Review Article |
Corresponding author: Yanli Che ( shirleyche2000@126.com ) Academic editor: Pavel Stoev
© 2019 Shuran Liao, 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:
Liao S, Wang Z, Che Y (2019) A new genus and a new species in the subfamily Polyzosteriinae (Blattodea, Blattidae) from China. ZooKeys 852: 85-100. https://doi.org/10.3897/zookeys.852.33325
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Laevifacies quadrialata gen. et sp. nov. is described from Hainan Province, China based on morphological data. COI data (DNA barcodes) is utilized to confirm the sexual dimorphism occurring in Laevifacies quadrialata gen. et sp. nov. Melanozosteria nitida Brunner von Wattenwyl, 1865, is reported from Guangxi Province, China. A key to the Chinese Polyzosteriinae is provided.
Blattaria, cockroaches, Laevifacies, Melanozosteria, molecular identification, morphology
Polyzosteriinae is a relatively species-abundant subfamily in the Blattidae. The subfamily is flightless (except Methanini), having lobiform vestigial tegmina or being totally apterous. Some species have pits or tubercles scattered on the pronotum (
The genus Melanozosteria was established with Polyzosteria nitida Brunner von Wattenwyl, 1865 as type species (
DNA barcodes have been proven to be a helpful method to identify species and to successfully match male and female. Barcoding has been applied to resolve the problems of sexual dimorphism and even to identify nymphs in cockroaches (
Morphological terminology used in this paper mainly follows
We used two cockroach specimens for COI sequencing in this study in order to resolve the sexual dimorphism. Both sequences are deposited in GenBank with the accession numbers: MK798103, MK798104 (Table
Family | Species | Accession number | Reference | |
Outgroups | Mantidae | Mantis religiosa | KM529415 | Hebert et al. 2015 (Unpublished) |
Mantis religiosa | KR148854 |
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Ingroups | Blattidae | Laevifacies quadrialata sp. nov. | MK798103 | |
Laevifacies quadrialata sp. nov. | MK798104 | |||
Periplaneta australiasiae | KX640825 |
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Shelfordella lateralis | KU684413 |
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Neostylopyga rhombifolia | KP986425 |
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Hebardina concinna | KF640073 |
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Methana parva | KP986422 |
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Angustonicus lifou | KP986393 |
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A total of ten COI sequences were analyzed (two sequences of Laevifacies species from our study, six sequences of Blattidae, and two sequences of a mantid outgroup downloaded from GenBank) (Table
In this study, we acquired two COI sequences, whose length, excluding primers, was 658 bp each. The genetic divergence value between male and female of Laevifacies quadrialata sp. nov. is 0.9%; however, the interspecific K2P genetic divergence among Laevifacies quadrialata sp. nov. and other species ranged from 10.4 to 13.1%.
The ML phylogenetic tree (Figure
Polyzosteriinae Tepper, 1893: 32;
1 | Sexual dimorphism present. Body small; tegmina and hind wings vestigial in male; tegmina vestigial and hind wings absent in female | Laevifacies quadrialata sp. nov. |
– | Sexes similar. Body large; tegmina vestigial and hind wings absent | 2 |
2 | Terga and abdomen uniformly dark reddish brown to black | Melanozosteria nitida |
– | Margin of terga with continuous and broad yellow stripes, the middle black; sometimes abdomen with continuous or discontinuous yellow stripes | Melanozosteria soror |
Laevifacies quadrialata sp. nov. here designated.
Body small to medium, thinner in male, thorax slightly broader than abdomen. Surface smooth and shining. Pronotum slightly semicircular, vertex barely exposed. Male with vestigial tegmina and hind wings on mesonotum and metanotum respectively, both nearly triangular; female only with vestigial tegmina, its shape similar to that of male, without hind wings. Legs strong but short, coxae with punctation, front femora Type A2. Mid and hind metatarsus with strong spines, claws symmetrical. Cerci strong, short and symmetrical. Styli long and symmetrical. Supra-anal plate in male short, triangular; subgenital plate broad and short, slightly quadrilateral and symmetrical. L1 divided into two parts, L3 bifurcated, one branch short, the other one long, R1 nearly claw-like and R2 large, hooked.
The name Laevifacies is derived from two Latin words laevis and facies, referring to the smooth and shining surface of terga. The gender of Laevifacies is feminine.
Based on former studies (
A–K Laevifacies quadrialata sp. nov. A–F, I–K male holotype A in dorsal view B in ventral view C pronotum, in dorsal view D head, in ventral view E femur, in ventral view F tibia, in ventral view I fore tarsus, in ventral view J mid tarsus, in ventral view K hind tarsus, in ventral view. G–H female paratype G in dorsal view H in ventral view. Scale bars: 5 mm (A–B, G–H); 1 mm (C–F, I–K).
A–G male genitalia features from holotype A left phallomere, in dorsal view B L1 of left phallomere, in dorsal view C right phallomere, in dorsal view D subgenital plate, in ventral view E supra-anal plate, in dorsal view F–G female genitalia features from paratype F subgenital plate, in ventral view G supra-anal plate and genitalia, in dorsal view. Abbreviations: a.a., anterior arch; acc.pr., accessory process; bsv., basivalvula; c.a., central apodeme; i.p.p., inner posterior process; lat.st.IX–X, laterosternal of the ninth-tenth segment; L.ph., left phallomere; L1–L3, parts of left phallomere; o.p.p., outer posterior process; R.ph., right phallomere; R1–R3, parts of right phallomere; v.I–III, first-third valve; v.ph., ventral phallomere. Scale bars: 1 mm (A, C–G); 0.5 mm (B).
A–E, I–L Melanozosteria nitida from Guangxi, male A in dorsal view B in ventral view C pronotum, in dorsal view D head, in ventral view E tarsus, in ventral view F–H Lectotype of Melanozosteria nitida, male F in dorsal view G in ventral view H labels I subgenital plate, in ventral view J left phallomere, in dorsal view K right phallomere, in dorsal view L supra-anal plate, in dorsal view M genitalia of Melanozosteria nitida in
China (Hainan).
Sexual dimorphism. Body small and black. Surface smooth and shining except last two terga with punctation. Tegmina and hind wings vestigial in male, tegmina vestigial and hind wings absent in female. Angles of T2–T7 sharp and protruded. Legs strong. Supra-anal plate short and triangular. Styli long and symmetrical.
Measurements. Male, pronotum: length × width 5.5–6.2 × 7.9–8.0 mm, overall length: 15.6–17.7 mm. Female, pronotum: length × width 7.0–7.1 × 10.5–10.7 mm, overall length: 17.0–21.0 mm.
Body black, smooth, shining. Vertex and frons black. Clypeus to part of labrum brown to dark brown, maxillary palpi and labial palpi dark brown to black. Eyes black when the specimens are fresh, fading after a long time (Figure
Size small to medium, female larger than male. Body oval, vertex nearly unexposed (Figure
Left phallomere consisting of three parts: L1, L2, and L3. L1 with two parts L1a and L1b, L1a with membranous finger-like projection; L1b with sclerotized projection. L2 consisting of L2d and L2v, L2d strongly sclerotized in anterior part, the posterior part with finger-like and with more small spines; L2va simple and broad, L2vb sclerotized and the posterior with a spinous projection. L3 with a simple hook, elongate to the right and bifurcated (Figure
The first valve (v.I) long, slightly broad and crescent-shaped, terminal membranous; the second valve (v.II) small, flaky and obscured by the v.I; the third valve (v.III) broader than v.I, terminal membranous; paraprocts (pp.) symmetrical and the middle concave; the middle of anterior arch (a.a.) concave; basivalvula (bsv.) trapezoidal (Figure
HOLOTYPE: male, CHINA, Hainan Prov., Baisha, Yinggeling Nature Reserve, 20-VIII-2010, Guo Zheng leg. PARATYPES: 1 male, same data as holotype; 1 male, Hainan Prov., Mt. Wuzhishan, 18-21-V-2014, Shunhua Gui, Xinran Li & Jianyue Qiu leg.; 1 male, Hainan Prov., Diaoluoshan, 18-IV-2015, Lu Qiu & Qikun Bai leg. (GenBank accession number: MK798103); 2 females, Hainan Prov., Lingshui, Mt. Diaoluoshan, 22-V-2014, Jianyue Qiu, Xinran Li & Shunhua Gui leg. (GenBank accession number: MK798104).
The species epithet comes from the Latin word quadrialata in reference to the male having four triangular vestigial wings.
In our study the interspecific K2P genetic divergence among L. quadrialata sp. nov. and other cockroach species ranged from 10.4 to 13.1%. But the genetic divergence value between male and female of L. quadrialata sp. nov. is only 0.9%, so we pair them based on their similar morphology combined with this COI data. Sexual dimorphism occurs in L. quadrialata sp. nov.: 1) females without hind wings, but males with vestigial hind wings (Figure
China (Hainan)
Melanozosteria Stål, 1874: 13;
Cutilia Stål, 1877: 36;
Symtomaptera Tepper, 1893: 106 (as a subgenus of Periplaneta);
Body broad oval and reddish brown to black. Pronotum slightly arched, surface with punctation. Vestigial tegmina sectorial with punctation, separated from mesonotum for nearly whole length, hind wings absent. Surface with punctation. Angles of T2–T7 protruded and sharp. The medial aspects to the styli with stubby and sharp spines.
Male, pronotum: length × width 7.4 × 12.5 mm, overall length: 26.1 mm.
Body uniformly deep reddish brown to black (Figure
Body large, broad oval and convex, surface shining. Pronotum slightly arched, surface with punctation. Anterior margin of pronotum roundly protruded, and posterior margin straight (Figure
Left phallomere includes L1, L2, and L3. L1 with three parts (a, b, c). L1a slightly sclerotized, posterior not sclerotized, membranous and blunt. L1b more sclerotized and posterior sharp. L1c anterior slightly sclerotized and posterior blunt membrane. L2 includes L2d and L2v. L2d with a well-sclerotized, strongly denticulate in anterior margin, while the posterior of the sclerite becomes more delicate and ends in a sharp point; L2v usually single, L3 is a simple hook, but the posterior divides into two small forks which resemble an elephant’s nose (Figure
1 male, CHINA, Guangxi Prov., Shangsi, Nadang, 15-XI-1958, Dexiang Gu & Jinting Liang leg.
Lectotype of Polyzosteria nitida, male, Ternate (Natural History Museum Vienna), “Ternate Jeynalle CoII. Br. V. W.”, “LECTOTYPE”, “LECTOTYPE of Polyzosteria nitida Brunn. Selected by KHL Key, 1963.”; holotype of Periplaneta polita, male, Taiwan (Natural History Museum), “Holotype”, “Periplaneta polita Walker”, “BMNH (E) #878036”, presented by
We compared the lectotype of M. nitida (from Ternate, Indonesia) with the specimen from Guangxi and found there are minor differences between them: the styli are straight in the Guangxi individual (Figure
Australia, Philippines, Malaysia, New Guinea, New Caledonia, New Zealand, China, Thailand.
Almost all members in the Polyzosteriinae are brachypterous or apterous (excepting the tribe Methanini), and display high developmental stochasticity (
We thank Dr Guo Zheng (Shenyang Normal University, China), Dr Shunhua Gui, Dr Xinran Li, Dr Lu Qiu, Dr Jianyue Qiu, and Mr Qikun Bai (all SWU) for collecting the valuable specimens. We also thank Harald Bruckner (Natural History Museum Vienna) for providing us with lectotype photos of M. nitida. And we are grateful to Prof John Richard Schrock (Department of Biological Sciences, Emporia State University) for revising the manuscript before submission. Sonia Lopes (Federal University of Rio de Janeiro, Brazil) and one anonymous reviewer are acknowledged for their valuable comments and suggestions to the manuscript. This study is supported by the National Natural Sciences Foundation of China (No. 31772506), the Program of the Ministry of Science and Technology of the People’s Republic of China (2015FY210300) and the Fundamental Research Funds for the Central Universities (XDJK2017B032).