Brephallus fruhstorferi (Shelford, 1910), comb. n., Brephallus tramlapensis (Anisyutkin, 1999), comb. n.

Pseudophoraspis fruhstorferi Shelford, 1910, by present designation.

Coloration brownish yellow. Pronotum smooth, completely covering vertex, anterior margin curved and posterior margin obtusely produced. Tegmina and wings fully developed in both sexes, entirely covering abdomen, tegmina about twice as long as broad, apices rounded (Figure 2A–F). Hind metatarsus shorter than succeeding tarsal segments combined, with two equal rows of spines along most of its length, 2^nd–4^th segments with large euplantulae. Supra-anal plate and hypandrium nearly symmetrical, posterior margin emarginate near mid-line (Figure 4G).

Figure 2. 

A–D Brephallus fruhstorferi (Shelford, 1910) comb. nov. (male A–B female C–D) E–F Brephallus tramlapensis (Anisyutkin, 1999) comb. nov. (male) G–J P. recurvata (male G–H female I–J) K–N P. clavellata (male K–L female M–N) O–R P. kabakovi (male O–P female Q–R). Scale bars = 10 mm (A–F) Scale bars = 5 mm (G–R).

Male genitalia (Figure 4D–E). Right phallomere similar to that in Morphna, Opisthoplatia, and Rhabdoblatta with well-developed caudal part of sclerite R1T subrectangular in shape, R2 rounded, R3 well developed, widened caudally and fused with R5. Sclerite L2D similar to Rhabdoblatta, divided into basal and apical parts, basal part rod-like, apical part more or less rounded, cap-shaped, but with more bristles. Sclerite L3 with terminal rectangular apex pointed and folded, scattered with bristles.

The new genus differs from other genera of Epilamprinae as follows: 1) male tegmina about twice as long as broad (Figure 2A, E); 2) facial part of head with large brown spot from ocellus to clypeus, basal margin of ocellus with brown spot (Figure 2B, D, F); 3) one third of radius vein of tegmen from base yellowish white (Figure 2A, C, E); 4) sclerite L3 with terminal rectangular, apex pointed (Figure 4D–E).

We propose the name Brephallus, based on the composition of two Latin words (“brevis” and “phallus”) meaning “short phallomere”, in reference to the short L2D sclerite of the male genitalia.

This genus differs from Pseudophoraspis in the apical part of sclerite L2D without a well-developed apical outgrowth. Meanwhile, the mean sequence divergence among species of Brephallus and Pseudophoraspis ranged from 15.2% to 18.8%, larger than that of congeners (Table 4). Although Brephallus fruhstorferi and B. tramlapensis only have the mean interspecific genetic distance of 4.1% (Table 4) between them, they show distinct morphological differences as follows: 1) mid-abdomen of B. tramlapensis (Anisyutkin, 1999) has two brown stripes while B. fruhstorferi (Shelford, 1910) lacks stripes (Figure 2A–F); and 2) the apical part of sclerite L2D of B. tramlapensis (Anisyutkin, 1999) is large and long, with a protrusion in the middle (Figure 4E) while in B. fruhstorferi (Shelford, 1910) it is short, without a protrusion in the middle (Figure 4D).

Epilampra nebulosa Burmeister, 1838.

The species P. clavellata and P. recurvata exhibit sexual dimorphism (male with developed tegmina and wings, females with tegmina reduced to lateral scales and wings absent) (Figure 2G–N); therefore, we provide below supplementary information on the nymphs and females.

Body slender, general color yellowish brown, head entirely covered by pronotum. Pronotum with numerous brown spots, smooth, without or with scattered punctuation. Male with fully-developed tegmina and wings, female with tegmina reduced to lateral scales without wings or with fully-developed tegmina and wings (Figure 2G–R). Hind metatarsus shorter than other tarsal segments combined, with small apical euplantulae along its lower margin, with spinules, euplantulae occupying less than half of its length, with two equal rows of spines along most of its length. Tarsal claws symmetrical and unspecialized. Supra-anal plate semicircular, meso-posterior margin emarginate (Figure 4F).

Male genitalia (Figure 4A–C). Right phallomere with well-developed caudal sclerite, R1T subrectangular in shape (Figure 4A–C “c.p.R1T”), R2 rounded, R3 weakly sclerotized, without branch, narrowed caudally. Sclerite L2D divided into basal and apical parts, basal part rod-like, apical part with well-developed apical outgrowth (Figure 4A–C “a.L2D”), with bristles. Sclerite L3 with apex pointed and folded structure scattered with bristles (Figure 4A–C “f.s.”).

Wang et al. (2013) subdivided the Chinese Pseudophoraspis into two species groups: the fruhstorferi group and the gorochovi group, but the latter lacked information on females. The fruhstorferi group currently includes three species: P. fruhstorferi Shelford, 1910, P. tramlapensis Anisyutkin, 1999 and P. kabakovi Anisyutkin, 1999. Because we have transferred the former two species to the new genus, Brephallus Wang et al., gen. n., the fruhstorferi group is renamed as nebulosa group. Some diagnostic characters between the nebulosa group and the gorochovi group are shown in Table 3.

The mean interspecific sequence divergence among the three Pseudophoraspis members ranged from 4.1% to 9.0% (Table 4), but there are distinguishing differences among them, as described below.

Wang et al. (2013) described the male of P. recurvata including detailed information on male genital structures (Figures 2G–H, 4A, 4F, 5B). The description of the female is provided here.

China: Hainan: five males and one female, Baoting County, 2013.V.2, coll. Yan Shi and Shun-Hua Gui; three males, Changjiang County, Qicha Township, 2015.IV.28, coll. Lu Qiu and Qi-Kun Bai; one male and two females, Sanya City, Liupan Village, 2015.IV.8, coll. Lu Qiu and Qi-Kun Bai; two males (holotype and paratype), Baoting County, 1959.VII.10, coll. Yi-Chuan Hu. China: Guangxi: one male (paratype), Mt. Daqingshan, 1958.IX, coll. Yi-Xin Xu.

(Figures 2I–J, 5A). Body brownish-yellow. Vertex, eyes and between the antennal sockets black-brown. Ocellar spots pale yellowish. Antennae, legs, thorax and abdomen brown. Maxillary palp with 1^st–2^nd segments pale yellowish and 3^rd–5^th segments brown. Cerci brown with apical segment yellow.

Head longer than wide. Interocular space slightly less wide than interocellar space, ocellar spots rather small, eyes elongate. Antennae short, not reaching to half length of body, first segment of flagellum twice length of next segment; interantennal portion of frons concave. Frons moderately punctuated; clypeus and labrum unmarked. Pronotum covering vertex of head, anterior margin arcuate, posterior margin truncate, with scattered punctuation and a pair of impressions on disc. Thoracic and abdominal tergites with small tubercles and longitudinal inflations along posterior margins. Tegmina reduced to lateral scales, with nearly indistinct venation, veins reduced, wings absent. Anterior margin of fore femur type B, with six large spines and one single apical spine. Tibial spines well developed. 3^rd–7^th abdominal tergites with paired rounded impressions. Hind metatarsus with spines along most of its length, equal to remaining joints, tarsal spines absent. Tarsal claws symmetrical, simple, arolia very small. Supra-anal plate transverse, beyond the subgenital plate, hind margin with a medial V-shaped excavation. Hypandrium widely rounded, caudal margin arcuate. Cerci abbreviated, apex blunt.

Morphology of paratypes is same as female type described above, but with following variation: five to six large spines scattered along anterior margin of fore femora; color of clypeus, labrum and abdomen tergites brown or yellow. Overall length: 20.1 ± 0.2 mm; head length × width: 3.6 ± 0.1 mm × 2.9 ± 0.1 mm; pronotum length × width: 6.2 ± 0.1 mm × 10.7 ± 0.1 mm.

China (Hainan, Guangxi).

Wang et al. (2013) described the male of P. clavellata including the male genitalic structures (Figures 2K–L, 4B and 5C). Description of the female and nymph is provided here.

China: Yunnan: Thirty males and one female, Pu’er City, Meizi Lake, 2016.V.20, coll. Lu Qiu and Zhi-Wei Qiu; two males, Jinhong City, Dadugang, 2014.VI.29, coll. Conlin McCat (= Xin-Ran Li) and Hong-Guang Liu; one nymph, Xishuangbanna, Menghai County, Bulong Natural Reserve, 2017.I.31, coll. Jian-Yue Qiu and Hao Xu; male (holotype), Xishuangbanna, 1981.V.27-30, coll. Zhi-Gang Zheng.

(Figures 2M–N, 3E). Identical to the female of P. recurvata but body larger; in addition, legs, venter of thorax and abdomen yellow.

Overall length 28.1 mm; head length × width: 3.8 mm × 3.7 mm; pronotum length × width: 7.0 mm × 12.5 mm.

(Figure 3C–D). Body flattened. Identical to adult female but lacking wings.

Figure 3. 

A–B P. kabakovi (nymph) scale bars = 5 mm C–D P. clavellata (nymph) E abdomen of female of P. clavellata F–G Brephallus fruhstorferi (Shelford, 1910) comb. nov. (nymph).

China (Yunnan).

The male of P. kabakovi was described (Figures 2O–P, 4C, 5E) by Anisyutkin (1999) and Wang et al. (2013), but little was known about the female and nymph until now.

Figure 4. 

Male genitalia of Pseudophoraspis and Brephallus Wang et al., gen. nov. A P. recurvata B P. clavellata C P. kabakovi D Brephallus fruhstorferi (Shelford, 1910) comb. nov. E Brephallus tramlapensis (Anisyutkin, 1999) comb. nov. F supra-anal plate of P. recurvata G supra-anal plate of Brephallus tramlapensis (Anisyutkin, 1999) comb. nov. (Scale bars = 1 mm).

China: Yunnan: One male, Xishuangbanna, 1974.IV.13, coll. Yao Zhou and Feng Yuan; twenty males, five females and one nymph, Xishuangbanna, Menglun Town, 2016.V.27, coll. Lu Qiu and Zhi-Wei Qiu; one male and two females, Xishuangbanna, Mengla County, Wangtianshu, 2016.V.23, coll. Lu Qiu and Zhi-Wei Qiu.

(Figures 2Q–R, 5F). Body yellowish brown. Eyes and antennae black, ocellar spots pale yellow. Pronotum with dense small brown spots. Tegmina with scattered large black spots. Abdominal sterna with small and fewer large black dots, large black dots along the hind margins of the segments. Cerci brown.

Figure 5. 

A female of P. recurvata from Hainan Province. This specimen, collected as a nymph in 8 April 2015, was reared at Southwest University by Lu Qiu and adult emergence occurred in 21 May 2015 B male of P. recurvata from Hainan Province C male of P. clavellata from Yunnan Province D habitat of P. clavellata E male of P. kabakovi from Yunnan Province F female of P. kabakovi from Yunnan Province. A–F Photographed by Lu Qiu G–H Brephallus fruhstorferi (Shelford, 1910) comb. nov. from Hainan Province (Photographed by Xin-Ran Li).

Similar to male in general appearance, but shorter and convex. Tegmina and wings shorter than in males. Fore femur with six spines along anterior margin and one single apical spine. Hind metatarsus with two rows of spines along most of its length. Claws symmetrical, simple; arolium well developed. Abdominal terga unspecialized. Supra-anal plate caudal margin with a medial V-shaped excavation. Hypandrium posterior margin emarginate near mid-line.

Overall length 32 ± 0.2 mm; head length × width: 4.2 ± 0.1 mm × 3.6 ± 0.1 mm; pronotum length × width: 8.3 ± 0.2 mm × 12.1 ± 0.2 mm; tegmina length × width: 25.4 ± 0.1 mm × 10.3 ± 0.2 mm.

Identical to adult females of P. recurvata and P. clavellata except for undeveloped wing (Figure 3A–B).

China (Yunnan); Vietnam.