Abstract
The Shanwang Fauna has been one of the well-known Miocene Konservat-Lagerstätten for a long time, but the high-fidelity preservation of the fossils has been often ignored. This paper illustrates the exceptional preservation displayed by some fossil insects from the Shanwang Formation. The microstructures of the preserved exoskeletons were analyzed using a scanning electron microscope and a transmission electron microscope. The epicuticle, procuticle, and some fine details such as cuticular scales could be recognized from the structural remains. These details could enable the identification of valued taxonomic and phylogenetic features and contribute to understanding the diagenetic environment.
References
Binnington, K.C. (1993) Ultrastructure of the attachment of Serratia entomophila to scarab larval cuticle and a review of nomenclature for insect epicuticular layers. International Journal of Insect Morphology and Embryology, 22(2–4), 145–155.
https://doi.org/10.1016/0020-7322(93)90006-M
Binnington, K. & Retnakaran, A. (1991) Physiology of the Insect Epidermis. CSIRO Publications, Melbourne, 252 pp.
Briggs, D.E.G. (1999) Molecular taphonomy of animal and plant cuticles: selective preservation and diagenesis. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 354, 7–17.
https://doi.org/10.1098/rstb.1999.0356
Chen, P.F. (2000) Using fish taphonomy to reconstruct the environment of ancient Shanwang Lake. Advances in Ecological Research, 31, 483–496.
https://doi.org/10.1016/S0065-2504(00)31026-1
Deng, T., Wei, W.M. & Yue, L.P. (2003) Recent advances of the establishment of the Shanwang Stage in the Chinese Neogene. Vertebrata PalAsiatica, 41(4), 314–323.
Duncan, I.J. (1997) The taphonomy of insects. Unpublished Ph. D. thesis, Department of Geology, University of Bristol. Bristol, United Kingdom, xvi + 328 pp.
Esperante, R., Brand, L.R., Chadwick, A.V., Poma, O. (2015) Taphonomy and paleoenvironmental conditions of deposition of fossil whales in the diatomaceous sediments of the Miocene/Pliocene Pisco Formation, southern Peru—A new fossil–Lagerstätte. Palaeogeography, Palaeoclimatology, Palaeoecology, 417, 337–370.
https://doi.org/10.1016/j.palaeo.2014.09.029
Evans, H.E. (1984) Insect Biology – A Textbook of Entomology. Addison-Wesley Publishing Company, London, pp. 70–76.
Guo, Z.F., Liu, J.Q. & Chen, X.Y. (2007) Effect of Miocene basaltic volcanism in Shanwang (Shandong Province, China) on environmental changes. Science in China Series D: Earth Sciences, 50 (12), 1823–1827.
https://doi.org/10.1007/s11430-007-0119-4
Gupta, N.S. & Briggs, D.E.G. (2011) Taphonomy of animal organic skeletons through time. In: Allison, P.A. and Bottjer, D.J. (eds) Taphonomy. Vol. 32. Aims & Scope Topics in Geobiology Book Series. Springer, Dordrecht, pp. 199–221.
Gupta, N.S., Michels, R., Briggs, D.E.G., Evershed, R.P. & Pancost, R.D. (2006) The organic preservation of fossil arthropods: an experimental study. Proceedings of the Royal Society B: Biological Sciences, 273, 2777–2783.
https://doi.org/10.1098/rspb.2006.3646
Hadley, N.F. (1986) The arthropod cuticle. Scientific American, 255, 98–106.
https://doi.org/10.1038/scientificamerican0786-104
Harding, I.C & Chant, L.S. (2000) Self-sedimented diatom mats as agents of exceptional fossil preservation in the Oligocene Florissant lake beds, Colorado, Unites States. Geology, 28 (3), 195–198.
https://doi.org/10.1130/0091-7613(2000)28<195:SDMAAO>2.0.CO;2
He, H.Y., Deng, C.L., Pan, Y.X., Deng, T., Luo, Z.H., Sun, J.M. & Zhu, R.X. (2011) New 40Ar/39Ar dating results from the Shanwang Basin, eastern China: Constraints on the age of the Shanwang Formation and associated biota. Physics of the Earth and Planetary Interiors, 187 (1–2), 66–75.
https://doi.org/10.1016/j.pepi.2011.05.002
Hepburn, H.R. (1985) Structure of the integument. In: Kerkut, G.A. & Gilbert, L.I. (Eds.) Comprehensive Insect Physiology, Biochemistry and Pharmacology, vol. 3, Integument, Respiration and Circulation. Pergamon Press, Oxford, pp. 1–58.
https://doi.org/10.1016/B978-0-08-030804-3.50007-8
Hong, Y.C. (1983) Fossil insect in the Diatoms of Shanwang. Bulletin of the Tianjin Institute of Geology and Mineral Resources, 8, 1–15.
Hong, Y.C. (1985) Fossil Insects, Scorpionids and Araneids in the Diatoms of Shanwang. Geological Publishing House, Beijing, 113 pp.
Hou, L.H., Zhou., Z.H., Zhang, F.C. & Li, J.D. (2000) A new vulture from the Miocene of Shandong, Eastern China. Vertebrata PalAsiatica, 38(2), 104–110.
Li, H.M. (1982) The age of the Shanwang flora, Shandong. In: Selected Papers from the 12th Annual Convention of Palaeontological Society of China. Scientific Publishing House, Beijing, pp. 159–162. [In Chinese].
Li, Y.M., Ferguson, D.K., Wang, Y.F. & Li, C.S. (2010) Paleoenvironmental inferences from diatom assemblages of the middle Miocene Shanwang Formation, Shandong, China. Journal of Paleolimnology, 43, 799–814.
https://doi.org/10.1007/s10933-009-9369-9
Liang, M.M., Yan, J.X., Song, S.Y. & Wang, Q. (1998) Advances of research on Miocene Flora from Shanwang in Shandong Province. Chinese Bulletin of Botany, 15(suppl.), 32–40.
Liu, H.F. & Shi, N. (1989) Paleomagnetic study of the Shanwang Formation, Shandong Province. Acta Scientiarum Naturalium Universitatis Pekinensis, 25,76–593. [In Chinese with English summary].
Liu, L.P., Fortelius, M. & Pickford, M. (2002) New fossil Suidae from Shanwang, Shandong, China. Journal of Vertebrate Paleontology, 22, 152–163.
https://doi.org/10.1671/0272-4634(2002)022[0152:NFSFSS]2.0.CO;2
Métais, G & Sen, S. (2018) The late Miocene mammals from the Konservat-Lagerstätte of Saint-Bauzile (Ardèche, France). Comptes Rendus Palevol, 17(7), 479–493.
https://doi.org/10.1016/j.crpv.2018.05.001
Neville, A.C. (1975) Biology of Arthropod Cuticle. Springer, Berlin, 448 pp.
O’Brien, N.R., Meyer, H.W., Reilly, K., Ross, A.M. & Maguire, S. (2002) Microbial taphonomic processes in the fossilization of insects and plants in the late Eocene Florissant Formation, Colorado. Rocky Mountain Geology, 37 (1), 1–11.
https://doi.org/10.2113/gsrocky.37.1.1
Pan, Y.H., Zheng, W.X., Moyer, A.E., O’Connor, J.K., Wang, M., Zheng, X.T., Wang, X.L., Schroeter, E.R., Zhou, Z.H. & Schweitzer, M.H. (2016) Molecular evidence of keratin and melanosomes in feathers of the Early Cretaceous bird Eoconfuciusornis. Proceedings of the National Academy of Sciences, 113 (49), E7900–E7907.
https://doi.org/10.1073/pnas.1617168113
Qiu, Z.D. & Yan, C.L. (2005) New Sciurids from the Miocene Shanwang Formation, Linqu, Shandong. Vertebrata PalAsiatica, 43, 196–207.
Riou, B. (1995) Les fossiles des diatomites du Miocène supérieur de la montagne d’Andance (Ardèche, France). Géologie Méditerranéenne, 22 (1), 1–15.
https://doi.org/10.3406/geolm.1995.1565
Roček, Z., Dong, L.P., Přikryl, T., Sun, C.K., Tan, J. & Wang, Y. (2011) Fossil frogs (Anura) from Shanwang (Middle Miocene; Shandong Province, China). Geobios, 44 (5), 499–518.
https://doi.org/10.1016/j.geobios.2010.11.009
Schaefer, J., Kramer, K.J., Garbow, J.R., Jacob, G.S., Stejskal, E.O., Hopkins, T.L., Speirs, R.D. (1987) Aromatic cross-links in insect cuticle: detection by solid-state 13C and 15N NMR. Science, 235, 1200–1204.
https://doi.org/10.1126/science.3823880
Shi, L. (1990) Miocene diatoms from the Shanwang basin of shandong province and analysis by fuzzy mathematics of the paleoenvironment. Acta Botanica Sinica, 32 (11), 888–895.
Singh, S.S., Jansen, M.A., Franz, N.M. & Chawla, N. (2016) Microstructure and nanoindentation of the rostrum of Curculio longinasus Chittenden, 1927 (Coleoptera: Curculionidae). Materials Characterization, 118, 206–211.
https://doi.org/10.1016/j.matchar.2016.05.022
Stankiewicz, B.A., Mastalerz, M., Hof, C.H.J., Bierstedt, A., Flannery, M.B., Briggs, D.E.G. & Evershed, R.P. (1998) Biodegradation of the chitin-protein complex in crustacean cuticle. Organic Geochemistry, 28, 67–76.
https://doi.org/10.1016/S0146-6380(97)00113-7
Storch, G. & Qiu, Z.D. (2004) First complete heterosoricine shrew: A new genus and species from the Miocene of China. Acta Palaeontologica Polonica, 49 (3), 357–363.
Sun, A.L. (1961) Notes on fossil snakes from Shanwang, Shantung. Vertebrata PalAsiatica, 4, 310–312.
Tian, H.S. Van Loon, A.J., Zhang, Z.Q., Zhang, S.H., Zhang, B.H., Lu, M.Y., Li, F.C. & Ma, X.M. (2015) Neogene paleoseismic events and the Shanwang Biota’s burial in the Linqu area, Shandong Province, China. Acta Geologica Sinica-English Edition, 89 (4), 1103–1119.
https://doi.org/10.1111/1755-6724.12517
Van de Kamp, T., Riedel, A. & Greven, H.G. (2016) Micromorphology of the elytral cuticle of beetles, with an emphasis on weevils (Coleoptera: Curculionoidea). Arthropod Structure & Development, 45 (1), 14–22.
https://doi.org/10.1016/j.asd.2015.10.002
Wang, Y.F., Ferguson, D.K., Li, C.S. & Herendeen, P.S. (2005) Ceratophyllum (Ceratophyllaceae) from the Miocene of eastern China and its paleoecological significance. Systematic Botany, 30 (4), 705–711.
https://doi.org/10.1600/036364405775097752
Yang, H. & Yang, S.P. (1994) The Shanwang fossil biota in eastern China: a Miocene Konservat-Lagerstätte in lacustrine deposits. Lethaia, 27 (4), 345–354.
https://doi.org/10.1111/j.1502-3931.1994.tb01585.x
Yeh, H.K. (1977) First discovery of Miocene bird in China. Vertebrata PalAsiatica, 15 (3), 244–248.
Zhang, J.F. (1986) Ecological and biogeographical analysis on Miocene Shanwang insect fauna. In: Selected Papers from the 13th and 14th Annual Conventions of Palaeontological Society of China. Anhui Sciennce and Technology Publishing House, Hefei, pp. 237–248.
Zhang, J.F. (1989) Fossil Insects From Shanwang, Shandong, China. Shandong Science & Technology Press, Jinan, 459 pp. [in Chinese with English summary]
Zhang, J.F. (1990) New fossil species of Apoidea (Insecta: Hymenoptera). Zoological Systematics, 15 (1), 83–91.
Zhang, J.F., Sun, B. & Zhang, X.Y. (1994) Miocene Insects and Spiders from Shanwang, Shandong. Science Press, Beijing, 298 pp.