Research Article |
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Corresponding author: Haoyu Liu ( liuhy@aliyun.com ) Corresponding author: Yuxia Yang ( yuxia0305@126.com ) Academic editor: Vinicius S. Ferreira
© 2023 Younan Wang, Haoyu Liu, Xingke Yang, Yuxia Yang.
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:
Wang Y, Liu H, Yang X, Yang Y (2023) Review of the Lycocerus pallidulus group (Coleoptera, Cantharidae), with descriptions of six new species from China. ZooKeys 1176: 243-285. https://doi.org/10.3897/zookeys.1176.107858
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The Lycocerus pallidulus subgroup, originally placed in the L. maculicollis group, is suggested as an independent species group herein and its diagnosis is redefined. Ten previously known species of Lycocerus are attributed to this group, including L. centrochinensis (Švihla, 2004), L. genaemaculatus (Wittmer, 1951), L. hubeiensis (Švihla, 2004), L. kubani (Švihla, 2004), L. zdeneki (Švihla, 2004), L. bilineatus (Wittmer, 1995), L. jelineki (Švihla, 2004), L. putzi Švihla, 2011, L. pictipennis (Wittmer, 1995), and L. curvatus (Wittmer, 1995). Additionally, six new species of this group are described from China, including L. laterophysus sp. nov., L. flavipennis sp. nov., L. putzimimus sp. nov., L. maoershanensis sp. nov., L. chongqingensis sp. nov., and L. bispermathecus sp. nov. These species are illustrated with photographs of habitus, aedeagi, abdominal sternites VIII, and reproductive systems of female. In addition, an identification key and a distribution map of the L. pallidulus group are provided.
Alpha taxonomy, Lycocerus, Oriental region, soldier beetles
Lycocerus Gorham, 1889 sensu
Prior to this study, three taxa were included in L. pallidulus subgroup, including L. pallidulus (Wittmer, 1995), L. guerryi (Pic, 1906), and L. guerryi atroapicipennis (Pic, 1914) (
The studied material is preserved in the following collections:
CWNU China West Normal University, Nanchong, China;
MHBU Museum of Hebei University, Baoding, China;
NWAFU Northwest Agriculture & Forestry University, Yangling, China;
ZFMK Zoological Research Museum Alexander Koenig, Bonn, Germany.
We identified specimens based on examination of the type material and reference to the relevant publications (
Genitalia of both sexes and abdominal sternites VIII of females were dissected and cleared in 10% NaOH solution, and female genitalia was dyed with hematoxylin. Habitus photos were taken by a Leica M205A stereomicroscope. Line drawings were made using a camera lucida attached to a Nikon SMZ1500 stereomicroscope, then edited in CorelDraw 12 and Adobe Photoshop CS6.13.0.
The label of the specimens in Chinese are transliterated, and the complete label data are cited for the type specimens. The distribution information was collected from the publications (
Common characteristics. Body middle-sized (8.0–11.5 mm), slender. Antennae filiform, present or absent with impressions on middle antennomeres in male. Pronotum subquadrate, longer than or nearly as long as wide. Elytra pale yellow or even transparent, sometimes with black longitudinal bands or markings on disc, subparallel-sided. Tarsal claws diverse: if all simple in male, fore and mid-anterior and or posterior claws each with a digitiform tooth at base in female; when fore and mid-anterior claws each with a tooth in male, same in female or both anterior and posterior claws each with a tooth; otherwise, fore and mid-anterior and posterior claws each with a tooth in both sexes; hind claws always simple. Aedeagus (Fig.
Aedeagus of Lycocerus curvatus (Wittmer, 1995): A ventral view B dorsal view C lateral view. Abbreviations: bp – basal piece; dp – dorsal plate of each paramere; lp – laterophyse; ml – median lobe; vp – ventral process of each paramere; is – inner sac. Scale bar: 1.0 mm. The red hues show basal pieces (with middle nodule in ventral part missing), green for parameres, and yellow represents median lobe exhibiting with inner sac and laterophyses.
Included species. L. pallidulus (Wittmer, 1995), L. guerryi (Pic, 1906), L. guerryi atroapicipennis (Pic, 1914), L. centrochinensis (Švihla, 2004), L. genaemaculatus (Wittmer, 1951), L. hubeiensis (Švihla, 2004), L. jelineki (Švihla, 2004), L. putzi Švihla, 2011, L. bilineatus (Wittmer, 1995), L. zdeneki (Švihla, 2004), L. kubani (Švihla, 2004), L. curvatus (Wittmer, 1995), L. pictipennis (Wittmer, 1995), L. laterophysus sp. nov., L. flavipennis sp. nov., L. putzimimus sp. nov., L. maoershanensis sp. nov., L. chongqingensis sp. nov. and L. bispermathecus sp. nov.
Distribution (Fig.
Athemus (Isathemus) pallidulus Wittmer, 1995: 261, figs 119, 189.
Lycocerus pallidulus:
Holotype
: 1♂ (
China (Yunnan).
This species was omitted from the Palaearctic Catalogue by
Cantharis guerryi Pic, 1906: 83.
Athemus (Isathemus) guerryi:
Lycocerus guerryi:
Holotype
: 1♂ (
China: 4♂1♀ (
Male. Aedeagus: basal piece nearly as long as dorsal plate of each paramere (Fig.
Female. Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
China (Yunnan).
Cantharis jeanvoinei Pic, 1927 was listed as a synonym of L. guerryi by
Cantharis atroapicipennis Pic, 1914:8.
Athemus (Isathemus) guerryi atroapicipennis:
Cantharis jeanvoinei
Pic, 1927: 2. Synonymized by
Lycocerus guerryi atroapicipennis:
Holotype
of Cantharis atroapicipennis: 1♂ (
Holotype
of Cantharis jeanvoinei: 1♀ (
China: 1♂ (
China (new record: Yunnan); Vietnam.
The aedeagus of this subspecies is very similar to that of L. guerryi guerryi, but its elytra are darkened at apices enabling external identification. The body coloration is constant, and the elytra of the individuals from northern Yunnan are uniformly pale yellow (L. guerryi guerryi), while those from southern Yunnan and adjacent area (northern Vietnam) are always darkened at apices (L. guerryi atroapicipennis), so they are kept as two subspecies at the moment. Here, L. guerryi atroapicipennis is recorded in the Chinese fauna for the first time.
Athemus (s.str.) centrochinensis Švihla, 2004: 182, figs 83–85, 202.
Lycocerus centrochinensis:
Holotype
: 1♂ (
China: 2♂1♀ (MHBU), Sichuan, Wolong, 6–7.VIII.2004, leg. X. J. Yang & H. R. Hua; 1♂1♀ (MHBU), Gansu, Wenxian, Huangtuling, 2350 m, 8.VII.2003, leg. Y. B. Ba & Y. Yu; 1♂ (MHBU), Hubei, Dalaoling Nature Reserve, 1200 m, 9.VII.2011, leg. X. L. Liang; 1♀ (MHBU), same locality as the preceding, 11.VII.2011, leg. X. Liao; 1♂ (MHBU), Hubei, Shennongjia, Muyuzhen, 1200 m, 12.VII.2004, leg. S. Q. Xu; 1♀ (MHBU), same data as the preceding, leg. F. L. Zou; 1♀ (MHBU), Hubei, Shennongjia, Wenshui Forestry, 1700–2000 m, 16.VII.2003, leg. C. Gui; 1♂2♀ (MHBU), Shaanxi, Fengxian, Jialing, Jiangyuan, 13.VII.2012, leg. G. D. Ren; 1♂4♀ (MHBU), Shaanxi, Ningshaan, Huoditang, 1505 m, 33.434126°N, 108.448091°E (DD), 15.VIII.2013, leg. X. C. Zhu & Y. Tian.
Male. Aedeagus: basal piece slightly longer than dorsal plate of each paramere (Fig.
Female. Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
China (Shaanxi, Hubei, Sichuan, Gansu).
Athemus genaemaculatus Wittmer, 1951: 100.
Athemus (Isathemus) genaemaculatus:
Lycocerus genaemaculatus:
Holotype
: 1♂ (ZFMK), Kuatun (Fukien), 2300 m, 27.40n, Br. 117.40ö, 20.V.1938, L. J. Klapperich. Paratypes: 1♀ (
China: 1♂ (
Male. Aedeagus: basal piece distinctly longer than dorsal plate of each paramere (Fig.
Female. Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
China (Fujian).
The aedeagus, abdominal sternite VIII, and internal genitalia of female are illustrated for the first time for this species herein.
Athemus (Isathemus) hubeiensis Švihla, 2004: 191, figs 124, 125.
Lycocerus hubeiensis:
Holotype
: 1♂ (
China: 1♂ (MHBU), Shaanxi, Yangxian, Huayanghanba, 1014 m, 33°32'52"N, 107°35'5"E (DMS), 6.V.2017, leg. H. Y. L iu; 1♀ (MHBU), Shaanxi, Yangxian, Huayangzhen, Yantou, 1206 m, 33°38'29"N, 107°31'6.9"E (DMS), 7.VIII.2017, leg. H. Y. Liu & X. D. Zhang; 1♀ (MHBU), Hubei, Chaoshuihe, 23.V. 2019, leg. P. Wang.
Male. Aedeagus: basal piece distinctly longer than dorsal plate of each paramere (Fig.
Female. Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
China (Hubei, Shaanxi).
The aedeagus of this species was illustrated only in ventral and lateral views by
Athemus (Isathemus) kubani Švihla, 2004: 190, figs 122, 123, 207.
Lycocerus kubani:
Holotype
: 1♂ (
China: 1♂, 1♀ (
Male. Aedeagus: basal piece obviously longer than dorsal plate of each paramere (Fig.
Female. Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
China (Shaanxi, Hubei).
The aedeagus of this species was illustrated only in ventral and lateral views by
Athemus (Isathemus) zdeneki Švihla, 2004: 192, figs 126–128.
Lycocerus zdeneki:
Holotype
: 1♂ (
China: 1♂ (
Male. Aedeagus: basal piece slightly longer than dorsal plate of each paramere (Fig.
Female. Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
China (Sichuan).
Only the apical parts of the aedeagus of this species was illustrated by
Athemus (Isathemus) bilineatus Wittmer, 1995: 275, figs 140, 141.
Athemus (s.str.) amplus Wittmer, 1995: 278, figs 146, 147, 203.
Lycocerus amplus:
Lycocerus bilineatus:
See
China: 2♂ , 1♀ (
Male. Aedeagus: basal piece feebly longer than dorsal plate of each paramere (Fig.
Female. Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
China (Jiangsu, Shanghai, Jiangxi, Hubei).
The aedeagus of this species was illustrated only in ventral and lateral views by
Athemus (Andrathemus) jelineki Švihla, 2004: 189, figs 109–111, 204.
Lycocerus jelineki:
Holotype
: 1♂ (
China: 1♂, W Hubei prov. Dashennongjia Nat. Res. Muyu, E slope, 2000 m, 12–15.VI.1997, leg. Bolm; 1♂ (MHBU), Hubei, Shennongjia, Pingqian, 1576 m, 31°28'08.0"N, 110°02'23.4"E (DMS), 4–7.VII.2014, leg. Y.B. Ba & S. Y. Tang; 1♂ (NWAFU), Shaanxi, Taibaishan, Haopingsi, 1200 m, 31.V.1981, collector unknown; 1♂ (NWAFU), Shaanxi, Taibaishan, Zhongshansi, 1500 m, 9.VI.1981, collector unknown; 1♀ (NWAFU), Shaanxi, Taibaishan, Haopingsi, 25.VI.1982, collector unknown; 1♀ (NWAFU), Shaanxi, Taibaishan, Zhongshansi, 400 m, 11.VI.1981, collector unknown.
Male. Aedeagus: basal piece feebly longer than dorsal plate of each paramere (Fig.
Female. Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
China (Shaanxi, Hubei).
Sometimes the pronotum and vertex have dark brown spots, in both sexes. Only the apical parts of the aedeagus were illustrated by
Lycocerus putzi Švihla, 2011: 11, figs 12, 60, 61–63.
China: 1♂1♀(
Male. Aedeagus: basal piece feebly longer than dorsal plate of each paramere (Fig.
Female. Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
China (Sichuan).
The aedeagus of this species was illustrated only for apical parts by
Athemus (Isathemus) pictipennis Wittmer, 1995: 276, figs 142, 143.
Holotype
: 1♂ (
1♂ (MHBU), Myanmar, Prov. Kachin Mt Emaw Bum, 2358 m road of Kanphant, 26°09'N, 98°31'E (DDM), 28.V.2006, leg. L. Langer.
Male. Aedeagus: basal piece as long as dorsal plate of each paramere (Fig.
China (Yunnan); Myanmar (new record).
This species was missing in the Palaearctic Catalogue by
Athemus (Isathemus) curvatus Wittmer, 1995: 260, figs 117, 118, 188.
Lycocerus curvatus:
Holotype
: 1♂ (
China: 1♂ (
Male. Aedeagus: basal piece nearly as long as dorsal plate of each paramere (Fig.
China (Sichuan).
The aedeagus of this species was illustrated only in ventral and lateral views by
Holotype : ♂ (MHBU), China, Guangxi, Wuming, Damingshan, 1230–1423 m, 20.V.2011, leg. H. Y. Liu. Paratypes: China: 4♀1♂ (MHBU), same data as holotype; 2♀ (MHBU), same locality as holotype, 1100 m, 27.V.2011, leg. H. Y. Liu; 1♀ (MHBU), same locality as holotype, 600–900 m, 25.V.2011, leg. H. Y. Liu.
The new species can be easily distinguished from all others by its body coloration, head and pronotum bicolored, with vertex black and clypeus yellow, pronotum with a black wide longitudinal median band (Fig.
Male (Fig.
Head feebly narrowed behind eyes, surface densely and finely punctate; eyes moderately large and protruding, head width across eyes feebly wider than anterior margin of pronotum; antennae filiform, extending to elytral mid-length when reclined, antennomere II shortest, ~ 2.5× longer than wide at apex, III–V feebly widened apically, IV–X each with a short smooth impression near apical part of outer margin, V longest, VI–XI nearly parallel-sided, XI acute at apex.
Pronotum distinctly longer than wide, anterior margin feebly arcuate, lateral margins subparallel, posterior margin nearly straight, anterior angles rounded, posterior angles right-angled, disc convex on posterolateral parts, surface finely and feebly sparsely punctate than that on head.
Elytra ~ 4.5× longer than pronotum, 5.4× longer than width across humeri, outer margins nearly parallel, disc semi-lustrous, coarsely and densely punctate.
Legs slender, all tarsal claws simple.
Aedeagus: basal piece nearly as long as dorsal plate of each paramere (Fig.
Female (Fig.
Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
Body length: 9.3–9.7 mm (9.3 mm in holotype); width: 1.8–2.0 mm (1.8 mm in holotype).
China (Guangxi).
The new species is named after the quite wide laterophyse of its aedeagus.
Holotype : ♂ (MHBU), China, Zhejiang, Lin’an, Qingliangfeng, 16–22.V.2012, leg. G. L. Xie. Paratypes: China: 2♀ (MHBU), same data as holotype; 1♀ (MHBU), Zhejiang, Lin’an, Qingliangfeng, Shunxiwu, 15–18.V.2012, leg. J. S. Xu & L. X. Chang; 1♀ (MHBU), Zhejiang, Lin’an, Qingliangfeng, Longtangshan, 19.V.2011, leg. G. L. Xie.
The new species resembles L. pictipennis in the coloration and characteristics of tarsal claws, but differs from the latter in the aedeagus: basal piece very large, ~ 3× longer than dorsal plate of each paramere in lateral view (Fig.
Also, it is similar to L. kubani in the structure of tarsal claws, genitalia of both sexes and female abdominal sternite VIII, but which are different in each component part in detail, and can be easily distinguished from the latter in the uniformly yellow elytra, while elytra black at apices in L. kubani. The coloration of elytra is stable and a good character to recognize the species in L. pallidulus group.
Male (Fig.
Head feebly narrowed behind eyes, surface densely and finely punctate; eyes moderately large and protruding, head width across eyes distinctly wider than anterior margin of pronotum; antennae filiform, extending to elytral mid-length when reclined, antennomere II shortest, ~ 2× longer than wide at apex, III–IX feebly widened apically, IV–IX each with a short smooth impression near basal part of outer margin (X–XI missing), VII longest.
Pronotum distinctly longer than wide, anterior margin feebly arcuate, lateral margins subparallel, posterior margin nearly straight, anterior angles rounded, posterior angles right-angled, disc convex on posterolateral parts, surface finely and feebly sparsely punctate than that on head.
Elytra ~ 3.5× longer than pronotum, 4.4× longer than width across humeri, outer margins nearly parallel, disc semi-lustrous, coarsely and densely punctate.
Legs slender, fore and middle legs with a digitiform tooth on each anterior and posterior claw, and hind claws simple.
Aedeagus: basal piece obviously longer than dorsal plate of each paramere (Fig.
Female (Fig.
Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
Body length: 11.0–12.0 mm (11.0 mm in holotype); width: 2.3–3.0 mm (2.3 mm in holotype).
China (Zhejiang).
The specific name is derived from the Latin flavus (golden-yellow) and pinna (wing), referring to its yellow elytra.
Holotype
: ♂ (
Although the new species is similar to L. laterophysus sp. nov. in the bicolored head, it is more related to L. zdeneki on basis of the structure of aedeagus. Further, L. maoershanensis sp. nov. can be distinguished from the latter in the following characters: scutellum dark brown; female abdominal sternite VIII (Fig.
Male (Fig.
Head feebly narrowed behind eyes, surface densely and finely punctate; eyes moderately large and protruding, head width across eyes slightly wider than anterior margin of pronotum; antennae filiform, extending to three-fifths of elytra when reclined, antennomere II shortest, ~ 1.5× longer than wide at apex, III–VI feebly widened apically, IV–XI each with a short smooth impression near basal part of outer margin, VII–XI nearly parallel-sided, VIII longest.
Pronotum subquadrate, slightly longer than wide, anterior margin feebly arcuate, lateral margins subparallel, posterior margin nearly straight, anterior angles rounded, posterior angles obtuse-angled, disc convex on posterolateral parts, surface finely and feebly sparsely punctate than that on head.
Elytra ~ 3.75× longer than pronotum, 5.0× longer than width across humeri, outer margins nearly parallel, disc semi-lustrous, coarsely and densely punctate.
Legs slender, fore and middle legs with a digitiform tooth on each anterior claw, and hind claws simple.
Aedeagus: basal piece slightly longer than dorsal plate of each paramere (Fig.
Female (Fig.
Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
Body length: 9.0–10.0 mm (9.1 mm in holotype); width: 1.7–2.3 mm (1.8 mm in holotype).
China (Guangxi).
The specific name is derived from the name of the type locality, Maoershan, Guangxi, China.
Holotype : ♂ (MHBU), China, Yunnan, Pingbian, Daweishan, 1900 m, 24.V.1996, leg. L. Y. Zheng.
The new species seems similar to L. putzi in the coloration, but differs in the following characters: fore and mid legs with a digitiform tooth on each anterior claw in male, while on both anterior and posterior claws in the latter; aedeagus with dorsal plate of each paramere wide (Fig.
Male (Fig.
Head feebly narrowed behind eyes, surface densely and finely punctate; eyes moderately large and protruding, head width across eyes slightly wider than anterior margin of pronotum; antennae filiform, extending to apical third length of elytra when reclined, antennomeres II shortest, ~ 2× longer than wide at apices, IV–XI nearly parallel-sided, each with a short smooth impression near apical part of outer margin, IV longest.
Pronotum subquadrate, feebly longer than wide, anterior margin feebly arcuate, lateral margins subparallel, posterior margin nearly straight, anterior angles obtuse-rounded, posterior angles nearly right-angled, disc convex on posterolateral parts, surface finely and feebly sparsely punctate than that on head.
Elytra ~ 4.4× longer than pronotum, 3.12× longer than width across humeri, outer margins nearly parallel, disc semi-lustrous, coarsely and densely punctate.
Legs slender, fore and middle legs with a digitiform tooth on each anterior claw, and hind claws simple.
Aedeagus: basal piece slightly longer than dorsal plate of each paramere (Fig.
Female. Unknown.
Body length: 10.2 mm; width: 2.2 mm.
China (Yunnan).
The specific name is derived from the Latin mimus (imitator), referring to its similarity to L. putzi Švihla, 2011.
Holotype : ♂ (MHBU), China, Chongqing, Nanchuan, Jinfoshan, 23–24.VII.2003, leg. Y. S. Liu & C. X. Yuan. Paratypes: 1♂2♀ (MHBU), same data as holotype.
The new species seems most similar to L. centrochinensis in both coloration and characteristics of tarsal claws, but differs in the following characters: body size is smaller; aedeagus: dorsal plates of parameres with inner margins feebly protuberant near base (Fig.
Male (Fig.
Head feebly narrowed behind eyes, surface densely and finely punctate; eyes moderately large and protruding, head width across eyes wider than anterior margin of pronotum; antennae filiform, extending to apical third of elytra when reclined, antennomere II shortest, ~ 2× longer than wide at apex, III–IV feebly expanded at apex, IV–X each with a short smooth impression near apical part of outer margin, V–XI nearly parallel-sided, VIII longest.
Pronotum subquadrate, feebly longer than wide, anterior margin feebly arcuate, lateral margins subparallel, posterior margin nearly straight, anterior angles obtuse-rounded, posterior angles nearly right-angled, disc convex on posterolateral parts, surface finely and feebly sparsely punctate than that on head.
Elytra ~ 4.6× longer than pronotum, 3.3× longer than width across humeri, outer margins nearly parallel, disc semi-lustrous, coarsely and densely punctate.
Legs slender, all claws simple.
Aedeagus: basal piece nearly as long as dorsal plate of each paramere (Fig.
Female (Fig.
Internal organ of reproductive system (Fig.
Abdominal sternite VIII (Fig.
Body length: 7.8–9.0 mm (8.0 mm in holotype); width: 1.7–2.1 mm (1.8 mm in holotype).
China (Chongqing).
The specific name is derived from the type locality, Chongqing, China.
Lycocerus centrochinensis
(Švihla, 2004):
Holotype : ♂ (MHBU), China, Ningxia, Kongtongshan, 6.VI.1992, leg. J. L. Ding. Paratypes: China: 1♀ (MHBU), Ningxia, Jingyuan, Liupanshan, 13.VI.1995, Collectors Group III of Forestry; 1♀ (MHBU), same data as the preceding, 8.VI.1995, Collectors Group III of Forestry; 1♀ (MHBU), same data as the preceding, 17.VI.1995, Collectors Group III of Forestry.
The new species seems similar to L. hubeiensis in the coloration, but differs in the following characters: tarsal claws simple in males, while fore and mid-anterior and posterior claws each with a digitiform tooth at base in the latter; aedeagus: dorsal plates of parameres triangular at apices (Fig.
Male (Fig.
Head feebly narrowed behind eyes, surface densely and finely punctate; eyes moderately large and protruding, head width across eyes nearly wider than anterior margin of pronotum; antennae filiform, extending to apical third length of elytra when reclined, antennomere II shortest, ~ 2× longer than wide at apices, IV–XI nearly parallel-sided, each with a short smooth impression near apical part of outer margin, IV longest.
Pronotum subquadrate, slightly longer than wide, anterior margin feebly arcuate, lateral margins subparallel, posterior margin nearly straight, anterior angles obtuse-rounded, posterior angles nearly right-angled, disc convex on posterolateral parts, surface finely and feebly sparsely punctate than that on head.
Elytra ~ 4.4× longer than pronotum, 3.12× longer than width across humeri, outer margins nearly parallel, disc semi-lustrous, coarsely and densely punctate.
Legs slender, all claws simple.
Aedeagus: basal piece obviously longer than dorsal plate of each paramere (Fig.
Female (Fig.
Internal organ of reproductive system (Fig.
Abdominal sterite VIII of female, ventral view A Lycocerus hubeiensis (Švihla, 2004) B L. centrochinensis (Švihla, 2004) C L. guerryi (Pic, 1906) D L. genaemaculatus (Wittmer, 1951) E L. flavipennis sp. nov. F L. zdeneki (Švihla, 2004) G L. kubani (Švihla, 2004) H L. laterophysus sp. nov. Scale bars: 0.5 mm.
Abdominal sternite VIII (Fig.
Body length: 9.0–10.0 mm (9.3 mm in holotype); width: 2.0–2.3 mm (2.0 mm in holotype).
China (Ningxia).
The specific name is derived from the Latin spermatike (sperm-carrying), referring to its distinctive spermatheca, which has two spiral tubes.
The characters of tarsal claws, which was emphasized by
| Species | Male fore and mid-legs | Female fore and mid-legs | ||
|---|---|---|---|---|
| anterior claws | posterior claws | anterior claws | posterior claws | |
| Lycocerus pallidulus | ||||
| L. guerryi | ||||
| L. guerryi atroapicipennis | ||||
| L. centrochinensis | ||||
| L. genaemaculatus | ||||
| L. hubeiensis | ||||
| L. jelineki | ||||
| L. bilineatus | ||||
| L. zdeneki | ||||
| L. kubani | ||||
| L. curvatus | ||||
| L. pictipennis | ||||
| L. putzi | ||||
| L. laterophysus sp. nov. | ||||
| L. flavipennis sp. nov. | ||||
| L. putzimimus sp. nov. | Unknown | Unknown | ||
| L. maoershanensis sp. nov. | ||||
| L. chongqingensis sp. nov. | ||||
| L. bispermathecus sp. nov. | ||||
At the beginning,
All species of L. pallidulus group are distributed in the southern China, located between 21.94–36.60°N and 98.31–21.60°E (Fig.
Originally, L. pallidulus group was regarded as a subgroup and placed in L. maculicollis group (
Within the L. pallidulus group, the species can be distinguished by their claws (Table
| 1 | Elytra bicolored, mixed black with light yellow | 2 |
| – | Elytra uniformly light yellow | 9 |
| 2 | Elytra with black median longitudinal bands | 3 |
| – | Elytra with black apices | 4 |
| 3 | Pronotum uniformly orange; aedeagus: ventral process of each paramere stout and nearly truncated apically in lateral view (Fig. |
L. bilineatus (Wittmer, 1995) |
| – | Pronotum yellow, dark brown in middle; aedeagus: ventral process of each paramere slender and nearly rounded apically in lateral view (Fig. |
L. curvatus (Wittmer, 1995) |
| 4 | Elytra black at apical 2/3, with a long triangular area pale yellow along suture | 5 |
| – | Elytra black at most at apical 1/5 | 6 |
| 5 | Pronotum with a small dark brown rounded marking; aedeagus: ventral process of each paramere moderately shorter than dorsal plate, laterophyse furcate at apex (Fig. |
L. putzimimus sp. nov. |
| – | Pronotum uniformly yellow; aedeagus: ventral process of each paramere much shorter than dorsal plate, laterophyse acute at apex (Fig. |
L. putzi Švihla, 2011 |
| 6 | All claws simple in male; spermatheca with 2 spiral tubes (Fig. |
L. bispermathecus sp. nov. |
| – | Fore and mid- anterior and or posterior claws legs each with a digitiform tooth at base in male; spermatheca with 1 spiral tube | 7 |
| 7 | Fore and mid- anterior and posterior claws legs each with a digitiform tooth at base in male; aedeagus: ventral process of each paramere nearly straight in ventral view (Figs |
8 |
| – | Fore and mid-anterior claws each with a digitiform tooth at base in male; aedeagus: ventral process of each paramere distinctly bent inwards in ventral view (Fig. |
L. guerryi atroapicipennis (Pic, 1914) |
| 8 | Aedeagus: ventral process of each paramere obviously longer than laterophyse (Fig. |
L. hubeiensis (Švihla, 2004) |
| – | Aedeagus: ventral process of each paramere nearly as long as laterophyse (Fig. |
L. kubani (Švihla, 2004) |
| 9 | Head bicolored, mixed yellow or orange with black | 10 |
| – | Head uniformly yellow or orange | 12 |
| 10 | Vertex yellow, each side with a black marking around eye; aedeagus: the distance between ventral processes distinctly wider than that between lateral margins of dorsal plates (Fig. |
L. genaemaculatus (Wittmer, 1951) |
| – | Vertex unlike above, never with markings around eyes; aedeagus: the distance between ventral processes narrower than that between lateral margins of dorsal plates (Figs |
11 |
| 11 | Head and pronotum yellow, vertex with a triangular dark brown marking in middle, pronotum with two irregular black markings near middle of anterior and posterior margins (Fig. |
L. maoershanensis sp. nov. |
| – | Vertex black, clypeus yellow, pronotum yellow, with a black wide median longitudinal band (Fig. |
L. laterophysus sp. nov. |
| 12 | All claws simple in male | 13 |
| – | Fore and mid-anterior and posterior claws each with a digitiform tooth at base in male | 14 |
| 13 | Body larger, 8.0–10.0 mm in length; aedeagus: ventral process of each paramere nearly vertical in lateral view (Fig. |
L. centrochinensis (Švihla, 2004) |
| – | Body smaller, 7.8–9.0 mm in length; aedeagus: ventral process of each paramere slightly bent ventrally in lateral view (Fig. |
L. chongqingensis sp. nov. |
| 14 | Fore and mid-anterior and posterior claws with a digitiform tooth at base in male | 15 |
| – | Fore and mid-anterior claws with a digitiform tooth at base in male | 16 |
| 15 | Aedeagus: basal piece very large, ~ 3× longer than dorsal plate of each paramere in lateral view (Fig. |
L. flavipennis sp. nov. |
| – | Aedeagus: basal piece nearly as long as dorsal plate of each paramere in lateral view (Fig. |
L. pictipennis (Wittmer, 1995) |
| 16 | Fore and mid-anterior and posterior claws each with a digitiform tooth at base in female | 17 |
| – | Fore and mid-anterior claws each with a digitiform tooth at base in female | L. jelineki (Švihla, 2004) |
| 17 | aedeagus: ventral process of each paramere obviously shorter than laterophyse in ventral view (Fig. |
L. zdeneki (Švihla, 2004) |
| – | aedeagus: ventral process of each paramere obviously longer than laterophyse in ventral view (Fig. |
18 |
| 18 | Female abdominal sternite VIII with the portions between middle and lateral emarginations wide and rounded at apices (Okushima, 2005: fig. 22d); aedeagus: ventral process of each paramere slightly bent inwards in ventral view (Okushima, 2005: fig. 22a), dorsal plate of each paramere narrow and obviously separate apically (Okushima, 2005: fig. 22c) | L. pallidulus (Wittmer, 1995) |
| – | Female abdominal sternite VIII with the portions between middle and lateral emarginations narrower and right-angled at apices (Fig. |
L. guerryi (Pic, 1906) |
The Lycocerus pallidulus subgroup originally placed in L. maculicollis group is suggested as an independent species-group herein. This group is mainly distributed in the southern China and easily recognized by the middle-sized body and pale yellow or even transparent elytra, sometimes with black longitudinal bands or markings, as well as the genitalia of both sexes. In total 19 species are currently attributed to this group, including, L. laterophysus sp. nov., L. flavipennis sp. nov., L. putzimimus sp. nov., L. maoershanensis sp. nov., L. chongqingensis sp. nov. and L. bispermathecus sp. nov., discovered from China. These species can be distinguished from one another by the body coloration, structures of tarsal claws in both sexes, shapes of aedeagus and abdominal sternite VIII of female. The results of this study provide a better understanding about the morphological and specific diversities of Lycocerus, to improve the classification of this speciose genus.
We are grateful to late Dr. Michel Brancucci (
The authors have declared that no competing interests exist.
No ethical statement was reported.
The present study was supported by the National Natural Science Foundation of China (Nos. 32270491, 31772507), the Natural Science Foundation of Hebei Province (No. C2022201005), the Interdisciplinary Research Program of Natural Science of Hebei University (No. DXK202103), and the Youth Scientific Research and Innovation Team of Hebei University (No. 605020521005).
Conceptualization: HL, YW. Data curation: YW. Formal analysis: YW. Funding acquisition: HL, YY. Investigation: YY. Methodology: YW. Project administration: YY. Resources: XY. Software: YW. Supervision: YY, HL, XY. Visualization: YW. Writing - original draft: YW. Writing - review and editing: YY.
Haoyu Liu https://orcid.org/0000-0003-1383-5560
Xingke Yang https://orcid.org/0000-0003-3676-6828
Yuxia Yang https://orcid.org/0000-0002-3118-6659
All of the data that support the findings of this study are available in the main text.