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New genus and species of Empheriidae (Psocodea: Trogiomorpha) from mid-Cretaceous amber of northern Myanmar
Journal Pre-proof New genus and species of Empheriidae (Psocodea: Trogiomorpha) from midCretaceous amber of northern Myanmar Sheng Li, Qiuzhu Wang, Dong Ren, Yunzhi Yao PII:
S0195-6671(19)30388-X
DOI:
https://doi.org/10.1016/j.cretres.2020.104421
Reference:
YCRES 104421
To appear in:
Cretaceous Research
Received Date: 5 September 2019 Revised Date:
30 December 2019
Accepted Date: 11 February 2020
Please cite this article as: Li, S., Wang, Q., Ren, D., Yao, Y., New genus and species of Empheriidae (Psocodea: Trogiomorpha) from mid-Cretaceous amber of northern Myanmar, Cretaceous Research, https://doi.org/10.1016/j.cretres.2020.104421. This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. © 2020 Elsevier Ltd. All rights reserved.
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New genus and species of Empheriidae (Psocodea: Trogiomorpha) from
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mid-Cretaceous amber of northern Myanmar
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Sheng Li, Qiuzhu Wang, Dong Ren, Yunzhi Yao*
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College of Life Sciences and Academy for multidisciplinary Studies, Capital Normal University, 105
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Xisanhuanbeilu, Haidian District, Beijing 100048, China
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*Corresponding author.
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E-mail address: [email protected]
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ABSTRACT
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A new genus, Burmempheria Li, Wang & Yao, gen. nov., with two new species, Burmempheria
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densuschaetae Li, Wang & Yao, sp. nov. and Burmempheria raruschaetae Li, Wang & Yao, sp.
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nov., are assigned to the Empheriidae (Trogiomorpha). This is the first finding of Empheriidae
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from Myanmar amber which enriches the geographical record. The rows of forewing vein setae
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cannot distinguish Empheriidae from Archaeatropidae. Empheriidae and Archaeatropidae may
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synonym.
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Keywords: Myanmar amber; Mesozoic; new taxon; fossil insects
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1. Introduction
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Atropetae belongs to Trogiomorpha, consists of five families, Empheriidae and Archaeatropidae
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are two extinct families with high similarity in morphological characteristics, living period and
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geographical distribution. Currently, six genera with eight species from the early Cretaceous to
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Paleogene have been assigned to Empheriidae, mostly known from Europe and North America.
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Four Cretaceous empheriids have been described: Empheropsocus arilloin Baz & Ortuño, 2001,
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Empheropsocus margineglabrus Baz & Ortuño, 2001, and Preempheria antiqua Baz & Ortuño,
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2001 from Cretaceous amber of Alava, northern Spain; and Jerseyempheria grimaldii Azar, Nel &
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Petrulevičius, 2010 from the Upper Cretaceous of New Jersey. All Cenozoic records are from
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Paleogene: Eoempheria intermedia Nel, Prokop, Ploëg & Millet, 2005 from Oise amber, France;
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Empheria reticulata Hagen, 1856，Empheria pertinens Enderlein, 1911 and Trichempheria villosa
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Hagen, 1882 from Baltic amber.
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In this research, all ambers were collected from the Hukawng Valley, Myitkyina District, Kachin
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State, Myanmar. There has been more research on Myanmar amber over the past few decades (Du
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et al., 2018; Gao et al., 2019; Lin et al., 2019). However, the records of Psocodea are rare, only 9
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families, 16 species within 10 genera have been reported (Ross, 2018; Wang et al., 2019). We
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collected six specimens from Myanmar amber, which preserved many intact characters, such as
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antennae, claws, and gonapophyses. Based on these specimens, we erected a new genus in
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Empheriidae.
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2. Materials and methods
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The amber specimens were collected from Kachin (Hukawng Valley) of the Northern Myanmar,
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and specifically at the north end of Noije Bum which is approximately at 26°15'N, 96°34'E, some
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18 km south-west of the town of Tanai (Shi et al., 2012, Zhang et al., 2018; Fig. 1). The age of
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Myanmar amber is attributed to the earliest Cenomanian (98.79 ± 0.62 Ma) (Shi et al., 2012). The
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specimens are housed in the Key Lab of Insect Evolution and Environmental Changes, College of
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Life Sciences, Capital Normal University, Beijing, China (CNU, Curator: Yunzhi Yao).
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Specimens were examined and photographed under Nikon SMZ25 microscope, with an attached
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Nikon DS-Ri2 digital camera system. Line drawings were edited by Adobe Illustrator CC and
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Adobe Photoshop CC. The morphological terminology mainly follows Smithers (1990) and
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Mockford (1993). Abbreviations: mx, maxillary palp; lc, lacinia; la, labium; Sc, subcostal vein;
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Sc’, first subcostal vein ; R, radius vein; Rs, radial sector; M, median vein; Cu, cubital vein; A,
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anal vein; ep, epiproct; p, paraproct; pa, parameres; s, subgenital plate; V3, external valve.
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3. Systematic palaeontology
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Suborder: Trogiomorpha Roesler, 1940
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Infraorder: Atropetae Pearman, 1936
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Family: Empheriidae Kolbe, 1884
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Included genera: Empheropsocus Baz & Ortuño, 2001; Preempheria Baz & Ortuño, 2001;
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Jerseyempheria Azar, Nel & Petrulevičius, 2010; Eoempheria Nel, Prokop, Ploëg & Millet, 2005;
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Empheria Hagen, 1856; Trichempheria Enderlein, 1911.
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Type genus: Empheria Enderlein, 1911
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Burmempheria Li, Wang & Yao, gen. nov.
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Etymology. The generic name is a combination of Latin words “Burma” (Myanmar) and
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“empheria” (type genera of Empheriidae). This designation refers to the locality of new genus.
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The gender is female.
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Type species. Burmempheria densuschaetae Li, Wang & Yao, sp. nov.
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Diagnosis: flagellum more than thirty segments; forewing Sc long, and reaching to the anterior
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margin; Cu1a and Cu1b almost parallel; Cu2 about 2/3 length of Cu1, Cu2 and 1A fused before
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reaching to posterior margin; veins with long setae except Cu2; distal part of tibiae with two apical
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spurs; lack of pulvillus.
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Remarks. Burmempheria Li, Wang & Yao, gen. nov. share series of characters with Trogiomorpha:
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(1) antennae with more than twenty segments; (2) labial palps two-segmented, with minute basal
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segment and rounded or elongated distal segment; (3) tarsi with three segments; (4) forewing
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pterostigma slightly opaque; (5) ventral and dorsal valves of gonapophyses strongly reduced (or
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absent), external valves well-developed and setose; (6) subgenital plate short, covering at most
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basal part of external valves (Yoshizawa et al., 2006). In addition, the Burmempheria Li, Wang &
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Yao, gen. nov. can be clearly classified into Atropetae by the following characters: (1) forewing
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basal segment of Sc well developed; (2) hind wing A simple; (3) paraproct with anal spine; (4)
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external valves of gonapophyses elongated and partially joined together on midline by membrane
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(Yoshizawa et al., 2006). Atropetae include five families, the Burmempheria Li, Wang & Yao, gen.
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nov. can be assigned to Empheriidae according to the following characters: (1) wings
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well-developed, rounded at apex; (2) forewing Sc generally bend to R; (3) venation covered with
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setae; (4) hind wing margin glabrous; (5) claws without preapical tooth (Baz and Ortuño, 2000,
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2001). Besides Burmempheria Li, Wang & Yao, gen. nov. resembles the genera Empheropsocus
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Baz & Ortuño, 2001 from the antennae without secondary annulations, shape of four wings and
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similar venation, thus Burmempheria Li, Wang & Yao, gen. nov. should assigned into
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Empheriidae.
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The main difference between Burmempheria Li, Wang & Yao, gen. nov. and Empheropsocus Baz
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& Ortuño, 2001 are: antennae more than 30 segments (vs. antennae 17-22 segments); forewing
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margin glabrous (vs. forewing margin glabrous in E. margineglabrus Baz & Ortuño, 2001 or
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forewing margin setose in E. arilloi Baz & Ortuño, 2001); Sc well-developed, with a short vein
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reaching to anterior margin (vs. a short vein on the basal sector of Sc, this vein can reaching to
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anterior margin or the distal sector of Sc forming a polygonal cell); with two apical spurs (vs. with
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three apical spurs); pulvillus absent (vs. pulvillus broadened at the tip).
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Burmempheria Li, Wang & Yao, gen. nov. can be distinguished from other genera of Empheriidae
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by following characters: antennae more than 30 segments (vs. antennae 19-28 segments);
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forewing Sc curved, joined to R, a short vein arises from Sc ending on the anterior margin (vs.
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forewing Sc curved back to reach R and fused for a distance); forewing veins with Sc, Cu and A
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with long setae arranged along both edges of vein, and other veins with a row of setae (vs.
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forewing veins with setae along both sides of the vein or forewing membrane densely setose).
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There are obvious differences between Burmempheria Li, Wang & Yao, gen. nov. and the genus in
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Archaeatropidae, based on: antennae without secondarily annulated (vs. antennae with secondarily
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annulated); the distal sector of Sc is directed backwards and a short vein arises from Sc reaching
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to the anterior margin (vs. the distal sector of Sc is directed backwards); the areola postica narrow
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and long (vs. the areola postica short); forewing veins with Sc, Cu and A with long setae arranged
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along both edges of vein, other veins with a row of setae (vs. forewing veins with a single row of
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setae).
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Burmempheria densuchaetae Li, Wang & Yao, sp. nov. (Figs. 1, 2)
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Etymology: A combination of Latin words densus (dense) and chaetae (setae), indicating that tibia
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covered with dense setae.
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Material. Holotype, CNU-PSO-MA-2019001, female. Paratype, CNU-PSO-MA-2019002, male
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with well-persevered wings and mouthparts, body almost broken; CNU-PSO-MA-2019003, sex
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unknown, wings well preserved, abdomen damaged.
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Locality and horizon. Hukawng Valley, Kachin State, Northern Myanmar; mid-Cretaceous,
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lowermost Cenomanian
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Diagnosis: vertex covered with long and dense setae; flagellum basal nine segments with long
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setae in distal part; forewing R1 slightly curved; tibia covered with long and dense setae.
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Description. Female (based on holotype specimen CNU-PSO-MA-2019001). Complete insect
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with well-preserved wings and body (Fig. 1).
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Head: Vertex broad, covered with long and dense setae (Fig. 2A, B); compound eyes small,
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diameter less than the 1/2 length of interorbital distance; three ocelli present, arranged in inverted
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triangle. Antennae long, left flagellum with 32 flagellomeres, right flagellum with 30
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flagellomeres, basal 9 segments covered with long setae, secondary annulations absent (Fig. 2C,
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D). Maxillary palps with four segments, covered with setae, terminal article hatchet-shaped, the
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second article longest, the second and fourth articles covered with setae; labrum and labium
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well-preserved; labium palps with two articles, terminal segment rounded (Fig. 2A, B).
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Thorax: Prothorax broad, prosternum with wrinkle; mesothorax well developed, mesonotum
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triangular, arthropleure extruded.
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Legs: Three legs covered with setae; all tibiae with densely setae, with two apical spurs (Fig. 2E,
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F); tarsi three-segmented, terminal segment with two claws, without pulvillus.
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Wings: Macropterous, forewing oval (Fig.1D), margin glabrous, veins with long setae except Cu2,
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membrane glabrous except vannal region. Sc, Cu and A with long setae arranged along both edges
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of vein, other veins with a row of setae; Sc long and curved and fused with R for a short distance,
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a short vein arises from distal part of Sc reaching to the anterior margin; Sc’ long, pterostigma
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quadrilateral, not thicken; R1 slightly curved, basal part with a crossvein joined to Rs, Rs and M
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fused together at base; Rs forking distal to M1+2; Cu1a and Cu1b fused together at base, Cu1a
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slightly curved, longer than Cu1b; Cu2 bare; A not branch, slightly curved; Cu2 and 1A merged for
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a distance (Fig. 2G).
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Hind wing glabrous (Fig. 1E), Sc short, reaching to wing margin nearly 2/3 length; basi-radial cell
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triangular, slender; R1 and Rs+M connected by a short crossvein; Rs with two branches, Rs
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forking distal to M forking; M with two branches, M1 and M2 fused together at base; Cu1 single,
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Cu2 faked; A single.
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Abdomen: Female epiproct and paraproct covered with strong setae; dorsal valves degraded,
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external valves elongated and partially joined together on midline by membrane (Fig. 2H). Male
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genitalia absent.
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Burmempheria raruschaetae Li, Wang & Yao, sp. nov. (Figs. 3, 4)
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Etymology: A combination of Latin words rarus (sparse) and chaetae (setae), indicating that tibia
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lack of long setae.
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Material. Holotype, CNU-PSO-MA-2019004, male. Paratype, CNU-PSO-MA-2019005, sex
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unknown, the specimen with impurities, thorax broken, abdomen covered with impurities.
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CNU-PSO-MA-2019006, male, antennae not completely preserved, wings preserved well, around
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the body with crack in amber.
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Diagnosis: vertex sparse covered with long setae; the basal 5 segments of flagellum with long
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setae at apical; forewing R1 strongly curved; tibia lack of long spurs.
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Description. Male (based on holotype specimen CNU-PSO-MA-2019004). Complete insect with
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well-preserved wings and body (Fig. 3).
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Head: vertex broad, covered with sparse setae; compound eyes large, diameter almost nearly 1/2
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length of interorbital distance, three ocelli present, arranged in inverted triangle (Fig. 3A); ecdysial
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suture well-developed; antennae long, left flagellum with 27 flagellomeres, right with 31
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flagellomeres, basal five segments with long setae, secondary annulations absent (Fig. 4A, B);
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mouthpart well-persevered, maxillary palps with four segments, covered with setae, mx2 longer
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than mx4 , the fourth segment hatchet-shaped (Fig. 4C). Labial palps with two segments, terminal
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segment rounded, covered with sparse setae (Fig. 4D); laciniae absent; cervical region with
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sculptures (Fig. 4E).
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Thorax: prothorax broad, pronotum with wrinkle; mesothorax well-developed, mesonotum
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inverted triangle, arthropleure extruded.
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Legs covered with sparse setae, all tibiae with two apical spurs (Fig. 4F, G); tarsi three segments,
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distal segment bearing claws.
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Wings: Forewing oval (Fig. 3D), margin glabrous, veins covered with long setae except Cu2,
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membrane glabrous except vannal region; Sc, Cu and A covered with long setae arranged along
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both sides of vein, other veins with a row of setae; Sc well-developed, curved back, fused with R
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for a short distance, a short vein arises from distal part of Sc reaching to the anterior margin; Sc’
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long, pterostigma quadrilateral, not thicken; R1 strongly curved, connect with Rs by a crossvein at
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base; Rs and M fused for a short distance; Rs forking distal to the M; Cu1a and Cu1b fused for a
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short distance, Cu1a slightly curved; Cu2 fade; Anal vein not branched; Cu2 and 1A fused for a
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distance and reaching to wing margin together (Fig. 4H).
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Hind wing glabrous (Fig. 3E); Sc short, reaching to margin nearly 2/3 length; basi-radial cell
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triangular; R1 and Rs+M fused for a distance; Rs with two branches, Rs forking distal to M
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forking; M with two branches, basal part of M1 and M2 fused together; Cu1 single; Cu2 faded; Anal
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vein with two branches, basal part fused.
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Abdomen: Male genitalia epiproct and paraproct covered with strong setae, epiproct with anal
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spine; subgential plate covered with short setae, with two triangular projections at apical (Fig. 4I,
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J). Female genitalia shares the same structure with Burmempheria densuchaetae Li, Wang & Yao,
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sp. nov.
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4. Discussion
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Based on the result of Yoshizawa et al., (2006), Atropetae contains three extant families,
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Lepidopsocidae, Trogiidae and Psoquillidae, the monophyly of Atropetae is supported by two
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synapomorphic characters: (1) external valves of gonapophyses elongated and partially joined
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together on midline by membrane, composing the ovipositor; (2) spermathecal sac with one or two
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glandular accessory bodies. Smithers mentioned that the reduction of female genitalia to setose,
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lobar external valve is a main apomorph of Atropetae (Smithers, 1972). Empheriidae and
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Archaeatropidae are two extinct families that were classified by Baz & Ortuño into Atropetae
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based on the reduction of female genitalia (Baz & Ortuño, 2000, 2001). Burmempheria Li, Wang
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& Yao, gen. nov. contains the synapomorphic characters of Atropetae: external valves of
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gonapophyses elongated and partially joined together on midline by membrane (spermathecal sac
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cannot be observed in fossil specimens) (Fig. 2H), also fitted the diagnose characters of Atropetae:
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(1) forewing basal segment of Sc well developed (Figs. 1, 3); (2) hind wing vein A simple (Figs. 1,
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3); (3) wing pilosity well developed (Figs. 1, 3); (4) paraproct with anal spine (Fig. 4I, J). Based
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on our specimens, we agree with Baz & Ortuño (2000, 2001) that there is no dispute about
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Empheriidae assigned to Atropetae.
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In the classification of extant psocids, venation usually not considered as diagnosis feature of
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family (Smithers, 1972; Mockford, 1993; Li, 2002; New et al., 2007). At the same time, different
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environmental conditions have different effect on wing venation development of extant
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Trogiomorpha (Kucerova, 1997). For extinct species, even one individual can express differences
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in venation of left and right wings (Baz & Ortuño, 2000; Wang et al., 2019). In Atropetae and
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Psyllipsocetae, some species showed that the forewing venation has asymmetry (Archaeatropos
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alavensis Baz & Ortuño, 2000; Concavapsocus parallelus Wang, Li, Ren & Yao., 2019) (Baz &
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Ortuño, 2000, Fig. 1; Wang et al., 2019, Fig. 4). Recent research indicates that venation of psocids
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show polymorphism, the difference within Trogiomorpha venation may be intraspecific difference
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(Wang et al., 2019). The results of Baz & Ortuño (2001) indicate that the main difference between
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Empheriidae and Archaeatropidae are the forewing characters: venation and vein setae
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(Empheriidae with two rows of setae along both side of vein vs. Archaeatropidae with only one
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row setae on the vein). The previous researches indicated that the forewing of Prospeleketor
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albianensis Perrichot, Azar, Neraudeau & Nel., 2003 and Proprionoglaris axioperierga Azar, Nel
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& Perrichot, 2014 (Archaeatropidae) with two rows of setae. Moreover, the forewing of
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Jerseyempheria grimaldii Azar, Nel & Petrulevičius, 2010 (Empheriidae) has setae arranged on
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membrane area. However, through our research characters unsteadiness arises not merely on
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venation but also on the vein setae of forewing. In the specimens of CNU-PSO-MA2019001,
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CNU-PSO-MA2019002, CNU-PSO-MA2019004, CNU-PSO-MA2019006, the setae covered on
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forewing Sc, Cu and A arranged in two rows, but distal vein with only one row of setae. Therefore,
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the venation and rows of setae in forewing may not be a family diagnostic character. According to
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the taxonomic about Empheriidae and Archaeatropidae, nodulus was considered as a vital
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character (Baz & Ortuño, 2000, 2001; Mockford et al., 2013). Nevertheless, among the recent
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research, nodulus in forewing is an unstable character (Wang et al., 2019). At the same time, the
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characters of both family has high similarity: (1) the number of antennae; (2) shape of the
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maxillary palpus; (3) wing shape and the venation; (4) absence of denticle on the claw (Hagen,
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1882, 1856; Enderlein, 1911; Baz & Ortuño, 2000, 2001; Perrichot et al., 2003; Nel et al., 2005;
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Azar & Nel, 2010; Azar et al., 2014). Based on the above evidence, Empheriidae and
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Archaeatropidae are closely related, it is possible that these two families are likely to be synonym,
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and the relationship between them needs further study.
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Conclusions
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Two new species of a new genus Burmempheria Li, Wang & Yao, gen. nov. from Myanmar amber
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are assigned to Empheriidae. It’s difficult to define some psocids only based on wing characters.
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Empheriidae and Archaeatropidae may synonym.
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Acknowledgements
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We are grateful to the editor and anonymous reviewers for constructive criticism and valuable
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comments on the manuscript. This project is supported by Joint Fund of the Beijing Municipal
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Natural Science Foundation and Beijing Municipal Education Commission (KZ201810028046),
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grants from the National Natural Science Foundation of China (No. 31970436, 31730087 and
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41688103), the Program for Changjiang Scholars and Innovative Research Team in University
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(IRT-17R75), the Support Project of High-level Teachers in Beijing Municipal Universities in the
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Period of 13th Five-year Plan (IDHT20180518) and Capacity Building for Sci-Tech Innovation -
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Fundamental Scientific Research Funds (No. 19530050144).
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References
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Azar, D., Nel, A., & Petrulevicius, V., 2010. First Psocodean (Psocadea, Empheriidae) from the
241
Cretaceous Amber of New Jersey. Acta Geologica Sinica English Edition 84(4):763
242
Azar, D., Nel, A., & Perrichot, V., 2014. Diverse barklice (Psocodea) from Late Cretaceous
243
Vendean amber. Paleontological Contributions 10:11
244
Baz, A. & Ortuño, V.M. 2000. Archaeatropidae, a new family of Psocoptera from the Cretaceous
245
amber of Alava, northern Spain. Annals of the Entomological Society of America 93, 367–373.
246
Baz, A., and Ortuño, V.M., 2001. New genera and species of empheriids (Psocoptera: Empheriidae)
247
from the Cretaceous amber of Alava, northern Spain. Cretaceous Research 22, 575-584
248
Du, S.L., Yao, Y.Z., 2018. A new genus and species of Tingidae (Heteroptera: Cimicomorpha)
249
from Myanmar, with the analysis of the evolution of hood, carinae and paranota. Zoological
250
Systematics 43(3), 283–293.
251
Enderlein, G., 1911. Die Fossilen Copeognathen und ihre Phylogenie. Palaeontographica 58:
252
279–360.
253
Gao T.P., Yin X.C., Shih C.K., Rasnitsyn A.P., Xu X., Chen S., Wang C. and Ren, D., 2019. New
254
insects feeding on dinosaur feathers in mid-Cretaceous amber. Nature Communications 10, 5424.
255
https://doi: 10.1038/s41467-019-13516-4
256
Hagen, H. 1882. Beitrage zur Monographie der Psociden. über Psociden im Bernstein. Stettiner
257
Entomologische Zeitung 43: 217–237 (in German).
258
Hagen, H.A. 1856. Die im Bernstein befindlichen Neuropteren der Vorwelt. Pp. 1–122 in Berendt,
259
G. C., Pictet de la Rive, F. J. & Hagen, H. A. (eds) Die im Bernstein befindlichen organischen
260
Reste der Vorwelt. Gerhard, Berlin.
261
Kolbe, H.J., 1884. Der Entwickelungsgang der Psociden im Individuum und in der Zeit. Berliner
262
Entomologische Zeitschrift 28, 35-38.
263
Kucerova, Z., 1997. Macropterous form of Dorypteryx domestica (Psocoptera: Psyllipsocidae).
264
European Journal of Entomology 94, 567–573.
265
Li, F.S., 2002. Psocoptera of China. National Natural Science Foundation of China, Science Press,
266
Beijing xlvi + 1976 pp. (two volumes). (in Chinese, with English summary).
267
Lin, X.D., Labandeira, C.C., Shih, C.K., Hotton, C.L., and Ren, D. (2019). Life habits and
268
evolutionary biology of new two-winged long-proboscid scorpionflies from mid-Cretaceous
269
Myanmar amber. Nature Communications 10: 1235.
270
Mockford, E.L., 1993. North American Psocoptera (Insecta). Flora & Fauna Handbook 10, 455pp.
271
Mockford, E.L., Lienhard, C., Yoshizawa, K., 2013. Revised classification of “Psocoptera” from
272
Cretaceous amber, a reassessment of published information. Insecta Matsumurana 69, 1-26.
273
Nel. A., Prokop, J., De Ploeg, G., and Millet, J., 2005. New Psocoptera (Insecta) from the
274
lowermost Eocene amber of Oise, France. Journal of Systematic Palaeonotology, 3(4): 371-391
275
New, T.R., Lienhard, C., 2007. The Psocoptera of Tropical South-east Asia. Fauna Malesina
276
Handbook 6. National Museum of Natural History Naturalis, Leiden, 48–168.
277
Pearman, J.V., 1936. The taxonomy of the Psocoptera: preliminary sketch. Proceedings of the
278
Royal Entomological Society, London (B) 5: 58–62.
279
Prerrichot, V., Azar, D., Néraudeau, D., Nel, A., 2003. New Psocoptera in the Early Cretaceous
280
amber of SW France and Lebanon (Insecta: Psocoptera: Trogiomorpha). Geological Magazine 140
281
(6), 669–683.
282
Ross, A.J., 2018. Burmese (Myanmar) amber taxa, on-line checklist v.2018.2 from.
283
https://www.nms.ac.uk/explore-our-cllections/stories/natural-would/burmese-amber.
284
Roesler, R., 1940. Neue und wenig bekannte Copeognathengattungen. I. Zoologischer Anzeiger
285
129 (9/10), 225-243.
286
Smithers, C.N., 1990. Keys to the family and genera of Psocoptera (Arthropoda: Insecta).
287
Technical Reports of Australian Museum, 1–82.
288
Smithers, C.N., 1972. The classification and phylogeny of the Psocoptera. Memoirs of the
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Australian Museum 14, 1–349.
290
Shi, G., Grimaldi, D.A., Harlow, G.E., Wang, J., Wang, J., Wang, M., Lei, W., Li, Q., Li, X., 2012.
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Age constraint on Burmese amber based on U-Pb dating of zicons. Cretaceous Research 37, 155–
292
163
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Wang, R., Li, S., Ren, D., Yao, Y., New genus and species of the Psyllipsocidae (Psocodea:
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Trogiomorpha)
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https://doi.org/10.1016/j.cretres.2019.07.008.
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Yoshizawa, K., Lienhard, C., Johnson, K.P., 2006. Molecular systematics of the suborder
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Trogiomorpha (Insecta: Psocodea: “Psocoptera”). Zoological Journal of the Linnean Society 146,
298
287–299.
299
Zhang, X., Ren, D., Yao, Y.Z., 2018. A new genus and species of Mimarachnidae (Hemiptera:
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Fulgoromprpha: Fulgoroidea) from mid-Cretaceous Burmese amber. Cretaceous Research 90,
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168–173.
from
mid-Cretaceous
Burmese
amber,
Cretaceous
Research,
302
303
FIGURE CAPTIONS
304
Fig.
305
CNU-PSO-MA2019001. (A) Photograph in dorsal view; (B) Photograph in ventral view; (C) Line
1.
Habitus
of
Burmempheria
densuschaetae
Li,
Wang
&
Yao,
sp.
nov.,
306
drawing in dorsal view. (D) Line drawing of forewing; (E) Line drawing of hind wing; Scale bars:
307
0.5mm.
308
Fig. 2. Detailed photographs of Burmempheria densuschaetae Li, Wang & Yao, sp. nov. (A)
309
Photograph of mouthpart; (B) Line drawing of mouthpart; (C) Photograph of left antenna; (D)
310
Line drawing of left antenna; (E) Photograph of left mid tibia; (F) Line drawing of left mid tibia;
311
(G) Photograph of Cu2 and 1A in left forewing; (H) Photograph of female genitalia; Scale bars: (A,
312
B, C, D, E, F) 0.1mm; (G, H) 0.5mm.
313
Fig. 3. Habitus of Burmempheria raruschaetae Li, Wang & Yao, sp. nov., CNU-PSO-MA2019004.
314
(A) Photograph in dorsal view; (B) Photograph in ventral view; (C) Line drawing in dorsal view.
315
(D) Line drawing of forewing; (E) Line drawing of hind wing; Scale bars: 0.5mm.
316
Fig. 4. Detailed photographs of Burmempheria raruschaetae Li, Wang & Yao, sp. nov. (A)
317
Photograph of left antenna; (B) Line drawing of left antenna; (C) Line drawing of right maxillary
318
palpus in ventral view; (D) Line drawing of labial palps; (E) Photograph of mouth part; (F)
319
Photograph of right hind tibia; (G) Line drawing of right hind tibia; (H) Photograph of Cu2 and 1A
320
in left forewing ; (I) Photograph of male genitalia; (J) Line drawing of male genitalia; Scale bars:
321
(A, B, C, D, H) 0.1mm; (I, J) 0.25mm; (F, G) 0.5mm.
S.L., Q.Z.W and Y.Z.Y designed the research. S.L. and Q.Z.W took the photographs and prepared the line drawings. S.L., D.R. and Y.Z.Y. performed morphological, and drafted the paper.
The authors declare no competing interests.
S0195-6671(19)30388-X
DOI:
https://doi.org/10.1016/j.cretres.2020.104421
Reference:
YCRES 104421
To appear in:
Cretaceous Research
Received Date: 5 September 2019 Revised Date:
30 December 2019
Accepted Date: 11 February 2020
Please cite this article as: Li, S., Wang, Q., Ren, D., Yao, Y., New genus and species of Empheriidae (Psocodea: Trogiomorpha) from mid-Cretaceous amber of northern Myanmar, Cretaceous Research, https://doi.org/10.1016/j.cretres.2020.104421. This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. © 2020 Elsevier Ltd. All rights reserved.
1
New genus and species of Empheriidae (Psocodea: Trogiomorpha) from
2
mid-Cretaceous amber of northern Myanmar
3 4
Sheng Li, Qiuzhu Wang, Dong Ren, Yunzhi Yao*
5
College of Life Sciences and Academy for multidisciplinary Studies, Capital Normal University, 105
6
Xisanhuanbeilu, Haidian District, Beijing 100048, China
7
*Corresponding author.
8
E-mail address: [email protected]
9
ABSTRACT
10
A new genus, Burmempheria Li, Wang & Yao, gen. nov., with two new species, Burmempheria
11
densuschaetae Li, Wang & Yao, sp. nov. and Burmempheria raruschaetae Li, Wang & Yao, sp.
12
nov., are assigned to the Empheriidae (Trogiomorpha). This is the first finding of Empheriidae
13
from Myanmar amber which enriches the geographical record. The rows of forewing vein setae
14
cannot distinguish Empheriidae from Archaeatropidae. Empheriidae and Archaeatropidae may
15
synonym.
16
Keywords: Myanmar amber; Mesozoic; new taxon; fossil insects
17
1. Introduction
18
Atropetae belongs to Trogiomorpha, consists of five families, Empheriidae and Archaeatropidae
19
are two extinct families with high similarity in morphological characteristics, living period and
20
geographical distribution. Currently, six genera with eight species from the early Cretaceous to
21
Paleogene have been assigned to Empheriidae, mostly known from Europe and North America.
22
Four Cretaceous empheriids have been described: Empheropsocus arilloin Baz & Ortuño, 2001,
23
Empheropsocus margineglabrus Baz & Ortuño, 2001, and Preempheria antiqua Baz & Ortuño,
24
2001 from Cretaceous amber of Alava, northern Spain; and Jerseyempheria grimaldii Azar, Nel &
25
Petrulevičius, 2010 from the Upper Cretaceous of New Jersey. All Cenozoic records are from
26
Paleogene: Eoempheria intermedia Nel, Prokop, Ploëg & Millet, 2005 from Oise amber, France;
27
Empheria reticulata Hagen, 1856，Empheria pertinens Enderlein, 1911 and Trichempheria villosa
28
Hagen, 1882 from Baltic amber.
29
In this research, all ambers were collected from the Hukawng Valley, Myitkyina District, Kachin
30
State, Myanmar. There has been more research on Myanmar amber over the past few decades (Du
31
et al., 2018; Gao et al., 2019; Lin et al., 2019). However, the records of Psocodea are rare, only 9
32
families, 16 species within 10 genera have been reported (Ross, 2018; Wang et al., 2019). We
33
collected six specimens from Myanmar amber, which preserved many intact characters, such as
34
antennae, claws, and gonapophyses. Based on these specimens, we erected a new genus in
35
Empheriidae.
36
2. Materials and methods
37
The amber specimens were collected from Kachin (Hukawng Valley) of the Northern Myanmar,
38
and specifically at the north end of Noije Bum which is approximately at 26°15'N, 96°34'E, some
39
18 km south-west of the town of Tanai (Shi et al., 2012, Zhang et al., 2018; Fig. 1). The age of
40
Myanmar amber is attributed to the earliest Cenomanian (98.79 ± 0.62 Ma) (Shi et al., 2012). The
41
specimens are housed in the Key Lab of Insect Evolution and Environmental Changes, College of
42
Life Sciences, Capital Normal University, Beijing, China (CNU, Curator: Yunzhi Yao).
43
Specimens were examined and photographed under Nikon SMZ25 microscope, with an attached
44
Nikon DS-Ri2 digital camera system. Line drawings were edited by Adobe Illustrator CC and
45
Adobe Photoshop CC. The morphological terminology mainly follows Smithers (1990) and
46
Mockford (1993). Abbreviations: mx, maxillary palp; lc, lacinia; la, labium; Sc, subcostal vein;
47
Sc’, first subcostal vein ; R, radius vein; Rs, radial sector; M, median vein; Cu, cubital vein; A,
48
anal vein; ep, epiproct; p, paraproct; pa, parameres; s, subgenital plate; V3, external valve.
49
3. Systematic palaeontology
50
Suborder: Trogiomorpha Roesler, 1940
51
Infraorder: Atropetae Pearman, 1936
52
Family: Empheriidae Kolbe, 1884
53
Included genera: Empheropsocus Baz & Ortuño, 2001; Preempheria Baz & Ortuño, 2001;
54
Jerseyempheria Azar, Nel & Petrulevičius, 2010; Eoempheria Nel, Prokop, Ploëg & Millet, 2005;
55
Empheria Hagen, 1856; Trichempheria Enderlein, 1911.
56
Type genus: Empheria Enderlein, 1911
57
Burmempheria Li, Wang & Yao, gen. nov.
58
Etymology. The generic name is a combination of Latin words “Burma” (Myanmar) and
59
“empheria” (type genera of Empheriidae). This designation refers to the locality of new genus.
60
The gender is female.
61
Type species. Burmempheria densuschaetae Li, Wang & Yao, sp. nov.
62
Diagnosis: flagellum more than thirty segments; forewing Sc long, and reaching to the anterior
63
margin; Cu1a and Cu1b almost parallel; Cu2 about 2/3 length of Cu1, Cu2 and 1A fused before
64
reaching to posterior margin; veins with long setae except Cu2; distal part of tibiae with two apical
65
spurs; lack of pulvillus.
66
Remarks. Burmempheria Li, Wang & Yao, gen. nov. share series of characters with Trogiomorpha:
67
(1) antennae with more than twenty segments; (2) labial palps two-segmented, with minute basal
68
segment and rounded or elongated distal segment; (3) tarsi with three segments; (4) forewing
69
pterostigma slightly opaque; (5) ventral and dorsal valves of gonapophyses strongly reduced (or
70
absent), external valves well-developed and setose; (6) subgenital plate short, covering at most
71
basal part of external valves (Yoshizawa et al., 2006). In addition, the Burmempheria Li, Wang &
72
Yao, gen. nov. can be clearly classified into Atropetae by the following characters: (1) forewing
73
basal segment of Sc well developed; (2) hind wing A simple; (3) paraproct with anal spine; (4)
74
external valves of gonapophyses elongated and partially joined together on midline by membrane
75
(Yoshizawa et al., 2006). Atropetae include five families, the Burmempheria Li, Wang & Yao, gen.
76
nov. can be assigned to Empheriidae according to the following characters: (1) wings
77
well-developed, rounded at apex; (2) forewing Sc generally bend to R; (3) venation covered with
78
setae; (4) hind wing margin glabrous; (5) claws without preapical tooth (Baz and Ortuño, 2000,
79
2001). Besides Burmempheria Li, Wang & Yao, gen. nov. resembles the genera Empheropsocus
80
Baz & Ortuño, 2001 from the antennae without secondary annulations, shape of four wings and
81
similar venation, thus Burmempheria Li, Wang & Yao, gen. nov. should assigned into
82
Empheriidae.
83
The main difference between Burmempheria Li, Wang & Yao, gen. nov. and Empheropsocus Baz
84
& Ortuño, 2001 are: antennae more than 30 segments (vs. antennae 17-22 segments); forewing
85
margin glabrous (vs. forewing margin glabrous in E. margineglabrus Baz & Ortuño, 2001 or
86
forewing margin setose in E. arilloi Baz & Ortuño, 2001); Sc well-developed, with a short vein
87
reaching to anterior margin (vs. a short vein on the basal sector of Sc, this vein can reaching to
88
anterior margin or the distal sector of Sc forming a polygonal cell); with two apical spurs (vs. with
89
three apical spurs); pulvillus absent (vs. pulvillus broadened at the tip).
90
Burmempheria Li, Wang & Yao, gen. nov. can be distinguished from other genera of Empheriidae
91
by following characters: antennae more than 30 segments (vs. antennae 19-28 segments);
92
forewing Sc curved, joined to R, a short vein arises from Sc ending on the anterior margin (vs.
93
forewing Sc curved back to reach R and fused for a distance); forewing veins with Sc, Cu and A
94
with long setae arranged along both edges of vein, and other veins with a row of setae (vs.
95
forewing veins with setae along both sides of the vein or forewing membrane densely setose).
96
There are obvious differences between Burmempheria Li, Wang & Yao, gen. nov. and the genus in
97
Archaeatropidae, based on: antennae without secondarily annulated (vs. antennae with secondarily
98
annulated); the distal sector of Sc is directed backwards and a short vein arises from Sc reaching
99
to the anterior margin (vs. the distal sector of Sc is directed backwards); the areola postica narrow
100
and long (vs. the areola postica short); forewing veins with Sc, Cu and A with long setae arranged
101
along both edges of vein, other veins with a row of setae (vs. forewing veins with a single row of
102
setae).
103
Burmempheria densuchaetae Li, Wang & Yao, sp. nov. (Figs. 1, 2)
104
Etymology: A combination of Latin words densus (dense) and chaetae (setae), indicating that tibia
105
covered with dense setae.
106
Material. Holotype, CNU-PSO-MA-2019001, female. Paratype, CNU-PSO-MA-2019002, male
107
with well-persevered wings and mouthparts, body almost broken; CNU-PSO-MA-2019003, sex
108
unknown, wings well preserved, abdomen damaged.
109
Locality and horizon. Hukawng Valley, Kachin State, Northern Myanmar; mid-Cretaceous,
110
lowermost Cenomanian
111
Diagnosis: vertex covered with long and dense setae; flagellum basal nine segments with long
112
setae in distal part; forewing R1 slightly curved; tibia covered with long and dense setae.
113
Description. Female (based on holotype specimen CNU-PSO-MA-2019001). Complete insect
114
with well-preserved wings and body (Fig. 1).
115
Head: Vertex broad, covered with long and dense setae (Fig. 2A, B); compound eyes small,
116
diameter less than the 1/2 length of interorbital distance; three ocelli present, arranged in inverted
117
triangle. Antennae long, left flagellum with 32 flagellomeres, right flagellum with 30
118
flagellomeres, basal 9 segments covered with long setae, secondary annulations absent (Fig. 2C,
119
D). Maxillary palps with four segments, covered with setae, terminal article hatchet-shaped, the
120
second article longest, the second and fourth articles covered with setae; labrum and labium
121
well-preserved; labium palps with two articles, terminal segment rounded (Fig. 2A, B).
122
Thorax: Prothorax broad, prosternum with wrinkle; mesothorax well developed, mesonotum
123
triangular, arthropleure extruded.
124
Legs: Three legs covered with setae; all tibiae with densely setae, with two apical spurs (Fig. 2E,
125
F); tarsi three-segmented, terminal segment with two claws, without pulvillus.
126
Wings: Macropterous, forewing oval (Fig.1D), margin glabrous, veins with long setae except Cu2,
127
membrane glabrous except vannal region. Sc, Cu and A with long setae arranged along both edges
128
of vein, other veins with a row of setae; Sc long and curved and fused with R for a short distance,
129
a short vein arises from distal part of Sc reaching to the anterior margin; Sc’ long, pterostigma
130
quadrilateral, not thicken; R1 slightly curved, basal part with a crossvein joined to Rs, Rs and M
131
fused together at base; Rs forking distal to M1+2; Cu1a and Cu1b fused together at base, Cu1a
132
slightly curved, longer than Cu1b; Cu2 bare; A not branch, slightly curved; Cu2 and 1A merged for
133
a distance (Fig. 2G).
134
Hind wing glabrous (Fig. 1E), Sc short, reaching to wing margin nearly 2/3 length; basi-radial cell
135
triangular, slender; R1 and Rs+M connected by a short crossvein; Rs with two branches, Rs
136
forking distal to M forking; M with two branches, M1 and M2 fused together at base; Cu1 single,
137
Cu2 faked; A single.
138
Abdomen: Female epiproct and paraproct covered with strong setae; dorsal valves degraded,
139
external valves elongated and partially joined together on midline by membrane (Fig. 2H). Male
140
genitalia absent.
141
Burmempheria raruschaetae Li, Wang & Yao, sp. nov. (Figs. 3, 4)
142
Etymology: A combination of Latin words rarus (sparse) and chaetae (setae), indicating that tibia
143
lack of long setae.
144
Material. Holotype, CNU-PSO-MA-2019004, male. Paratype, CNU-PSO-MA-2019005, sex
145
unknown, the specimen with impurities, thorax broken, abdomen covered with impurities.
146
CNU-PSO-MA-2019006, male, antennae not completely preserved, wings preserved well, around
147
the body with crack in amber.
148
Diagnosis: vertex sparse covered with long setae; the basal 5 segments of flagellum with long
149
setae at apical; forewing R1 strongly curved; tibia lack of long spurs.
150
Description. Male (based on holotype specimen CNU-PSO-MA-2019004). Complete insect with
151
well-preserved wings and body (Fig. 3).
152
Head: vertex broad, covered with sparse setae; compound eyes large, diameter almost nearly 1/2
153
length of interorbital distance, three ocelli present, arranged in inverted triangle (Fig. 3A); ecdysial
154
suture well-developed; antennae long, left flagellum with 27 flagellomeres, right with 31
155
flagellomeres, basal five segments with long setae, secondary annulations absent (Fig. 4A, B);
156
mouthpart well-persevered, maxillary palps with four segments, covered with setae, mx2 longer
157
than mx4 , the fourth segment hatchet-shaped (Fig. 4C). Labial palps with two segments, terminal
158
segment rounded, covered with sparse setae (Fig. 4D); laciniae absent; cervical region with
159
sculptures (Fig. 4E).
160
Thorax: prothorax broad, pronotum with wrinkle; mesothorax well-developed, mesonotum
161
inverted triangle, arthropleure extruded.
162
Legs covered with sparse setae, all tibiae with two apical spurs (Fig. 4F, G); tarsi three segments,
163
distal segment bearing claws.
164
Wings: Forewing oval (Fig. 3D), margin glabrous, veins covered with long setae except Cu2,
165
membrane glabrous except vannal region; Sc, Cu and A covered with long setae arranged along
166
both sides of vein, other veins with a row of setae; Sc well-developed, curved back, fused with R
167
for a short distance, a short vein arises from distal part of Sc reaching to the anterior margin; Sc’
168
long, pterostigma quadrilateral, not thicken; R1 strongly curved, connect with Rs by a crossvein at
169
base; Rs and M fused for a short distance; Rs forking distal to the M; Cu1a and Cu1b fused for a
170
short distance, Cu1a slightly curved; Cu2 fade; Anal vein not branched; Cu2 and 1A fused for a
171
distance and reaching to wing margin together (Fig. 4H).
172
Hind wing glabrous (Fig. 3E); Sc short, reaching to margin nearly 2/3 length; basi-radial cell
173
triangular; R1 and Rs+M fused for a distance; Rs with two branches, Rs forking distal to M
174
forking; M with two branches, basal part of M1 and M2 fused together; Cu1 single; Cu2 faded; Anal
175
vein with two branches, basal part fused.
176
Abdomen: Male genitalia epiproct and paraproct covered with strong setae, epiproct with anal
177
spine; subgential plate covered with short setae, with two triangular projections at apical (Fig. 4I,
178
J). Female genitalia shares the same structure with Burmempheria densuchaetae Li, Wang & Yao,
179
sp. nov.
180
4. Discussion
181
Based on the result of Yoshizawa et al., (2006), Atropetae contains three extant families,
182
Lepidopsocidae, Trogiidae and Psoquillidae, the monophyly of Atropetae is supported by two
183
synapomorphic characters: (1) external valves of gonapophyses elongated and partially joined
184
together on midline by membrane, composing the ovipositor; (2) spermathecal sac with one or two
185
glandular accessory bodies. Smithers mentioned that the reduction of female genitalia to setose,
186
lobar external valve is a main apomorph of Atropetae (Smithers, 1972). Empheriidae and
187
Archaeatropidae are two extinct families that were classified by Baz & Ortuño into Atropetae
188
based on the reduction of female genitalia (Baz & Ortuño, 2000, 2001). Burmempheria Li, Wang
189
& Yao, gen. nov. contains the synapomorphic characters of Atropetae: external valves of
190
gonapophyses elongated and partially joined together on midline by membrane (spermathecal sac
191
cannot be observed in fossil specimens) (Fig. 2H), also fitted the diagnose characters of Atropetae:
192
(1) forewing basal segment of Sc well developed (Figs. 1, 3); (2) hind wing vein A simple (Figs. 1,
193
3); (3) wing pilosity well developed (Figs. 1, 3); (4) paraproct with anal spine (Fig. 4I, J). Based
194
on our specimens, we agree with Baz & Ortuño (2000, 2001) that there is no dispute about
195
Empheriidae assigned to Atropetae.
196
In the classification of extant psocids, venation usually not considered as diagnosis feature of
197
family (Smithers, 1972; Mockford, 1993; Li, 2002; New et al., 2007). At the same time, different
198
environmental conditions have different effect on wing venation development of extant
199
Trogiomorpha (Kucerova, 1997). For extinct species, even one individual can express differences
200
in venation of left and right wings (Baz & Ortuño, 2000; Wang et al., 2019). In Atropetae and
201
Psyllipsocetae, some species showed that the forewing venation has asymmetry (Archaeatropos
202
alavensis Baz & Ortuño, 2000; Concavapsocus parallelus Wang, Li, Ren & Yao., 2019) (Baz &
203
Ortuño, 2000, Fig. 1; Wang et al., 2019, Fig. 4). Recent research indicates that venation of psocids
204
show polymorphism, the difference within Trogiomorpha venation may be intraspecific difference
205
(Wang et al., 2019). The results of Baz & Ortuño (2001) indicate that the main difference between
206
Empheriidae and Archaeatropidae are the forewing characters: venation and vein setae
207
(Empheriidae with two rows of setae along both side of vein vs. Archaeatropidae with only one
208
row setae on the vein). The previous researches indicated that the forewing of Prospeleketor
209
albianensis Perrichot, Azar, Neraudeau & Nel., 2003 and Proprionoglaris axioperierga Azar, Nel
210
& Perrichot, 2014 (Archaeatropidae) with two rows of setae. Moreover, the forewing of
211
Jerseyempheria grimaldii Azar, Nel & Petrulevičius, 2010 (Empheriidae) has setae arranged on
212
membrane area. However, through our research characters unsteadiness arises not merely on
213
venation but also on the vein setae of forewing. In the specimens of CNU-PSO-MA2019001,
214
CNU-PSO-MA2019002, CNU-PSO-MA2019004, CNU-PSO-MA2019006, the setae covered on
215
forewing Sc, Cu and A arranged in two rows, but distal vein with only one row of setae. Therefore,
216
the venation and rows of setae in forewing may not be a family diagnostic character. According to
217
the taxonomic about Empheriidae and Archaeatropidae, nodulus was considered as a vital
218
character (Baz & Ortuño, 2000, 2001; Mockford et al., 2013). Nevertheless, among the recent
219
research, nodulus in forewing is an unstable character (Wang et al., 2019). At the same time, the
220
characters of both family has high similarity: (1) the number of antennae; (2) shape of the
221
maxillary palpus; (3) wing shape and the venation; (4) absence of denticle on the claw (Hagen,
222
1882, 1856; Enderlein, 1911; Baz & Ortuño, 2000, 2001; Perrichot et al., 2003; Nel et al., 2005;
223
Azar & Nel, 2010; Azar et al., 2014). Based on the above evidence, Empheriidae and
224
Archaeatropidae are closely related, it is possible that these two families are likely to be synonym,
225
and the relationship between them needs further study.
226
Conclusions
227
Two new species of a new genus Burmempheria Li, Wang & Yao, gen. nov. from Myanmar amber
228
are assigned to Empheriidae. It’s difficult to define some psocids only based on wing characters.
229
Empheriidae and Archaeatropidae may synonym.
230
Acknowledgements
231
We are grateful to the editor and anonymous reviewers for constructive criticism and valuable
232
comments on the manuscript. This project is supported by Joint Fund of the Beijing Municipal
233
Natural Science Foundation and Beijing Municipal Education Commission (KZ201810028046),
234
grants from the National Natural Science Foundation of China (No. 31970436, 31730087 and
235
41688103), the Program for Changjiang Scholars and Innovative Research Team in University
236
(IRT-17R75), the Support Project of High-level Teachers in Beijing Municipal Universities in the
237
Period of 13th Five-year Plan (IDHT20180518) and Capacity Building for Sci-Tech Innovation -
238
Fundamental Scientific Research Funds (No. 19530050144).
239
References
240
Azar, D., Nel, A., & Petrulevicius, V., 2010. First Psocodean (Psocadea, Empheriidae) from the
241
Cretaceous Amber of New Jersey. Acta Geologica Sinica English Edition 84(4):763
242
Azar, D., Nel, A., & Perrichot, V., 2014. Diverse barklice (Psocodea) from Late Cretaceous
243
Vendean amber. Paleontological Contributions 10:11
244
Baz, A. & Ortuño, V.M. 2000. Archaeatropidae, a new family of Psocoptera from the Cretaceous
245
amber of Alava, northern Spain. Annals of the Entomological Society of America 93, 367–373.
246
Baz, A., and Ortuño, V.M., 2001. New genera and species of empheriids (Psocoptera: Empheriidae)
247
from the Cretaceous amber of Alava, northern Spain. Cretaceous Research 22, 575-584
248
Du, S.L., Yao, Y.Z., 2018. A new genus and species of Tingidae (Heteroptera: Cimicomorpha)
249
from Myanmar, with the analysis of the evolution of hood, carinae and paranota. Zoological
250
Systematics 43(3), 283–293.
251
Enderlein, G., 1911. Die Fossilen Copeognathen und ihre Phylogenie. Palaeontographica 58:
252
279–360.
253
Gao T.P., Yin X.C., Shih C.K., Rasnitsyn A.P., Xu X., Chen S., Wang C. and Ren, D., 2019. New
254
insects feeding on dinosaur feathers in mid-Cretaceous amber. Nature Communications 10, 5424.
255
https://doi: 10.1038/s41467-019-13516-4
256
Hagen, H. 1882. Beitrage zur Monographie der Psociden. über Psociden im Bernstein. Stettiner
257
Entomologische Zeitung 43: 217–237 (in German).
258
Hagen, H.A. 1856. Die im Bernstein befindlichen Neuropteren der Vorwelt. Pp. 1–122 in Berendt,
259
G. C., Pictet de la Rive, F. J. & Hagen, H. A. (eds) Die im Bernstein befindlichen organischen
260
Reste der Vorwelt. Gerhard, Berlin.
261
Kolbe, H.J., 1884. Der Entwickelungsgang der Psociden im Individuum und in der Zeit. Berliner
262
Entomologische Zeitschrift 28, 35-38.
263
Kucerova, Z., 1997. Macropterous form of Dorypteryx domestica (Psocoptera: Psyllipsocidae).
264
European Journal of Entomology 94, 567–573.
265
Li, F.S., 2002. Psocoptera of China. National Natural Science Foundation of China, Science Press,
266
Beijing xlvi + 1976 pp. (two volumes). (in Chinese, with English summary).
267
Lin, X.D., Labandeira, C.C., Shih, C.K., Hotton, C.L., and Ren, D. (2019). Life habits and
268
evolutionary biology of new two-winged long-proboscid scorpionflies from mid-Cretaceous
269
Myanmar amber. Nature Communications 10: 1235.
270
Mockford, E.L., 1993. North American Psocoptera (Insecta). Flora & Fauna Handbook 10, 455pp.
271
Mockford, E.L., Lienhard, C., Yoshizawa, K., 2013. Revised classification of “Psocoptera” from
272
Cretaceous amber, a reassessment of published information. Insecta Matsumurana 69, 1-26.
273
Nel. A., Prokop, J., De Ploeg, G., and Millet, J., 2005. New Psocoptera (Insecta) from the
274
lowermost Eocene amber of Oise, France. Journal of Systematic Palaeonotology, 3(4): 371-391
275
New, T.R., Lienhard, C., 2007. The Psocoptera of Tropical South-east Asia. Fauna Malesina
276
Handbook 6. National Museum of Natural History Naturalis, Leiden, 48–168.
277
Pearman, J.V., 1936. The taxonomy of the Psocoptera: preliminary sketch. Proceedings of the
278
Royal Entomological Society, London (B) 5: 58–62.
279
Prerrichot, V., Azar, D., Néraudeau, D., Nel, A., 2003. New Psocoptera in the Early Cretaceous
280
amber of SW France and Lebanon (Insecta: Psocoptera: Trogiomorpha). Geological Magazine 140
281
(6), 669–683.
282
Ross, A.J., 2018. Burmese (Myanmar) amber taxa, on-line checklist v.2018.2 from.
283
https://www.nms.ac.uk/explore-our-cllections/stories/natural-would/burmese-amber.
284
Roesler, R., 1940. Neue und wenig bekannte Copeognathengattungen. I. Zoologischer Anzeiger
285
129 (9/10), 225-243.
286
Smithers, C.N., 1990. Keys to the family and genera of Psocoptera (Arthropoda: Insecta).
287
Technical Reports of Australian Museum, 1–82.
288
Smithers, C.N., 1972. The classification and phylogeny of the Psocoptera. Memoirs of the
289
Australian Museum 14, 1–349.
290
Shi, G., Grimaldi, D.A., Harlow, G.E., Wang, J., Wang, J., Wang, M., Lei, W., Li, Q., Li, X., 2012.
291
Age constraint on Burmese amber based on U-Pb dating of zicons. Cretaceous Research 37, 155–
292
163
293
Wang, R., Li, S., Ren, D., Yao, Y., New genus and species of the Psyllipsocidae (Psocodea:
294
Trogiomorpha)
295
https://doi.org/10.1016/j.cretres.2019.07.008.
296
Yoshizawa, K., Lienhard, C., Johnson, K.P., 2006. Molecular systematics of the suborder
297
Trogiomorpha (Insecta: Psocodea: “Psocoptera”). Zoological Journal of the Linnean Society 146,
298
287–299.
299
Zhang, X., Ren, D., Yao, Y.Z., 2018. A new genus and species of Mimarachnidae (Hemiptera:
300
Fulgoromprpha: Fulgoroidea) from mid-Cretaceous Burmese amber. Cretaceous Research 90,
301
168–173.
from
mid-Cretaceous
Burmese
amber,
Cretaceous
Research,
302
303
FIGURE CAPTIONS
304
Fig.
305
CNU-PSO-MA2019001. (A) Photograph in dorsal view; (B) Photograph in ventral view; (C) Line
1.
Habitus
of
Burmempheria
densuschaetae
Li,
Wang
&
Yao,
sp.
nov.,
306
drawing in dorsal view. (D) Line drawing of forewing; (E) Line drawing of hind wing; Scale bars:
307
0.5mm.
308
Fig. 2. Detailed photographs of Burmempheria densuschaetae Li, Wang & Yao, sp. nov. (A)
309
Photograph of mouthpart; (B) Line drawing of mouthpart; (C) Photograph of left antenna; (D)
310
Line drawing of left antenna; (E) Photograph of left mid tibia; (F) Line drawing of left mid tibia;
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(G) Photograph of Cu2 and 1A in left forewing; (H) Photograph of female genitalia; Scale bars: (A,
312
B, C, D, E, F) 0.1mm; (G, H) 0.5mm.
313
Fig. 3. Habitus of Burmempheria raruschaetae Li, Wang & Yao, sp. nov., CNU-PSO-MA2019004.
314
(A) Photograph in dorsal view; (B) Photograph in ventral view; (C) Line drawing in dorsal view.
315
(D) Line drawing of forewing; (E) Line drawing of hind wing; Scale bars: 0.5mm.
316
Fig. 4. Detailed photographs of Burmempheria raruschaetae Li, Wang & Yao, sp. nov. (A)
317
Photograph of left antenna; (B) Line drawing of left antenna; (C) Line drawing of right maxillary
318
palpus in ventral view; (D) Line drawing of labial palps; (E) Photograph of mouth part; (F)
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Photograph of right hind tibia; (G) Line drawing of right hind tibia; (H) Photograph of Cu2 and 1A
320
in left forewing ; (I) Photograph of male genitalia; (J) Line drawing of male genitalia; Scale bars:
321
(A, B, C, D, H) 0.1mm; (I, J) 0.25mm; (F, G) 0.5mm.
S.L., Q.Z.W and Y.Z.Y designed the research. S.L. and Q.Z.W took the photographs and prepared the line drawings. S.L., D.R. and Y.Z.Y. performed morphological, and drafted the paper.
The authors declare no competing interests.