Research Article |
Corresponding author: Xueyan Li ( lixy@mail.kiz.ac.cn ) Academic editor: Hans Pohl
© 2022 Zhiwei Dong, Xingyue Liu, Chuyang Mao, Jinwu He, Xueyan Li.
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:
Dong Z, Liu X, Mao C, He J, Li X (2022) Xenos yangi sp. nov.: A new twisted-wing parasite species (Strepsiptera, Xenidae) from Gaoligong Mountains, Southwest China. ZooKeys 1085: 11-27. https://doi.org/10.3897/zookeys.1085.76484
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Here we report a new twisted-wing parasite species of the family Xenidae based on both morphological and molecular evidence. By using nearly complete mitogenomes, we confirmed the twisted-wing parasites on two wasps (Vespa velutina and Vespa bicolor) (China: Yunnan) as the same species, and associated its neotenic females and alate males. Combining the mitogenomic data (COI) and morphological traits, this species was identified to be a new species of the genus Xenos, namely Xenos yangi Dong, Liu & Li, sp. nov. Detailed descriptions and illustrations are provided for the new species.
Mitogenome, morphology, new species, taxonomy, wasp endoparasite
Strepsiptera are obligate endoparasites of silverfish, cockroaches, mantids, orthopterans, hemipterans, wasps, bees and flies, and they comprise about 630 species in 15 families (
In December 2019, some wasps (Vespa velutina Lepeletier, 1836 and Vespa bicolor Fabricius, 1787) were collected by local villagers in southern Gaoligong Mountains (Yunnan, China). We checked these wasp individuals and found some of them parasitized by twisted-wing parasites. We collected male adults (Figs
The male and neotenic female specimens of the new species Xenos yangi Dong, Liu & Li, sp. nov. were collected from the nests of both V. velutina and V. bicolor in Gaoligong Mountains, Xiangda Township, Longling County, Yunnan Province in December, 2019. The type materials of the new species described in this paper are deposited in the Insect Collection of Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China (
Images of the living adults were taken using a Canon 70D camera in conjunction with a Canon EF 100 mm f/2.8L IS USM. The habitus images were taken using a stereomicroscope Nikon, SMZ18 equipped with NIS-Elements (Nikon, Japan). Scanning electron microscopes (SEM) images were taken using TM4000 II (Hitachi, Japan). The specimens used for SEM were directly fixed in 70% ethanol, and then dried at the room temperature. Morphological terminology follows those of
Total genomic DNA of one male collected from V. velutina nest and one neotenic female collected from V. bicolor nest was extracted using a TIANamp Genomic DNA Kit (TIANGEN, China) based on manual instruction. Library construction, sequencing, mitogenome assembly follows those in our previous work (
COI is an useful molecular marker for species identification in many insects, including twisted-wing parasites (
Combined with 45 COI sequences of Xenos published by others (
We assembled nearly complete mitogenomes of one male adult collected from a V. velutina nest (15324 bp) (GenBank accession number: OK329871) and one neotenic female collected from a V. bicolor nest (14670 bp) (GenBank accession number: OK32987). The mitogenomes of these two individuals in this study and one neotenic female in our previous work (
We further extracted a major fragment (1518 bp) of COI sequences from three Chinese Xenos individuals (one male and two neotenic females), and combined 45 COI sequences of identified species of Xenos reported by others to make the dataset for the phylogenetic analyses using ML and MP methods. All phylogenetic trees show that the three Chinese Xenos individuals (one male individual from the V. velutina nest and two female individuals from the V. velutina and V. bicolor nests) cluster together with high bootstrap values (Fig.
Xenidae Saunders, 1872
China, Yunnan, Longling County, Xiangda Township.
Holotype
: male (KIZ0130767), “Gaoligong Mountains, Xiangda Township, Longling County, Yunnan Province, 24.4441083 N, 98.7239194 E, 1666 m, 20.XII.2019, local villagers leg.”, kept in 75% ethanol, [red label]. (
One neotenic female, “Gaoligong Mountains, Xiangda Township, Longling County, Yunnan Province, 20. XII. 2019, local villagers leg.”, partially used for extracting genomic DNA (accession number MW222190;
Male. Head transverse. Antenna (Fig.
Male (Fig.
Hind wing
sector with nine veins (Fig.
Proventrite
laterally with anepisternum angulately curved at middle, and posteromedially with a small U-shaped notch, forming a pair of short lobes (Fig.
Cephalotheca of male puparium (Fig.
Neotenic female (Fig.
Xenos yangi Dong, Liu & Li sp. nov. A male cephalotheca frontal view (CP, Clypeus; AN, Antenna; EYE, Eye; MD, Mandible; MX, Maxillae) B female ventral view (BC, brood canal; BOR, birth organs) C, D female cephalothorax ventral view (BO, birth opening; SBHP, segmental border between head and prothorax; OS, mouth opening; MD, mandible). Scale bar: 0.5 mm.
Considering the geographic distance and host association of those species of Africa and Americas, we mainly compared the male adult, the cephalotheca of the male papurium, and the neotenic female of this new species with ten described known species distributed in Asia (nine species) and Europe (one species) (Table
Distribution and described stages of 11 Xenos species from Asia and Europe. Literature in which the species was originally described is highlighted in bold.
Species | Distribution | Male | Cephalotheca of male puparium | Neotenic female | Primary larvae |
---|---|---|---|---|---|
Xenos yangi Dong, Liu & Li sp. nov. | China: Yunnan | This study | This study | This study | NA |
Xenos moutoni (Buysson, 1903) | China: Yunnan, Anhui, Taiwan | Kifune & Maeta, 1985 | Buysson, 1904 | Buysson, 1903 | NA |
Xenos circularis Kifune & Maeta 1985 | China: Taiwan | NA | NA | Kifune & Maeta, 1985 | NA |
Xenos yamaneorum Kifune & Maeta, 1985 | China: Taiwan | NA | Kifune & Maeta, 1985 | Kifune & Maeta, 1985 | NA |
Xenos formosanus Kifune & Maeta, 1985 | China: Taiwan | Kifune & Maeta, 1985 | Kifune & Maeta, 1985 | Kifune & Maeta, 1985 | NA |
Xenos dianshuiwengi Yang, 1999 | China: Fujian | Yang, 1999 | NA | NA | NA |
Xenos oxyodontes Yuta & Makoto 2013 | Japan | Yuta & Makoto, 2013 | Yuta & Makoto, 2013 | Yuta & Makoto, 2013 | NA |
Xenos vespularum Kifune & Maeta, 1975 | Japan | Kifune & Maeta, 1975 | Kifune & Maeta, 1975 | Kifune & Maeta, 1975 | NA |
Xenos hebraei Kinzelbach, 1978 | India | NA | Kinzelbach, 1978 | Kinzelbach, 1978 | NA |
Xenos provesparum Kifune,1986 | Indonesia | Kifune, 1986 | Kifune, 1986 | Kifune, 1986 | NA |
Xenos vesparum Rossius, 1793 | Europe; Northern Africa | Rossius, 1793 | Rossius, 1793 | Rossius, 1793 | Pohl & Beutel, 2005 |
The new species can be distinguished from X. circularis (China: Taiwan), X. yamaneorum (China: Taiwan), X. vespularum (Japan), X. hebraei (India) and X. vesparum (Europe; Northern Africa) by the female cephalothorax. It is almost circular or ovoid in X. yamaneorum, X. circularis, X. vespularum and X. vesparum (
China (Yunnan).
The hosts of this new species are Vespa velutina (Fig.
The specific epithet is dedicated to the late famous Chinese entomologist Chi-Kun Yang, who made significant contributions to the studies on Strepsiptera in China.
Phylogeny tree of Xenos species inferred from mitochondrial cytochrome c oxidase subunit 1 (COI) using Maximum parsimony method. In total, 48 COI sequences of different Xenos species were used to investigate their phylogenetic relationships. two sequences (str6-b-female and Xenos-male) were sequenced in this study, and that of Xenos cf. moutoni (MW222190.2) was sequenced in
Due to the discovery of X. yangi sp. nov., the number of Chinese Xenos species increases to six (Fig.
Among the 10 Asian Xenos species, six species (including the new species here) are described based on both males and neotenic females, one species solely based on males, two species are based on neotenic females and the cepholotheca of the male puparium, and one species is solely based on neotenic females (Table
Xenos moutoni was originally described by
In the molecular data analysis, we noticed that different populations of five monophylic species (X. moutoni, X. oxyodontes, X. yangi, X. pecki, and X. vespularum) show genetic divergence of less than 0.036 (Table
Summary of pairwise distances based on COI sequences among different Xenos species.
Group | Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 |
---|---|---|---|---|---|---|---|---|---|---|---|---|
1 | X. moutoni | 0–0.014 | – | – | – | – | – | – | – | – | – | – |
2 | X. oxyodontes | 0.111–0.123 | 0–0.071 | – | – | – | – | – | – | – | – | – |
3 | X. yangi sp. nov. | 0.191–0.208 | 0.269–0.287 | 0–0.014 | – | – | – | – | – | – | – | – |
4 | X. pecki | 0.322–0.344 | 0.369–0.382 | 0.338–0.361 | 0–0.036 | – | – | – | – | – | – | – |
5 | X. ropalidiae | 0.330–0.348 | 0.308–0.341 | 0.302–0.333 | 0.326–0.358 | 0.089 | – | – | – | – | – | – |
6 | X. minor | 0.425–0.432 | 0.451–0.465 | 0.497–0.508 | 0.544–0.560 | 0.367–0.377 | 0 | – | – | – | – | – |
7 | X. vesparum | 0.425–0.440 | 0.451–0.473 | 0.497–0.516 | 0.544–0.568 | 0.367–0.385 | 0–0.003 | 0–0.003 | – | – | – | – |
8 | X. minor | 0.425–0.432 | 0.451–0.465 | 0.497–0.508 | 0.544–0.560 | 0.367–0.377 | 0 | 0–0.003 | NA | – | – | – |
9 | X. vesparum | 0.411–0.434 | 0.439–0.475 | 0.494–0.507 | 0.526–0.588 | 0.358–0.393 | 0.001–0.003 | 0.001–0.007 | 0.001–0.003 | 0.005 | – | – |
10 | X. ropalidiae | 0.275–0.383 | 0.329–0.404 | 0.382–0.433 | 0.452–0.474 | 0.368–0.421 | 0.435–0.517 | 0.435–0.525 | 0.435–0.517 | 0.437–0.552 | 0.2 | – |
11 | X. vespularum | 0.481–0.505 | 0.451–0.478 | 0.4472–0.478 | 0.549–0.577 | 0.458–0.472 | 0.608–0.612 | 0.608–0.616 | 0.608–0.612 | 0.590–0.608 | 0.519–0.523 | 0.001 |
We give thanks to those anonymous villagers for helping to collect wasps harboring parasites, Dr Jiangli Tan for helping in wasp identification, Dr Guichun Liu and Ms Ruoping Zhao for helping in DNA extraction, Dr Ru Zhang and Jinwu He for helping in data analysis, Dr Tian-Tian Yu for reading the manuscript. Special thanks are given to Prof. Wen Wang for his courage to perform insect taxonomic work. This work was supported by grants from Yunnan Provincial Science and Technology Department (202105AC160039) and Chinese Academy of Sciences (CAS “Light of West China” Program) to X.L.
Table S1
Data type: xlsx file
Explanation note: Known species of Xenos and their geographical distribution, type depository and host.