title: The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies.
abstract: Mid-Mesozoic kalligrammatid lacewings (Neuroptera) entered the fossil record 165 million years ago (Ma) and disappeared 45 Ma later. Extant papilionoid butterflies (Lepidoptera) probably originated 80-70 Ma, long after kalligrammatids became extinct. Although poor preservation of kalligrammatid fossils previously prevented their detailed morphological and ecological characterization, we examine new, well-preserved, kalligrammatid fossils from Middle Jurassic and Early Cretaceous sites in northeastern China to unravel a surprising array of similar morphological and ecological features in these two, unrelated clades. We used polarized light and epifluorescence photography, SEM imaging, energy dispersive spectrometry and time-of-flight secondary ion mass spectrometry to examine kalligrammatid fossils and their environment. We mapped the evolution of specific traits onto a kalligrammatid phylogeny and discovered that these extinct lacewings convergently evolved wing eyespots that possibly contained melanin, and wing scales, elongate tubular proboscides, similar feeding styles, and seed-plant associations, similar to butterflies. Long-proboscid kalligrammatid lacewings lived in ecosystems with gymnosperm-insect relationships and likely accessed bennettitalean pollination drops and pollen. This system later was replaced by mid-Cretaceous angiosperms and their insect pollinators.
authors: College of Life Sciences, Capital Normal University, Beijing 100048, People's Republic of China Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA Department of Entomology and BEES Program, University of Maryland, College Park, MD 20742, USA rendong@mail.cnu.edu.cn.
authors: Conrad C
authors: CC
authors: Labandeira
authors: College of Life Sciences, Capital Normal University, Beijing 100048, People's Republic of China State Key Laboratory of Biocontrol, Key Laboratory of Biodiversity Dynamics and Conservation of Guangdong Higher Education Institute, College of Ecology and Evolution, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, People's Republic of China Geoscience Museum, Shijiazhuang University of Economics, Shijiazhuang 050031, People's Republic of China.
authors: Qiang
authors: Q
authors: Yang
authors: Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA Department of Crop and Agroenvironmental Sciences, University of Puerto Rico, Mayagüez, PR 00681, USA.
authors: Jorge A
authors: JA
authors: Santiago-Blay
authors: Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA National Centre for Biotechnology Information, National Library of Medicine, Bethesda, MD 20892, USA.
authors: Carol L
authors: CL
authors: Hotton
authors: Department of Ecology and Evolutionary Biology, Yale University, New Haven, CT 06511, USA Department of Biological Sciences, National University of Singapore, Singapore 117543, Singapore Yale-NUS College, Singapore 138614, Singapore.
authors: Antónia
authors: A
authors: Monteiro
authors: College of Life Sciences, Capital Normal University, Beijing 100048, People's Republic of China.
authors: Yong-Jie
authors: YJ
authors: Wang
authors: Department of Mineral Sciences, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA Jet Propulsion Laboratory, National Aeronautics and Space Administration, Pasadena, CA 91125, USA.
authors: Yulia
authors: Y
authors: Goreva
authors: College of Life Sciences, Capital Normal University, Beijing 100048, People's Republic of China Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA.
authors: ChungKun
authors: C
authors: Shih
authors: Department of Mineral Sciences, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA Department of Chemistry, Materials and Surfaces, SP Technical Research Institute of Sweden, Borås 51115, Sweden Geophysical Laboratory, Carnegie Institution of Washington, Washington, DC 20015, USA.
authors: Sandra
authors: S
authors: Siljeström
authors: Department of Mineral Sciences, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA.
authors: Tim R
authors: TR
authors: Rose
authors: Departments of Geology and Biology, Indiana University, Bloomington, IN 47405, USA.
authors: David L
authors: DL
authors: Dilcher
authors: College of Life Sciences, Capital Normal University, Beijing 100048, People's Republic of China labandec@si.edu.
authors: Dong
authors: D
authors: Ren
mesh_terms: D000818:Animals; D005075:Biological Evolution; D002080:Butterflies / Q000033:anatomy & histology; D005580:Fossils / Q000033:anatomy & histology*; D007313:Insecta / Q000033:anatomy & histology*; D014921:Wings, Animal / Q000033:anatomy & histology*
keywords: Kalligrammatidae; Papilionoidea; angiosperms; gymnosperms; tubular proboscis; wing eyespots