Research Article |
Corresponding author: Yanli Che ( shirleyche2000@126.com ) Academic editor: Dominic Evangelista
© 2016 Chenchen 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 C, Wang Z, Che Y (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: 57-73. https://doi.org/10.3897/zookeys.574.7111
|
The blattid genus Protagonista Shelford, 1908, which is tentatively assigned to the subfamily Archiblattinae (= Planeticinae), is reported from China for the first time with illustrations and description of P. lugubris Shelford, 1908. It is a wood-dwelling and potentially a wood-feeding species. The male and female genitalia of Protagonista are described and illustrated for the first time. The species Protagonista pertristis Hanitsch, 1923 is revived from the synonymy of P. lugubris, and the remaining three nominal species that were also considered as synonyms of P. lugubris are now recognized as synonyms of P. pertristis. In agreement with
Planeticinae , new record, synonym, taxonomy, Eroblatta
The cockroach subfamily Archiblattinae (= Planeticinae) belongs to the family Blattidae and is distributed in southeast Asia. Archiblattinae has a controversial taxonomic history. Planeticidae was erected by
The genus Protagonista was established by
Previous studies of the subfamily Archiblattinae are fairly limited. In this paper, we report one known species P. lugubris newly discovered from China, distributed in Hainan and Guangxi, of which the male and female genitalia are described in detail for the first time. A key to all species of Protagonista and the related genus Eroblatta is given. The taxonomic status of this subfamily and the genus Eroblatta, as well as the validity of the name Archiblattinae, are discussed. We also deal with the synonymy of Protagonista lugubris based on the examination of holotypes, geographical distribution and original descriptions.
The terminology for the body, male and female genitalia used in this paper mainly follows
The terms of veins (abbreviations given in parentheses) are: subcosta (Sc), radius (R), radius anterior (RA), radius posterior (RP), media (M), cubitus anterior (CuA), cubitus posterior (CuP), anal (A), anal anterior (AA), anal posterior (AP). The terms of female genitalia (abbreviations given in parentheses) are: paraprocts (pp.), anterior arch (a.a.), paratergites (pt.), first valve (v.I), second valve (v.II), third valve (v.III), laterosternite IX (ltst.IX), basivalvula (bsv.), laterosternal shelf (ltst.sh), common oviduct opening (c.o.o.), spermathecal opening (sp.o.), and vestibular sclerite (vst.s.).
The terminology of
The standard barcoding sequences of the mitochondrial COI gene (658 bp) of Protagonista lugubris from Hainan and Guangxi are approved, which are deposited in GenBank under the accession numbers KU511283, KU511284, KU511285 and KU511286.
Planeticidae Walker, 1868: 25;
Archiblattinae Kirby, 1904: 148, as a substitute name based on the synonymy of Planetica with Archiblatta;
Archiblattidae:
Protagonista Shelford, 1908: 158;
The genus Protagonista is remarkable on account of the shape of the pronotum (as long as broad, quadrangular, with rounded angles, sides not deflexed), and the pubescence on its pronotum and tegmina (after
Antennae slightly moniliform. Ocelli present. Pronotum as long as broad, quadrangular, with rounded angles, sides not deflexed and not covering vertex (Figs
Vietnam; Malaysia (Malacca State); Sumatra; Borneo; China (new record; Hainan, Guangxi).
Protagonista lugubris Shelford, 1908: 158;
Male. Body slender, dark brown to black (Figs
Head with vertex punctate, with three smooth longitudinal stripes (Figs
Supra-anal plate in ventral view symmetrical, hind margin convex with a weak medial indentation, two paraprocts large and nearly symmetrical (Figs
Female. Body black (Figs
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.
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
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.
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).
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).
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.
China (new recored; Hainan, Guangxi); Vietnam.
The adult P. 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.
Validity of the name Archiblattinae. As explained in the introduction, the scientific names Archiblattinae and Planeticinae are synonyms. According the Principle of Priority in ICZN, Archiblattinae should be abandoned and the earlier name Planeticinae is valid although its type genus is no longer valid. But the name Planeticinae/–idae has been ignored for a long time and the substitute name Archiblattinae/–idae is in prevailing usage since
Taxonomic status of subfamily Archiblattinae and its genera. The subfamily Archiblattinae has a controversial taxonomic history since it was established. Although
Synonymy of Protagonista lugubris.
Taxonomic status of the genus Eroblatta. The genus Eroblatta was erected based on Protagonista borneensis Shelford, 1908 by
Protagonista | |
P. lugubris Shelford, 1908 | Vietnam (Tonkin, type locality), South China |
P. pertristis Hanitsch, 1923, stat. rev. | Sunda Shelf including Malay Peninsula (type locality) |
syn. P. fusca Hanitsch, 1925 | Sarawak |
syn. P. aterrima Hanitsch, 1929 | Sumatra |
syn. P. laeta Hanitsch, 1931 | Singapore |
Eroblatta | |
E. borneensis Shelford, 1908 | Borneo (Sarawak, type locality) |
1 | Tibia with 3 rows of spines along outer margin | Eroblatta borneensis |
– | Tibia with 2 rows of spines along outer margin | (Protagonista) 2 |
2 | Coxae of all legs with the distal half and the base of femora orange yellow | Protagonista pertristis |
– | All legs uniform in colour | Protagonista lugubris |
We are sincerely grateful to reviewers Marie Djernæs, Michael Kotyl and Sonia Lopes for providing helpful comments on this paper. We also express our thanks to Amoret Spooner (Oxford University Museum of Natural History, Hope Entomological Collections) and Gunvi Lindberg (Naturhistoriska Riksmuseet) for providing photographs of the holotypes. We thank colleagues Lu Qiu, Qikun Bai, Xinran Li (= Conlin McCat), Zhiwei Qiu, Shunhua Gui and Jianyue Qiu for collecting specimens, among them Xinran Li also took the photos of Protagonista lugubris in the wild.We also thank Prof. John Richard Schrock (Department of Biological Sciences, Emporia State University, USA) for revising this manuscript.
This project was supported by a Program of Ministry of Science and Technology of the People’s Republic of China (2015FY210300), the National Natural Sciences Foundation of China (31472026, 31093430 and 31493021) and The Fundamental Research Funds for the Central Universities, China (XDJK2013B013, 2362015XK04).