Research Article |
Corresponding author: Zong-Qing Wang ( zqwang2006@126.com ) Academic editor: Fred Legendre
© 2024 Bian-Lun Li, Peng-Hui Hu, Lin Guo, Yan-Li Che, Zong-Qing Wang.
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:
Li B-L, Hu P-H, Guo L, Che Y-L, Wang Z-Q (2024) Discovery of five new species of Allacta from Yunnan and Hainan, China (Blattodea, Pseudophyllodromiidae). ZooKeys 1191: 1-21. https://doi.org/10.3897/zookeys.1191.113043
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We examined new Allacta materials from Yunnan and Hainan Province, China, and discovered new species using both morphological and molecular species delimitation (ABGD) methods. Five new species are described: A. bifolium Li & Wang, sp. nov., A. hemiptera Li & Wang, sp. nov., A. lunulara Li & Wang, sp. nov., A. redacta Li & Wang, sp. nov., and A. unicaudata Li & Wang, sp. nov. All five species are placed under the hamifera species group. An updated key and checklist of Allacta species from China are provided.
ABGD, checklist, cockroach, cryptic species, DNA barcodes, female genitalia, identification key
The genus Allacta Saussure & Zehntner, 1895 belongs to Pseudophyllodromiidae, with 47 known species mainly distributed in the Oriental and Australasian regions (
Eight species were recorded from China according to
DNA barcoding has been widely used in cockroach identification in recent years, but is considered more reliable when used in combination with morphological evidence (
In this study, newly collected Allacta materials from Yunnan and Hainan Provinces were examined, and they were found morphologically different from the known species. Based on morphological characters as well as the ABGD approach, five new species were confirmed, including the establishment of Allacta hemiptera Li & Wang, sp. nov. for the nomen nudum Temnopteryx hainanensis
Twenty-six studied specimens of Allacta were collected from Yunnan, Hainan and Xizang Provinces and were deposited in College of Plant Protection, Southwest University, Chongqing, China (SWU) and Shanghai Entomology Museum, Shanghai, China (SEM). Morphological terminology used in this paper follows
CuA cubitus anterior;
CuP cubitus posterior;
M media;
R radius;
RA radius anterior;
RP radius posterior;
ScP subcosta posterio;
V vannal;
Pcu postcubitus.
All materials are preserved in absolute ethanol and stored at -20 °C. The genitalia were handled based on a standard procedure in which terminal segments of the abdomen were cut off, heated in tap-water and rinsed with sterile water to remove trace amounts of NaOH, and then immersed in glycerol for further photography, dissection and preservation. Photos were taken with Leica DFC camera through a Leica M205A stereomicroscope; dissection and observation were performed under a Motic K400 stereomicroscope. All photos and images were edited with Adobe Photoshop CC 2019.
Total DNA was extracted from hindleg tissues by the Hipure Tissue DNA Mini Kit (Magen Biotech, Guangzhou), and the remaining body parts were stored in absolute ethanol as voucher specimens. Primers for PCR amplification were LCO1490 (5’-GGTCAACAAATCATAAGATATTGG-3’) and HCO2198 (5’-TAAACTTCAGGGTGACCAAAAAATCA-3’) (
A total of 30 COI sequences were analyzed, including 18 newly obtained sequences from this study; eight sequences representing eight Allacta species downloaded from GenBank; and four sequences represent the outgroup from four genera (Margattea Shelford, 1911, Sorineuchora Caudell, 1927, Balta Tepper, 1893 and Shelfordina Hebard, 1929) of Pseudophyllodromiidae (Table
Species | Voucher ID | GenBank accession number | Collection information |
---|---|---|---|
ingroup | |||
A. bimaculata | OQ736904 | Menglun, Yunnan, China | |
5002287 | PP133869 | Menglun, Yunnan, China | |
5002288 | PP133870 | Menglun, Yunnan, China | |
5002329, F | PP133873 | Menglun, Yunnan, China | |
5002286 | PP133874 | Menglun, Yunnan, China | |
A. transversa | OQ736996 | Wuzhishan, Hainan, China | |
5002314, F | PP133872 | Jianfengling, Hainan, China | |
A. bruna | OQ736905 | Puer, Yunnan, China | |
5002343, F | PP133875 | Jianfengling, Hainan, China | |
5002342, F | PP133876 | Jianfengling, Hainan, China | |
A. xizangensis | OQ736995 | Linzhi, Xizang, China | |
5002302, F | PP133871 | Linzhi, Xizang, China | |
A. robusta | OQ736903 | Limushan, Hainan, China | |
5002282, F | PP133867 | Puer, Yunnan, China | |
5002308, F | PP133868 | Puer, Yunnan, China | |
A. ornata | KY349665 | ||
A. australiensis | MG882127 | ||
A. redacta sp. nov. | 5002334 | PP133862 | Honghe, Yunnan, China |
A. unicaudata sp. nov. | 5002289, F | PP133863 | Honghe, Yunnan, China |
5002291, F | PP133866 | Honghe, Yunnan, China | |
5002290 | PP133865 | Honghe, Yunnan, China | |
A. lunulara sp. nov. | 5015272, F | PP133864 | Chuxiong, Yunnan, China |
A. bifolium sp. nov. | 5002309 | PP133860 | Baoshan, Yunnan, China |
A. hemiptera sp. nov. | 5002310 | PP133861 | Baoshan, Yunnan, China |
OQ736902 | Jianfengling, Hainan, China | ||
5013913 | PP133877 | Jianfengling, Hainan, China | |
outgroup | |||
Margattea concava | MW970256 | ||
Balta vilis | KT279743.1 | ||
Sorineuchora nigra | KY349516 | ||
Shelfordina volubilis | KY349562 |
ML analysis clustered females together with morphologically similar males. We identified 11 morphospecies of Allacta on the basis of morphological characters, mainly body color, pronotum pattern, head features, legs, wing venation and male genitalia (Fig.
Maximum likelihood (ML) tree based on COI sequence. Branch node labels are support values. Colored bars indicate different species delimitation by different methods A morphology (green) B ABGD results (pink). The colored clades (red, yellow, green and blue highlights) on the tree correspond to four new morphological species.
The characteristics of the external structure and male genitalia were given in full in
Species of Allacta from China were listed by
polygrapha species group:
Allacta alba He, Zheng, Qiu, Che & Wang, 2019: 6. China (Zhejiang).
Allacta bimaculata Bey-Bienko, 1969: 858. China (Yunnan, Guangxi).
Allacta bruna He, Zheng, Qiu, Che & Wang, 2019: 4. China (Hainan).
Allacta xizangensis Wang, Gui, Che & Wang, 2014: 449. China (Xizang).
hamifera species group:
Allacta ornata Bey-Bienko, 1969: 859. China (Yunnan, Hainan).
Allacta robusta Bey-Bienko, 1969: 860. China (Yunnan).
Allacta transversa Bey-Bienko, 1969: 859. China (Hainan); Vietnam.
Allacta bifolium Li & Wang, sp. nov. China (Yunnan).
Allacta lunulara Li & Wang, sp. nov. China (Yunnan).
Allacta redacta Li & Wang, sp. nov. China (Yunnan).
Allacta unicaudata Li & Wang, sp. nov. China (Yunnan).
Allacta hemiptera Li & Wang, sp. nov. China (Hainan).
1 | Male interstylar margin broadly V-shaped | 2 (hamifera-group) |
– | Male interstylar margin without a keel-like ridge | 9 (polygrapha-group) |
2 | Tegmina and hind wings reduced, not reaching end of abdomen | 3 |
– | Tegmina and hind wings fully developed, both extending beyond end of abdomen | 5 |
3 | hind wings absent | A. bifolium sp. nov. |
– | hind wings present | 4 |
4 | Body broad, disc of pronotum dark brown with a yellowish vertical stripe in the middle | A. hemiptera sp. nov. |
– | Body narrow, disc of pronotum dark brown without vertical stripe | A. redacta sp. nov. |
5 | Head with two dark brown longitudinal stripes reaching from vertex to frons between the antennal sockets, and subgenital plate with dissimilar styli | A. robusta Bey-Bienko, 1969 |
– | Head with one dark brown longitudinal stripe reaching from vertex to clypeus or not, and subgenital plate with similar styli | 6 |
6 | Pronotal disk with an inverted triangular yellowish spot in the middle | 7 |
– | Pronotal disk without an inverted triangular yellowish spot in the middle | 8 |
7 | Face with one vertical wide dark brown stripe | A. ornata Bey-Bienko, 1969 |
– | Face with three narrow horizontal dark brown stripes | A. transversa Bey-Bienko, 1969 |
8 | Female genitalia with third valves asymmetrical and slender rod-shaped | A. unicaudata sp. nov. |
– | Female genitalia with third valves symmetrical and broad crescent-shaped | A. lunulara sp. nov. |
9 | Subgenital plate symmetrical | A. bruna He, Zheng, Qiu, Che & Wang, 2019 |
– | Subgenital plate asymmetrical | 10 |
10 | Pronotal disc brown without maculae | A. bimaculata Bey-Bienko, 1969 |
– | Pronotal disc with maculae | 11 |
11 | Pronotal disc with trapezoidal symmetrical white maculae | A. alba He, Zheng, Qiu, Che & Wang, 2019 |
– | Pronotal disc without large trapezoidal shaped white maculae posteriorly | A. xizangensis Wang, Gui, Che & Wang, 2014 |
(All in SWU). Holotype: China • male; Yunnan Prov., Baoshan City, Baihualing; 1253 m; 24 Aug., 2015; Xin-Ran Li, Zhi-Wei Qiu leg. Paratypes: China • 3 males; same data as holotype; 1 male, Yuxi City, Xinping County, Mount Ailao, 1933 m, 12 May, 2016, Lu Qiu, Zhi-Wei Qiu leg.
This species can be easily distinguished from its congeners by the small leaf-shaped tegmina, the absence of hind wings as well as the right side of the right stylus with a long, finger-like protrusion.
(mm). Male, pronotum length × width: 3.1–3.2 × 4.9–5.2, tegmina length: 1.9–2.3, overall length: 11.2–12.5.
Male. Body dark brown (Fig.
Allacta bifolium sp. nov., male holotype A dorsal view B ventral view C head, ventral view D pronotum, dorsal view E maxillary palpi, ventral view F front femur, ventral view G tarsus and tarsal claws of front leg H tegmen, dorsal view I supra-anal plate and paraprocts, ventral view J subgenital plate, dorsal view K left phallomere, dorsal view L median phallomere, dorsal view M hook-like phallomere, dorsal view. Scale bars: 5 mm (A, B); 1 mm (C–M).
Vertex with interocular space greater than the distance between antennal sockets. The third, and fourth maxillary palpi of approximately the same length, slightly longer than the fifth (Fig.
Male abdomen and genitalia. Abdominal terga unspecialized. Supra-anal plate short, symmetrical, and hind margin arc-shaped. Paraprocts simple and plate-like (Fig.
The Latin words bi- means pair, double, and folium means leaf, referring to the tegmina being degenerated into small leaf-like structures and hind wings absent.
China (Yunnan).
Holotype : China • male (SWU); Yunnan Prov., Pingbian County, Mount Dawei; 1496 m; 15 May, 2016; Lu Qiu, Zhi-Wei Qiu leg.
This species can be easily distinguished from its congeners by the wings being reduced and the pronotal disk with a brownish mushroom-shape marking.
(mm). Male, pronotum length × width: 3.3 × 4.5, tegmina length: 4.5, overall length: 13.5.
Male. Body medium-sized, yellowish brown (Fig.
Allacta redacta sp. nov., male holotype A dorsal view B ventral view C head, ventral view D pronotum, dorsal view E maxillary palpi, ventral view F front femur, ventral view G tarsus and tarsal claws of front leg H tegmen, dorsal view I hind wing, dorsal view J supra-anal plate and paraprocts, ventral view K subgenital plate and median phallomere, dorsal view L left phallomere, dorsal view M hook-like phallomere, dorsal view. Scale bars: 5 mm (A, B); 1 mm (C–M).
Vertex with interocular space greater than distance between antennal sockets. Pronotum subparabolic with hind margins nearly straight (Fig.
Male abdomen and genitalia. Abdominal terga unspecialized. Supra-anal plate short, nearly triangular, symmetrical, with hind margin blunt round. Paraprocts simple and plate-like, with scattered setae on distal margin (Fig.
The specific name redacta derived from Latin, refers to both the tegmina and hind wings being reduced, which do not reach half the length of the normal wings of its congeners.
China (Yunnan).
(All in SWU). Holotype: China • male; Yunnan Prov., Pingbian County, Mount Dawei; 1496 m; 15 May, 2016; Lu Qiu, Zhi-Wei Qiu leg. Paratypes: China • 1 male and 1 female, same data as holotype.
This species can be easily distinguished from all congeners by the absence of the left stylus in males, except for Allacta lunulara sp. nov., of which males are unknown (see below for females). This species shares a similar appearance with A. lunulara sp. nov., but it can be differentiated from the latter mainly by the following characters of the female genitalia: 1) third valves asymmetrical and slender rod-shaped, while symmetrical and broad crescent-shaped in A. lunulara; 2) posterior half of basivalvula narrower than the basal half, while basivalvula oval-shaped in A. lunulara; 3) spermatheca plate rounded with a sharp protrusion in the middle of the spermatheca plate, while front margin of spermatheca plate truncated in A. lunulara; and 4) laterosternal shelf asymmetrical, narrow, long and slightly curved, while symmetrical, broad and triangular in A. lunulara.
(mm). Male, pronotum length × width: 3.7–4.0 × 5.5–5.9, tegmina length: 10.9–12.6, overall length: 14.9–16.0; female, pronotum length × width: 3.7–4.4 × 4.6–5.7, tegmina length: 9.2–11.4, overall length: 11.5–14.7.
Male. Body yellowish brown (Fig.
Allacta unicaudata sp. nov., male holotype A dorsal view B ventral view C paratype (female), dorsal view D paratype (female), ventral view E head, ventral view F pronotum, dorsal view G maxillary palpi, ventral view H front femur, ventral view I tarsus and tarsal claws of front leg J tegmen, dorsal view K hind wing, dorsal view L supra-anal plate and paraprocts, ventral view M subgenital plate, dorsal view N left phallomere, dorsal view O median phallomere, dorsal view P hook-like phallomere, dorsal view Q female subgenital plate, dorsal view. Scale bars: 5 mm (A–D); 1 mm (E–Q). Abbreviations: a.a. anterior arch, bsv. basivalvula, intc.s. intercalary sclerite, ltst.sh. laterosternal shelf, pp. paraprocts, pt. paratergites, sp.pl. spermathecal plate, vlf.I first valvifer, v.I first valves, v.II second valves, v.III third valves. For vein abbreviations see Material and methods.
Vertex with interocular space narrower than distance between antennal sockets. The third and fourth maxillary palpi slightly longer than the fifth (Fig.
Male abdomen and genitalia. Abdominal terga unspecialized. Supra-anal plate short, symmetrical, with hind margin slightly concave. Paraprocts simple and plate-like (Fig.
Female genitalia. Supra-anal plate nearly symmetrical. Paraprocts broad. Intercalary sclerite irregular plate-shaped and translucent. First valves robust, with inward protrusions. Second valves small. Third valves asymmetrical, slender rod-shaped, and the left branch obviously curved outward. The anterior margin of anterior arch slightly sclerotized. First valvifer long and narrow plate-like with setae on the inside. Basivalvula spindle-shaped, wide in the middle and tapers at both ends. Laterosternal shelf asymmetrical, narrow, long and slightly curved. Front margin of spermathecal plate rounded with a sharp protrusion in the middle. The spermatheca lobe forked, the end of one spermatheca branch enlarged, and the other long and tubular (Fig.
This species resembles A. alba, but it can be differentiated from the latter by the following characters: 1) pronotal disc with an inverted triangular yellowish spot, while with subtrapezoidal symmetrical white markings in A. alba; 2) tegmina and hind wings slightly extending beyond the end of the abdomen, while extending far beyond the end of the abdomen in A. alba; and 3) subgenital plate without left stylus, while with two styli in A. alba. This species is placed in the hamifera species group by having the male interstylar margin broadly V-shaped.
The Latin words uni- meaning one, single, and caudata meaning tailed, referring to subgenital plate with only one stylus.
China (Yunnan).
(All in SWU). Holotype: China • female; Yunnan Prov., Chuxiong City, Mount Zixi; 2239 m; 31 Jul., 2022; Lin Guo, Wei Han leg. Paratype: China • 1 female, same data as holotype.
This species resembles A. unicaudata, but it can be differentiated from A. unicaudata mainly by the symmetrical and crescent-shaped third valves.
(mm). Female, pronotum length × width: 3.3–3.4 × 4.8–5.2, tegmina length: 8.5–9.0, overall length: 11.6–12.2.
Female. Body yellowish brown (Fig.
Allacta lunulara sp. nov., female holotype A dorsal view B ventral view C head, ventral view D pronotum, dorsal view E maxillary palpi, ventral view F front femur, ventral view G tarsus and tarsal claws of front leg H tegmen, dorsal view I hind wing, dorsal view J subgenital plate, dorsal view K supra-anal plate, dorsal view L supra-anal plate, ventral view. Scale bars: 5 mm (A, B); 1 mm (C–L). Abbreviations: a.a. anterior arch, bsv. basivalvula,intc.s. intercalary sclerite, ltst.sh. laterosternal shelf, pp. paraprocts, pt. paratergites, sp.pl. spermathecal plate, vlf.I first valvifer, v.I first valves, v.II second valves, v.III third valves. For vein abbreviations see Materials and methods.
Vertex with interocular space narrower than distance between antennal sockets (Fig.
Female genitalia. Supra-anal plate nearly symmetrical. Paraprocts broad, not extending to the posterior margin of supra-anal plate. Intercalary sclerite irregular plate-shaped and translucent. First valves robust, with inward protrusions. Second valves small. Third valves symmetrical and broad crescent-shaped. The anterior margin of anterior arch slightly sclerotized. First valvifer long and narrow plate-like with setae on the inside. Basivalvula oval-shaped. Laterosternal shelf symmetrical, broad and triangle. Front margin of spermathecal plate truncated. The spermatheca lobe forked, the end of one spermatheca branch enlarged, and the other long and tubular (Fig.
This species was found to be a cryptic species, very similar to A. unicaudata Li & Wang, sp. nov. but it can be differentiated from A. unicaudata by the female genitalia characters aforementioned. In this study, after comparing the female genitalia in five Allacta species, it is confirmed that the variation in female genitalia can be applied to identify the species of Allacta. The COI divergence between them (6.6%) is significantly larger than the usual intraspecies distance in Allacta (Suppl. material
The specific epithet is derived from the Latin word lunulara which means moon-like or relating to a crescent shape, referring to its crescent-shaped third valves.
China (Yunnan).
Temnopteryx hainanensis
Allacta hainanensis:
Holotype : China • male (SWU); Hainan Prov., Ledong County, Mount Jianfeng; 997 m; 16 Apr., 2015; Lu Qiu leg. Paratypes: China • 1 male (SEM); Hainan Prov., Changjiang County, Mount Bawang; 1495 m; 22 Sep., 2011; Xian-Wei Liu leg • 1 male (SWU); Hainan Prov., Ledong County, Mount Jianfeng; 1050 m; 6 Jul., 2007; Wei-Wei Zhang leg • 1 female (SWU); Hainan Prov., Ledong County, Mount Jianfeng; 997 m; 16 Apr., 2015; Lu Qiu leg • 3 males & 5 females (SWU); Hainan Prov., Qiongzhong County, Limushan Stone Forest; 585 m; 12 Jul., 2023; Wen-Bo Deng leg • 3 females (SWU); Hainan Prov., Qiongzhong County, Quling Valley; 662 m; 11 Jul., 2023; Yi-Shu Wang leg.
This species resembles A. redacta, but it can be differentiated from A. redacta mainly by the pronotal disk with a nib-shaped yellowish spot.
(mm). Male, pronotum length × width: 4.3–4.6 × 6.9–7.4, tegmina length: 5.5–5.7, overall length: 17.0–17.2; female, pronotum length × width: 4.0–4.4 × 6.5–7.4, tegmina length: 5.3–5.8, overall length: 16.9–17.2.
Male. Body dark brown (Fig.
Allacta hemiptera sp. nov., male holotype A dorsal view B ventral view C pronotum, dorsal view D head, ventral view E maxillary palpi, ventral view F front femur, ventral view G tarsus and tarsal claws of front leg H female subgenital plate, dorsal view I tegmen, dorsal view J hind wing, dorsal view K subgenital plate, dorsal view L supra-anal plate and paraprocts, ventral view M left phallomere, dorsal view N median phallomere, dorsal view O hook-like phallomere, dorsal view. Scale bars: 5 mm (A, B); 2 mm (C–I); 1 mm (J–O). Abbreviations: a.a. anterior arch, bsv. basivalvula, intc.s. intercalary sclerite, ltst.sh. laterosternal shelf, pp. paraprocts, pt. paratergites, sp.pl. spermathecal plate, vlf.I first valvifer, v.I first valves, v.II second valves, v.III third valves.
Vertex with interocular space obviously narrower than distance between antennal sockets (Fig.
Male abdomen and genitalia. Abdominal terga unspecialized. Supra-anal plate short, nearly triangular, symmetrical, with incision at the middle of hind margin. Paraprocts simple and plate-like, with scattered setae on distal margin (Fig.
Female genitalia. Supra-anal plate nearly symmetrical. Paraprocts broad, not extending to the posterior margin of supra-anal plate. Intercalary sclerite irregular plate-shaped and translucent. First valves robust, with inward protrusions. Second valves small. Third valves symmetrical and broad rod-shaped. The anterior margin of anterior arch slightly sclerotized. First valvifer irregular swollen and oval with short setae on the inside. Basivalvula oval-shaped. Laterosternal shelf symmetrical, broad and trapezoid. Front margin of spermathecal plate truncated. The spermatheca lobe forked, the end of one spermatheca branch enlarged, and the other long and tubular (Fig.
According to the International Code of Zoological Nomenclature (Article 16.4.2) (ICZN 1999), Temnopteryx hainanensis
The Latin terms hemi- means half, ptera means wing, and hemiptera means that the tegmina is half the normal wing length.
China (Hainan).
External characteristics and male genitalia have been traditionally used to define species of Allacta (
A Allacta transversa Bey-Bienko, 1969. Female B Allacta bimaculata Bey-Bienko, 1969. Female C Allacta robusta Bey-Bienko, 1969. Female D Allacta bruna He, Zheng, Qiu, Che & Wang, 2019. Female A–D supra-anal plate, dorsal view. Scale bars: 1 mm. Abbreviations: a.a. anterior arch, bsv. basivalvula, intc.s. intercalary sclerite, ltst.sh. laterosternal shelf, pp. paraprocts, pt. paratergites, sp.pl. spermathecal plate, vlf.I first valvifer, v.I first valves, v.II second valves, v.III third valves.
The clade highlighted in blue (Fig.
In future research, morphology specific to females should be taken into greater consideration, as it played a key role in the discovery of new species in this study. Sometimes female genitalia can even be used to discover cryptic species, for example,
In this study, the absence of one stylus in the family Pseudophyllodromiidae was observed for the first time (A. unicaudata Li & Wang, sp. nov.), but this is not a unique case in Blattodea (
We are deeply grateful to the following collectors of specimens: Zhi-Wei Qiu, Lu Qiu, Yong Li, Wei Han, and Xin-Ran Li. We also extend our sincere thanks to Dominic Evangelista and the anonymous reviewer for their valuable suggestions on our manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This study is supported by a Program of the Ministry of Science and Technology of the People’s Republic of China (2022FY202100) and the National Natural Sciences Foundation of China (nos. 32170458, 32070468 and 31872271).
Bianlun Li: Data curation (equal); methodology (lead); visualization (equal); writing – original draft (lead); writing – review and editing (equal). Penghui Hu: Data curation (equal); visualization (supporting); Lin Guo: Data curation (equal); visualization (supporting); writing – review and editing (supporting). Yanli Che: Funding acquisition (equal); project administration (equal); supervision (equal); writing – review and editing (equal). Zongqing Wang: Funding acquisition (equal); methodology (supporting); project administration (equal); supervision (equal); writing – review and editing (equal).
Bian-Lun Li https://orcid.org/0009-0009-1773-0900
Yan-Li Che https://orcid.org/0000-0003-3214-9494
Zong-Qing Wang https://orcid.org/0000-0001-9413-1105
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Interspecific and intraspecific genetic distances of Allacta
Data type: xls