Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2026-04-29
Page range: 125-138
Abstract views: 169
PDF downloaded: 89

Cretolala kachinensis gen. et sp. nov. (Hemiptera: Fulgoromorpha: Delphacoidea), the first lalacid from mid-Cretaceous Kachin amber: morphological and taphonomic implications

Yunnan Key Laboratory for Palaeobiology, Institute of Palaeontology, Yunnan University, South Waihuan Road, Chenggong District, Kunming 650500, China, MEC International Joint Laboratory for Palaeobiology and Palaeoenvironment, Yunnan University, Kunming 650500, China, Bonn Institute for Organismic Biology (BIOB), Department for Paleontology, University of Bonn, Bonn 53115, Germany
Institute of Rock Structure and Mechanics of the Czech Academy of Sciences, V Holešovičkách 41, 18200, Prague, Czech Republic
Institute of Physiology II, University Hospital Bonn, University of Bonn, 53115 Bonn, Germany
Institut de Systématique, Évolution, Biodiversité (UMR 7205), Muséum national d’Histoire naturelle, CNRS, Sorbonne Université, EPHE-PSL, Université des Antilles, Paris F-75005, France
The Steinhardt Museum of Natural History and The New Environmental School, Tel Aviv University, 69978 Tel Aviv, Israel
Bonn Institute for Organismic Biology (BIOB), Department for Paleontology, University of Bonn, Bonn 53115, Germany
Natural History Museum of Denmark, University of Copenhagen, Copenhagen 1350, Denmark, Institute of Environmental Sciences, Faculty of Biology, Jagiellonian University, Kraków, Poland
Department of Geological Survey and Mineral Exploration, Ministry of Natural Resources and Environmental Conservation, Myanmar Gems Museum, Nay Pyi Taw 15011, Myanmar
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 21008, China
Oxford University Museum of Natural History, University of Oxford, Parks Road, Oxford OX1 3PW, UK
Institute of Geosciences, University of Bonn, Bonn 53115, Germany
Laboratory of Evolutionary Entomology and Museum of Amber Inclusions, Department of Invertebrate Zoology and Parasitology, University of Gdańsk, Gdańsk, PL80–308, Poland
Bonn Institute for Organismic Biology (BIOB), Department for Paleontology, University of Bonn, Bonn 53115, Germany, State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 21008, China, Institute of Geology and Paleontology, Charles University, Prague 12843, Czech Republic
Hemiptera Fulgoromorpha Delphacoidea Mesozoic Burmese amber Planthopper taxonomy taphonomy mineral infilling wing maculation

Abstract

We report Cretolala kachinensis gen. et sp. nov., the first formally described representative of the family Lalacidae from mid-Cretaceous (~99 Ma) Kachin amber. The morphology and taphonomy of the specimen were studied using optical microscopy, micro-computed tomography (micro-CT), confocal laser scanning microscopy (CLSM), and scanning electron microscopy coupled with energy-dispersive X-ray spectroscopy (SEM‒EDS). Cretolala kachinensis gen. et sp. nov. exhibits a unique combination of forewing characters, including a strengthened, broad, and wrinkled costal margin; ScP+R fork proximal to both the Pcu+A1 fusion and the CuA fork; MP with five terminals; and CuA1 and CuA2 each with two terminals. Crossvein patterns differ between the left and right forewings, indicating that crossvein expression alone is not diagnostic. The forewing bears distinct maculation, with a face-like pattern dorsally and an extensive lateral pattern that may indicate aposematism, mimicry, or plant-like camouflage in an arboreal herbivore. Taphonomic analyses based on optical micrographs and material-dependent X-ray attenuation in micro-CT show that minerals are distributed throughout the fossil, coating anatomical surfaces and infilling pre-existing cavities and voids. These phases help stabilize morphology and influence the micro-CT reconstruction of soft-bodied structures. SEM–EDS analyses indicate that the body-coating and void-filling minerals are dominated by quartz and pyrite, with feldspars and aluminosilicate mixtures also present, locally enriched in K, Na, Mg, or Ca. This assemblage and its textures point to a polyphase paragenetic sequence, with early detrital infill and later authigenic mineralization. Our results suggest that minerals and some chemicals derived from the host sediments can be transported into and, in some cases, precipitated within amber inclusions, infilling voids and providing structural support that influences fossil preservation. Collectively, these phases occlude porosity, stabilize morphology, and enhance X-ray contrast, although they locally obscured primary tissue boundaries. Our results highlight multistage diagenesis in Kachin amber and underscore variability among inclusions in both mineral assemblages and the timing of mineralization/diagenetic events.

References

  1. Boderau, M., Nel, A. & Fu, Y.Z. (2025a) A new Lapicixius planthopper species (Fulgoromorpha: Lalacidae) from the Lower Cretaceous Yixian Formation of northeastern China. Journal of Insect Biodiversity, 77 (1), 85–90. https://doi.org//10.12976/jib/2025.77.1.12
  2. Boderau, M., Fu, Y.Z., Jiang, H., Guan, S., Peng, A., Nel, A. & Jouault, C. (2025b) Bayesian modelling of the fossil record enlightens the evolutionary history of Hemiptera. Proceedings of the Royal Society B, 292, 20251133. https://doi.org//10.1098/rspb.2025.1133
  3. Boderau, M., Nel, A. & Jouault, C. (2025c) Diversification and extinction of Hemiptera in deep time. Communications Biology, 8 (1), 352. https://doi.org//10.1038/s42003-025-07773-x
  4. Bourgoin, T. (1993) Female genitalia in Hemiptera Fulgoromorpha, morphological and phylogenetic data. Annales de la Société entomologique de France, (N.S.), 29 (3), 225–244. https://doi.org//10.1080/21686351.1993.12277686
  5. Bourgoin, T. (2025) FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to Fulgoromorpha. Available from: https://flow.hemiptera-databases.org/flow/?&lang=en (accessed 30 Oct 2025).
  6. Bourgoin, T. & Szwedo, J. (2022) Toward a new classification of planthoppers Hemiptera Fulgoromorpha: 1. What do Fulgoridiidae really cover? Annales Zoologici, 72 (4), 951–962. https://doi.org//10.3161/00034541ANZ2022.72.4.011
  7. Bourgoin, T. & Szwedo, J. (2023) Toward a new classification of planthoppers Hemiptera Fulgoromorpha: 2. Higher taxa, their names and their composition. Zootaxa, 5297 (4), 562–568. https://doi.org//10.11646/zootaxa.5297.4.5
  8. Bourgoin, T., Wang, R.R., Asche, M., Hoch, H., Soulier-Perkins, A., Stroiński, A., Yap, S. & Szwedo, J. (2015) From micropterism to hyperpterism: recognition strategy and standardized homology-driven terminology of the forewing venation patterns in planthoppers (Hemiptera: Fulgoromorpha). Zoomorphology, 134, 63–77. https://doi.org//10.1007/s00435-014-0243-6
  9. Brożek, J., Stroiński, A., Romaniak, A. & Bourgoin, T. (2024) Disparity of metatibial and metatarsal cuticular and sensory structures in Cixiidae (Hemiptera: Fulgoromorpha) with a metatibiotarsal diagnosis for the tribes. Zoological Letters, 10 (1), 16. https://doi.org//10.1186/s40851-024-00239-8
  10. Bucher, M. (2024) Les grandes extinctions de masse ont-elles impacté l’évolution des insectes? Exemple de l’histoire macroévolutive des Hémiptères Fulgoromorpha. Systématique, phylogénie et taxonomie. Thèse de doctorat, Museum national d’histoire naturelle (MNHN), Paris, 409 pp.
  11. Bucher, M., Gignoux, G., Szwedo, J. & Bourgoin, T. (2024) Time-traveling through fossil planthopper tegmina in the Paleozoic and Mesozoic eras (Insecta: Hemiptera: Fulgoromorpha). Palaeoentomology, 7 (1), 1–67. https://doi.org//10.11646/palaeoentomology.7.1.1
  12. Cruickshank, R.D. & Ko, K.O. (2003) Geology of an amber locality in the Hukawng Valley, Northern Myanmar. Journal of Asian Earth Sciences, 21 (5), 441–455. https://doi.org//10.1016/S1367-9120(02)00044-5
  13. Hamilton, K.G.A. (1990) Insecta from the Santana Formation, Lower Cretaceous, of Brazil. Chapter 6. Homoptera. Bulletin of the American Museum of Natural History, 195, 82–122.
  14. Haug, J.T., Azar, D., Ross, A.J., Szwedo, J., Wang, B., Arillo, A., Baranov, V., Bechteler, J., Beutel, R.G., Blagoderov, V., Delclòs, X., Feldberg, K., Feldmann, R., Foth, C., Fraaije, R.H.B., Gehler, A., Harms, D., Hedenäs, L., Hyžný, M., Jagt, J.W.M., Jagt-Yazykova, E.A., Jarzembowski, E.A., Kerp, H., Khine, P.K., Kirejtshuk, A.G., Klug, C., Kopylov, D.S., Kotthoff, U., Kriwet, J., McKellar, R.C., Nel, A., Nützel, A., Peñalver, E., Perrichot, V., Pint, A., Ragazzi, E., Regalado, L., Reich, M., Rikkinen, J., Schmidt, A.R., Schneider, H., Schram, F.R., Schweigert, G., Selden, P.A., Solórzano-Kraemer, M.M., Stilwell, J.D., van-Bakel, B.W.M., Vega, F.J., Wang, Y., Xing, L.D. & Haug, C. (2020) Comment on the letter of the Society of Vertebrate Paleontology (SVP) dated April 21, 2020, regarding “Fossils from conflict zones and reproducibility of fossil-based scientific data”: Myanmar amber. Paläontologische Zeitschrift, 94, 431–437. https://doi.org//10.1007/s12542-020-00522-x
  15. International Commission on Zoological Nomenclature (ICZN) (1999) International code of zoological nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, xxix + 336 pp.
  16. Jiang, H., Tomaschek, F., Muscente, A. D., Niu, C., Nyunt, T. T., Fang, Y., Schmidt, U., Chen, J., Lönartz, M., Mähler, B., Wappler, T., Jarzembowski, E.A., Szwedo, J., Zhang, H., Rust, J. & Wang, B. (2022) Widespread mineralization of soft-bodied insects in Cretaceous amber. Geobiology, 20, 363–376. https://doi.org//10.1111/gbi.12488
  17. Jiang, H., Szwedo, J., Labandeira, C.C., Chen, J., Moulds, M.S., Mähler, B., Muscente, A.D., Zhuo, D., Nyunt, T.T., Zhang, H.C., Wei, C., Rust, J. & Wang, B. (2024) Mesozoic evolution of cicadas and their origins of vocalization and root feeding. Nature Communications, 15, 376. https://doi.org//10.1038/s41467-023-44446-x
  18. Lozano, R.P., López Del Valle, R., Baeza, E., Delvene, G., Barrón, E., Peñalver, E., Rodrigo, A. & Pérez-de la Fuente, R. (2025) Preventive conservation of amber: some preliminary investigations. Pyrite decay in amber: deterioration of collections and conservation guidelines. Geoheritage, 17 (4), 168. https://doi.org//10.1007/s12371-025-01219-w
  19. Nel, A., Prokop, J., Nel, P., Grandcolas, P., Huang, D.Y., Roques, P., Guilbert, E., Dostál, O. & Szwedo, J. (2012) Traits and evolution of wing venation pattern in paraneopteran insects. Journal of Morphology, 273 (5), 480–506. https://doi.org//10.1002/jmor.11036
  20. Parchem, R.J., Perry, M.W. & Patel, N.H. (2007) Patterns on the insect wing. Current Opinion in Genetics and Development, 17 (4), 300–308. https://doi.org//10.1016/j.gde.2007.05.006
  21. Ren, D., Lu, L.W., Guo, Z.G. & Ji, S.A. (1995) Faunae and stratigraphy of Jurassic-Cretaceous in Beijing and the adjacent areas. Seismic Publishing House, Beijing, viii+222 pp.
  22. Ren, D., Yin, J. & Dou, W. (1998) New planthoppers and froghoppers from the Late Jurassic of northeast China (Homoptera: Auchenorrhyncha). Acta Zootaxonomica Sinica, 23 (3), 281–288.
  23. Schubnel, T., Desutter-Grandcolas, L., Legendre, F., Prokop, J., Mazurier, A., Garrouste, R., Grandcolas, P. & Nel, A. (2019) To be or not to be: postcubital vein in insects revealed by microtomography. Systematic Entomology, 45 (2), 327–336. https://doi.org//10.1111/syen.12399
  24. Scotese, C.R., Vérard, C., Burgener, L., Elling, R.P. & Kocsis, A.T. (2025) The Cretaceous world: plate tectonics, palaeogeography and palaeoclimate. Geological Society, London, Special Publications, 544, 31–202. https://doi.org//10.1144/SP544-2024-28
  25. Shi, G.H., Grimaldi, D.A., Harlow, G.E., Wang, J., Wang, J., Yang, M.C., Lei, W.Y., Li, Q.L. & Li, X.H. (2012) Age constraint on Burmese amber based on U-Pb dating of zircons. Cretaceous Research, 37, 155–163. https://doi.org//10.1016/j.cretres.2012.03.014
  26. Stagg, N.A., Jiang, H., Tomaschek, F., Staniczek, A.H., Mähler, B., Bruthansová, J., Nyunt, T.T. & Godunko, R.J. (2026). Hidden taxonomic and taphonomic diversity revealed by mayflies (Ephemeroptera: Hexagenitidae) from mid-Cretaceous Kachin amber. Scientific Reports (in press). https://doi.org//10.1038/s41598-026-46621-8
  27. Szwedo, J. (2007) Fulgoromorpha: planthoppers. In: Martill D.M., Bechly, G. & Loveridge, R.F. (Eds), The Crato fossil beds of Brazil: window into an ancient world. Cambridge University Press, Cambridge, pp. 297–313. https://doi.org//10.1017/CBO9780511535512.012
  28. Szwedo, J., Bourgoin, T. & Lefebvre, F. (2004) Fossil planthoppers (Hemiptera: Fulgoromorpha) of the world. An annotated catalogue with notes on Hemiptera classification. Studio 1, Warsaw, 199 pp.
  29. Szwedo, J., Wang, B., Soszyńska-Maj, A., Azar, D. & Ross, A.J. (2020) International Palaeoentomological Society statement. Palaeoentomology, 3 (3), 221–222. https://doi.org//10.11646/palaeoentomology.3.3.1
  30. Thu, K. & Zaw, K. (2017) Gem deposits of Myanmar. Geological Society, London Memoirs, 48, 497–529. https://doi.org//10.1144/m48.23
  31. Wang, L.N., Szwedo, J., Zhuo, D., Xiao, C.T. & Luo, C.H. (2025) Adding to the diversity of Katlasidae (Hemiptera: Fulgoromorpha: Fulgoridoidea)—a new genus and species from mid-Cretaceous Kachin amber of northern Myanmar. Palaeontologia Electronica, 28 (3), 1–15. https://doi.org//10.26879/1575
  32. Wittkopp, P.J., Carroll, S.B. & Kopp, A. (2003) Evolution in black and white: genetic control of pigment patterns in Drosophila. Trends in Genetics, 19 (9), 495–504. https://doi.org//10.1016/S0168-9525(03)00194-X
  33. Wittkopp, P.J. & Beldade, P. (2009) Development and evolution of insect pigmentation: genetic mechanisms and the potential consequences of pleiotropy. Seminars in Cell & Developmental Biology, 20 (1), 65–71. https://doi.org//10.1016/j.semcdb.2008.10.002
  34. Zhang, Z. (2002) New early Cretaceous lalacid from Jingxi Basin of Beijing, China (Homoptera: Fulgoroidea). Acta Zootaxonomica Sinica, 27 (1), 20–23.