Protagonista lugubris Shelford, 1908

Wang, Chenchen, Wang, Zongqing & Che, Yanli, 2016, Protagonista lugubris, a cockroach species new to China and its contribution to the revision of genus Protagonista, with notes on the taxonomy of Archiblattinae (Blattodea, Blattidae), ZooKeys 574, pp. 57-73 : 59-64

Protagonista lugubris Shelford, 1908 new record from China Figs 1-6 , 7-12 , 13-22 , 23-27 , 28-33 , 34-43 , 45 , 48-49

Protagonista lugubris Shelford, 1908: 158; Shelford 1910: 22; Hanitsch 1927: 40; Hanitsch 1929: 17; Bruijning 1948: 117.

Description.

Male. Body slender, dark brown to black (Figs 1-2 , 5-6 , 49 ). Eyes black, ocelli yellowish white. Vertex and face dark brown to black. Clypeus yellowish or dark brown and the base of labrum pale, labial palpi and maxillary palpomeres brown. Antennae brown or black, apical joints creamy-white. Pronotum and tegmina dark brown or black and apex of tegmina brownish yellow or brown. Legs brown or black. Abdomen reddish brown with dark brown margins or uniformly black. Cerci brown or yellowish brown.

Head with vertex punctate, with three smooth longitudinal stripes (Figs 13 , 34 ). Eyes reniform and closer together than antennal sockets. Ocelli elliptical with distinct border. Face punctate, with some smooth interspaces and lines (Figs 13 , 34 ). Antennae with numerous bristles, shorter than the body. Pronotum elongate, trapezoidal, with punctations and pubescence, margins thickened and raised, with three smooth longitudinal impressions and some smooth interspaces, disk not flat (Figs 7 , 16 , 28 , 37 ). Both tegmina and hind wings fully developed, extending beyond the end of abdomen. Tegmina narrow with scattered erect pubescence; basal half sclerotized (Figs 8 , 29 ), veins indistinct other than Sc, R and CuP; CuP ending at the middle of the hind margin, A almost invisible (Figs 17 , 38 ). Hind wing with indistinct Sc, M bifurcated; CuA with 9 branches, of which four branch again (Figs 18 , 39 ). Legs slender with dense pubescence. Tarsal claws symmetrical and unspecialized. First abdominal tergum specialized, with dense setae medially (Figs 9 , 19 , 30 , 40 ).

Supra-anal plate in ventral view symmetrical, hind margin convex with a weak medial indentation, two paraprocts large and nearly symmetrical (Figs 10 , 20 ). Cerci conical and segmented (Figs 10 , 31 ). Subgenital plate in dorsal view nearly symmetrical; styli modest, cylindroid (Figs 11 , 21 , 32 , 42 ). Left phallomere consisting of three parts: L1, L2 and L3; sclerite L1 folded with a narrow and fingerlike terminus; sclerite L2 large, flat, and folded over posteriorly so that L2v lies on the ventral surface of the phallomere and L2d on the dorsal, sclerite L2d with a rough and curved margin, terminus acute, sclerite L2v with two acute ends posteriorly, of which (in dorsal view) the left one tapers towards the right and the other towards the left; sclerite L3 forming an elongate hook of which the curved part has a small spinous protuberance (Figs 12 , 22 , 33 , 43 ) that is inconspicuous due to the observation angle. Right phallomere consisting of three parts: R1, R2 and R3; sclerite R1 expanding downward towards the left and with a serrate edge; R2, hook-like, expanding towards the left; the basal sclerite of R3 broad and slightly curved, joining with R2. The ventral phallomere (v.ph.) under the right phallomere, flat, posteriorly rounded, with a more or less sclerotized ventral surface (Figs 12 , 22 , 33 , 43 ).

Female. Body black (Figs 3-4 , 48 ). Eyes, ocelli and antennae similar to those of male. Vertex and face reddish brown. Labial palpi and maxillary palpomeres brown. Pronotum black. Abdominal terga black, but with the last segment brown. Abdominal sterna black and center reddish brown. Legs and cerci brown.

Vertex exposed, with 3 longitudinal shining stripes. Face punctuated. Tegmina short, just exceeding the metanotum, with punctures and scattered erect pubescence, heavily sclerotized with metallic shine. Hind wings much reduced. Legs slender, front femur Type A2. Each hind tibia with 2 rows of spines along outer margin. Hind metatarsus exceeding the remaining joints in length.

Supra-anal plate nearly symmetrical, roof shaped, the hind margin nearly blunt and round; paraprocts (pp.) broad and similar (Fig. 23 ). The juncture between the spermatheca plate and the anterior arch (a.a.) membranous and somewhat extensible (Fig. 23 ). First valve (v.I) falciform, sclerotized, with slender base and weakly sclerotized terminus (Figs 23 , 24 , 25 ); laterosternite IX (ltst.IX) large, fused to paratergites (pt.); paratergites (pt.) slender (Fig. 23 ). Second valve (v.II) small and slender, basally fused, connecting to third valve (v.III) by membrane (Figs 23 , 24 , 26 ). Third valve larger than second valve but smaller than first valve, with weakly sclerotized and curved apex, basal portion fused and slightly raised (Figs 23 , 24 , 27 ). Anterior arch (a.a.) claviform, with tapering terminus (Fig. 23 ). Well developed basivalvula (bsv.) strongly sclerotized, fused with the anterolateral deflections of the spermatheca plate. Laterosternal shelf (ltst.sh.) flat, divided by common oviduct opening (c.o.o.). Subgenital plate in dorsal view symmetrical (Fig. 23 ).

Nymph. Body color, characters of pronotum and antennae similar to those of adults. Legs light brown. Cerci reddish brown.

Infraspecific variation. The individual differences in morphological characters mainly involve: 1) the number and shape of smooth areas of pronotum (Figs 7 , 16 , 28 , 37 ); 2) the dentate tine close to the largest tine of the serrated edge of sclerite R1 sclerotized (Figs 12 , 22 ) or not sclerotized (only one case, Figs 33 , 43 ); 3) body color (Figs 1-2 , 5-6 ). We provide pictures for detailed comparison (one male from Hainan, one male from Guangxi) illustrating the appearance of individual differences (Figs 1-2 , 5-6 , 7-12 , 13-22 , 28-33 , 34-43 ). These infraspecific variations cannot separate the populations from each other into different species and the key morphological characters strongly suggest they are conspecific. However, their COI genes show a great genetic divergence among them: the standard barcoding sequence of one (Baisha) of the three Hainan populations has a distance of 3.0% and 3.1% from the other two (Wuzhishan, Baoting) respectively, and it is far distant (4.6%) from the Guangxi population, which in turn is very distinct from the two remaining Hainan populations (Wuzhishan, 6.1%; Baoting, 6.2%).

Male measurements (mm). Body length: 17.5-21.0. Total length including tegmen: 19.0-24.5. Pronotum length × width: 4.9-5.5 × 5.5-6.5. Tegmen length × width: 15.0-20.0 × 5.0-6.5.

Female measurements (mm). Body length: 19.0-20.0. Pronotum length × width: 5.5-6.0 × 6.0-6.5. Tegmen length × width: 4.7-5.0 × 4.7-5.0.

Material examined.

One male and one nymph, China: Hainan Prov., Baisha County, Yinggeling Natural Reserve , Nankai Station , in rotten wood, 21 April 2015, coll. Xinran Li (= Conlin McCat) and Zhiwei Qiu; one male, China: Hainan Prov., Mt. Diaoluoshan , 275m, 18°40.080'N, 109°53.998'E, 25 May 2014, coll. Shunhua Gui and Xinran Li (= Conlin McCat) ; one male, China: Hainan Prov., Baoting County, Maogan Township , 11-12 April 2015, coll. Qikun Bai ; one female, China: Hainan Prov., Mt. Wuzhishan , 795m, 18-21 May 2014, coll. Shunhua Gui, Xinran Li (= Conlin McCat) and Jianyue Qiu; one male, China: Guangxi Aut. Reg., Guiping City , Longtan Park , 386 m, 23°31.140'N, 109°59. 510'E, 31 May-2 June 2014, coll. Shunhua Gui and Xinran Li (= Conlin McCat) ; one male and two females, China: Guangxi Aut. Reg., Fangchenggang City , Shangsi County, Shiwandashan Forest Park, 296 m, 28 June 2015, coll. Lu Qiu and Qikun Bai.

Distribution.

China (new recored; Hainan, Guangxi); Vietnam.

Habitat.

The adult Protagonista lugubris were observed in shrubs at night by the collectors who also found the nymphs and adults in rotten wood. Their rugged pronotum with thickened and raised margins, which resembles that of Cryptocercinae and Panesthiinae is conducive to moving about in rotten wood. However, if Protagonista utilizes the wood tunneled by other organisms or if they bore the wood themselves is yet to be seen. Additionally, wood feeding has not been observed but is still a possibility in Protagonista and the other morphologically similar Archiblattinae .

Figures

Figures 1 - 6. Protagonista lugubris Shelford. 1 - 2 Male from Hainan: 1 dorsal view 2 ventral view 3 - 4 Female from Hainan: 3 dorsal view 4 ventral view 5 - 6 Male from Guangxi: 5 dorsal view 6 ventral view. Scale bars: 1.0 cm.

Figures 7 - 12. Protagonista lugubris Shelford, male from Hainan: 7 pronotum 8 tegmen 9 abdominal tergum 1, dorsal view 10 supra-anal plate and paraprocts, ventral view 11 subgenital plate, dorsal view 12 left phallomere and right phallomere. Scale bars: 1.0 mm.

Figures 13 - 22. Protagonista lugubris Shelford, male from Hainan: 13 head, frontal view 14 maxillary palps 3 - 5 15 front femur 16 pronotum 17 tegmen 18 hind wing 19 abdominal tergum 1, dorsal view 20 supra-anal plate and paraprocts, ventral view 21 subgenital plate, dorsal view 22 left phallomere and right phallomere. Scale bars: 1.0 mm (13, 15 - 16, 19 - 21), 0.5 mm (14, 22), 2.0 mm (17 - 18).

Figures 23 - 27. Protagonista lugubris Shelford, female from Hainan, female genitalia: 23 posterior view and dorsal view of subgenital plate 24 valves and accessory sclerites, dorsal view 25 first valve, ventral view 26 second valve, ventral view 27 third valve, ventral view. Scale bars = 1.0 mm.

Figures 28 - 33. Protagonista lugubris Shelford, male from Guangxi: 28 pronotum 29 tegmen 30 abdominal tergum 1, dorsal view 31 supra-anal plate and paraprocts broken, ventral view 32 subgenital plate, dorsal view 33 left phallomere and right phallomere. Scale bars: 1.0 mm (28, 30 - 33), 2.0 mm (29).

Figures 34 - 43. Protagonista lugubris Shelford, male from Guangxi: 34 head, frontal view 35 maxillary palps 3 - 5 36 front femur 37 pronotum 38 tegmen 39 hind wing 40 abdominal tergum 1, dorsal view 41 supra-anal plate and paraprocts broken, ventral view 42 subgenital plate, dorsal view 43 left phallomere and right phallomere. Scale bars: 1.0 mm (34, 36 - 37, 40), 0.5 mm (35, 41 - 42), 2.0 mm (38 - 39).

Figures 44 - 47. Protagonista species, holotypes and labels. 44 Protagonista lugubris, male 45 Protagonista pertristis stat. rev., female 46 Protagonista fusca, now synonym of Protagonista pertristis, male 47 Protagonista aterrima, now synonym of Protagonista pertristis, female. Scale bars: 5.0 mm.

Figures 48 - 49. 48 Protagonista lugubris Shelford, adult female from Longtan Park in Guiping, Guangxi, 2011. VIII. 4, eating a wasp body near a light trap 49 Protagonista lugubris Shelford, adult male from Diaoluo Mt. in Lingshui, Hainan. Both photographed by XinRan Li (= Conlin McCat).