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Corresponding author: Yanli Che ( shirleyche2000@126.com ) Academic editor: Steffen Pauls
© 2022 Duting Jin, Qiongyao Zhao, Wei Han, Jinxiang Li, Zongqing Wang, Yanli Che.
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Members of the morphologically and ecologically diverse Blattellidae provide a dilemma: their systematic assignment, whether morphologically similar or different, is uncertain. We pay special attention to several taxa: the morphologically similar Episymploce Bey-Bienko, 1950 and Symploce Hebard, 1916, which were strongly disputed because of their extremely similar generic diagnosis in the past century, and one brachypterous species of Blattella Caudell, 1903, which can be easily distinguished from other macropterous members, but is at risk of being misassigned to other genera on the basis of morphological characters. We address the phylogeny of Blattellidae using DNA sequences (mitochondrial 12S rRNA, 16S rRNA, COII, nuclear 28S rRNA, histone H3) from a broad sample of taxa. A new genus (Centrocolumna gen. nov.) and four new species (Centrocolumna ericea sp. nov., Symploce nigra sp. nov., Symploce tubercularis sp. nov., Blattella foliolata sp. nov.) are established on the basis of morphological characters combined with the molecular data. The phylogenetic results indicate the relationships ((Episymploce + Blattella) + Centrocolumna gen. nov.) + Symploce. Furthermore, we delimited the taxonomic status of 12 new combinations, which involve Centrocolumna gen. nov., Episymploce, Symploce, and Blattella.
Bayesian Inference, interspecific competition, key, Maximum Likelihood, mimicry, new species
Blattellidae is an extremely species-diverse family in Blaberoidea, which contains approximately 1070 species from 79 genera (
Because of the notorious German cockroach, Blattella germanica, people have a relatively good understanding of the genus Blattella, which is characterized by its generally small size, the tongue-shaped tergite 10 (supra-anal plate) and the tergal glands on abdominal tergites VII and VIII. Developed tegmina and hindwings are very common in Blattella males with the exception of Blattella kevani Roth, 1985, whose tegmina and hindwings are missing or possibly reduced (
During the last decade, a number of well-accepted molecular phylogenetic studies obtained excellent results in cockroach systematics, including
Here we used targeted taxon sampling (mainly from Blattellidae) and generated molecular data (mitochondrial 12S rRNA (12S), mitochondrial 16S rRNA (16S), cytochrome c oxidase subunit II (COII), nuclear 28S rRNA (28S) and histone H3 (H3)), in order to infer the relationships of the species of the morphologically similar genera Episymploce and Symploce, and the morphologically different brachypterous Blattellidae species, and clarify the taxonomic status of these problematic groups. Finally, we present one new genus Centrocolumna gen. nov., four new species of Symploce, Blattella and Centrocolumna gen. nov., and 12 new combinations based on morphological characters and molecular data.
Samples of Blattellidae were collected from southeast and southwest localities in China, mainly in Yunnan, Hainan, and Guangdong (supplementary material Table S1). Specimens were preserved in analytical pure ethanol and stored at –80°C until processing. All voucher specimens (more information in Table S1) are kept at the Institute of Entomology, College of Plant Protection, Southwest University, Chongqing, China.
Morphological terminology used mainly follows
Measurements are based on specimens examined by vernier caliper as follows: the body length: from the anterior to the posterior; the body length including tegmina: from the anterior to the tip of tegmina; and pronotum length × width: the length at the longest and the width at the widest part. Genital segments of examined specimens were placed in centrifuge tubes and soaked with 10% NaOH, then the centrifuge tube was immersed in hot water for 15–20 minutes and rinsed with distilled water, and then observed in glycerin under a SZ780 Continuous Zoom Stereo Microscope. Specimen and genital photographs were taken by a Leica M205A stereomicroscope with a Leica DFC Camera. All photos provided here were edited and assembled into plates in Adobe Photoshop CS6. Male postabdominal structures are shown at greater detail in supplementary material Fig. S2 than in the figures included in the main article. Type material of new taxa is deposited in the Institute of Entomology, Southwest University, Chongqing, China.
We sequenced five genes in 48 samples across 26 species (Table S1) of Blattellidae: three mitochondrial genes: 12S, 16S, COII, and two nuclear genes: 28S, H3. Total genomic DNA was extracted with the HiPure Tissue DNA Mini Kit (Magen, Guangzhou) from the hindleg tissue of the specimens according to the manufacturer’s protocol. The legs were retained afterwards and stored in 100% ethanol.
All gene fragments were amplified by PCR. The primers used for amplification are given in supplementary material Table S2. For PCR amplification, the 25 μL reaction system included 22 μL of 1.1 × T3 Super PCR Mix (Tsingke Biological Technology, Beijing), 2 μL of each primer and 1 μL of DNA template. Amplification conditions were as follows: initial denaturation at 98°C for 2 min, followed by 35 cycles of 10 s at 98°C, 10 s at annealing temperature (depending on the amplified gene fragment; see Table S2 for more details), and 10 s at 72°C, and with a final extension of 2 min at 72°C.
PCR products were checked by agarose gel electrophoresis and then sent to Tsingke Biological Technology for purification and sequencing. We used an unrestricted BLAST search to check for contamination in all new sequences. All sequences were deposited in GenBank (accession numbers given in Table S1).
In our study, the lengths of five genes excluding the primer were 422 nt (12S), 464 nt (16S), 681 nt (COII), 726 nt (28S), and 330 nt (H3). Sequences were aligned by MAFFT online version 7 (https://mafft.cbrc.jp/alignment/server). For the alignment of the ribosomal sequences (12S, 16S and 28S), an adjusted direction according to the first sequence was selected to check if there was the reverse sequence. The Q-INS-i algorithm was selected for the ribosomal sequences, the G-INS-i algorithm for protein-coding genes COII and H3, while other parameters were set to default values. Alignments of 12S, 16S and 28S were checked visually in MEGA 7 (
We assembled a data set with 93 samples from 42 Blattellidae species combined with some samples sequenced in previous studies, and 17 outgroup species (ten from other families of Blaberoidea Blattodea except Blaberidae, three from Mantodea, and one each from Phasmatodea, Mantophasmatodea, Grylloblattodea and Orthoptera) downloaded from GenBank (https://www.ncbi.nlm.nih.gov) to infer the position of the root in our phylogenetic analysis (supplementary material Table S1). Phylogenetic analysis was performed based on the combined data set, using maximum likelihood (ML) and Bayesian inference (BI). We performed ML using RAxML version 7.7.1 (
Our ML and BI phylogenetic analyses yielded almost identical topologies with generally high support values (Fig.
In Blattellidae, the topology derived from both analyses showed that Symploce ligulata (Bey-Bienko, 1957), the brachypterous Blattellidae species, and Blattella samples formed a clade with high support values (ML bootstrap support (bs) 100%, posterior probability (pp) 1.0). Then this clade was placed as sister to the monophyletic Episymploce (bs 100%, pp 1.0), followed by the clade Symploce evidens Wang and Che, 2013 + one unknown species (bs 100%, pp 1.0). Other species of the genus Symploce were grouped together to form a clade in the ML analysis, which was recovered to be the sister of clade comprised of the three groupings listed above (bs 100%) (Fig.
Maximum likelihood (ML) tree based on three mitochondrial genes 12S rRNA, 16S rRNA, COII and two nuclear genes 28S rRNA, histone H3. The topologies yielded by Bayesian inference (BI) are almost identical with ML tree (see supplementary material Fig. S1). Branch labels provide bootstrap support (bs) of nodes in ML and posterior probability (pp) (%) of nodes in BI; (*) indicates the branch label of the corresponding analysis is 100 or 1.0, respectively; (-) means the node is absent for the given analysis.
Our phylogenetic inference and subsequent critical examination of morphology led us to propose one new genus, four new species and 12 new combinations as follows.
Both the ML analyses and BI highly support that the clade Symploce evidens Wang and Che, 2013 plus one unknown species is the sister to the clade of Blattella + Episymploce (bs 100%, pp 1.0). Hind margin of subgenital plate with an obvious protrusion and with the right stylus located on it make them obviously distinguishable from other related genera. Therefore, we establish for them and some other taxonomic units the new genus Centrocolumna Jin and Che gen. nov. and also propose one new species and eight new combinations.
Symploce evidens Wang and Che, 2013, by present designation.
Male. Tegmina and hindwings fully developed or slightly reduced. Hindwings with RP branched near the middle, M simple and unbranched, CuA curved with 1–4 complete and 0–5 incomplete branches; apical triangle small or absent. Anteroventral margin of front femur usually Type A. Tergal specialization and medial setal tuft always present on T1, T7 and T10. Subgenital plate asymmetrical, hind margin with a prominent protrusion in the middle, and the right stylus arising upon the protrusion. Bases of two styli obtuse, apices tapering.
Centrocolumna, the combination of the Latin words centr- and columna, refers to the right stylus arising upon the protrusion. The gender of Centrocolumna is feminine.
This genus is similar to Symploce Hebard, 1916, but it can be distinguished from Symploce by the following characters: 1) T10 usually specialized with setal tuft at disk (absent in Symploce); 2) hind margin of subgenital plate with a prominent protrusion (absent in Symploce).
According to the generic diagnosis of Centrocolumna gen. nov., we transferred the following species of Symploce to this genus after our critical examination of the original literature, especially the grouping of species of Symploce collected from Taiwan and Japan in
(1) Centrocolumna evidens (Wang and Che, 2013) comb. nov.
Symploce evidens Wang and Che, 2013: 13.
(2a) Centrocolumna striata striata (Shiraki, 1906) comb. nov.
Periplanela striata Shiraki, 1906: 27.
Ischnoptera striata: Shiraki, 1931: 194.
Symploce striata striata: Asahina, 1979b: 223; Roth, 1987: 158.
(2b) Centrocolumna striata wulaii (Asahina, 1979b) comb. nov.
Symploce striata wulaii Asahina, 1979b: 225; Roth, 1987: 159.
(3) Centrocolumna yayeyamana (Asahina, 1979b) comb. nov.
Symploce yayeyamana Asahina, 1979b: 225–226; Roth, 1987: 159.
(4a) Centrocolumna gigas gigas (Asahina, 1979b) comb. nov.
Symploce gigas gigas Asahina, 1979b: 226; Roth, 1987: 160.
(4b) Centrocolumna gigas okinawana (Asahina, 1979b) comb. nov.
Symploce gigas okinawana Asahina, 1979b: 228; Roth, 1987: 160.
(5) Centrocolumna japonica (Shelford, 1907) comb. nov.
Phyllodromia (?) japonica Shelford, 1907: 33–34.
Symploce japonica: Asahina, 1979b: 229; Roth, 1987: 157.
(6) Centrocolumna okinoerabuensis (Asahina, 1974) comb. nov.
Symploce okinoerabuensis Asahina, 1974: 151, 1979b: 230; Roth, 1987: 158.
(7) Centrocolumna furcata (Shiraki, 1931) comb. nov.
Ischnoptera furcata Shiraki, 1931: 190–191.
Symploce furcata: Asahina, 1979b: 230; Roth, 1987: 160.
(8) Centrocolumna acuminata (Shiraki, 1931) comb. nov.
Ischnoptera acuminata Shiraki, 1931: 195.
Symploce acuminata: Asahina, 1979b: 231; Roth, 1987: 157.
(9) Centrocolumna ericea Jin and Che sp. nov.
1 | Tegmina and hindwings reduced, length about 0.5 times body length | C. acuminata comb. nov. |
1’ | Tegmina and hindwings fully developed, length about 1.2 times body length | 2 |
2 | T1 without a medial setal tuft | 3 |
2’ | T1 with a medial setal tuft | 4 |
3 | T10 without a setal tuft | C. striata striata comb. nov. |
3’ | T10 with an obvious setal tuft | C. striata wulai comb. nov. |
4 | Styli elongate, slender and straight, apically acute | C. yayeyamana comb. nov. |
4’ | Styli relatively short and slightly curved | 5 |
5 | CuA branches of hindwings all reaching the wing margin | 6 |
5’ | CuA branches of hindwings partly not reaching the wing margin | 7 |
6 | T10 with a setal tuft in the middle | C. japonica comb. nov. |
6’ | T10 apparently without setal tuft in the middle | C. furcata comb. nov. |
7 | Right paraproct with a cockscomb-like process | 8 |
7’ | Right paraproct without such a process | 9 |
8 | T7 with a setal tuft | Centrocolumna ericea sp. nov . |
8’ | T7 without a setal tuft | C. evidens comb. nov. |
9 | Left phallomere with one hook-like structure and an additional irregular sclerite | C. okinoerabuensis comb. nov. |
9’ | Left phallomere with only one hook-like structure | 10 |
10 | CuA of hindwings with 3 complete and 4–5 small incomplete branches | C. gigas gigas comb. nov. |
10’ | CuA of hindwings with 2 complete and 1 long incomplete branches | C. gigas okinawana comb. nov. |
Holotype: CHINA • ♂; Chongqing, Qianjiang County, Zhuoshui Town; 19-26 June 2013; Shunhua Gui, Yan Shi leg. Paratypes: CHINA • 1 ♂, Fujian Prov., Mt. Wuyi, Tongmu Village; 8 July 2013; Shunhua Gui, Yan Shi leg. • 1 ♂, 1 ♀; Guangxi Prov., Jinxiu County, Mt. Shengtang; 4–5 June 2014; Shunhua Gui, Xinran Li leg. • 2 ♂♂; Hunan Prov., Changde City, Taoyuan County, Wuyunjie, Tujindong; 19 June 2019; Rong Chen, Hao Xu leg.
Overall length: ♂ 20.6, ♀ 19.6; pronotum length × width: ♂ 4.3 × 4.9, ♀ 4.2 × 5.1; tegmina: ♂ 17.5, ♀ 16.4.
Male. Coloration: Body yellowish brown or brown (Fig.
Centrocolumna ericea gen. et sp. nov., male: A holotype, dorsal view; B holotype, ventral view; C head, ventral view; D pronotum; E maxillary palp; F front femur, ventral view; G seventh abdominal tergite; H tegmen; I wing; J supra-anal plate, ventral view; K left phallomere; L median phallomere; M right phallomere; N subgenital plate, dorsal view. Scale bars: 10 mm (A, B), 1 mm (C, D), 2 mm (H, I), 0.5 mm (E–G, J–N).
The specific epithet is derived from the Latin word ericeus, referring to the right paraproct scattered with spines.
Centrocolumna ericea Jin and Che sp. nov. resembles Centrocolumna striata striata comb. nov., but differs from it as follows: 1) the former with a setal tuft on T1, but the latter without; 2) the former with the left stylus larger than right one, but the latter with the right stylus larger than left one. Furthermore, the general color differs greatly among specimens of the new species collected in different provinces: specimens from Chongqing and Fujian are generally yellowish brown, specimens from Hunan are brown, and specimens from Guangxi are dark brown. In terms of paraprocts, there are some minor differences exhibited in the cockscomb-like process of right paraproct, the teeth usually ranging from 2 to 8.
China (Chongqing, Fujian, Guizhou, Jiangxi, Guangxi, Guangdong, Yunnan, Hunan).
In this genus, with the type species S. hebardi Princis, 1969 (not included in our analyses), we establish two new species mainly according to the morphological characters. In addition, inferred from our phylogenetic analyses (Fig.
Episymploce maxima Li, Liu, Qiu et Yue, 2020: 40.
CHINA • 1 ♂, 1 ♀; Guangxi Prov, Guiping City, Longtan National Forest Park; 386 m; 2 June 2014; Shunhua Gui, Xinran Li leg. • 1 ♂, Guizhou Prov., Qiannan Dushan County, Mawei Town; 9 June 2019; Lu Qiu, Wenbo Deng leg. • 1 ♂, Guizhou, Prov., Leishan County, Changbei Village; 8 June 2013; Guiqiang Huang, Xiangxiang Zhang leg.
Symploce maxima (
China (Guangxi, Guizhou).
Holotype: CHINA • ♂; Hainan Prov., Ledong County, Mt. Jianfengling; 26–28 June 2014; Bing Liu leg.; light traps. Paratypes: CHINA • 2 ♂♂, same data as for holotype.
Overall length: ♂ 21.5; pronotum length × width: ♂ 4.6 × 6.0; tegmina: ♂ 18.5.
Male. Coloration: Body chestnut brown (Fig.
Symploce nigra sp. nov., male: A holotype, dorsal view; B holotype, ventral view; C head, ventral view; D pronotum; E maxillary palp; F front femur, ventral view; G seventh abdominal tergite; H tegmen; I wing; J supra-anal plate, ventral view; K left phallomere; L median phallomere; M right phallomere; N subgenital plate, dorsal view. Scale bars: 10 mm (A, B), 1 mm (C, D), 2 mm (H, I), 0.5 mm (E–G, J–N).
The Latin word niger, meaning black, refers to the posterior and lateral border areas of pronotum being black.
Symploce nigra Jin and Che sp. nov. is similar to Centrocolumna evidens comb. nov. in general morphology, however, they can be distinguished by the following characteristics: 1) pronotum of the former chestnut brown, posterior and narrow lateral borders shiny black, but lateral region of the pronotum of the latter species yellowish brown with dark U-shaped maculae at disc; 2) supra-anal plate of the former specialized with setal tuft near the posterior margin, but the latter without; 3) two styli similar and cylindrical, both situated at the posterior margin of subgenital plate, while the right stylus of the latter, arising at the protrusion of subgenital plate.
China (Hainan).
Holotype: CHINA • ♂, Guangxi Prov., Jinxiu County, Mt. Shengtang; 4–5 June 2014; Shunhua Gui, Xinran Li leg. Paratypes: CHINA • 2 ♂♂, Guangxi Prov., Wuming County, Mt. Daming; 3 July 2015, Lu Qiu and Qikun Bai leg. • 1 ♀, Guangxi Prov., Jinxiu County, Yinshan Park; 16–17 July 2015; Lu Qiu, Qikun Bai leg.
Overall length: ♂ 25.0, ♀ 23.0; pronotum length × width: ♂ 5.4 × 6.2, ♀ 5.5 × 6.4; tegmina: ♂ 22.5, ♀ 18.7.
Male. Coloration: Large body, yellowish-brown to dark brown (Fig.
Symploce tubercularis sp. nov., male: A holotype, dorsal view; B holotype, ventral view; C head, ventral view; D pronotum; E maxillary palp; F front femur, ventral view; G seventh abdominal tergite; H tegmen; I wing; J supra-anal plate, ventral view; K left phallomere; L median phallomere; M right phallomere; N subgenital plate, dorsal view. Scale bars: 10 mm (A, B), 1 mm (C, D), 2 mm (H, I), 0.5 mm (E–G, J–N).
The Latin word tubercularis indicates T7 with a tubercular process in the middle.
Symploce tubercularis Jin and Che sp. nov. is closely related to Symploce maxima (
China (Guangxi).
In our phylogenetic analysis (the type species Episymploce paradoxura Bey-Bienko, 1950 not included), Symploce torchaceus Feng and Woo, 1999 and Symploce wulingensis Feng and Woo, 1993 were recovered as sister taxa and deeply embedded within the clade of Episymploce. 2 new combinations were set up accordingly, and 1 synonymy was proposed as follows.
Symploce torchaceus Feng and Woo, 1999: 51 (Type locality: Fujian, China); Wang and Che, 2013: 4.
CHINA • 1 ♂ (holotype of S. torchaceus); Fujian Prov., Mt. Wuyi; 10 July 1982; Feng Xia leg. • 1 ♂ (paratype of S. torchaceus); Hainan Prov., Mt. Jianfengling; 21 March 1983; Shaoying Liang leg. • 2 ♂♂; Hainan Prov., Linshui County, Mt. Diaoluo; 3 May 2013; Shunhua Gui leg.
Symploce torchaceus Feng and Woo, 1999 was established based on the samples form Mt. Wuyi and Mt. Jianfengling, which was redescribed by
Symploce wulingensis Feng and Woo, 1993: 40 (Type locality: Sichuan, China).
Symploce stellatus
Feng and Woo, 1999: 52 (Type locality: Fujian, China).
CHINA • 1 ♂ (holotype of S. stellatus); Fujian Prov., Mt. Wuyi; 10–18 June 1980; Shiyang Xia leg. • 1 ♂ (paratype of S. stellatus); Fujian Prov., Sangang; 15 June 1981; Jingying Liu leg. • 1 ♂, 1 ♀; Hainan Prov., Lingshi, Mt. Diaoluo; 3 May 2013; Yan Shi leg. • 1 ♂; Guangxi Prov., Jinxiu County, Mt. Shengtang; 900–1200 m; 28 June 2000; Jian Yao leg.
Overall length: ♂ 17.7–18.0; pronotum length × width: ♂ 3.8 × 4.2; tegmina: ♂ 15.0–15.2.
Male. Coloration: Medium body, yellowish brown (Fig.
Episymploce wulingensis (Feng et Woo, 1993) comb. nov., male: A dorsal view; B ventral view; C head, ventral view; D pronotum; E maxillary palp; F front femur, ventral view; G seventh abdominal tergite; H tegmen; I wing; J supra-anal plate, ventral view; K left phallomere; L median phallomere; M right phallomere; N subgenital plate, dorsal view. Scale bars: 10 mm (A, B), 1 mm (C, D), 2 mm (H, I), 0.5 mm (E–G, J–N).
China (Fujian, Guangdong, Hainan, Guangxi, Sichuan, Guizhou, Chongqing).
Symploce kunmingi Bey-Bienko, 1969: 855 (Type locality: Yunnan, China).
Episymploce kunmingi: Roth, 1985b, 215.
Symploce quadrispinis Woo and Feng, 1992: 54 (Type locality: Yunnan, China) syn. nov.
CHINA • 1 ♂ (holotype of S. quadrispinis); Yunnan Prov., Lushui City, Liuku Town, 12 June 1981, Jianmin Zhao leg. • 1 ♂, 1 ♀; Yunnan Prov., Kunming City; 13 June 1986; Pingzhang Feng leg. • 1 ♂, 1 ♀; Sichuan Prov., Leshan City, Jinhekou County; 26–27 May 2011, Keliang Wu leg.
Episymploce kunmingi was published by Bey-Bienko in Symploce and was transferred to Episymploce by
In our molecular analyses, Symploce ligulata (Bey-Bienko, 1957) and the brachypterous Blattellidae species are clustered together with Blattella samples to form a monophyly with high support values. Therefore, we proposed one new combination and assign the brachypterous species into the genus Blattella.
Episymploce ligulata Bey-Bienko, 1957: 911 (Type locality: Yunnan, China); Princis, 1969: 877; Roth, 1997: 107.
Symploce ligulata: Roth, 1985c: 391.
CHINA • 1 ♂, 3 ♀; Yunnan Prov., Xishuangbanna, Dadugang; 3 May 2013; Zongqing Wang leg. • 2 ♂♂; Yunnan Prov., Xishuangbanna, Naban River Reserve; 4 July 2020; Yishu Wang leg. • 3 ♂♂; Yunnan Prov., Xishuangbanna, Mengban Town; 8 July 2020; Duting Jin leg.
Blattella ligulata (Bey-Bienko, 1957) comb. nov. (Fig.
Blattella ligulata (Bey-Bienko, 1957) comb. nov., male: A dorsal view; B ventral view; C head, ventral view; D pronotum; E maxillary palp; F front femur, ventral view; G seventh abdominal tergite; H tegmen; I wing; J supra-anal plate, ventral view; K left phallomere L median phallomere; M right phallomere; N subgenital plate, dorsal view. Scale bars: 10 mm (A, B), 1 mm (C, D), 2 mm (H, I), 0.5 mm (E–G, J–N).
China (Yunnan).
Holotype. CHINA • ♂; Sichuan Prov., Panzhihua City, Wuben Town; 8 June 2014; Xinran Li leg. Paratypes: CHINA • 2 ♂, 2 ♀, same data as for holotype.
Overall length: ♂ 7.7–7.8, ♀ 7.7–10.2; pronotum length × width: ♂ 1.9–2.1 × 2.9–3.2, ♀ 2.0–2.5 × 3.4–3.7; tegmina: ♂ 1.6–1.7, ♀ 1.7–1.9.
Male. Coloration: Body blackish brown with yellowish brown lateral border (Fig.
Blattella foliolata sp. nov., A, B male, adult: A holotype, dorsal view; B holotype, ventral view; C, D female, adult: C paratype, dorsal view; D paratype, ventral view; E, F male, nymph: E dorsal view; F ventral view; G, H female, nymph: G dorsal view; H ventral view; I maxillary palp; J front femur, ventral view; K seventh abdominal tergite; L pronotum; M head, ventral view; N tegmen; O wing; P supra-anal plate, ventral view; Q left phallomere; R median phallomere; S right phallomere; T subgenital plate, dorsal view. Scale bars: 5 mm (A–H), 1 mm (K–O), 0.5 mm (I, J, P–T).
Blattella foliolata Jin and Che sp. nov. is similar to Ignabolivaria bilobata Chopard, 1929 because of nearly identical scale-shaped tegmina and wings. However, they can be distinguished by the following: 1) the former with black markings on the face, but the latter entirely dark brown; 2) abdominal T7 longer than the previous tergites in the former, but shorter than most of previous tergites in the latter; 3) styli close to each other in the former, but separated by a distance of about the length of one stylus in the latter. Meanwhile, this species is easily recognized and distinguished from other macropterous Blattella species owing to the strongly reduced wings, but it is very challenging to distinguish them from the nymphs of Blattella bisignata (Brunner von Wattenwyl, 1893) in the wild. Like most species of Blattella, the brachypterous Blattella foliolata Jin and Che sp. nov. has a small body, pronotum blackish brown with a pair of large black stripes, a symmetrical T10, and an asymmetrical subgenital plate with a concave excavation on the left posterior margin. Moreover, Blattella foliolata Jin and Che sp. nov. was deeply embedded within the clade of Blattella in our phylogenetic analysis, and is retrieved as the sister group of B. bisignata + B. nipponica + B. germanica + B. parilis comb. nov.
The Latin word foliolatus refers to the tegmina and hindwings strongly reduced as a lobule.
China (Sichuan).
For the similar genera Episymploce, Symploce and Centrocolumna Jin and Che gen. nov., a morphology-based placement and delimitation is currently difficult because of the absence of obvious discontinuities in diagnostic features and the large structural diversity in Blattodea and in Blaberoidea, with considerable homoplasy (see e.g.
Episymploce and Symploce are morphologically similar, but they were proposed to be distinguishable by a few diagnostic features: the specialized shape of T7 and T9, the symmetry or asymmetry of T10, and the posterior margin of the subgenital plate being incrassate and setaceous or not (Roth, 1984; 1985a; 1985b; 1997). Previous research about Episymploce and Symploce based only on a non-phylogenetic evaluation of morphological characters revealed that the taxonomic status of some species was controversial, such as
In our study, the new genus Centrocolumna Jin and Che gen. nov. is established, with Symploce evidens Wang and Che, 2013 as type species. At present, there are nine species included in Centrocolumna Jin and Che gen. nov., of which eight are transferred from Symploce except for the new species, Centrocolumna ericea sp. nov. Herein, all of them consistently share generic diagnostic features: their T1, T7, T10 always specialized with medial setal tufts, subgenital plate asymmetrical with a prominent protrusion in the middle of hind margin, and the right stylus arising on it; and their grouping is also supported by our ML and BI analyses with high support values (bs 100%, pp 1.0). A tuft on tergite X (T10) was present at the base of clade Centrocolumna Jin and Che gen. nov., but absent at the base of clade Symploce + (Centrocolumna gen. nov. + (Episymploce + Blattella)), which means the unspecialized T10 was their common ancestral state. Symploce was once thought to be an unnatural unit containing several natural groups (
Another point worth noting is that the different geographical populations of the new species Centrocolumna ericea sp. nov. exhibit some differences in coloration: they are overall dark brown from Guangxi, light-yellow brown from Chongqing and Fujian, while those from Hunan range between dark brown and light-yellow brown. After checking the male genitalia of different geographical samples combined with the molecular clustering, we confirmed these color differences as intraspecific variation.
As in the case of the jumping cockroach, the morphology-based placement of brachypterous species with a high divergence in external morphology can also be difficult.
The small brachypterous species in our study has a blackish brown pronotum with a pair of large black stripes, a symmetrical supra-anal plate, the subgenital plate distinctly asymmetrical with a concave excavation on the left posterior margin, and the ootheca rotates prior to desposition, like most species of Blattella. But all known species of Blattella usually have fully developed tegmina and hindwings, or the hind wings are micropterous: female of Blatta lamotteana (Princis, 1963), Blatta rossi Roth, 1985, male and female of Blatta portalensis (Giglio-Tos, 1907), except for Blatta kevani Roth, 1985, whose tegmina are missing (possibly reduced) and hindwings reduced (
In Blattodea, the reduction of wings usually manifests itself in forewings being larger than the hindwings, or in micropterous forewings and completely absent or extremely reduced hindwings (
Samples of Blattella foliolata sp. nov. (Fig.
We would like to express gratitude to all specimen collectors, as well as Keliang Wu and Lu Qiu who provided photos of habitat and information about field observations. We also thank John Richard Schrock for the revision of manuscript. This study is supported by the National Natural Science Foundation of China (Nos. 31772506, 32070468).
Table S1
Data type: .xlsx
Explanation note: The species and details used in the study.
Table S2
Data type: .docx
Explanation note: Primers used in the study.
Figure S1
Data type: .pdf
Explanation note: Bayesian inference (BI) tree.
Figure S2
Data type: .pdf
Explanation note: Paraprocts, subgenital plate, and male genitalia shown in greater detail.