Research Article |
Corresponding author: Yuxia Yang ( yuxia0305@126.com ) Corresponding author: Haoyu Liu ( liuhy@aliyun.com ) Academic editor: Hume Douglas
© 2019 Yuxia Yang, Huacong Xi, Xingke Yang, Haoyu Liu.
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:
Yang Y, Xi H, Yang X, Liu H (2019) Taxonomic review of the Themus (Telephorops) nepalensis species-group (Coleoptera, Cantharidae). ZooKeys 884: 81-106. https://doi.org/10.3897/zookeys.884.32550
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The diagnosis of the Themus (Telephorops) nepalensis species-group is summarized. A catalogue, a key and a distribution map of all world species are provided. Two synonymies are proposed: Themus (Telephorops) subcaeruleiformis Wittmer, 1983, syn. nov. = T. (Telephorops) crassimargo Champion, 1926; T. (Telephorops) separandus Wittmer, 1975, syn. nov. = T. (Telephorops) laboissierei (Pic, 1929). The female internal genitalia are photographed and described in this species-group for the first time, the aedeagi of T. (Telephorops) crassipes Pic, 1929 and T. (Telephorops) impressipennis (Fairmaire, 1886) are illustrated and described for the first time, and some additional distribution information is provided for the species. Themus (Telephorops) cavipennis (Fairmaire, 1897) is a new record for the Chinese fauna.
female internal genitalia, new faunistic record, new synonym, soldier beetles, taxonomy
Themus Motschulsky, 1858 is one of the largest cantharid genera and comprises about 250 species in total (
The subgenus Telephorops Fairmaire, 1886 for T. impressipennis (by original and monotypic designation) was subdivided into two species groups, which however were not given names (
Most species of the nepalensis species-group were described by early taxonomists, such as
Up to now, 15 species were included in the nepalensis species-group. This group has not previously been reviewed globally, and some sibling species remain difficult to diagnose from others due to there being few characters known for the females when males are unavailable. Furthermore, attribution of species to the groups is difficult because species diagnoses are often imprecise. For example, T. minor Wittmer, 1997, T. subcaeruleus (Pic, 1911) and T. crassimargo Champion, 1926, whose elytra are enlarged and laterophyses well developed, give contradictory information about their placement in the species-group defined by
The material is deposited in the following collections:
MHBU Museum of Hebei University, Baoding, China;
Genitalia of both sexes and abdominal sternites VIII of females were dissected and cleared in 10% KOH solution, and female genitalia were stained with hematoxylin. The female internal genitalia is attached to the ventral side of abdominal tergite IX and the vulva opens between the coxites. The dorsal or ventral side of vagina is established according to the tergite IX. The situation of median oviduct opening is on the opposite side of tergite IX and established as the ventral side of vagina. The diverticulum and spermatheca arise from apex of vagina.
Habitus photos were taken using a Leica M205 A stereomicroscope, multiple image layers were stacked using Combine ZM (Helicon Focus 5.3). Line drawings were made using a camera lucida attached to a Nikon SMZ1500 stereomicroscope, then edited in CorelDRAW 12 and Adobe Photoshop 8.0.1. Body length was measured from the anterior edge of the clypeus to the elytral apex and body width across the humeri of elytra. Morphological terminology of female genitalia followed
In the checklist, valid scientific names and original sources, synonyms and publications for the taxonomical changes, type localities and depositories, additional material information and all distributions were included, as well as additional description or remarks were added if necessary. Complete label data were cited for type specimens, using square brackets “ []” for our remarks and comments, [p] indicating that the following data were machine printed and [h] that they were handwritten, quotation marks to separate data from different labels. A distribution map was prepared using the geographic information system software ArcGIS (ver. 10.2), based literature records and the author's databases of specimens examined for this study.
The specimens were identified based on examination of types if available and original literature. In practice, species were determined mainly by the aedeagus of male, and the females were associated with males based on evidence that they were collected at the same locality and date. Also, the female could be identified by the structure of abdominal sternite VIII, which was useful in species’ recognition and illustrated in the literature by cantharid specialists. For each species, compared with males, the females have smaller eyes, shorter and narrower antennae, simple middle antennomeres, without smooth narrow impressions along the outer edges, wider pronotum and elytra, and only seven abdominal ventrites.
Elytra enlarged posteriorly and widest near apical third. Aedeagus: conjoint dorsal plate of parameres narrowed apically in dorsal view, emarginate at middle of apical edge; laterophyses flattened dorsoventrally, reduced and not reaching apices of conjoint dorsal plate except in a few species. Female internal genitalia: diverticulum situated at end of vagina, presenting with a sclerotized ring around at base, confluent in middle and extending to median oviduct; spermatheca arising from middle of the sclerotized ring.
Distribution. Most species are restricted in their distribution (Figs
The diagnosis is developed from the definition of the species-group by
(characters based on illustrations in the present study or those from
1 | Aedeagus: laterophyses reaching apices of conjoint dorsal plate of parameres | 2 |
– | Aedeagus: laterophyses reduced, not reaching apices of conjoint dorsal plate of parameres | 4 |
2 | Antennae, mid and hind legs uniformly black; aedeagus ( |
T. minor Wittmer, 1997 |
– | Antennae and legs mixed yellow and black; aedeagus: laterophyses with a narrow process each side | 3 |
3 | Aedeagus ( |
T. subcaeruleus (Pic, 1911) |
– | Aedeagus ( |
T. crassimargo Champion, 1926 |
4 | Aedeagus: ventral process of each paramere hooked at apex in lateral view | 5 |
– | Aedeagus: ventral process of each paramere not hooked at apex | 9 |
5 | Aedeagus: ventral process of each paramere expanded and obtusely hooked dorsally at apex (e.g. Fig. |
6 |
– | Aedeagus: ventral process of each paramere narrowed and acutely hooked ventrally at apex | 8 |
6 | Elytra dark green or blue, strongly metallic | T. nepalensis (Hope, 1831) |
– | Elytra purple-black, weakly metallic | 7 |
7 | Body longer than 16.0 mm; female internal genitalia (Fig. |
T. crassipes Pic, 1929 |
– | Body 13.0–15.0 mm in length; female internal genitalia (Fig. |
T. masatakai Okushima, 2003 |
8 | Aedeagus (Fig. |
T. cavipennis (Fairmaire, 1897) |
– | Aedeagus ( |
T. uncinatus Wittmer, 1983 |
9 | Elytra no more than 1.5 times as long as maximal width; aedeagus ( |
T. laboissierei Pic, 1929 |
– | Elytra about twice as long as maximal width; aedeagus: ventral process of each paramere with apex unlike above, not bent inwards in ventral view, longer than conjoint dorsal plate in lateral view | 10 |
10 | Aedeagus ( |
T. sauteri (Pic, 1912) |
– | Aedeagus (Fig. |
11 |
11 | Tibiae mixed yellow and black; aedeagus ( |
T. coelestis (Gorham, 1889) |
– | Tibiae uniformly black or yellow; aedeagus: ventral process of each paramere almost even in width in ventral view, conjoint dorsal plate narrowly emarginate medially at apical edge in dorsal view | 12 |
12 | Femora mixed yellow and black, tibiae black; female abdominal sternite VIII (Fig. |
T. impressipennis (Fairmaire, 1886) |
– | Femora and tibiae uniformly yellow; female abdominal sternite VIII (Fig. |
T. bicoloricornis Wittmer, 1983 |
Themus (Telephorops) bicoloricornis Wittmer, 1983a: 153, figs 48 (aedeagus illustration), 51 (female abdominal sternite VIII illustration).
1♂ (paratype,
Other material examined. 1♂, 1♀ (
Supplementary description. Female. Like male, but antennomeres V–X without impressions along outer edges (while present with smooth narrow longitudinal or oblong impressions in male), terminal abdominal ventrite wide (while narrow and triangular in male) (Fig.
Taiwan.
Tryblius cavipennis Fairmaire, 1897: 228.
Themus ancoralis
Champion, 1926: 128. Synonymized by
Themus (Tryblius) cavipennis:
Themus (Telephorops) cavipennis:
1♂ (holotype,
Other material examined. CHINA: Xizang: 1♂, 1♀ (
Male (Fig.
China (new record: Xizang), Bhutan, Nepal, northern India.
Telephorus coelestis Gorham, 1889: 104, t.10, fig. 7.
Themus (Telephorops) coelestis:
Themus rugosus
Pic, 1929b: 8. Synonymized by
Themus violetipennis
Wang & Yang, 1992: 265, fig. 2 (habitus illustration). Synonymized by
1♂ (
1♀ (
1♀ (
CHINA, Shaanxi: 2♂ (MHBU), Chushui, Niubeiliang, 1056 m, 2011.VIII.22–29, leg. X.C. Zhu & Y. Zhao. Hubei: 1♀ (MHBU), Badong, Lvcongpo, 1700 m, 2006.VII.14, leg. M. Li; 1♀ (MHBU), same data, leg. J. H. Wan; 1♀(MHBU), Yuan’an, Hehua, 2009.VII.12, leg. X.M. Sun; 1♀ (MHBU), same data, leg. Y. Dong; 1♀ (MHBU), Yichang, Xianrenxi, 2009.VI.25, leg. G.L. Xie; 1♀ (MHBU), Yichang, Dalaoling Forestry, 2009.VI.26, leg. Y. Tian; 1♀ (MHBU), Yidu, Niejiahe, 2008.VI.16, leg. G. L. Xie; 1♀ (MHBU), Yichang, Hejiaping, Qinggangping, 2013.VII–XI, leg. T.H. Du; 1♀ (MHBU), Changyang, Langping, Changfeng, 900 m, 2013.VII.11, leg. Y.Q. Wu; 1♂ (MHBU), Jingshan, Huzhuashan Forestry, 2007.VII.15, leg. G.L. Xie; 1♂ (MHBU), Wufeng, Houhe, 2002.VII.16, leg. F.Y. Wang; 1♂ (MHBU), same locality, 2002.VII.21, C.H. Shi. Guangxi: 1♀ (MHBU), Luocheng, Pingying, 2004.V.29, leg. J.M. Zhang. Guangdong: 1♀ (MHBU), Nanling, 2010.VIII.8–18, leg. H.Y. Liu; 1♀ (MHBU), same locality and collector, 2010.VIII.17; 1♀ (MHBU), same locality and collector, 2010.VIII.8–11. Hebei: 1♂ (MHBU), Changli, Huangjin seaside, 1999.VIII.18, leg. H.Z. Liang; 1♂ (MHBU), same data, leg. Z.J. Ma; 1♂(MHBU), Zushan, 1998.VII.14, leg. X.J. Li. Zhejiang: 1♀ (MHBU), Longquan, Fengyangshan, 2007.VII.25, leg. L.K. Tan; 1♀ (MHBU), same locality and collector, 2007.VII.30; 1♀ (MHBU), same locality, 2007.VII.26, leg. G.L. Xie; 1♀ (MHBU), same locality and collector, 2007.VII.27; 1♀ (MHBU), same locality and collector, 2007.VII.31; 3♂, 1♀ (MHBU), same locality, 2007.VII.25–VIII.1, leg. H.Y. Liu & Z.H. Gao; 1♂ (MHBU), same locality, 2012.VII.18, leg. G.L. Xie & J. Jiao; 2♀ (MHBU), Lin’an, Tianmushan, 2013.VI.26–VII.2, leg. J.Y. Su; 2♀ (MHBU), Hangzhou, Lin’an, Dajingwu, 2012.VI.10, leg. H. Xu; 1♂ (MHBU), Qingyuan, Baishanzu, 2012.VII.24, G.L. Xie & X. Wang. Yunnan: 2♂ (MHBU), Dali, 2008.VIII.18, leg. G. L. Xie.
Female. Like male, but antennomeres IV–X without impressions along outer edges (while present with smooth narrow longitudinal or oblong impressions in male), terminal abdominal ventrite wide (while narrow and triangular in male) (Fig.
China (Shaanxi, Gansu, Henan, Anhui, Zhejiang, Hubei, Jiangxi, Hunan, Fujian, Hainan, Guangxi, Sichuan, Guizhou).
Themus crassimargo Champion, 1926: 127.
Themus (Tryblius) crassimargo:
Themus (Telephorops) subcaeruleiformis Wittmer, 1983b: 199, fig. 3, 3a (aedeagus illustration), syn. nov.
Themus (Telephorops) crassimargo:
1♂ (
1♂ (
CHINA, Xizang: 1♂ (
China (Xizang, Sichuan), N. India, Bhutan, Nepal.
Female (Fig.
Themus (T.) subcaeruleiformis Wittmer, 1983 was originally described based on a single male type, from China, Szechwan, Kuanshien Umg. (now in Sichuan, Dujiangyan). Here a female (Fig.
Themus crassipes Pic, 1929b: 8.
Themus (Telephorops) crassipes:
1♂ (
CHINA, Guangxi: 1♂, 1♀ (
Male. Aedeagus (Fig.
Female. Like male, but antennomeres V–X without impressions along outer edges (while present with smooth narrow longitudinal or oblong impressions in male), terminal abdominal ventrite wide (while narrow and triangular in male) (Fig.
China (Guangxi, Yunnan); Vietnam.
Telephorops impressipennis Fairmaire, 1886: 339.
Telephorops violaceipennis
Gorham, 1889: 105. Synonymized by
Themus (Telephorops) impressipennis:
1♂ (
1♀ (
CHINA, Guizhou: 2♀ (MHBU), Daozhen, Xiannvhe, 2004.VIII.24–26, leg. X.J. Yang & H.R. Hua; 1♀ (MHBU), Suiyang, Baishaogou, 2010.VIII.14, leg. L.Y. Guo; 1♂ (
Male. Aedeagus (Figs
Female. Like male, but antennomeres V–X without impressions along outer edges(while present with smooth narrow longitudinal or oblong impressions in male), terminal abdominal ventrite wide (while narrow and triangular in male) (Fig.
China (Gansu, Shaanxi, Henan, Jiangsu, Anhui, Zhejiang, Hubei, Jiangxi, Hunan, Fujian, Taiwan, Guangxi, Sichuan, Guizhou, Yunnan).
Triblius laboissierei Pic, 1929a: 195, 196.
Themus (Tryblius) separandus Wittmer, 1975: 252, fig. 3 (aedeagus illustration). syn. n.
Themus (Telephorops) laboissierei:
Themus (Telephorops) separandus:
1♂ (
1♂ (
CHINA, Yunnan: 1♂ (
Female. Like male, but antennomeres V–X without impressions along outer edges (while present with smooth narrow longitudinal or oblong impressions in male), terminal abdominal ventrite wide (narrower and triangular in male) (Fig.
China (Yunnan, Guangxi); northern Laos, northern Vietnam, northern India.
Themus (Tryblius) separandus was described based on a single male type and its aedeagus was illustrated by
In the present study, the habitus (Fig.
Themus (Telephorops) masatakai
Okushima, 2003: 280, figs 1–4 (habitus photo, aedeagus illustrations);
LAOS: 1♂, 1♀ (
CHINA,Yunnan: 1♂ (
Female. Like male, but antennomeres VII–XI without impressions along outer edges (while present with smooth narrow longitudinal or oblong impressions in male), terminal abdominal ventrite wide (while narrow and triangular in male) (Fig.
China (Yunnan, Guangxi); Laos, northernVietnam.
Themus (Telephorops) minor
Wittmer, 1997: 272, fig. 104 (aedeagus illustration);
1♂ (holotype,
China (Yunnan).
Telephorus nepalensis Hope, 1831: 26.
Themus (Tryblius) nepalensis:
Themus (Telephorops) nepalensis:
Northern India, Nepal.
Cantharis sauteri Pic, 1912: 46.
Themus sauteri:
Themus (Telephorops) sauteri:
Taiwan: 1♂ (
Female. Like male, but antennomeres VI–X without impressions along outer edges (while present with smooth narrow longitudinal or oblong impressions in male), terminal abdominal ventrite wide (while narrow and triangular in male) (Fig.
Tryblius cavipennis var. subcaeruleus Pic, 1911: 132.
Themus (Tryblius) subcaeruleus:
Themus (Telephorops) subcaeruleus:
1♂ (
CHINA: 1♀ (
China (Yunnan), northern Vietnam.
Themus (Telephorops) uncinatus Wittmer, 1983b: 200, figs 5 (aedeagus illustration), 66 (female abdominal sternite VIII illustration).
1♂ (holotype,
CHINA, Sichuan: 1♂ (
Female. Like male, but antennomeres V–X without impressions along outer edges (while present with smooth narrow longitudinal or oblong impressions in male), terminal abdominal ventrite wide (while narrow and triangular in male) (Fig.
China (Sichuan, Yunnan), northern Vietnam.
Female abdominal sternite VIII, ventral view A Themus bicoloricornis Wittmer, 1983 B T. cavipennis Champion, 1926 C T. coelestis (Gorham, 1889) D T. crassimargo Champion, 1926 E T. crassipes Pic, 1929 F T. impressipennis (Fairmaire, 1886) G T. laboissierei (Pic, 1921) H T. masatakai Okushima, 2003 I T. sauteri (Pic, 1912) J T. uncinatus Wittmer, 1983. Scale bars: 1.0 mm.
We are grateful to Dr Yûichi Okushima (Kurashiki Museum of Natural History, Japan) for proving some material to us, and to the late Dr Michel Brancucci (