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The hatching mechanism of 130‐million‐year‐old insects: an association of neonates,egg shells and egg bursters in Lebanese amber
Authors:Ricardo Pérez‐de la Fuente  Michael S. Engel  Dany Azar  Enrique Peñalver
Affiliation:1.

https://orcid.org/0000-0002-2830-2639;2. Oxford University Museum of Natural History, Parks Road, Oxford, OX1 3PW UK;3.

https://orcid.org/0000-0003-3067-077X;4. Division of Entomology, Natural History Museum, and Department of Ecology & Evolutionary Biology, University of Kansas, 1501 Crestline Drive – Suite 140, Lawrence, KS, 66045 USA;5. Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY, 10024 USA;6. Department of Biology, Faculty of Sciences II, Lebanese University, Fanar Matn, PO Box 7. 26110217, Beirut, Lebanon;8. State Key Laboratory of Palaeobiology & Stratigraphy, Nanjing Institute of Geology & Palaeontology, Chinese Academy of Sciences, Nanjing, 210008 China;9. Instituto Geológico y Minero de Espa?a (Museo Geominero), C/Cirilo Amorós 42, 46004 Valencia, Spain

Abstract:Hatching is a pivotal moment in the life of most animals. Diverse chemical, behavioural and mechanical methods have evolved in metazoans to break the egg membranes. Among them, many arthropod and vertebrate embryos hatch using ephemeral, frequently convergent structures known as egg bursters. However, the evolutionary processes by which hatching mechanisms and related embryonic structures became established in deep time are poorly understood due to a nearly complete absence from the fossil record. Herein we describe an exceptional c. 130‐million‐year‐old association in Lebanese amber composed of multiple neonate green lacewing larvae, Tragichrysa ovoruptora gen. et sp. nov. (Neuroptera, Chrysopoidea), and conspecific egg remains. Egg bursters with a serrated blade bearing a short process are attached to three longitudinally split egg shells. Embryos of extant green lacewing relatives (Chrysopidae) utilize this egg burster morphotype to open a vertical slit on the egg, after which the burster is moulted and left joined to the empty egg shell. Additionally, the new larval species has extremely elongate dorsal tubercles, an adaptation to carry exogenous debris for protection and camouflage also known from other Cretaceous chrysopoids but absent in modern relatives. The present discovery demonstrates that the hatching mechanism of modern green lacewings was established in the chrysopoid lineage by the Early Cretaceous and proves through direct fossil evidence how some morphological traits related to hatching and linked behaviours, at least in insect embryos, have been subject to a high degree of evolutionary conservatism.
Keywords:hatching  egg burster  fossil insects  Neuroptera  Cretaceous  amber
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