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.

Pollinator Diversity: Distribution, Ecological Function, and Conservation

“…Fossil insects from China, Spain and Russia have revealed ancient groups of insects that appear, based on interpretations of their mouthparts and associated pollen grains, to have been pollinators; examples include mid-Mesozoic thrips (Thysanoptera), flies (Diptera), lacewings (Neuroptera), scorpionflies (Mecoptera) and beetles (Coleoptera) (Ren 1998;Ren et al 2009;Labandeira 2010;Labandeira et al, 2007Labandeira et al, , 2016Peñalver et al 2012Peñalver et al , 2015Peris et al in press). It is notable that lacewings and scorpionflies are no longer significant pollinators (in terms of diversity) compared to groups such as the bees (Hymenoptera) and butterflies and moths (Lepidoptera) that have replaced them over time (see section 2.1).…”

The natural history of oviposition on a ginkgophyte fruit from the Middle Jurassic of northeastern China

“…and Ren, D., unpublished data). Based on these occurrences and knowledge of the ovipositor structure of the younger kalligrammatid lacewing Oregramma illecebrosa of the Yixian Formation that probably was associated with bennettitalean pollinators (Yang et al, 2014a(Yang et al, , 2014bLabandeira et al, 2016b), we infer that this type ovipositor structure originated for slashing deep into fleshy tissues. Such an ovipositional habit may have been tracking a particular, fleshy tissue type throughout the Jiulongshan flora among unrelated plant hosts.…”

“…The most likely maker of DT272 damage was an early, Callovian‐age species of the subfamily Oregrammatinae (Neuroptera: Kalligrammatidae), based on the ovipositor structure of Oregramma illecebrosa from the younger Yixian Formation of northeastern China (Labandeira et al ., ; Meng et al ., unpublished data).…”

“…The most likely maker of DT272 damage was an early, Callovian-age species of the subfamily Oregrammatinae (Neuroptera: Kalligrammatidae), based on the ovipositor structure of Oregramma illecebrosa from the younger Yixian Formation of northeastern China (Labandeira et al, 2016b;Meng et al, unpublished data). Other material DT272 has not been found on other specimens of Yimaia capituliformis, but it does occur on several of the basal attachment pads of Cycadolepis bracts on a specimen of Williamsonia sp.…”

“…Such an ovipositional habit may have been tracking a particular, fleshy tissue type throughout the Jiulongshan flora among unrelated plant hosts. Kalligrammatid lacewings also were long-proboscid pollinators that likely pollinated gymnosperms such as Williamsonia that provided rewards of pollination drops and pollen (Labandeira et al, 2016b). Additional understanding of the relationships between the fleshy tissues of ginkgoalean and bennettitalean plants, kalligrammatid ovipositors, and the pollination biology of kalligrammatids and bennettitaleans and possibly ginkgoalean hosts await additional plant-insect studies of the Jiulongshan biota.…”