Mesozoic lacewings from China provide phylogenetic insight into evolution of the Kalligrammatidae (Neuroptera)

Abstract

BackgroundThe Kalligrammatidae are distinctive, large, conspicuous, lacewings found in Eurasia from the Middle Jurassic to mid Early Cretaceous. Because of incomplete and often inadequate fossil preservation, an absence of detailed morphology, unclear relationships, and unknown evolutionary trends, the Kalligrammatidae are poorly understood.ResultsWe describe three new subfamilies, four new genera, twelve new species and four unassigned species from the late Middle Jurassic Jiulongshan and mid Early Cretaceous Yixian Formations of China. These kalligrammatid taxa exhibit diverse morphological characters, such as mandibulate mouthparts in one major clade and siphonate mouthparts in the remaining four major clades, the presence or absence of a variety of distinctive wing markings such as stripes, wing spots and eyespots, as well as multiple major wing shapes. Based on phylogenetic analyses, the Kalligrammatidae are divided into five principal clades: Kalligrammatinae Handlirsch, 1906, Kallihemerobiinae Ren & Engel, 2008, Meioneurinae subfam. nov., Oregrammatinae subfam. nov. and Sophogrammatinae subfam. nov., each of which is accorded subfamily-level status. Our results show significant morphological and evolutionary differentiation of the Kalligrammatidae family during a 40 million-year-interval of the mid Mesozoic.ConclusionA new phylogeny and classification of five subfamilies and their constituent genera is proposed for the Kalligrammatidae. These diverse, yet highly specialized taxa from northeastern China suggest that eastern Eurasia likely was an important diversification center for the Kalligrammatidae. Kalligrammatids possess an extraordinary morphological breadth and panoply of adaptations during the mid-Mesozoic that highlight our conclusion that their evolutionary biology is much more complex than heretofore realized.

Phylogenetic analyses elucidate the inter‐relationships of Pamphilioidea (Hymenoptera, Symphyta)

“…Notwithstanding, the bias in character sampling is always a serious issue owing to limited available characters preserved on fossils (Prevosti and Chemisquy, ; Pattinson et al., ). As a result, the phylogenetic inference and affinity are mostly dependent on the wing venation characters (Li et al., ; Yang et al., ). Our present study also has a similar issue mentioned above, i.e.…”

New Aetheogrammatidae (Insecta: Neuroptera) from the Lower Cretaceous Yixian Formation, China

“…Recently, the family Kalligrammatidae was divided into five subfamilies: Kalligrammatinae, Kallihemerobiinae, Meioneuritinae, Oregrammatinae and Sophogrammatinae (Yang et al, 2014b).…”

“…Yang et al (2011) considered this genus in Kalligrammatidae, and thought that similar character states of Kalligrammina Panfilov, 1980 andAetheogramma Ren andEngel, 2008 "may have arisen independently, although this seems to be less plausible than they are indeed nearly related" (Yang et al, 2011, p. 65). Recently, the genus was tentatively assigned to the kalligrammatid subfamily Kalligrammatinae (Yang et al, 2014b). Based on the data presented here from the new fossils, we believe that Kalligrammina areolata belongs to Aetheogrammatidae.…”

“…It is closely related to another highly specialized Mesozoic family, the Kalligrammatidae. Adults of both families have long-proboscid siphonate mouthparts allowing them to presumptively feed on gymnosperm fluids and pollen (Zherikhin, 1978;Rasnitsyn, 1980;Labandeira, 2010;Yang et al, 2012Yang et al, , 2014b. Hitherto, aetheogrammatids were known from only three species from the Middle Jurassic to the Lower Cretaceous of China and Kazakhstan: Aetheogramma speciosum Ren and Engel, 2008, Ectopogramma kalligrammoides Engel et al, 2011and Kalligrammina areolata Panfilov, 1980(Ren and Engel, 2008Engel et al, 2011).…”

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 ., , ; Labandeira et al ., ), 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.…”