-798. Academic Press, San Diego, London, Burlington.

Tauber C.A., Tauber M.J. & Albuquerque G.S. (2006) - Berchmansus elegans (Neuroptera: Chrysopidae): larval and adult characteristics and new tribal affiliation. European Journal of Entomology, 103(1), 221-231.

Tillyard R.J. (1918) - Studies in Australian Neuroptera. No. 7. The life-history of Psychopsis elegans (Guérin). Proceedings of the Linnean Society of New South Wales, 43: 787-818.

Vasilikopoulos A., Misof B., Meusemann K., Lieberz D., Flouri T., Beutel R.G., Niehuis O., Wappler T., Rust J., Peters R.S., Donath A., Podsiadlowski L., Mayer C., Bartel D., Böhm A., Liu S., Kapli P., Greve C., Jepson J.E., Liu X., Zhou X., Aspöck H. & Aspöck U. (2020) - An integrative phylogenomic approach to elucidate the evolutionary history and divergence times of Neuropterida (Insecta: Holometabola). BMC Evolutionary Biology, 20(1): 133.

Winterton S.L., Hardy N.B. & Wiegmann, B.M. (2010) - On wings of lace: Phylogeny and Bayesian divergence time estimates of Neuropterida (Insecta) based on morphological and molecular data. Systematic Entomology, 35(3): 349-378.

Winterton S.L., Lemmon A.R., Gillung J.P., Garzon I.J., Badano D., Bakkes D.K., Breitkreuz L.C. V., Engel M.S., Lemmon E.M., Liu X., Machado R.J.P., Skevington J.H. & Oswald J.D. (2018) - Evolution of lacewings and allied orders using anchored phylogenomics (Neuroptera, Megaloptera, Raphidioptera). Systematic Entomology, 43(2): 330-354.

Yu T., Kelly R., Mu L., Ross A., Kennedy J., Broly P., Xia F., Zhang H., Wang B. & Dilcher D. (2019) - An ammonite trapped in Burmese amber. Proceedings of the National Academy of Sciences, 116: 11345-11350.

Zachos F.E. (2011) - Linnean ranks, temporal banding, and time-clipping: why not slaughter the sacred cow? Biological Journal of the Linnean Society, 103(3): 732-734.

Zachos F.E. (2014) - Paraphyly—again!? A plea against the dissociation of taxonomy and phylogenetics. Zootaxa, 3764(5): 594-596.

Zimmermann D., Randolf S. & Aspöck U. (2019) - From chewing to sucking via phylogeny—from sucking to chewing via ontogeny: mouthparts of Neuroptera. In: Krenn H.W. (Ed.) - Insect Mouthparts. Form, Function, Development and Performance. Zoological Monographs: 361-385.

Zippel A., Kiesmüller C., Haug G.T., Müller P., Weiterschan T., Haug C., Hörnig M.K. & Haug J.T. (2021) - Long-headed predators in Cretaceous amber - fossil findings of an unusual type of lacewing larva. Palaeoentomology, 4(5): 475-498.

Zwick P. (1967) - Beschreibung der aquatischen Larve von Neurorthus fallax (Rambur) und Errichtung der neuen Planipennierfamilie Neurorthidae fam. nov. Gewässer und Abwässer, 44: 65-86.






Received 2023-08-09
Accepted 2024-01-23
Published 2024-02-22