Research Article |
Corresponding author: Yanli Che ( shirleyche@sohu.com ) Academic editor: Gavin Svenson
© 2014 Xiudan Wang, Zongqing Wang, Yanli Che.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Wang X, Wang Z, Che Y (2014) A taxonomic study of the genus Panesthia (Blattodea, Blaberidae, Panesthiinae) from China with descriptions of one new species, one new subspecies and the male of Panesthia antennata. ZooKeys 466: 53-75. https://doi.org/10.3897/zookeys.466.8111
|
One new species Panesthia guizhouensis sp. n. and one new subspecies Panesthia stellata concava ssp. n. are described and illustrated. The male of Panesthia antennata Brunner von Wattenwyl, 1893 and its brachypterous form are described and illustrated for the first time. Panesthia strelkovi Bey-Bienko, 1969 is redescribed and illustrated. Three known species, Panesthia birmanica Brunner von Wattenwyl, 1893, Panesthia sinuata Saussure, 1839 and Panesthia angustipennis cognata Bey-Bienko, 1969 are illustrated. In addition, a key to all species of the genus Panesthia from China is presented.
Wing polymorphism, brachypterism, illustration, key, Salganea
The wood-feeding cockroach genus Panesthia was established by
In this paper, we report one new species and one new subspecies, and also provide a key including nine species and three subspecies of the Panesthia from China. We also take this opportunity to report the male and the brachypterous form of P. antennata for the first time.
The terminology of the head, body and male genitalia used in this paper mainly follows
Panesthia Serville, 1831: 38;
Proterodia Costa, 1866: 5. Type species: Proterodia punctatissima Costa, 1866. Synonymized by
Dicellonotus Butler, 1882: 387. Type species: Dicellonotus lucanoides Butler, 1882. Synonymized by
(mainly following
Anterolateral corners of terga rarely with holes and without setae, or just tergum six (T6) and tergum seven (T7) with holes. Lateral margins of T6 smooth, and laterocaudal angles not produced, or with a spine and directed caudally. Lateral margins of T7 straight and not crenulate, laterocaudal angles sometimes produced and usually directed caudally. Lateral margins of sternite seven (S7) with a feeble and short ridge or without ridge. In the male, the hind margin of the last sternite is truncate or concave, and the subgenital fig is slightly exposed. In the female, the hind margin of the last sternite is convex and rounded. Both sexes are without styli. Supra-anal fig punctate, with uneven or rounded hind margin, and cerci are short and broad basally. Paraprocts are asymmetrical, the left one in ventral view with a finger-like projection lacking in the right one. Anterior margin and lateral margins of subgenital fig concave and the hind margin is rounded. Four genital phallomeres as follow: first sclerite of the left phallomere (L1) figd; second ventromedial sclerite of left phallomere (L2vm) rod-like; second dorsal sclerite of the left phallomere (L2d) variable; second sclerite of the right phallomere (R2) well developed or reduced, if developed, it is often hook-like and curved to right side in dorsal view.
The genus Panesthia is recognized by both T6 and T7 having smooth lateral margins, their hind margins without tubercles; the hind angles of T7 spine-like, but T6 not. Some species in this genus may have individuals with mixed characters resembling the genus Salganea Stål, 1877, Ancaudellia Shaw, 1925 or Miopanesthia Saussure, 1895 (
Oriental Region, Australian Region, a few locations in the Palaearctic Region (China, Japan).
1 | Tegmina and wings absent | P. larvata |
– | Tegmina and wings present, or tegmina present and wings absent | 2 |
2 | Tegmina reduced (Fig. |
P. strelkovi |
– | Tegmina and wings well developed or reduced | 3 |
3 | Tegmina with more than one colour in the form of spots | P. transversa |
– | Tegmina unicoloured without spots | 4 |
4 | Laterocaudal angles of T6 acute and posteriorly directed | P. birmanica |
– | Laterocaudal angles of T6 not acute | 5 |
5 | Anterior margin of pronotum broadly excavated and with a projection mesally in male, slightly concave and without projection in female | 6 |
– | Anterior margin of pronotum slightly concave and without middle projection in both sexes | 8 |
6 | Body length < 30 mm, anteroventral margin of front femur with 0–1 spine | P. sinuata |
– | Body length > 30 mm, anteroventral margin of front femur with two spines or more (P. angustipennis complex) | 7 |
7 | Hind margin of supra-anal fig entire dorsally or weakly undulate; median phallomere L2d elongate, tapering to a round apex ( |
P. angustipennis spadica |
– | Hind margin of supra-anal fig undulate dorsally; median phallomere L2d short with variable apex (Fig. |
P. angustipennis cognata |
8 | Hind margin of supra-anal fig slightly crenate in ventral view (Fig. |
P. guizhouensis sp. n. |
– | Hind margin of supra-anal fig crenulate in ventral and dorsal view | 9 |
9 | Vertex with a foveola in dorsal view (Fig. |
P. antennata |
– | Vertex without foveola (Fig. |
P. stellata concava ssp. n. |
1–2 Panesthia antennata Brunner von Wattenwyl, 1893, male: 1 dorsal view 2 ventral view 3–4 Panesthia antennata Brunner von Wattenwyl, 1893, nymph: 3 dorsal view 4 ventral view 5–6 brachypterous form of Panesthia antennata Brunner von Wattenwyl, 1893, male: 5 dorsal view 6 ventral view 7–8 brachypterous form of Panesthia antennata Brunner von Wattenwyl, 1893, nymph: 7 dorsal view 8 ventral view 9–10 Panesthia stellata concava ssp. n., male: 9 holotype, dorsal view 10 same, ventral view 11–12 Panesthia stellata concava ssp. n., nymph: 11 paratype, dorsal view 12 same, ventral view 13–14 Panesthia guizhouensis sp. n., male: 13 holotype, dorsal view 14 same, ventral view 15–16 Panesthia guizhouensis sp. n., nymph: 15 paratype, dorsal view 16 same, ventral view. Scale bars = 1.0 cm.
17–18 Panesthia strelkovi Bey-Bienko, 1969, male: 17 dorsal view 18 ventral view 19–20 Panesthia birmanica Brunner von Wattenwyl, 1893, male: 19 dorsal view 20 ventral view 21–22 Panesthia sinuata Saussure, 1895, male: 21 dorsal view 22 ventral view 23–24 Panesthia angustipennis cognata Bey-Bienko, 1969, female: 23 dorsal view 24 ventral view 25–26 Panesthia angustipennis cognata Bey-Bienko, 1969, nymph: 25 dorsal view 26 ventral view. Scale bars = 1.0 cm.
Panesthia antennata Brunner von Wattenwyl, 1893: 50;
Male. Body black or dark brown (Figs
Vertex punctate, with a small foveola in dorsal view, which is exposed to the excision of pronotum in anterior margin (Fig.
Panesthia antennata Brunner von Wattenwyl, 1893 27 vertex and pronotum 28 tegmen 29 hind wing 30 abdominal tergum 7, dorsal view 31 abdominal sternite 7, ventral view 32 supra-anal fig and paraprocts, ventral view 33 subgenital fig, dorsal view 34 left phallomere (L1) 35 median phallomere (L2vm and L2d) 36 right phallomere (R2). Scale bars = 1.0 mm (Figs 27, 30–33), 2.0 mm (Figs 28–29), 0.5 mm (Figs 34–36).
Male genitalia. Genital phallomere L1 well developed but slightly sclerotized (Fig.
Female. Similar to male, but hind margin of S7 rounded. In all specimens observed, the anterior margin of pronotum with an excision. In brachypterous form, tegmina and wings also reduced similar to males.
Nymph. Body black and punctate, with two yellowish brown marks on the mesonotum and metanotum without marks (Figs
Measurements. Male, body length: 29.1–32.0 mm; pronotum: length/width: 6.5–6.9/9.6–10.5 mm; width of excision of anterior margin of pronotum: 1.7–2.0 mm; distance between disc tubercles: 1.5–2.0 mm; tegmen: 22.7–26.4 mm. In brachypterous form, body length: 28.5–35.5 mm; pronotum: length/width: 6.8–7.1/9.8–11.3 mm; width of excision of anterior margin of pronotum: 2.0–2.5 mm; distance between disc tubercles: 1.9–2.5 mm; tegmen: 13.5–16.7 mm.
Female, body length: 29.7–32.6 mm; pronotum: length/width: 6.5–7.0/10.1–10.3 mm; width of excision of anterior margin of pronotum: 1.7–2.0 mm; distance between disc tubercles: 1.9–2.0 mm; tegmen: 23.0–26.8 mm. In brachypterous form, body length: 32.0–32.5 mm; pronotum: length/width: 6.1–7.1/9.6–11.1 mm; width of excision of anterior margin of pronotum: 1.6–2.5 mm; distance between disc tubercles: 2.0–2.4 mm; tegmen: 12.6–16.0 mm.
Two males, two females and one nymph, China: Yunnan Prov., Tengchong County, Mt. Gaoligong, 9 August 2005, coll. Benyong Mao (HBU); 13 males, six females and 32 nymphs, Yunnan Prov., Tengchong County, Lingjiapu, 13–14 August 2006, coll. Biao Liu (SWFU).
In brachypterous form, one male and one female, China: Yunnan Prov., Tengchong County, Mt. Gaoligong, Baihualing, 30 July 2012, coll. Jishan Xu and Lingxiao Chang (HBU); two males, two females and one nymph, Yunnan Prov., Longling County, Longxin Village, Mt. Hei, 2300m, 23–25 July 2008, coll. Jishan Xu and Zhenhua Gao (HBU); one male and one female, China: Yunnan Prov., Nanjian County, Mt. Wuliang, 8–9 July 2006, coll. Benyong Mao and Juntong Lang (HBU); one male and one nymph, China: Yunnan Prov., Nanjian County, Mt. Wuliang, 17 July 2003, coll. Benyong Mao (HBU); three males and five nymphs, China: Yunnan Prov., Longchuan County, Mt. Ping, 1800m, 6 November 2003, coll. Jinxin Song (SWFU); one male, China: Yunnan Prov., Lijiang, Snow Mt. Yulong, Yuanbinping, 3228m, 31 October 2007, coll. Biao Liu (SWFU); one male and one female, China: Yunnan Prov., Tengchong County, Dahaoping, 2000m, 3 May 2002, coll. Jinxin Song and Yingxian Situ (SWFU); one male, China: Yunnan Prov., Baoshan City, Baihualing, 1980m, 12 April 2002, coll. Yingxian Situ (SWFU); one male and one nymph, Yunnan Prov., Mt. Wuliang, 2000m, 17 July 2003, coll. Benyong Mao (DLU).
China (Yunnan); Myanmar.
Male. Body black (Fig.
Vertex and face punctate, the former exposed and without foveola (Fig.
Panesthia stellata concava ssp. n. 39 vertex and pronotum 40 tegmen 41 hind wing 42 abdominal tergum 7, dorsal view 43 abdominal sternite 7, ventral view 44 supra-anal fig and paraprocts, ventral view 35 subgenital fig, dorsal view 46 left phallomere (L1) 47 median phallomere (L2vm and L2d) 48 right phallomere (R2). Scale bars = 1.0 mm (Figs 39, 42–45), 2.0 mm (Figs 40–41), 0.5 mm (Figs 46–48).
Male genitalia. Genital phallomere L1 well developed (Fig.
Female. Essentially similar to male, differs with the anterior margin of pronotum weakly concave as well as S7 with rounded hind margin.
Nymph. Body black and punctate, with a broad yellow band on the mesonotum which extends to the middle of the metanotum, hind border of the mark concave (Figs
Measurements. Male, body length: 27.0–32.5 mm; pronotum: length/width: 5.9–7.3/9.4–11.2 mm; width of excision of anterior margin of pronotum: 1.3–2.0 mm; distance between disc tubercles: 1.6–2.1 mm; tegmen: 22.1–26.7 mm.
Female, body length: 28.5–32.5 mm; pronotum: length/width: 6.5–7.1/10.3–11.0 mm; width of excision of anterior margin of pronotum: 1.0–1.3 mm; distance between disc tubercles: 1.5–2.0 mm; tegmen: 23.8 mm.
Holotype, male, China: Xizang Prov., Chayu County, Shangchayu Town, 8 August 2013, coll. Xinglong Bai and Junsheng Wang (HBU). Paratypes, four males, one female and four nymphs, same data as holotype (HBU); one male, two females and two nymphs, Sichuan Prov., Dege County, Gengqing Town, 3270m, 19 July 2009, coll. Guodong Ren (HBU); one female, China: Xizang Prov., Chayu, 2000m, 24 August 1973, coll. Fusheng Huang (SWU).
This subspecies is close to P. stellata stellata Saussure, 1895, but can be distinguished by the following characteristics: 1) nymph with broad yellowish band on mesonotum and metanotum, nymph of P. stellata stellata with two yellowish markings on mesonotum and without markings on metanotum; 2) anterior margin of pronotum with an excision in both sexes and the corners of the excision upturned in male; anterior margin of pronotum entire or slightly concave in both sexes of P. stellata stellata.
The subspecific epithet is derived from the Latin word “concavus” which refers to the hind margin of the yellowish mark on nymphs being concave.
Male. Body dark brown or black (Fig.
Vertex slightly punctate, exposed (Fig.
Panesthia guizhouensis sp. n. 49 vertex and pronotum 50 tegmen 51 hind wing 52 abdominal tergum 7, dorsal view 53 abdominal sternite 7, ventral view 54 supra-anal fig and paraprocts, ventral view 55 subgenital fig, dorsal view 56 left phallomere (L1) 57 median phallomere (L2vm and L2d) 58 right phallomere (R2). Scale bars = 1.0 mm (Figs 49–55), 0.5 mm (Figs 56–58).
Male genitalia. Genital phallomere well developed, L1 well sclerotized (Fig.
Female. Differs from male as follows: the anterior margin of pronotum weakly concave and the corners not upturned; the tubercles on surface smaller than in male. Hind margin of S7 rounded.
Nymph. Body black and punctuate, with two yellowish brown marks on the mesonotum and metanotum without marks (Figs
Measurements. Male, body length: 24.5–27.0 mm; pronotum: length/width: 4.2–5.2/ 7.9–9.0 mm; width of excision of anterior margin of pronotum: 1.5–1.8 mm; distance between disc tubercles: 1.1–1.5 mm; tegmen: 22.0–25.6 mm.
Female, body length: 26.8–31.5 mm; pronotum: length/width: 4.5–5.0/8.5–9.0 mm; width of excision of anterior margin of pronotum: 1.1–1.3 mm; distance between disc tubercles: 1.3–1.5 mm; tegmen: 21.2–23.2 mm.
Holotype, male, China: Guizhou Prov., Zunyi City, Suiyang County, Kuankuoshui Natural Reserve, 14 August 2010, coll. Keliang Wu (SWU). Paratypes, five males, two females and five nymphs, same data as holotype (SWU); four males, 12 females and 13 nymphs, Guizhou Prov., Zunyi City, Suiyang County, Kuankuoshui Natural Reserve, 1–2 August 2013, coll. Xiudan Wang and Yuhong Zheng (SWU).
This species is similar to P. angustipennis spadica, but can be distinguished by the following characteristics: 1) anterior margin of pronotum weakly concave in male, and without mesal elevation, male of latter with anterior margin broadly excised and with mesal elevation; 2) body length < 30 mm in most, or few surpassing 30 mm, body length of P. angustipennis spadica > 30 mm.
The specific epithet is named after the locality of the holotype, Guizhou Province.
Panesthia strelkovi Bey-Bienko, 1969: 834;
Male. Body black (Fig.
Face punctulate, with weak ocelli; vertex slightly exposed (Fig.
Panesthia strelkovi Bey-Bienko, 1969 59 vertex and pronotum 60 tegmen 61 abdominal tergum 7, dorsal view 62 abdominal sternite 7, ventral view 63 supra-anal fig and paraprocts, ventral view 64 subgenital fig, dorsal view 65 left phallomere (L1) 66 median phallomere (L2vm and L2d) 67 right phallomere (R2). Scale bars = 1.0 mm (Figs 59, 61–62), 0.5 mm (Figs 60, 63–67).
Male genitalia.L1 well developed (Fig.
Measurements. Male, body length: 29.0–37.0 mm; pronotum: length/width: 7.2–8.2/10.0–13.0 mm; width of excision of anterior margin of pronotum: 1.0–1.7 mm; distance between disc tubercles: 2.0–2.5 mm; tegmen: 3.6–5.5 mm.
Two males, China: Hainan Prov., Mt. Diaoluo, 12 May 1965, coll. Sikong Liu; one male, China: Hainan Prov., Mt. Jianfengling, 17 April 1982, coll. Zhiqin Chen. (SWU)
China (Hainan).
Panesthia birmanica Brunner, 1893: 54;
This species is distinguished from other species by its small body length, ranging from 22.5 mm to 28 mm; and also by pronotum virtually flattened, anterior margin slightly thickened, entire or weakly indented in male, entire and not thickened in female (Fig.
One female, China: Hainan Prov., Mt. Jianfengling, Tianchi, 6 July 1981, collector unknown; one female, China: Hainan Prov., Changjiang, Mt. Bawangling, 8–13 July 2006, coll. Jiliang Wang and Chao Gao; three females, China: Yunnan Prov., Xishuangbanna, Dadugang, 27 April 2014, Xinran Li. (SWU).
China (Hainan, Yunnan); India; Myanmar; Vietnam; Thailand.
Panesthia sinuata Saussure, 1895: 318;
This species is similar to P. angustipennis spadica, but can be distinguished by body length (19–29 mm), which is commonly smaller than the latter (34–42 mm). In male of P. sinuata, the anterior margin of pronotum is broadly concave and the corners of the indentation upturned (Fig.
Panesthia sinuata Saussure, 1895 74 vertex and pronotum 75 tegmen 76 hind wing 77 abdominal tergum 7, dorsal view 78 abdominal sternite 7, ventral view 79 supra-anal fig and paraprocts, ventral view 80 subgenital fig, dorsal view 81 left phallomere (L1) 82 median phallomere (L2vm and L2d) 83 right phallomere (R2). Scale bars = 1.0 mm (Figs 74–78), 0.5 mm (Figs 79–83).
Two males and one female, China: Yunnan Prov., Xishuangbanna, Menghai, 1200–1600m, 26 July 1958, coll. Shuyong Wang; one male, China: Yunnan Prov., Xishuangbanna, 1050–1080m, 9 June 1958, coll. Fuji Pu. (SWU)
China (Yunnan, Guangdong); Vietnam; Laos; Malaysia.
Panesthia cognata Bey-Bienko, 1969: 833;
Panesthia angustipennis cognata,
This subspecies resembles P. angustipennis spadica, and it is difficult to distinguish adults.
Panesthia angustipennis cognata Bey-Bienko, 1969 84 vertex and pronotum 85 tegmen 86 hind wing 87 abdominal tergum 7, dorsal view 88 abdominal sternite 7, ventral view 89 supra-anal fig and paraprocts, ventral view 90 subgenital fig, dorsal view 91 left phallomere (L1) 92 median phallomere (L2vm and L2d) 93 right phallomere (R2). Scale bars = 2.0 mm (Figs 84–90), 0.5 mm (Figs 91–93).
One nymph, China: Hainan Prov., Mt. Jianfengling, 29 June 1981, coll. Kegang Hu; one nymph, China: Hainan Prov., Ledong County, 25 December 1963, coll. Yuliang Luo; one nymph, China: Hainan Prov., Mt. Jianfengling, Tianchi, 24 February 1982, Ruilin Pan; one female, China: Hainan Prov., Mt. Jianfengling, 24 November 1981, Ruilin Pan; one nymph, China: Hainan Prov., Mt. Jianfengling, 8 July 1981, collector unknown; one nymph, China: Hainan Prov., Mt. Jianfengling, Tianchi, 8–10 May 1964, coll. Hui Ren; one nymph, China: Hainan Prov., Mt. Jianfengling, 11 May 1981, collector unknown; one female, China: Hainan Prov., Ledong County, Mt. Jianfengling, 1050m, 6–7 December 2007, coll. Weiwei Zhang; one female, China: Hainan Prov., Mt. Jianfengling, Tianchi, 26 July 1983, coll. Lishen Hua; one nymph, China: Hainan Prov., Wanning City, Xinglong, 23 February 1964, coll. Sikong Liu; one female, China: Hainan Prov., Mt. Jianfengling, 9 May 1964, coll. Sikong Liu; one nymph, China: Yunnan Prov., Menglun, 1 August 2009, coll. Zongqing Wang; one nymph, China: Yunnan Prov., Yingjiang County, Tongbiguan, 1450m, 30 July 2009, coll. Benyong Mao; one male, China: Guizhou Prov., Liping County, Mt. Taiping, 27 July 2009, coll. Yang; two males, one female and one nymph, China: Xizang Prov., Motuo, 1100m, 5 January 1983, coll. Yinheng Han; one female, China: Xizang Prov., Motuo, 700-1050m, 23 June 1983, coll. Yinheng Han; one male, China: Xizang Prov., Chayu County, Songgu, 23 July 1972, coll. Fusheng Huang. (SWU). One male, one female and one nymph, China: Guangxi Prov., Hechi City, Tian'e County, 14–19 September 2002, coll. Ming Bai. (HBU)
China (Hainan, Guangxi, Guizhou, Yunnan, Xizang); India; Sikkim; Myanmar; Vietnam; Laos; Thailand.
Gregarious, xylophagous cockroaches of the blaberid genus Panesthia exhibit little variation in morphology. They have dark, hard, rigid and pitted exoskeletons. Body sizes range from 15 mm to over 50 mm. They usually live in decaying wood, fallen leaves, rubbish, cracks in rocks, or in some kind of debris, and feed on wood (
Nymphs of P. angustipennis spadica have two morphs. One has a pair of large reddish spots on mesonotum (
Panesthia is the only genus of the subfamily Panesthiinae in which several species and subspecies have tegmina and wings which are fully developed or variably reduced (
The reason for wing polymorphism in cockroaches is still unknown.
We are sincerely grateful to Prof. J. R. Schrock (Department of Biological Sciences, Emporia State University, USA) for revising the manuscript and also thanks to Prof. Guodong Ren (Hebei University, China), Prof. Xiaohong Ou (Southwest Forest University, China) and Prof. Benyong Mao (Dali University, China) for their kindness in loaning specimens to us. This study is supported by the National Natural Sciences Foundation of China (Nos. 30900146, 31093430, 31472026), and also partly by the Project-sponsored by SRF for ROCS, SEM.