Research Article |
Corresponding author: Hong Pang ( lsshpang@mail.sysu.edu.cn ) Corresponding author: Dong Ren ( rendong@cnu.edu.cn ) Academic editor: Shaun Winterton
© 2018 Qiang Yang, Chaofan Shi, Xiangchuan Li, Hong Pang, Dong Ren.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Yang Q, Shi C, Li X, Pang H, Ren D (2018) The first fossil brown lacewing from the Miocene of the Tibetan Plateau (Neuroptera, Hemerobiidae). ZooKeys 726: 145-154. https://doi.org/10.3897/zookeys.726.21086
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A new species of Hemerobiidae, Wesmaelius makarkini Yang, Pang & Ren, sp. n. is described from the Lower Miocene, Garang Formation of Zeku County, Qinghai Province (northeastern Tibetan Plateau), China. The species is assigned to the widely distributed extant genus Wesmaelius Krüger (Hemerobiinae). The species represents the first named fossil of this family from China, which sheds light on the historical distribution of Wesmaelius and early divergences within Hemerobiinae.
Cenozoic, China, Wesmaelius
Hemerobiidae, commonly known as brown lacewings, are the third largest family of Neuroptera, with about 520 species assigned to 27 genera (
Compared with its putative sister group Chrysopidae, Hemerobiidae have a relatively sparse and recent fossil record (
List of Named Fossil Hemerobiidae (updated from
Taxon | Deposit | Reference |
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Drepanepteryx oedobia Makarkin | Miocene, Russia |
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Drepanepteryx ramosa Makarkin | Miocene, Russia |
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Hemerobius incertus Makarkin | Miocene, Russia |
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Hemerobius prohumulinus Makarkin | Miocene, Russia |
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Megalomus caucasicus Makarkin | Miocene, Russia |
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Megalomus sikhotensis Makarkin | Miocene, Russia |
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Notiobiella thaumasta Oswald | Miocene, Dominican amber | Oswald 1999 |
Wesmaelius makarkini sp. n. | Miocene, China | This paper |
Bothromicromus lachlani Scudder | Oligocene, Canada |
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Drepanepteryx resinata (Krüger, 1923) | Eocene, Baltic amber |
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Megalomus tinctus (Jarzembowski) | Eocene, England |
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Prolachlanius resinatus (Hagen) | Eocene, Baltic amber |
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Proneuronema gradatum Makarkin, Wedmann et Weiterschan | Eocene, Baltic amber |
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Proneuronema minor Makarkin, Wedmann et Weiterschan | Eocene, Baltic amber |
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Proneuronema wehri (Makarkin, Archibald et Oswald) | Eocene, USA |
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Prospadobius moestus (Hagen) | Eocene, Baltic amber |
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Sympherobius completus Makarkin and Wedmann | Eocene, Baltic amber |
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Sympherobius siriae Jepson, Penney et Green 2010 | Eocene, Baltic amber |
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Wesmaelius mathewesi Makarkin, Archibald et Oswald 2003 | Eocene, Canada |
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Megalomus densistriatus Henriksen | Eocene, Denmark |
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Plesiorobius canadensis (Klimaszewski et Kevan, 1986) | Late Cretaceous, Canadian amber |
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Cretomerobius disjunctus Ponomarenko | Early Cretaceous, Mongolia |
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Purbemerobius medialis Jepson, Makarkin et Coram | Early Cretaceous, England |
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Promegalomus anomalus Panfilov | Late Jurassic, Kazakhstan |
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Fossil hemerobiids have never been described from China. Mesohemerobius jeholensis Ping, from the Lower Cretaceous, Yixian Formation of China, was previously placed in Hemerobiidae, but was later excluded from the family by
The specimen was collected from the Guide Group at Caergen Village, Duohemao Town, Zeku County, eastern Qinghai Province, China (34°56'N, 101°48'E, 3700 m a.s.l.) (fig. 1 in
The specimen was examined using a Zeiss Discovery V20 stereomicroscope and photographed with an AxioCam HRc digital camera attached to the Zeiss Discovery V20 stereomicroscope (both instruments Carl Zeiss Light Microscopy, Göttingen, Germany). Line drawings were prepared with the CorelDraw 12 graphics software and with the aid of Adobe Photoshop CS6. The vein terminology in general follows
Venation abbreviations used in the text and figures:
AA1–AA3 first to third anterior anal vein;
CuA anterior cubitus;
CuP posterior cubitus;
hv humeral veinlet;
fl flexion fold line;
MA anterior branche of media;
MP posterior branche of media;
ORB1, ORB2, ORB3 first to third oblique radial branches;
RA anterior radius;
RP posterior sector;
ScA subcosta anterior;
ScP subcosta posterior.
Forewing with transparent spots on veins, and dark spots on the graduate crossveins, darker pigmentation along wing margin, subcostal veinlets, and longitudinal veins with dark intervals or dots. MA and MP pectinately forked, 2m-cu located at the fork of MA and M, the crossveins of the third gradate series more oblique.
Holotype CNU-NEU-QZ2017001. Only forewing preserved.
Forewing oval, 8.31 mm long, 3.17 mm wide. Trichosors prominent, along the entire wing margin. Setae distinct, scarce on the veins and dense on the margin. Costal space relatively broad, dilated basally. Humeral veinlet recurrent, with two pectinate branches. Presumable ScA present. Majority of subcostal veinlets branched once, several basal veinlets branched twice, with no crossveins between them. Subcostal space moderately broad, with two prestigmal sc-r crossveins: basal 1sc-r and distal 3sc-r. Posterior trace of RA forked apically, with two distal branches. One RA branch forked once, the other twice. RP with three branches (ORBs) originated from RA; ORB1with two pectinate branches between 3r-m and 4r-m, all with distal forks; ORB2 dichotomously forked between the third and fourth gradate series of crossveins, each branch dichotomously forked; ORB3 forked between the second and third gradate series, with two dichotomously forked branches. M appear to be fused basally with R. M forked at 2m-cu; MA, MP configuration similar, parallel for a long distance, then each with two pectinate branches between the third and fourth gradate series. The second branch of MP dichotomously forked. Forewing with three m-cu crossveins. Crossvein 2r-m present and positioned distally to crossvein 2m-cu; 2m-cu at the fork of MA and MP. Cu divided into CuA and CuP proximal of the first gradate series, close to wing base; CuA with four pectinate branches distal to 2cua-cup, all branches with marginal forks; CuP simple, only with marginal fork. AA1 with three pectinate branches, all with marginal forks. AA2 with two simple branches, forked proximal to aa1-aa2. AA3 simple. Three flexion fold (line) distinct between RP and MA, MP and CuA, CuP and AA1. The third gradate series with nine crossveins and the fourth gradate with seven crossveins. Forewing with transparent spots on veins, and dark spots at the graduate crossveins; margined with darker pigmentation, and no other distinct maculation; wing margin, subcostal veinlets and longitudinal veins with dark intervals or dots.
The specific epithet is in honor of the entomologist Dr. Vladimir Nikolaevich Makarkin to acknowledge his great help to the first author in his study of Neuropteran.
Caergen Village, Zeku County, Qinghai Province, China; Garang Formation; The early Miocene.
The species can be easily attributed to the genus Wesmaelius due to the following characters: two prestigmal sc-r crossveins, three RP branches (ORBs), crossvein 2r-m present and positioned distally to crossvein 2m-cu; intersection of crossvein 2m-cu with M not more than the crossvein’s length distal to fork MA/MP (sometimes anterior to this fork), resulting in cell c2m-cu broad distally; forewing with three m-cu crossveins (Oswald, 1993).
In the genus, Wesmaelius makarkini sp. n. is most similar to the extant species of W. nervosus (Fabricius, 1793), W. subnebulosus (Stephens, 1836) and W. reisseri U. Aspöck & H. Aspöck, 1982. The new species with two ORB3 branches, 2m-cu located at the division of MA and MP; while W. nervosus with three ORB3 branches, 2m-cu distal to the division of MA and MP; and W. makarkini with a distinct large darker pigmentation at the apex of forewing. The new species differs from W. subnebulosus and W. reisseri in the pectinately forked MA and RP1, 2m-cua located at the division of MA and MP, instead of dichotomously forked MA and RP1, and 2m-cu distal to the division of MA and MP in W. subnebulosus and W. reisseri. Moreover, the new species has seven crossveins in the fourth series, while W. reisseri has four crossveins.
Named fossil Hemerobiidae have been described from the Late Jurassic to the Miocene, including four extinct genera from the Mesozoic and 11 genera from the Cenozoic (Table
Wesmaelius is an extant genus with approximately 62 extant species and two fossil species from the Eocene of Canada (
We thank Dr. Chungkun Shih (College of Life Sciences, Capital Normal University and National Museum of Natural History, Smithsonian Institution) for his helpful comments, fruitful suggestion and improvement of our manuscript. We are grateful to Dr. Shaun L. Winterton (California State Arthropod Collection, California Department of Food and Agriculture, Sacramento, USA) for his kind help and permitting us to check the extant hemerobiids specimens. We sincerely thank Drs Yunzhi Yao and Junjie Gu, Zhipeng Zhao, Lei Li, Yizi Cao, Yingnan Li, Siyuan Wu, and He Tian for collecting specimens. We appreciate the valuable comments and useful suggestions on our manuscript from reviewers and the editor. This research was funded by grants from key project of Science-technology basic condition platform from The Ministry of Science and Technology of the People’s Republic of China (grant no. 2005DKA21402), the specimen platform of China, teaching specimens sub-platform, Web, http://mnh.scu.edu.cn/, National Natural Science Foundation of China (grant nos. 41602014, 31501881, 41402009, 31730087), China Postdoctoral Science Foundation (grant no. 2016M592570), Natural Science Foundation of Hebei Province (grant no. C2015403012), Basal Research Fund of SYSU (grant nos. 32110-41030349) and Program for Changjiang Scholars and Innovative Research Team in University (grant no. IRT-17R75).