Research Article |
Corresponding author: Yanli Che ( shirleyche2000@126.com ) Academic editor: Fred Legendre
© 2022 Xinyi Deng, Jieyu Shan, Chengcheng Xiao, Jing Zhu, 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:
Deng X, Shan J, Xiao C, Zhu J, Wang Z, Che Y (2022) Establishment of two new Anaplecta species (Blattodea, Blattoidea, Anaplectidae) based on morphological and COI data with an additional description of Anaplecta furcata Deng & Che, 2020. ZooKeys 1130: 153-166. https://doi.org/10.3897/zookeys.1130.87810
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Based on morphological characteristics, including male and female genitalia, combined with DNA barcodes, two new species, Anaplecta circinalis Deng & Che, sp. nov. and Anaplecta bihamata Deng & Che, sp. nov., are described in detail. Additional information on the female genitalia of Anaplecta furcata Deng & Che, 2020 is also provided. Photographs of external morphology and caudal anatomy of these species, as well as a key to the Chinese Anaplecta species, are provided.
Cockroaches, COI, DNA barcodes, female genitalia, morphology, new species, taxonomy
At present, 112 species of Anaplecta have been recorded, widely distributed in Asia, North America, South America, Africa and Oceania (21 species in China) (
The effective discernment of species using male genitalia was verified in
Recently, after collecting cockroach specimens in Fujian, Yunnan, Guangdong and Hunan provinces in China, two new Anaplecta species were discovered based on morphological characters. This result was then further verified here by using DNA barcodes. In addition, an exhaustive description of the female genitalia of Anaplecta furcata was also provided herein.
Sixty-eight specimens from Fujian, Yunnan, Guangdong and Hunan provinces were examined in this study. The genitalia terminology used in this paper mainly follows
The measurements are based on the specimens examined. The genitalia were processed with 10% NaOH at 65 °C for 30–35 min for digestion of soft tissues. The genitalia segments were dissected and stored in glycerol, then observed with a Motic K400 stereomicroscope. These segments were then preserved with the remainder of the specimen in ethyl alcohol. Photographs of the genitalia and body were taken with a Leica M205A stereomicroscope and edited with Adobe Photoshop CS6. All type materials are deposited at the Institute of Entomology, College of Plant Protection, Southwest University, Chongqing, China (SWU).
Abbreviations in this paper are as follows:
L1, L2, L3 sclerites of the left phallomere;
L2d L2 dorsal;
L2v L2 ventral;
L2vm median sclerite;
M media veins;
R1, R2, R3 sclerites of the right phallomere;
CuA cubitus anterior;
CuP cubitus posterior.
Four specimens were used for cytochrome oxidase subunit I (COI) sequencing in this study. Total DNA was extracted from the muscle of legs according to the HiPure Tissue DNA Mini Kit (Magen Biotech, Guangzhou). Primers for polymerase chain reaction (PCR) were COI-F (5’-CAACYAATCATAAAGANATTGGAAC-3’) and COI-R (5’-TAAACTTCTGGRTGACCAAARAATCA-3’) (a simple adjustment based on
A total of 62 COI sequences were analyzed in this study: four newly-sequenced sequences of our newly described Anaplecta species and 55 published sequences of 20 Chinese Anaplecta species, and three sequences of Periplaneta Burmeister, 1838 (as outgroup) downloaded from GenBank (Table
Species | Location (voucher number, gender) | Literature source | GenBank Accession Number |
---|---|---|---|
A. circinalis sp. nov. | Pu’er, Yunnan (Anapcircm, ♂) | OP306078 | |
A. circinalis sp. nov. | Pu’er, Yunnan (Anapcircf, ♀) | OP306077 | |
A. bihamata sp. nov. | Shaoyang, Hunan (Anapbiham, ♂) | OP306076 | |
A. bihamata sp. nov. | Shaoyang, Hunan (Anapbihaf, ♀) | OP306075 | |
A. bicruris |
|
OL790029, OL790030, OL790036 | |
A. spinosa |
|
OL790028, OL790038 | |
A. ungulata |
|
OL790031, OL790053, OL790048 | |
A. anomala |
|
OL790032, OL790050 | |
A. serrata |
|
OL790033, OL790047, OL790046 | |
A. bombycina |
|
OL790037, OL790049, OL790034, OL790052 | |
A. longihamata |
|
OL790035, OL790051 | |
A. paraomei |
|
OL790039, OL790045, OL790041, OL790042 | |
A. condensa |
|
OL790040, OL790043, OL790044 | |
A. truncatula |
|
OL790054, OL790055 | |
A. omei |
|
OL790056, OL790057, OL790058 | |
A. omei |
|
MT800287 | |
A. corneola |
|
OL790063 | |
A. corneola |
|
MT800293, MT800296 | |
A. cruciata |
|
OL790061 | |
A. cruciata |
|
MT800303, MT800304 | |
A. basalis |
|
OL790060 | |
A. basalis |
|
MT800305, MT800309 | |
A. nigra |
|
MT800306 | |
A. staminiformis |
|
OL790062 | |
A. staminiformis |
|
MT800297, MT800299 | |
A. arcuata |
|
OL790065 | |
A. arcuata |
|
MT800307, MT800308 | |
A. strigata |
|
OL790064 | |
A. strigata |
|
MT800291, MT800292 | |
A. furcata |
|
MT800301,MT800302 | |
A. bicolor |
|
OL790059 | |
A. bicolor |
|
MT800310 | |
Periplaneta americana |
|
KC617846 | |
Periplaneta fuliginosa |
|
KM577133 | |
Periplaneta australasiae |
|
KF640069 |
After observing the external morphological and genital characteristics of 42 Anaplecta samples from Fujian, Yunnan, Guangdong and Hunan provinces, two new morphospecies and one known species, Anaplecta furcata, were identified. One morphospecies can be distinguished by its curled L2vm from other Chinese Anaplecta species; while the other is characterized by its hook-shaped L2vm and R1.
In this study, the sequenced length of COI excluding the primer was approximately 658bp. Four new COI sequences have been deposited in GenBank with accession numbers OP306075 to OP306078 (Table
Interspecific genetic distance calculated by the K2P model using COI sequences in MEGA.
species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1 | A. bihamata sp. nov. | |||||||||||||||||||||
2 | A. staminiformis | 0.1397 | ||||||||||||||||||||
3 | A. bicruris | 0.1686 | 0.1827 | |||||||||||||||||||
4 | A. spinosa | 0.1835 | 0.1764 | 0.1653 | ||||||||||||||||||
5 | A. longihamata | 0.2177 | 0.2460 | 0.2134 | 0.2222 | |||||||||||||||||
6 | A. condensa | 0.2121 | 0.2370 | 0.2160 | 0.2131 | 0.0554 | ||||||||||||||||
7 | A. paraomei | 0.2003 | 0.2220 | 0.2037 | 0.2169 | 0.0832 | 0.0881 | |||||||||||||||
8 | A. omei | 0.1966 | 0.2202 | 0.2192 | 0.2065 | 0.0779 | 0.0937 | 0.0699 | ||||||||||||||
9 | A. corneola | 0.2078 | 0.2115 | 0.1972 | 0.1975 | 0.1425 | 0.1447 | 0.1437 | 0.1513 | |||||||||||||
10 | A. nigra | 0.1899 | 0.1904 | 0.1943 | 0.2150 | 0.2318 | 0.2365 | 0.2146 | 0.2277 | 0.2108 | ||||||||||||
11 | A. basalis | 0.1734 | 0.1875 | 0.1918 | 0.1909 | 0.2417 | 0.2400 | 0.2176 | 0.2202 | 0.2133 | 0.2132 | |||||||||||
12 | A. bicolor | 0.2265 | 0.2215 | 0.1939 | 0.2025 | 0.2211 | 0.2209 | 0.2104 | 0.2207 | 0.2070 | 0.2399 | 0.2246 | ||||||||||
13 | A. anomala | 0.2123 | 0.2119 | 0.2159 | 0.2065 | 0.2106 | 0.2154 | 0.1981 | 0.1974 | 0.2157 | 0.2220 | 0.2342 | 0.1972 | |||||||||
14 | A. strigata | 0.2123 | 0.2236 | 0.2077 | 0.2012 | 0.2032 | 0.2100 | 0.2095 | 0.2035 | 0.2004 | 0.2094 | 0.2104 | 0.2071 | 0.0868 | ||||||||
15 | A. serrata | 0.2027 | 0.2287 | 0.2090 | 0.2198 | 0.2366 | 0.2526 | 0.2213 | 0.2144 | 0.2353 | 0.2141 | 0.2124 | 0.2438 | 0.1986 | 0.1852 | |||||||
16 | A. circinalis sp. nov. | 0.2014 | 0.2175 | 0.2179 | 0.1885 | 0.2427 | 0.2485 | 0.2236 | 0.2196 | 0.2416 | 0.2368 | 0.2063 | 0.2282 | 0.2145 | 0.2019 | 0.2051 | ||||||
17 | A. bombycina | 0.2487 | 0.2423 | 0.2201 | 0.2050 | 0.2269 | 0.2310 | 0.2247 | 0.2198 | 0.2141 | 0.2176 | 0.2530 | 0.2471 | 0.2263 | 0.2216 | 0.2373 | 0.2263 | |||||
18 | A. arcuata | 0.1970 | 0.2339 | 0.2367 | 0.2310 | 0.2258 | 0.2355 | 0.2197 | 0.2291 | 0.2442 | 0.2128 | 0.2364 | 0.2239 | 0.2089 | 0.2100 | 0.2495 | 0.2027 | 0.1925 | ||||
19 | A. furcata | 0.2201 | 0.2357 | 0.2133 | 0.2208 | 0.2199 | 0.2419 | 0.2376 | 0.2178 | 0.2284 | 0.2126 | 0.2275 | 0.2128 | 0.2124 | 0.2084 | 0.2059 | 0.2192 | 0.1887 | 0.1867 | |||
20 | A. truncatula | 0.2025 | 0.2161 | 0.2008 | 0.1961 | 0.2387 | 0.2274 | 0.2187 | 0.2219 | 0.2400 | 0.2395 | 0.2031 | 0.2393 | 0.2405 | 0.2398 | 0.2348 | 0.2099 | 0.2494 | 0.2456 | 0.2334 | ||
21 | A. cruciata | 0.2087 | 0.2167 | 0.2201 | 0.2258 | 0.2352 | 0.2233 | 0.2359 | 0.2107 | 0.2475 | 0.2291 | 0.2292 | 0.2354 | 0.2228 | 0.2370 | 0.2383 | 0.2230 | 0.2615 | 0.2342 | 0.2362 | 0.2546 | |
22 | A. ungulata | 0.2337 | 0.2644 | 0.2720 | 0.2411 | 0.2424 | 0.2503 | 0.2509 | 0.2536 | 0.2349 | 0.2284 | 0.2250 | 0.2569 | 0.2276 | 0.2068 | 0.2628 | 0.2435 | 0.2504 | 0.2374 | 0.2454 | 0.2753 | 0.2571 |
Based on the morphology and ML analysis, we confirmed two new species: Anaplecta circinalis Deng & Che, sp. nov. and Anaplecta bihamata Deng & Che, sp. nov.
[A. simplex Shiraki, 1931 is not included because we were unable to collect a sample and only wings were described by the author.]
1 | Disk of pronotum bicolored | 2 |
– | Disk of pronotum unicolored | 6 |
2 | Disk dark and faded gradually or sharply towards the posterior end | 3 |
– | Disk dark with some markings | 4 |
3 | Tegmina yellowish brown, 1/3 of the base black (except the lateral margins) | A. basalis Bey-Bienko, 1969 |
– | Tegmina completely yellowish brown (except the lateral margins) | A. bicolor Deng & Che, 2020 |
4 | Disk with a pair of blurred longitudinal darker areas | A. bicruris Zhu & Che, 2022 |
– | Disk with a lighter blurred centre | 5 |
5 | Tegmina unicolored | A. strigata Deng & Che, 2020 |
– | Tegmina bicolored, 1/3 of the base darker than remaining parts (except lateral margins and anal field) | A. anomala Zhu & Che, 2022 |
6 | Tegmina with obvious markings | 7 |
– | Tegmina without obvious markings | 9 |
7 | Tegmina yellowish brown, with a nearly oval brown spot at CuP | A. ungulata Zhu & Che, 2022 |
– | Tegmina yellowish brown, with a subrectangular black spot at base | 8 |
8 | R1 needle-shaped | A. truncatula Zhu & Che, 2022 |
– | R1 arc-shaped | A. nigra Deng & Che, 2020 |
9 | Male paraprocts with dense spines on curly posterior margin | 10 |
– | Male paraprocts not as above | 13 |
10 | Intercalary sclerite small, nearly filamentous | A. condensa Zhu & Che, 2022 |
– | Intercalary sclerite large, strip-shaped or sheet-like | 11 |
11 | Right first valvifer arm long, lateral edges folded up | A. longihamata Zhu & Che, 2022 |
– | Right first valvifer arm short, lateral edges not folded up | 12 |
12 | The posterior margin of anterior arch hip-shaped | A. paraomei Zhu & Che, 2022 |
– | The posterior margin of anterior arch smooth | A. omei Bey-Bienko, 1958 |
13 | L1 with a long and curved filamentary structure | 14 |
– | L1 with a short and robust uncinate structure | A. cruciata Deng & Che, 2020 |
14 | R1 degraded or merged with L2vm | 15 |
– | R1 well developed, not merged with L2vm | 18 |
15 | Male paraprocts specialized, stripe-shaped, with spines on posterior margin | A. spinosa Zhu & Che, 2022 |
– | Male paraprocts unspecialized | 16 |
16 | The apex of L2v bifurcated, sheet-like | 17 |
– | The apex of L2v not bifurcated, shaped like ‘3’ | A. bombycina Zhu & Che, 2022 |
17 | One sclerite of R2 serrated | A. serrata Zhu & Che, 2022 |
– | All sclerites of R2 without serration | 18 |
18 | L2vm slender | A. arcuata Deng & Che, 2020 |
– | L2vm broad | A. circinalis Deng & Che, sp. nov. |
19 | R1 curved | 20 |
– | R1 straight, cylindrical | A. staminiformis Deng & Che, 2020 |
20 | R1 highly sclerotized, horn-shaped | 21 |
– | R1 sightly sclerotized, arc-shaped | A. furcata Deng & Che, 2020 |
21 | Right phallomere without special horny structure | A. bihamata Deng & Che, sp. nov. |
– | Right phallomere with special horny structure | A. corneola Deng & Che, 2020 |
Holotype : China • male; Yunnan Prov., Pu’er County, Xiniuping Scenic Area of Pu’er National Park; 26°36.14'N, 101°5.53'E; 1602 m; 29 Jun. 2021; Jia-Wei Zhang & Jin-Lin Liu leg; SWU-B-AN-0175. Paratypes: China • 9 males & 14 females; same data as holotype; SWU-B-AN-0176 to 0198.
Diagnosis. This species can be easily separated from other Chinese Anaplecta species by its curled left phallomere (L2vm).
The specific epithet ‘circinalis’ is derived from the Latin word circinalis, referring to the curled L2vm.
Measurements (mm). Male: Pronotum length × width: 1.0–1.3 × 1.8–1.9, tegmina length: 5.0–5.2, overall length: 5.8–6.1. Female: Pronotum length × width: 1.2–1.4×1.7–2.0, tegmina length: 4.7–5.1, overall length: 5.6–6.2.
Coloration. Body dark yellowish brown, eyes black, antennae dark brown. Head yellowish brown (Fig.
Anaplecta circinalis Deng & Che, sp. nov. holotype, male SWU-B-AN-0175 A habitus, dorsal view B habitus, ventral view C pronotum, dorsal view D head, ventral view E tegmen F supra-anal plate, ventral view G front femur H wing I subgenital plate, dorsal view J left phallomere K right phallomere, dorsal view. Abbreviations: afd anal fold, A[1] anterior anal vein, cfd cubitus fold, CuA cubitus anterior, CuP cubitus posterior, L1, L2, L3 sclerites of the left phallomere, L2d L2 dorsal, L2v L2 ventral, L2vm median sclerite, M median, Pcu postcubitus, R radius, RA radius anterior, RP radius posterior, R2, R3 sclerites of the right phallomere, ScP subcostal posterior. Scale bars: 2 mm (A–E, G, H); 0.5 mm (F, I–K).
Head and thorax. Interocular space slightly greater than distance between antennal sockets. Third and fifth maxillary palpi equal in length, longer than fourth; fifth maxillary palpus subelliptical and thicker than other segments (Fig.
Tegmina with indistinct veins; the radius posterior veins of hind wings slightly indistinct, CuP and CuA merging into one venation (Fig.
Male abdomen and genitalia. Supra-anal plate nearly symmetrical, sheet-like (Fig.
Female abdomen and genitalia. Supra-anal plate nearly symmetrical. Paraprocts (pp.) broad, not extending to the posterior margin of supra-anal plate. Intercalary sclerite (intc.s) nearly stripe-shaped, slightly curved. First valvifer arm short. First valve (v.I) robust. Second valve (v.II) small, basally fused. Third valve (v.III) broad. The anterior margin of anterior arch (aa.) symmetrical, slightly sclerotized, extending forward in a flaky shape with a deep concavity in the middle. Basivalvula (bsv.) nearly flattened, elliptical (Fig.
China (Yunnan).
Holotype : China • male; Hunan Prov., Shaoyang City, Chengbu County, Ten Miles Flat Monitoring Station; 26°14.12'N, 110°25.52'E; 821 m; 22 May 2021; Jing Zhu leg; SWU-B-AN-0199. Paratypes: China • 7 males; SWU-B-AN-0200 to 206; same collection data as holotype • 1 male; SWU-B-AN-0207; Guangdong Prov., Shaoguan City • 2 males and 1 female; SWU-B-AN-0208 to 0210; Ruyuan County, Nanling Nature Reserve Xiaozhu Parking Lot; 24°54.10'N, 113°2.53'E; 695 m, 18 May 2021; Wei Han & Li-Min Qiao leg. • 1 male; SWU-B-AN-0211; Hunan Prov., Yongzhou City, Ningyuan County, Mt. Jiuyi, Yellow River Village; 25°9.8'N, 111°34.17'E; 629 m, 6 Jun. 2021; Jing Zhu leg. • 3 males; SWU-B-AN-0212 to 214; Fujian Prov., Wuyishan City, Sisin Integrated Observation Site; 27°35.30'N, 117°46.4'E; 450 m, 23 Jun. 2021; Wei Han & Jing Zhu leg.
This species can be easily separated from other Chinese Anaplecta species by its hook-shaped L2vm and R1.
The specific epithet is derived from the Latin word hamatus, referring to both L2vm and R1 being hook-like.
Measurements (mm). Male: Pronotum length × width: 1.2–1.4 × 1.9–2.0, tegmina length: 5.8–6.4, overall length: 6.8–7.3. Female: Pronotum length × width: 1.1–1.4 × 1.9–2.2, tegmina length: 5.4–5.6, overall length: 6.2–6.7.
Coloration. Body yellowish brown, eyes black, antennae dark brown. Head yellowish brown (Fig.
Anaplecta bihamata Deng & Che, sp. nov. holotype, male SWU-B-AN-0177 A habitus, dorsal view B habitus, ventral view C pronotum, dorsal view D head, ventral view E tegmen F supra-anal plate, ventral view G front femur H wing I subgenital plate, dorsal view J, K left phallomere L, M right phallomere, dorsal view. Abbreviations: afd anal fold, A[1] anterior anal vein, cfd cubitus fold, CuA cubitus anterior, CuP cubitus posterior, L1, L2, L3 sclerites of the left phallomere, L2v L2 ventral, L2vm median sclerite, M median, Pcu postcubitus, R radius, RA radius anterior, RP radius posterior, R1, R2, R3 sclerites of the right phallomere, ScP subcostal posterior. Scale bars: 2 mm (A–E, G, H); 0.5 mm (F, I–M).
Head and thorax. Interocular space slightly greater than distance between antennal sockets. Fourth and fifth maxillary palpi equal in length, shorter than third maxillary palpus; fifth maxillary palpus triangular and thicker than others (Fig.
Male abdomen and genitalia. Supra-anal plate symmetrical, sheet-like (Fig.
Female abdomen and genitalia. Supra-anal plate nearly symmetrical. Paraprocts (pp.) broad, extending to the posterior margin of supra-anal plate. Intercalary sclerite (intc.s) short, nearly stripe-shaped, slightly curved. First valvifer arm short. First valve (v.I) robust. Second valve (v.II) small, basally fused. Third valve (v.III) broad. The anterior margin of anterior arch (aa.) slightly sclerotized, with a near spine-shaped protrusion and dense tiny punctuations. Basivalvula (bsv.) nearly flat (Fig.
A–C Anaplecta circinalis Deng & Che, sp. nov. paratype, female SWU-B-AN-0176 D–F Anaplecta bihamata Deng & Che, sp. nov. paratype, female SWU-B-AN-0178 G–I Anaplecta furcata female SWU-B-B-A060471 A, D, G supra-anal plate, ventral view B, E, H supra-anal plate, dorsal view C, F, I subgenital plate, dorsal view. Abbreviations: a.a. anterior arch, bsv. basivalvula, cp. crosspiece, pt. paratergites, intc.s. intercalary sclerite, inst.f. Intersternal fold, ltst.sh. laterosternal shelf, pp. paraprocts, sp. spermatheca, v.I first valve, v.II second valve, v.III third valve. Scale bars: 2 mm.
China (Hunan, Fujian, Guangdong).
Anaplecta furcata
Deng & Che in
Measurements (mm). Female: pronotum length × width: 1.1–1.4 × 1.9–2.2, tegmina length: 5.4–5.6, overall length: 6.2–6.7.
Female abdomen and genitalia. Supra-anal plate nearly symmetrical. Paraprocts broad, extending to the posterior margin of supra-anal plate. Intercalary sclerite (intc.s) nearly stripe-shaped. First valve (v.I) robust. Second valve (v.II) small, basally fused. Third valve (v.III) broad. The anterior margin of anterior arch (aa.) slightly sclerotized, extending forward in a flaky shape with a deep concavity in the middle. Basivalvula (bsv.) broad, some areas with dense punctuations. Laterosternal shelf (ltst.sh) slightly sclerotized, lateral margin slightly curved.
China • 2 females; SWU-B-AN-0215 to 216; Guangxi Prov., Jinxiu County, Mt. Dayao; 24°8.43'N, 110°11.70'E; 944 m; 7 Jul. 2015; Lu Qiu & Qi-Kun Bai leg.
China (Guangxi).
We sincerely thank all collectors for their efforts in collection. We thank Prof. John Richard Schrock (Department of Biological Sciences, Emporia State University) for proof-reading the final draft. This study is supported by the National Natural Science Foundation of China (No. 32170458, 31772506, 32070468), and GDAS Special Project of Science and Technology Development (No. 2020GDASYL-20200102021, 2020GDASYL-20200301003).