Three new species of aneuretopsychids (Insecta: Mecoptera) from the Jehol Biota, China
Highlights
► Three new species of Aneuretopsychidae are described. ► The species with siphonate mouthparts might be pollinators of gymnosperm hosts. ► These proboscides imply insects carry pollination with their matched proboscides. ► The finding enriches our understanding and broadens the diversity of this family.
Introduction
The Aneuretopsychidae is a small, enigmatic family of long-proboscid insects with two genera, Aneuretopsyche Rasnitsyn and Kozlov, 1990 with three species (Aneuretopsyche rostrata, A. minima and A. vitimensis) and Jeholopsyche Ren, Shih and Labandeira, 2011 with one species (Jeholopsyche liaoningensis) (Rasnitsyn and Kozlov, 1990; Ren et al., 2009, 2011). Aneuretopsychids are moderate-sized insects, with a small head and a remarkably prolonged, siphonate proboscis, the exterior of which is covered with well-developed dense hairs or microtrichia arranged in distinct annulae; the terminus is surrounded by a distinctive, lobed, fleshy pseudolabellum. The antennae are distinctly longer than the proboscis and multiarticulate; the articles are covered with annulate hairs. The forewing is narrow and elongate, Rs diverges from R1 distally of an anastomosis of MP with CuA, both Rs and MA bifurcate, the MP 4 is branched and the CuA is single or probably bifurcating, CuA bends at about its midpoint, A1 reaches the middle of the posterior margin, there are several long, wide cells between Rs, MA and MP, and the veins are bent in places where crossveins abut on them. Hairs on the legs are arranged in distinctive rings (Rasnitsyn and Kozlov, 1990; Ren et al., 2011).
Aneuretopsychidae is one of the four families in Mecoptera, having prolonged siphonate mouthparts: Mesopsychidae (Ren et al., 2009, 2010a; Bashkuev, 2011a), Pseudopolycentropodidae (Ren et al., 2009, 2010b; Shih et al., 2011), Aneuretopsychidae (Rasnitsyn and Kozlov, 1990; Ren et al., 2009, 2011) and Nedubroviidae (Bashkuev, 2011b). These four families are members of a particularly interesting clade of the Aneuretopsychina, which lasted for 150 myr from the Late Permian to the latest Early Cretaceous (Ren et al., 2009, 2011). This essentially pre-angiospermous lineage exhibited a tubular siphonate proboscis, indicating that mid Mesozoic scorpionflies were accessing pollination drops from gymnospermous plants (Labandeira et al., 2007; Ren et al., 2009, 2011; Labandeira, 2010). Such a siphonate mouthpart structure, modified from the mecopteran labial structure, was convergent with and homologous to labial mouthpart elements of the brachycerous dipteran proboscis, but analogous to the maxillary glossae of lepidopterans, which evolved during the Early Cretaceous (Liu et al., 2005; Labandeira et al., 2007; Ren et al., 2009, 2011; Labandeira, 2010). Grimaldi et al. considered that Aneuretopsychidae, Pseudopolycentropodidae, Mesopsychidae are not true mecopterans but rather stem-group mecopteroids showing remarkable parallel evolution with the Diptera (Grimaldi and Engel, 2005; Grimaldi et al., 2005). Ren et al. regarded these three families to represent a major, early-derived lineage of basal Mecoptera (Ren et al., 2009, 2011).
Recently, we collected three fossils of Aneuretopsychidae, all with overlapped wings and relatively well-preserved body. These fossils were collected from the Early Cretaceous (late Barremian) Yixian Formation exposed at Huangbanjigou Village, Shangyuan Township, Beipiao City, Liaoning Province, China (Ren et al., 1997, 2010, 2011). Based on their different forewing characters (Sc with three crossveins to C; R1 single; Rs+MA bifurcation distinctly basal of the first forking of MP), we describe three new species of Jeholopsyche herein.
Section snippets
Material and methods
The three fossil specimens are housed in the Key Laboratory of Insect Evolution and Environmental Changes, College of Life Sciences, Capital Normal University, Beijing, China (CNUB; Dong Ren, Curator).
Photographs of whole specimens were taken using a Nikon D100 digital camera coupled to a Nikkor 105 mm macro lens. The specimens were examined using a Leica MZ12.5 dissecting microscope, and illustrated with the aid of a drawing tube attached to the microscope. Line drawings were made with
Systematic palaeontology
Order Mecoptera Packard, 1886
Family Aneuretopsychidae Rasnitsyn and Kozlov, 1990
Genus Jeholopsyche Ren, Shih and Labandeira, 2011
Type species. Jeholopsyche liaoningensis Ren, Shih and Labandeira, 2011
Emended diagnosis. Head, oviform with prolonged siphonate mouthparts, compound eyes big and oval and ocellus small and nearly round; mesothorax and metathorax almost the same size, but prothorax smaller; legs, long and slender with dense hairs, and all legs nearly the same shape, but hind legs
Discussion
Data on these three new species enhance our understanding of the morphological characters of Aneuretopsychidae, and provide a basis for future phylogenetic studies of this family. The new species, which have prolonged siphonate mouthparts, might have been feeders on the secretions and exudates of the reproductive structures of contemporaneous gymnosperms living near the site of deposition of the Yixian Formation. From the proboscis structure and inferred ecological relationships, we believe
Acknowledgements
We thank Yang Qiang, Cui Yingying, Yang Xiaoguang and Peng Yuanyuan in the CNU Key Lab for their detailed comments and fruitful suggestions. We are grateful to Dr Alexandr P. Rasnitsyn, Dr Alexei S. Bashkuev and an anonymous reviewer for their positive and useful comments. This research is supported by the National Basic Research Program of China (973 Program) (2012CB821906), National Natural Science Foundation of China (No. 40872022, 30811120038, 31071964) and PHR Project of Beijing Municipal
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