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Two new species of Panorpa (Mecoptera, Panorpidae) from the Hengduan Mountains in Yunnan, China
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S1226-8615(18)30836-7 https://doi.org/10.1016/j.aspen.2019.12.005 ASPEN 1482
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Journal of Asia-Pacific Entomology
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9 December 2018 7 December 2019 9 December 2019
Please cite this article as: N. Li, B-Z. Hua, Two new species of Panorpa (Mecoptera, Panorpidae) from the Hengduan Mountains in Yunnan, China, Journal of Asia-Pacific Entomology (2019), doi: https://doi.org/10.1016/j.aspen. 2019.12.005
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Two new species of Panorpa (Mecoptera, Panorpidae) from the Hengduan Mountains in Yunnan, China
Ning Li, Bao-Zhen Hua* Key Laboratory of Plant Protection Resources and Pest Management, Ministry of Education, Entomological Museum, Northwest A&F University, Yangling, Shaanxi 712100, China
*Corresponding
authors: Bao-Zhen Hua ([email protected])
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Abstract Two new species of the genus Panorpa Linnaeus, 1758 are described from the Hengduan Mountains in Yunnan, China. Panorpa diqingensis sp. n. can be distinguished from its congeners by vertex dark brown, and occiput yellow; pleura dark brown; gonocoxites bearing a cluster of stout black setae on inner apex; parameres S-shaped in male genitalia. Panorpa dispergens sp. n. can be readily differentiated from other Panorpa species by apical band with a hyaline spot in wing; gonocoxites with a cluster of stout dark brown setae and 3–5 long setae on inner apex; female medigynium with a pair of basal plates. The number of Panorpa species in Yunnan is raised to six. A key to Yunnan species of Panorpa is presented.
www.zoobank.org/urn:lsid:zoobank.org:pub:1CFD5277-EA5A-4C99-A7DA-EA26722F2C76
Keywords Mecoptera, Panorpa, Yunnan, Scorpionfly, Hengduan Mountains
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Introduction The Hengduan Mountains are a well-known biodiversity hotspot in the southwestern region of China, consisting of western Sichuan in the north, eastern Tibet in the west, and northern Yunnan in the south (Boufford, 2014; Xing and Ree, 2017). They are featured by extreme topographic relief, unique geology, dramatic climate, and are one of the richest faunas and floras of temperate regions in the northern hemisphere (Boufford, 2014; Sun et al., 2017; Wu et al., 2017). There have been several studies of the diversity of this region, especially of plants (Sun et al., 2017; Wang et al., 2018) and birds (Wu et al., 2013; Wu et al., 2017), but the diversity of insects, especially the Mecoptera, has received little attention. Panorpidae is the largest family in Mecoptera, consisting of approximately 500 extant species, which are widely distributed across Asia, North America and Europe (Penny and Byers, 1979; Byers and Thornhill, 1983; Wang et al., 2019). Panorpids are commonly called scorpionflies because their enlarged and recurved genital bulbs resemble the stinger of a scorpion (Byers and Thornhill, 1983). They are known from fossil records from Middle Jurassic (Ding et al., 2014) and from Eocene (Krzemiński and Soszyńska-Maj, 2012). Scorpionflies usually inhabit dense forests with high humidity and elevation (Byers and Thornhill, 1983). Panorpidae are saprophagous, mainly feeding on dead and dying arthropods (Byers and Thornhill, 1983; Palmer, 2010). The eruciform larvae are also saprophagous (Jiang and Hua, 2015) and bear two compound eyes that are rare in the Holometabola larvae (Chen et al., 2012). The Panorpidae currently consists of two fossil genera (Krzemiński and Soszyńska-Maj, 2012; Ding et al., 2014) and eight extant genera (Wang and Hua, 2019a). The largest extant genus Panorpa Linneaus, 1758 was confirmed paraphyletic based on morphological (Willmann, 1989; Ma et al., 2012) and molecular data (Whiting, 2002; Hu et al., 2015; Miao et al., 2019). The species of Panorpa are diverse and are assigned to several species groups based on morphological criteria for regional fauna (Issiki, 1933; Cheng, 1957; Willmann, 1977; Byers, 1993). Recently, four genera (Sinopanorpa Cai & Hua, 2008, Furcatopanorpa Ma & Hua, 2011, Dicerapanorpa Zhong & Hua, 2013, and Cerapanorpa Gao, Ma & Hua, 2016) have been erected from Panorpa. Even with the removal of the new genera, Panorpa is still considered to be paraphyletic (Miao et al., 2019). Thirty-five species of Panorpidae have been reported from the Hengduan Mountains in Yunnan (Byers, 1970; Fu and Hua, 2009; Wang and Hua, 2016, 2017, 2018, 2019b; Hu et al., 2019), and are 3
assigned to three genera: Panorpa Linneaus, 1758 (4 spp.), Neopanorpa Weele, 1909 (24 spp.), and Dicerapanorpa Zhong & Hua, 2013 (7 spp.). The genus Panorpa is differentiated from Dicerapanorpa by lacking two anal horns on tergum VI in males, and is mainly distinguishable from Neopanorpa by vein 1A terminating at the posterior margin beyond the origin of Rs and characters of the male genital bulb (Cheng, 1957; Ma et al., 2012; Hua et al., 2018). In this paper, two undescribed species of Panorpa are identified from Yunnan, and are described herein, raising the species number of Panorpa to six and Panorpidae to 37 in Yunnan. A key to the species of Yunnan Panorpa is presented.
Materials and methods Adults of the new species were collected using sweeping nets from the Hengduan Mountains, Haba Village, Shangri-La County, Diqing, Yunnan Province (Fig. 1) from early June to late June in 2014 and 2016. These specimens are preserved in 75% ethanol at the Entomological Museum, Northwest A&F University, Shaanxi, China (NWAU). Specimens were observed and dissected under a Nikon SMZ1500 Stereoscopic Zoom Microscope (Nikon, Tokyo, Japan). The photographs of adult habitus were taken with a Canon EOS 70D digital camera and the light micrographs of genitalia were taken using an advanced Stereo Microscope System (Discovery V20, Zeiss; an auto-montage imaging system Axio ICc5). The photographs were assembled with Adobe Photoshop CS6.
Results Key to species of Panorpa from Yunnan (males) 1. Paramere bifurcated subbasally........................................................................................................................2 - Paramere unbranched........................................................................................................................................4 2. Two branches of paramere stretched oppositely, 3-shaped (Yongde) ...............P. reflexa Wang & Hua, 2016 - Two branches of paramere stretched caudally, with a short ventral branch and a long dorsal branch.............3 3. Gonostylus slightly concave on outer margin, with a spoon-like basal process and an obtuse triangular median tooth (Kunming) ............................................................P. kunmingensis Fu & Hua, 2009 - Gonostylus not concave on outer margin, a long and acute apically basal process, without median tooth (Yunnan) ............................................................................................................P. issikiana Byers, 1970
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4. Parameres long stick-like, straight and parallel; nota brown with a broad yellowish longitudinal stripe (Yongde) ...............................................................................................P. parallela Wang & Hua, 2016 - Parameres slender, twisted in an S-shape; nota dark brown without any longitudinal stripe...........................5 5. Gonocoxites with a cluster of stout black setae on inner apex; hypovalves with long yellow setae at distal half on inner margin (Shangri-La) ...........................................................................P. diqingensis sp. n. - Gonocoxites with a cluster of stout dark brown setae and 3–5 long setae on inner apex; hypovalves with long thick black bristles along the apical half on inner margin (Shangri-La)............P. dispergens sp. n.
Panorpa diqingensis sp. n. Figs. 2–4 Type material. Holotype: ♂, CHINA: Yunnan: Diqing, Shangri-La, 2600–2700 m, 20 June 2014, leg. Chao Gao and Mei Liu. Paratypes: 33♂25♀, same data as holotype.
Diagnosis. The new species can be distinguished from other Panorpa species by the following characters: (1) head with vertex dark brown, occiput yellow; (2) pleura in thoracic region dark brown; (3) forewing with pterostigmal band brown, and apical band brown but faint; (4) hypovalves bearing a row of long yellow setae along inner margin; (5) gonocoxites with a cluster of stout black bristles on inner apex.
Description of male. Head. Head unevenly colored, with frons brown, vertex dark brown, and occiput yellow. Rostrum with two brown longitudinal stripes, maxillary and labial palps brown with distal segments dark brown (Fig. 2C). Antennae filiform, and yellowish brown (Fig. 2A, B). Thorax. Pronotum unevenly dark, bearing 8–12 setae along anterior margin. Mesonotum dark brown, and paler posteriorly. Metanotum similar to Mesonotum in coloration (Fig. 2D). Pleura dark brown and legs yellow. Wings. Forewing length 13.66 ± 0.45 mm, width 3.10 ± 0.09 mm (n = 5). Wing membrane hyaline, with markings brown. Pterostigmal band usually reduced and not reaching R2+3. Apical band indistinct and extremely reduced. Hindwing length 12.50 ± 0.39 mm, width 3.10 ± 0.07 mm (n = 5), similar to forewings in markings, but slightly smaller in shape (Fig. 2A). Abdomen. Terga I–V dark brown, pleural membrane pale. Notal organ on tergum III weaklydeveloped, and dorsal process of tergum IV slightly produced (Fig. 2E). Tergum VI with the basal third
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brown and yellow posteriorly, without anal horns on posterior margin, but slightly projected on the dorsal apex. Segments VII and VIII uniformly yellowish and constricted basally (Fig. 2E). Male genitalia. Genital bulb globular, yellowish (Fig. 3B, C). Epandrium broad basally and tapering toward apex, with a deep U-shaped terminal emargination bearing dense setae. Cerci clavate and expanded apically (Fig. 3C). Hypandrium with a greatly shortened basal stalk and a pair of elongate hypovalves. Hypovalves extending beyond half of gonocoxites, with a row of long yellow bristles at distal half along inner margin. Gonocoxites bearing a cluster of stout black setae on inner apex (Fig. 3A, B). Gonostylus roughly half length of gonocoxite, with an obtuse triangular median tooth and a spoon-like basal process on the inner margin (Fig. 3A, B). Parameres slender, twisted in an S-shape (Fig. 3D). Dorsal valves of aedeagus slender, with a bundle of long hairs ventrally and middle part a quadrilateral flap dorsally (Fig. 3E). A pair of short distinct lateral process from dorsal valves basally (Fig. 3D–F). Ventral valves greatly short and membranous (Fig. 3D). Description of female. Similar to male except for more extensive and prominent wing markings. Pterostigmal band extending beyond R4+5. Apical band indistinct but broader than in male (Fig. 2B). Forewing length 14.16 ± 0.55 mm, width 3.41 ± 0.09 mm; hindwing length 12.99 ± 0.56 mm, width 3.33 ± 0.09 mm (n = 5). Female genitalia. Subgenital plate ligulate, bearing long setae along caudal margins (Fig. 4A–C). Medigynium weakly sclerotized, with main plate three times as long as wide. Posterior arms curved, forming a broad U-shaped emargination. Median axis elongate, extending beyond main plate for a quarter length, with posterior slightly extruding (Fig. 4D, E).
Etymology. The specific epithet is derived from the type locality, Diqing in Yunnan Province, Shangri-La County.
Distribution. China (Yunnan Province). Remarks. In some individuals, wing markings more extensive and distinct, especially apical band of forewing in female. None or one cross-vein between veins R2a and R2b. One or two cross-vein between veins R2 and R3.
Panorpa dispergens sp. n. Figs. 5–7
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Type material. Holotype. ♂, CHINA: Yunnan: Diqing, Shangri-La (27.37°N, 100.13°E), 2700 m, 2 June 2016, leg. Wei Du and Gui-Lin Hu. Paratypes. 1♂3♀, same data as holotype; 13♂12♀, Diqing, Shangri-La, 2600–2700 m, 20 June 2014, leg. Chao Gao and Mei Liu. Diagnosis. Panorpa dispergens sp. n. can be separated from its congeners by the following characters: (1) vertex dark brown, occiput paler; (2) pleura in thoracic region yellowish; (3) forewing with pterostigmal band brown, with apical band brown with a hyaline spot; (4) apical half of hypovalves with long thick black bristles on inner margin (5) gonocoxites with a cluster of stout dark brown setae and 3–5 long setae on inner apex.
Description of male. Head. Vertex and ocellar triangle dark brown, occiput paler. Rostrum yellowish, with two brown submedian stripes along clypeus, maxillary palp brown with distal segments dark brown (Fig. 5C). Antennae long, filiform, dark brown (Fig. 5A, B). Thorax. Pronotum unevenly dark brown, with 10–12 setae along anterior margin. Mesonotum dark brown anteriorly, and paler posteriorly, with scutellum yellowish brown. Metanotum brown, and paler posteriorly (Fig. 5D). Pleura and legs yellowish. Wings. Forewing length 13.50 ± 0.87 mm, width 3.31 ± 0.18 mm (n = 5). Wing membrane with brown markings. Pterostigmal band reduced, consisting of an irregular discrete spot, extending from R1 to beyond R4+5. Apical band distinct with a hyaline spot innerly. Hindwing length 12.61 ± 0.78 mm, width 3.31 ± 0.26 mm (n = 5), similar to forewing in markings (Fig. 5A). Abdomen. Terga I–V dark brown, pleural membrane pale. Notal organ on tergum III flat triangular and an acute postnotal organ on tergum IV (Fig. 5E). Tergum VI with the basal two-thirds brown and yellowish distally and slightly projected on the dorsal apex. Segments VII and VIII elongate, uniformly yellowish, and constricted basally (Fig. 5E). Male genitalia. Male genital bulb long elliptical, yellowish (Fig. 6B, C). Epandrium broad basally, forming a square deep emargination bearing dense setae distally. Cerci clavate and expanded apically (Fig. 6C). Hypandrium with a shortened basal stalk and a pair of long strip-like hypovalves. Hypovalves with long thick black bristles along the apical half on inner margin, reaching apex of gonocoxite. Gonocoxites with a row of brown bristles of various lengths on inner margin apically (Fig. 6B). Gonostylus with a spoon-like basal process and an obtuse triangular median tooth (Fig. 6A, B). Parameres slender, twisted in an S-shape (Fig. 6D). The dorsal valves slender, with a bundle of long
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hairs ventrally and bearing a foot-shaped process on the distal part dorsally (Fig. 6E). Lateral process small, but distinct (Fig. 6D–F). Ventral valves greatly shortened and membranous (Fig. 6D). Description of female. Forewing length 14.06 ± 0.40 mm, width 3.39 ± 0.17 mm; hindwing length 12.99 ± 0.41 mm, width 3.31 ± 0.19 mm (n = 5). Similar to male in coloration and wing pattern. Pterostigmal band extremely reduced with a large spot not reaching R2+3 and a small spot between R4+5 and M1+2 (Fig. 5B). Female genitalia. Subgenital plate ligulate, with long setae on distal half, especially caudal and lateral margins (Fig. 7A–C). Medigynium with main plate oblong, narrow basally, medial part bearing a pair of basal plates laterally, distal margin with a U-shaped emargination formed by a pair of sclerotized and curved posterior arms. Median axis elongated and bifurcated anteriorly, extending beyond main plate for half its length, with posterior slightly extruding (Fig. 7D, E).
Etymology. The specific name is derived from Latin dispergens (disperse), referring to the disperse spots.
Distribution. China (Yunnan Province). Remarks. In some individuals, wing markings more reduced, without or with an extremely reduced pterostigmal band. The size of hyaline spot in apical band is distinctly variable. Apical band is absent in some individuals. The length of parameres is slightly variable.
Discussion Yunnan Province is located in the southwestern region of China, bordering Vietnam in the southeast, Laos in the south, and Myanmar in the west. It possesses a high degree of biodiversity and occupies a key position connecting the tropical Indo-China Peninsula and subtropical China (Boufford, 2014). Species of Panorpidae from Yunnan mainly inhabit the Hengduan Mountain region. The Hengduan Mountains are a group of several north-south mountain ranges, and are characterized by significant vertical relief originating from the Indian subcontinent’s collision with the Eurasian Plate, and further carved out by the major rivers draining the eastern Qinghai-Tibetan Plateau. Mount Gongga, the highest peak of the Hengduan Mountains reaches 7556 m above the see level, while the elevation of the Tiger Leaping Gorge is only 1900 m located between the Yulong Snow Mountain and the Haba Snow Mountain with elevations over 5000 m (Boufford, 2014). Low-altitude regions and mountain gorges of the Hengduan Mountains were considered as refugia in the glacial periods (Meng et al., 2015; 8
Wang and Hua, 2018). Species in this region responded to environmental changes by migrating north and south along the Hengduan Mountains with the change of elevation (Hewitt, 2004; Wang and Hua, 2018; Hu et al., 2019). Populations after migration and colonization were long-term isolated into glacial refugia by Pleistocene glaciers, therefore population differentiation and speciation would be promoted (Meng et al., 2015; Ye et al., 2016). It is reasonable that more species of Panorpidae await description from this region. The main parts of the Hengduan Mountains are located in Sichuan, Tibet, and Yunnan Provinces (Boufford, 2014; Xing and Ree, 2017). Based on the comparative morphology of Panorpa species from the Hengduan Mountains, the two new species from Yunnan have close relationship with the six species from Sichuan―P. curva, Carpenter, 1938, P. davidi Navás, 1908, P. difficilis Carpenter, 1938, P. fructa Cheng, 1949, P. typicoides Cheng, 1949, P. jinchuana Hua, Sun & Li, 2001. These eight species of Panorpa are characterized by tergum VI slightly projected on the dorsal apex; basal stalk extremely shortened; parameres twisted in an S-shape; gonocoxites bearing some bristles on inner margin apically; gonostylus with a spoon-like basal process and an undeveloped median tooth. Compared with the six species of Panorpa from Sichuan, two new species are unique for their pattern of wing markings, arrangement and quantity of bristles on the inner apex of male gonocoxites and the characters of female medigynium.
Acknowledgements We are indebted to Gui-Lin Hu, Wei Du, Chao Gao, and Mei Liu for collecting specimens, Ji-Shen Wang for help with the Latin names and Xin Tong and Kai Gao for assistance in taking photographs. We also thank Agnieszka Soszynska-Maj and the two anonymous reviewers for valuable comments and suggestions to the revision of the manuscript. This research was funded by the National Natural Science Foundation of China (grant no. 31672341).
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Figure legends Figure 1. Distribution map of the two new species of Panorpa in Yunnan. Figure 2. Adults of Panorpa diqingensis sp. n. (A) Habitus of male, dorsal view; (B) habitus of female, dorsal view; (C) head of male, frontal view; (D) dorsum of male; (E) abdomen of male, lateral view. no, notal organ; pno, postnotal organ. Scale bars: 5 mm (A and B); 0.5 mm (C); 1 mm (D and E). Figure 3. Male genitalia of Panorpa diqingensis sp. n. (A) Gonostylus, ventral view; (B) and (C) genital bulb in ventral and dorsal views; (D)−(F) aedeagal complex in ventral, lateral, and dorsal views. ae, aedeagus; bp, basal process; bst, basal stalk of hypandrium; ce, cercus; dv, dorsal valve; ep, epandrium; gcx, gonocoxite; gs, gonostylus; hv, hypovalve; lp, lateral process; mt, median tooth; pm, paramere; vv, ventral valve. Scale bars: 0.2 mm. Figure 4. Female of Panorpa diqingensis sp. n. (A) and (B) Abdominal segments VIII–X, lateral and ventral views; (C) subgenital plate, ventral view; (D) and (E) medigynium, ventral and dorsal views. ax, axis; ce, cercus; mp, main plate; pa, posterior arm; sgp, subgenital plate. Scale bars: 0.2 mm. Figure 5. Adults of Panorpa dispergens sp. n. (A) Habitus of male, dorsal view; (B) habitus of female, dorsal view; (C) head of male, frontal view; (D) dorsum of male; (E) abdomen of male, lateral view. no, notal organ; pno, postnotal organ. Scale bars: 5 mm (A and B); 0.5 mm (C); 1 mm (D and E). Figure 6. Male genitalia of Panorpa dispergens sp. n. (A) Gonostylus, ventral view; (B) and (C) genital bulb in ventral and dorsal views; (D)−(F) aedeagal complex in ventral, lateral, and dorsal views. ae, aedeagus; bp, basal process; bst, basal stalk of hypandrium; ce, cercus; dv, dorsal valve; ep, epandrium; gcx, gonocoxite; gs, gonostylus; hv, hypovalve; lp, lateral process; mt, median tooth; pm, paramere; vv, ventral valve. Scale bars: 0.2 mm. Figure 7. Female of Panorpa dispergens sp. n. (A) and (B) Abdominal segments VIII–X, lateral and ventral views; (C) subgenital plate, ventral view; (D) and (E) medigynium, ventral and dorsal view. ax, axis; bp, basal plate; ce, cercus; mp, main plate; pa, posterior arm; sgp, subgenital plate. Scale bars: 0.2 mm.
Conflict of Interest
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All authors have approved the manuscript and agree with submission to Journal of Asia-Pacific Entomology. The authors have no conflicts of interest to declare.
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Highlights: • Two new species of Panorpa are described from the Hengduan Mountains in Yunnan, China. • A key to Yunnan species of Panorpa is presented. • All new species of illustrations are provided.
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S1226-8615(18)30836-7 https://doi.org/10.1016/j.aspen.2019.12.005 ASPEN 1482
To appear in:
Journal of Asia-Pacific Entomology
Received Date: Revised Date: Accepted Date:
9 December 2018 7 December 2019 9 December 2019
Please cite this article as: N. Li, B-Z. Hua, Two new species of Panorpa (Mecoptera, Panorpidae) from the Hengduan Mountains in Yunnan, China, Journal of Asia-Pacific Entomology (2019), doi: https://doi.org/10.1016/j.aspen. 2019.12.005
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Two new species of Panorpa (Mecoptera, Panorpidae) from the Hengduan Mountains in Yunnan, China
Ning Li, Bao-Zhen Hua* Key Laboratory of Plant Protection Resources and Pest Management, Ministry of Education, Entomological Museum, Northwest A&F University, Yangling, Shaanxi 712100, China
*Corresponding
authors: Bao-Zhen Hua ([email protected])
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Abstract Two new species of the genus Panorpa Linnaeus, 1758 are described from the Hengduan Mountains in Yunnan, China. Panorpa diqingensis sp. n. can be distinguished from its congeners by vertex dark brown, and occiput yellow; pleura dark brown; gonocoxites bearing a cluster of stout black setae on inner apex; parameres S-shaped in male genitalia. Panorpa dispergens sp. n. can be readily differentiated from other Panorpa species by apical band with a hyaline spot in wing; gonocoxites with a cluster of stout dark brown setae and 3–5 long setae on inner apex; female medigynium with a pair of basal plates. The number of Panorpa species in Yunnan is raised to six. A key to Yunnan species of Panorpa is presented.
www.zoobank.org/urn:lsid:zoobank.org:pub:1CFD5277-EA5A-4C99-A7DA-EA26722F2C76
Keywords Mecoptera, Panorpa, Yunnan, Scorpionfly, Hengduan Mountains
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Introduction The Hengduan Mountains are a well-known biodiversity hotspot in the southwestern region of China, consisting of western Sichuan in the north, eastern Tibet in the west, and northern Yunnan in the south (Boufford, 2014; Xing and Ree, 2017). They are featured by extreme topographic relief, unique geology, dramatic climate, and are one of the richest faunas and floras of temperate regions in the northern hemisphere (Boufford, 2014; Sun et al., 2017; Wu et al., 2017). There have been several studies of the diversity of this region, especially of plants (Sun et al., 2017; Wang et al., 2018) and birds (Wu et al., 2013; Wu et al., 2017), but the diversity of insects, especially the Mecoptera, has received little attention. Panorpidae is the largest family in Mecoptera, consisting of approximately 500 extant species, which are widely distributed across Asia, North America and Europe (Penny and Byers, 1979; Byers and Thornhill, 1983; Wang et al., 2019). Panorpids are commonly called scorpionflies because their enlarged and recurved genital bulbs resemble the stinger of a scorpion (Byers and Thornhill, 1983). They are known from fossil records from Middle Jurassic (Ding et al., 2014) and from Eocene (Krzemiński and Soszyńska-Maj, 2012). Scorpionflies usually inhabit dense forests with high humidity and elevation (Byers and Thornhill, 1983). Panorpidae are saprophagous, mainly feeding on dead and dying arthropods (Byers and Thornhill, 1983; Palmer, 2010). The eruciform larvae are also saprophagous (Jiang and Hua, 2015) and bear two compound eyes that are rare in the Holometabola larvae (Chen et al., 2012). The Panorpidae currently consists of two fossil genera (Krzemiński and Soszyńska-Maj, 2012; Ding et al., 2014) and eight extant genera (Wang and Hua, 2019a). The largest extant genus Panorpa Linneaus, 1758 was confirmed paraphyletic based on morphological (Willmann, 1989; Ma et al., 2012) and molecular data (Whiting, 2002; Hu et al., 2015; Miao et al., 2019). The species of Panorpa are diverse and are assigned to several species groups based on morphological criteria for regional fauna (Issiki, 1933; Cheng, 1957; Willmann, 1977; Byers, 1993). Recently, four genera (Sinopanorpa Cai & Hua, 2008, Furcatopanorpa Ma & Hua, 2011, Dicerapanorpa Zhong & Hua, 2013, and Cerapanorpa Gao, Ma & Hua, 2016) have been erected from Panorpa. Even with the removal of the new genera, Panorpa is still considered to be paraphyletic (Miao et al., 2019). Thirty-five species of Panorpidae have been reported from the Hengduan Mountains in Yunnan (Byers, 1970; Fu and Hua, 2009; Wang and Hua, 2016, 2017, 2018, 2019b; Hu et al., 2019), and are 3
assigned to three genera: Panorpa Linneaus, 1758 (4 spp.), Neopanorpa Weele, 1909 (24 spp.), and Dicerapanorpa Zhong & Hua, 2013 (7 spp.). The genus Panorpa is differentiated from Dicerapanorpa by lacking two anal horns on tergum VI in males, and is mainly distinguishable from Neopanorpa by vein 1A terminating at the posterior margin beyond the origin of Rs and characters of the male genital bulb (Cheng, 1957; Ma et al., 2012; Hua et al., 2018). In this paper, two undescribed species of Panorpa are identified from Yunnan, and are described herein, raising the species number of Panorpa to six and Panorpidae to 37 in Yunnan. A key to the species of Yunnan Panorpa is presented.
Materials and methods Adults of the new species were collected using sweeping nets from the Hengduan Mountains, Haba Village, Shangri-La County, Diqing, Yunnan Province (Fig. 1) from early June to late June in 2014 and 2016. These specimens are preserved in 75% ethanol at the Entomological Museum, Northwest A&F University, Shaanxi, China (NWAU). Specimens were observed and dissected under a Nikon SMZ1500 Stereoscopic Zoom Microscope (Nikon, Tokyo, Japan). The photographs of adult habitus were taken with a Canon EOS 70D digital camera and the light micrographs of genitalia were taken using an advanced Stereo Microscope System (Discovery V20, Zeiss; an auto-montage imaging system Axio ICc5). The photographs were assembled with Adobe Photoshop CS6.
Results Key to species of Panorpa from Yunnan (males) 1. Paramere bifurcated subbasally........................................................................................................................2 - Paramere unbranched........................................................................................................................................4 2. Two branches of paramere stretched oppositely, 3-shaped (Yongde) ...............P. reflexa Wang & Hua, 2016 - Two branches of paramere stretched caudally, with a short ventral branch and a long dorsal branch.............3 3. Gonostylus slightly concave on outer margin, with a spoon-like basal process and an obtuse triangular median tooth (Kunming) ............................................................P. kunmingensis Fu & Hua, 2009 - Gonostylus not concave on outer margin, a long and acute apically basal process, without median tooth (Yunnan) ............................................................................................................P. issikiana Byers, 1970
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4. Parameres long stick-like, straight and parallel; nota brown with a broad yellowish longitudinal stripe (Yongde) ...............................................................................................P. parallela Wang & Hua, 2016 - Parameres slender, twisted in an S-shape; nota dark brown without any longitudinal stripe...........................5 5. Gonocoxites with a cluster of stout black setae on inner apex; hypovalves with long yellow setae at distal half on inner margin (Shangri-La) ...........................................................................P. diqingensis sp. n. - Gonocoxites with a cluster of stout dark brown setae and 3–5 long setae on inner apex; hypovalves with long thick black bristles along the apical half on inner margin (Shangri-La)............P. dispergens sp. n.
Panorpa diqingensis sp. n. Figs. 2–4 Type material. Holotype: ♂, CHINA: Yunnan: Diqing, Shangri-La, 2600–2700 m, 20 June 2014, leg. Chao Gao and Mei Liu. Paratypes: 33♂25♀, same data as holotype.
Diagnosis. The new species can be distinguished from other Panorpa species by the following characters: (1) head with vertex dark brown, occiput yellow; (2) pleura in thoracic region dark brown; (3) forewing with pterostigmal band brown, and apical band brown but faint; (4) hypovalves bearing a row of long yellow setae along inner margin; (5) gonocoxites with a cluster of stout black bristles on inner apex.
Description of male. Head. Head unevenly colored, with frons brown, vertex dark brown, and occiput yellow. Rostrum with two brown longitudinal stripes, maxillary and labial palps brown with distal segments dark brown (Fig. 2C). Antennae filiform, and yellowish brown (Fig. 2A, B). Thorax. Pronotum unevenly dark, bearing 8–12 setae along anterior margin. Mesonotum dark brown, and paler posteriorly. Metanotum similar to Mesonotum in coloration (Fig. 2D). Pleura dark brown and legs yellow. Wings. Forewing length 13.66 ± 0.45 mm, width 3.10 ± 0.09 mm (n = 5). Wing membrane hyaline, with markings brown. Pterostigmal band usually reduced and not reaching R2+3. Apical band indistinct and extremely reduced. Hindwing length 12.50 ± 0.39 mm, width 3.10 ± 0.07 mm (n = 5), similar to forewings in markings, but slightly smaller in shape (Fig. 2A). Abdomen. Terga I–V dark brown, pleural membrane pale. Notal organ on tergum III weaklydeveloped, and dorsal process of tergum IV slightly produced (Fig. 2E). Tergum VI with the basal third
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brown and yellow posteriorly, without anal horns on posterior margin, but slightly projected on the dorsal apex. Segments VII and VIII uniformly yellowish and constricted basally (Fig. 2E). Male genitalia. Genital bulb globular, yellowish (Fig. 3B, C). Epandrium broad basally and tapering toward apex, with a deep U-shaped terminal emargination bearing dense setae. Cerci clavate and expanded apically (Fig. 3C). Hypandrium with a greatly shortened basal stalk and a pair of elongate hypovalves. Hypovalves extending beyond half of gonocoxites, with a row of long yellow bristles at distal half along inner margin. Gonocoxites bearing a cluster of stout black setae on inner apex (Fig. 3A, B). Gonostylus roughly half length of gonocoxite, with an obtuse triangular median tooth and a spoon-like basal process on the inner margin (Fig. 3A, B). Parameres slender, twisted in an S-shape (Fig. 3D). Dorsal valves of aedeagus slender, with a bundle of long hairs ventrally and middle part a quadrilateral flap dorsally (Fig. 3E). A pair of short distinct lateral process from dorsal valves basally (Fig. 3D–F). Ventral valves greatly short and membranous (Fig. 3D). Description of female. Similar to male except for more extensive and prominent wing markings. Pterostigmal band extending beyond R4+5. Apical band indistinct but broader than in male (Fig. 2B). Forewing length 14.16 ± 0.55 mm, width 3.41 ± 0.09 mm; hindwing length 12.99 ± 0.56 mm, width 3.33 ± 0.09 mm (n = 5). Female genitalia. Subgenital plate ligulate, bearing long setae along caudal margins (Fig. 4A–C). Medigynium weakly sclerotized, with main plate three times as long as wide. Posterior arms curved, forming a broad U-shaped emargination. Median axis elongate, extending beyond main plate for a quarter length, with posterior slightly extruding (Fig. 4D, E).
Etymology. The specific epithet is derived from the type locality, Diqing in Yunnan Province, Shangri-La County.
Distribution. China (Yunnan Province). Remarks. In some individuals, wing markings more extensive and distinct, especially apical band of forewing in female. None or one cross-vein between veins R2a and R2b. One or two cross-vein between veins R2 and R3.
Panorpa dispergens sp. n. Figs. 5–7
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Type material. Holotype. ♂, CHINA: Yunnan: Diqing, Shangri-La (27.37°N, 100.13°E), 2700 m, 2 June 2016, leg. Wei Du and Gui-Lin Hu. Paratypes. 1♂3♀, same data as holotype; 13♂12♀, Diqing, Shangri-La, 2600–2700 m, 20 June 2014, leg. Chao Gao and Mei Liu. Diagnosis. Panorpa dispergens sp. n. can be separated from its congeners by the following characters: (1) vertex dark brown, occiput paler; (2) pleura in thoracic region yellowish; (3) forewing with pterostigmal band brown, with apical band brown with a hyaline spot; (4) apical half of hypovalves with long thick black bristles on inner margin (5) gonocoxites with a cluster of stout dark brown setae and 3–5 long setae on inner apex.
Description of male. Head. Vertex and ocellar triangle dark brown, occiput paler. Rostrum yellowish, with two brown submedian stripes along clypeus, maxillary palp brown with distal segments dark brown (Fig. 5C). Antennae long, filiform, dark brown (Fig. 5A, B). Thorax. Pronotum unevenly dark brown, with 10–12 setae along anterior margin. Mesonotum dark brown anteriorly, and paler posteriorly, with scutellum yellowish brown. Metanotum brown, and paler posteriorly (Fig. 5D). Pleura and legs yellowish. Wings. Forewing length 13.50 ± 0.87 mm, width 3.31 ± 0.18 mm (n = 5). Wing membrane with brown markings. Pterostigmal band reduced, consisting of an irregular discrete spot, extending from R1 to beyond R4+5. Apical band distinct with a hyaline spot innerly. Hindwing length 12.61 ± 0.78 mm, width 3.31 ± 0.26 mm (n = 5), similar to forewing in markings (Fig. 5A). Abdomen. Terga I–V dark brown, pleural membrane pale. Notal organ on tergum III flat triangular and an acute postnotal organ on tergum IV (Fig. 5E). Tergum VI with the basal two-thirds brown and yellowish distally and slightly projected on the dorsal apex. Segments VII and VIII elongate, uniformly yellowish, and constricted basally (Fig. 5E). Male genitalia. Male genital bulb long elliptical, yellowish (Fig. 6B, C). Epandrium broad basally, forming a square deep emargination bearing dense setae distally. Cerci clavate and expanded apically (Fig. 6C). Hypandrium with a shortened basal stalk and a pair of long strip-like hypovalves. Hypovalves with long thick black bristles along the apical half on inner margin, reaching apex of gonocoxite. Gonocoxites with a row of brown bristles of various lengths on inner margin apically (Fig. 6B). Gonostylus with a spoon-like basal process and an obtuse triangular median tooth (Fig. 6A, B). Parameres slender, twisted in an S-shape (Fig. 6D). The dorsal valves slender, with a bundle of long
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hairs ventrally and bearing a foot-shaped process on the distal part dorsally (Fig. 6E). Lateral process small, but distinct (Fig. 6D–F). Ventral valves greatly shortened and membranous (Fig. 6D). Description of female. Forewing length 14.06 ± 0.40 mm, width 3.39 ± 0.17 mm; hindwing length 12.99 ± 0.41 mm, width 3.31 ± 0.19 mm (n = 5). Similar to male in coloration and wing pattern. Pterostigmal band extremely reduced with a large spot not reaching R2+3 and a small spot between R4+5 and M1+2 (Fig. 5B). Female genitalia. Subgenital plate ligulate, with long setae on distal half, especially caudal and lateral margins (Fig. 7A–C). Medigynium with main plate oblong, narrow basally, medial part bearing a pair of basal plates laterally, distal margin with a U-shaped emargination formed by a pair of sclerotized and curved posterior arms. Median axis elongated and bifurcated anteriorly, extending beyond main plate for half its length, with posterior slightly extruding (Fig. 7D, E).
Etymology. The specific name is derived from Latin dispergens (disperse), referring to the disperse spots.
Distribution. China (Yunnan Province). Remarks. In some individuals, wing markings more reduced, without or with an extremely reduced pterostigmal band. The size of hyaline spot in apical band is distinctly variable. Apical band is absent in some individuals. The length of parameres is slightly variable.
Discussion Yunnan Province is located in the southwestern region of China, bordering Vietnam in the southeast, Laos in the south, and Myanmar in the west. It possesses a high degree of biodiversity and occupies a key position connecting the tropical Indo-China Peninsula and subtropical China (Boufford, 2014). Species of Panorpidae from Yunnan mainly inhabit the Hengduan Mountain region. The Hengduan Mountains are a group of several north-south mountain ranges, and are characterized by significant vertical relief originating from the Indian subcontinent’s collision with the Eurasian Plate, and further carved out by the major rivers draining the eastern Qinghai-Tibetan Plateau. Mount Gongga, the highest peak of the Hengduan Mountains reaches 7556 m above the see level, while the elevation of the Tiger Leaping Gorge is only 1900 m located between the Yulong Snow Mountain and the Haba Snow Mountain with elevations over 5000 m (Boufford, 2014). Low-altitude regions and mountain gorges of the Hengduan Mountains were considered as refugia in the glacial periods (Meng et al., 2015; 8
Wang and Hua, 2018). Species in this region responded to environmental changes by migrating north and south along the Hengduan Mountains with the change of elevation (Hewitt, 2004; Wang and Hua, 2018; Hu et al., 2019). Populations after migration and colonization were long-term isolated into glacial refugia by Pleistocene glaciers, therefore population differentiation and speciation would be promoted (Meng et al., 2015; Ye et al., 2016). It is reasonable that more species of Panorpidae await description from this region. The main parts of the Hengduan Mountains are located in Sichuan, Tibet, and Yunnan Provinces (Boufford, 2014; Xing and Ree, 2017). Based on the comparative morphology of Panorpa species from the Hengduan Mountains, the two new species from Yunnan have close relationship with the six species from Sichuan―P. curva, Carpenter, 1938, P. davidi Navás, 1908, P. difficilis Carpenter, 1938, P. fructa Cheng, 1949, P. typicoides Cheng, 1949, P. jinchuana Hua, Sun & Li, 2001. These eight species of Panorpa are characterized by tergum VI slightly projected on the dorsal apex; basal stalk extremely shortened; parameres twisted in an S-shape; gonocoxites bearing some bristles on inner margin apically; gonostylus with a spoon-like basal process and an undeveloped median tooth. Compared with the six species of Panorpa from Sichuan, two new species are unique for their pattern of wing markings, arrangement and quantity of bristles on the inner apex of male gonocoxites and the characters of female medigynium.
Acknowledgements We are indebted to Gui-Lin Hu, Wei Du, Chao Gao, and Mei Liu for collecting specimens, Ji-Shen Wang for help with the Latin names and Xin Tong and Kai Gao for assistance in taking photographs. We also thank Agnieszka Soszynska-Maj and the two anonymous reviewers for valuable comments and suggestions to the revision of the manuscript. This research was funded by the National Natural Science Foundation of China (grant no. 31672341).
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Figure legends Figure 1. Distribution map of the two new species of Panorpa in Yunnan. Figure 2. Adults of Panorpa diqingensis sp. n. (A) Habitus of male, dorsal view; (B) habitus of female, dorsal view; (C) head of male, frontal view; (D) dorsum of male; (E) abdomen of male, lateral view. no, notal organ; pno, postnotal organ. Scale bars: 5 mm (A and B); 0.5 mm (C); 1 mm (D and E). Figure 3. Male genitalia of Panorpa diqingensis sp. n. (A) Gonostylus, ventral view; (B) and (C) genital bulb in ventral and dorsal views; (D)−(F) aedeagal complex in ventral, lateral, and dorsal views. ae, aedeagus; bp, basal process; bst, basal stalk of hypandrium; ce, cercus; dv, dorsal valve; ep, epandrium; gcx, gonocoxite; gs, gonostylus; hv, hypovalve; lp, lateral process; mt, median tooth; pm, paramere; vv, ventral valve. Scale bars: 0.2 mm. Figure 4. Female of Panorpa diqingensis sp. n. (A) and (B) Abdominal segments VIII–X, lateral and ventral views; (C) subgenital plate, ventral view; (D) and (E) medigynium, ventral and dorsal views. ax, axis; ce, cercus; mp, main plate; pa, posterior arm; sgp, subgenital plate. Scale bars: 0.2 mm. Figure 5. Adults of Panorpa dispergens sp. n. (A) Habitus of male, dorsal view; (B) habitus of female, dorsal view; (C) head of male, frontal view; (D) dorsum of male; (E) abdomen of male, lateral view. no, notal organ; pno, postnotal organ. Scale bars: 5 mm (A and B); 0.5 mm (C); 1 mm (D and E). Figure 6. Male genitalia of Panorpa dispergens sp. n. (A) Gonostylus, ventral view; (B) and (C) genital bulb in ventral and dorsal views; (D)−(F) aedeagal complex in ventral, lateral, and dorsal views. ae, aedeagus; bp, basal process; bst, basal stalk of hypandrium; ce, cercus; dv, dorsal valve; ep, epandrium; gcx, gonocoxite; gs, gonostylus; hv, hypovalve; lp, lateral process; mt, median tooth; pm, paramere; vv, ventral valve. Scale bars: 0.2 mm. Figure 7. Female of Panorpa dispergens sp. n. (A) and (B) Abdominal segments VIII–X, lateral and ventral views; (C) subgenital plate, ventral view; (D) and (E) medigynium, ventral and dorsal view. ax, axis; bp, basal plate; ce, cercus; mp, main plate; pa, posterior arm; sgp, subgenital plate. Scale bars: 0.2 mm.
Conflict of Interest
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All authors have approved the manuscript and agree with submission to Journal of Asia-Pacific Entomology. The authors have no conflicts of interest to declare.
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Highlights: • Two new species of Panorpa are described from the Hengduan Mountains in Yunnan, China. • A key to Yunnan species of Panorpa is presented. • All new species of illustrations are provided.
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