﻿Review of the Lycoceruspallidulus group (Coleoptera, Cantharidae), with descriptions of six new species from China

﻿Abstract The Lycoceruspallidulus 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.laterophysussp. nov., L.flavipennissp. nov., L.putzimimussp. nov., L.maoershanensissp. nov., L.chongqingensissp. nov., and L.bispermathecussp. 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.

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. 1): dorsal plates of parameres separate, each with a keel near lateral margin and located on inner surface, laterophyses well-developed and nearly as long as ventral processes, with apices opposite to the keels of dorsal plates, inner sac of median lobe lengthened apically and nearly as long as tegmen, without dorsal process.Female internal genitalia: vagina stout and abruptly thinned at ventroapical portion into a stout tube, where diverticulum and spermathecal duct arising separately; diverticulum moderately long, thin, and spiral; spermathecal duct short and stout; spermatheca with a spiral tube, often abruptly thinned apically near base, basal portion of spermatheca extended into a short tube, where accessory gland opening, accessory gland longer than spermatheca.

Lycocerus pallidulus
Remarks.This species was omitted from the Palaearctic Catalogue by Kazantsev and Brancucci (2007).Both the aedeagus and female internal genitalia of this species have been well illustrated by Okushima (2005).
Abdominal sternite VIII (Fig. 16C): triangular emarginations on both sides and in middle of posterior margin, lateral emarginations slightly deeper and much wider than the middle one, the portions between lateral and middle emarginations wide and obtuse-triangular at apices, slightly extending over apices of latero-apical angles, which are nearly truncated.

Lycocerus guerryi atroapicipennis
Remarks.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.(Švihla, 2004)
Abdominal sternite VIII (Fig. 16B): rounded emargination in middle and triangular emarginations on both sides of posterior margin, lateral emarginations obviously deeper than and nearly as wide as the middle one, the portions between lateral and middle emarginations moderately wide and right-angled at apices, obviously extending over apices of latero-apical angles, which are narrowly triangular.Distribution.China (Shaanxi, Hubei, Sichuan, Gansu).Remarks.Li et al. (2015) provided an illustration of female internal genitalia for L. centrochinensis, which is of an unknown species.Additionally, the distribution range of this species is expanded, with geographic records added from Sichuan and Gansu provinces.(Wittmer, 1951 Descriptive notes.Male.Aedeagus: basal piece distinctly longer than dorsal plate of each paramere (Fig. 4A-C); ventral process of each paramere short and thin, nearly straight and approaching to each other in ventral view (Fig. 4A), slightly bent dorsally in lateral view (Fig. 4C); dorsal plates of parameres as long as ventral process (Fig. 4B, C), with outer margins abruptly converging apically, apical margins subrounded in dorsal view (Fig. 4B); laterophyse feebly shorter than ventral process, with apices acute and markedly directing dorso-outwards (Fig. 4C).
Abdominal sternite VIII (Fig. 16D): triangular emargination in middle and subrounded emarginations on both sides of posterior margin, lateral emarginations slightly deeper and obviously wider than the middle one, the portions between lateral and middle emarginations narrow and acute at apices, obviously extending over apices of latero-apical angles, which are widely triangular.
Remarks.The aedeagus, abdominal sternite VIII, and internal genitalia of female are illustrated for the first time for this species herein.(Švihla, 2004)
Abdominal sternite VIII (Fig. 16A): triangular emargination in middle and narrowly rounded emarginations on both sides of posterior margin, lateral emarginations obviously deeper and feebly wider than the middle one, the portions between lateral and middle emarginations moderately wide and right-angled at apices, obviously extending over apices of latero-apical angles, which are narrowly rounded.
Remarks.The aedeagus of this species was illustrated only in ventral and lateral views by Švihla (2004).Here, the aedeagus is illustrated in ventral, dorsal, and lateral views, and the abdominal sternite VIII and internal genitalia of the female are illustrated for the first time.Additionally, the distribution range of this species is expanded, with geographic records added from Shaanxi Province.
Abdominal sternite VIII (Fig. 16G): triangular emargination in middle and rounded emarginations on both sides of posterior margin, lateral emarginations obviously deeper and wider than the middle one, the portion between lateral and middle emarginations narrow and acute at apices, obviously extending over apices of latero-apical angles, which are rounded.
Remarks.The aedeagus of this species was illustrated only in ventral and lateral views by Švihla (2004).Here, the aedeagus is illustrated in ventral, dorsal, and lateral views, and female internal genitalia is illustrated for the first time.Additionally, the distribution range of this species is expanded, with geographic records added from Hubei province.
Descriptive notes.Male.Aedeagus: basal piece slightly longer than dorsal plate of each paramere (Fig. 5D-F); ventral process of each paramere slender and slightly expanded at apices, nearly straightly and approaching to each other in ventral view (Fig. 5D), inclining ventrally in lateral view (Fig. 5F); dorsal plates of parameres feebly longer than ventral process (Fig. 5D, F), with inner margins feebly emarginate at inner apical angles, outer margins slightly converging apically in dorsal view (Fig. 5E); laterophyse slightly longer than ventral process, with apices acute and appreciably directing dorso-outwards (Fig. 5F).
Abdominal sternite VIII (Fig. 16F): right-triangular emargination in middle and subrounded emarginations on both sides of posterior margin, lateral emarginations feebly deeper and wider than the middle one, the portions between lateral and middle emarginations wide and wide-triangular at apices, feebly extending over apices of latero-apical angles, which are truncated.
Remarks.Only the apical parts of the aedeagus of this species was illustrated by Švihla (2004).In this work, the aedeagus is illustrated in general views, and the abdominal sternite VIII and internal genitalia of the female are illustrated for the first time.Type material examined.See Yang et al. (2013).
Female.Internal organ of reproductive system (Fig. 13A): spermatheca feebly longer than diverticulum; accessory gland nearly as long as spermatheca.
Abdominal sternite VIII (Fig. 17A): triangular emarginations in middle and on both sides of posterior margin, lateral emarginations obviously deeper and wider than the middle one, the portion between lateral and middle emarginations wide and right-angled at apices, obviously extending over apices of latero-apical angles, which are narrowly rounded.
Remarks.The aedeagus of this species was illustrated only in ventral and lateral views by Wittmer (1995).Herein, the aedeagus is illustrated in ventral, dorsal, and lateral views, and the female internal genitalia is illustrated for the first time.Descriptive notes.Male.Aedeagus: basal piece feebly longer than dorsal plate of each paramere (Fig. 6D-F); ventral process of each paramere nearly slender and approaching to each other in ventral view (Fig. 6D), nearly vertical in lateral view (Fig. 6F); dorsal plates feebly longer than ventral process (Fig. 6D, F), with inner margins abruptly diverging near middle, outer margins slightly converging apically, apical margins subrounded (Fig. 6E); laterophyse slightly shorter than ventral process, with apices acute and appreciably directing dorso-outwards (Fig. 6D, F).
Abdominal sternite VIII (Fig. 17B): rounded emargination in middle and subtriangular emargination on both sides of posterior margin, lateral emarginations deeper and feebly narrower than the middle one, the portion between lateral and middle emarginations wide and widely triangular at apices, extending over apices of latero-apical angles, which are subrounded.
Remarks.Sometimes the pronotum and vertex have dark brown spots, in both sexes.Only the apical parts of the aedeagus were illustrated by Švihla (2004).Here, the aedeagus is illustrated in general views, and the female internal genitalia is illustrated for the first time.Descriptive notes.Male.Aedeagus: basal piece feebly longer than dorsal plate of each paramere (Fig. 7A-C); ventral process of each paramere nearly straight and approaching to each other in ventral view (Fig. 7A), nearly vertical in lateral view (Fig. 7C); dorsal plates of parameres obviously longer than ventral processes (Fig. 7A, C), outer margins obviously converging apically, apical margins rounded in dorsal view (Fig. 7B); laterophyse feebly longer than ventral process, with apices acute and appreciably directing dorso-outwards (Fig. 7A, C).
Abdominal sternite VIII (Fig. 17D): triangular emargination in middle and subrounded emarginations on both sides of posterior margin, lateral emarginations feebly deeper and obviously wider than the middle one, the portion between lateral and middle emarginations narrow and acute at apices, obviously extending over apices of latero-apical angles, which are rounded.
Remarks.The aedeagus of this species was illustrated only for apical parts by Švihla (2011).Here, the aedeagus is illustrated with general views, and the female internal genitalia is illustrated for the first time.Descriptive notes.Male.Aedeagus: basal piece as long as dorsal plate of each paramere (Fig. 7D-F); ventral process of each paramere thin and bent inwards in ventral view (Fig. 7D), inclining ventrally in lateral view (Fig. 7F); dorsal plates of parameres obviously longer than ventral processes (Fig. 7D,  F), with apical margins rounded in lateral view (Fig. 7E); laterophyse slightly shorter than ventral process, with apices acute and directing dorso-outwards (Fig. 7D, F).
Remarks.This species was missing in the Palaearctic Catalogue by Kazantsev and Brancucci (2007).It is recorded to Myanmar for the first time herein.The aedeagus of this species was illustrated only in ventral and lateral views by Wittmer (1995).Here, the aedeagus is illustrated in ventral, dorsal, and lateral views, and the abdominal sternite VIII and internal genitalia of female are illustrated for the first time.Descriptive notes.Male.Aedeagus: basal piece nearly as long as dorsal plate of each paramere (Fig. 11A-C); ventral process of each paramere slender and bent inwards apically in ventral view (Fig. 11A), inclining ventrally in lateral view (Fig. 11C); dorsal plates of parameres feebly longer than ventral processes (Fig. 11A, C), with apical margins rounded in dorsal view (Fig. 11B); laterophyse slightly shorter than ventral process, with apices acute and appreciably directing dorso-outwards (Fig. 11A, C).
Remarks.The aedeagus of this species was illustrated only in ventral and lateral views by Wittmer (1995).Here, the aedeagus is illustrated in ventral, dorsal, and lateral views, and the female internal genitalia is illustrated for the first time.Diagnosis.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. 18A, B).Also, its aedeagus is unique and differs from all others in the ventral process of each paramere expanded near base in lateral view (Fig. 8C), and broad laterophyse in ventral view (Fig. 8A).
Description.Male (Fig. 18A).Head, prothorax and legs orange, vertex black, mandibles dark brown, antennomeres III-XI black, pronotum with a wide black longitudinal median band extending from posterior to anterior margins, scutellum black, elytra pale yellow, tarsi darkened, meso-and metasterna and abdomen black.Body densely covered with yellow recumbent pubescence.
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.
Legs slender, all tarsal claws simple.Aedeagus: basal piece nearly as long as dorsal plate of each paramere (Fig. 8A-C); ventral process of each paramere slender, feebly bent inwards and approaching each other in ventral view (Fig. 8A), slightly expanded near base and bent ventrally in lateral view (Fig. 8C); dorsal plates of parameres obviously longer than ventral process (Fig. 8A, C), with apical margins arcuate and descending inwards in dorsal view (Fig. 8B); laterophyse nearly as long as ventral process and broad in ventral view, with apices acute and directing dorso-outwards (Fig. 8A, C).
Female (Fig. 18B).Similar to the males, but eyes less protruding, antennae thinner and shorter, extending to basal one-third length of elytra when reclined, middle antennomeres without impressions, fore and middle legs with a digitiform tooth on each anterior claw, pronotum feebly longer than wide.
Abdominal sternite VIII (Fig. 16H): hardly emarginate in middle and subtriangular emarginations on both sides of posterior margin, lateral emarginations obviously deeper than the middle one, the portion between lateral and middle emarginations wide and rounded at apices, obviously extending over apices of latero-apical angles, which are subrounded.
Etymology.The new species is named after the quite wide laterophyse of its aedeagus.Diagnosis.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. 8F), dorsal plate narrow (Fig. 8E), laterophyse slightly bent inwards in ventral view (Fig. 8D).In comparison, in the aedeagus of L. pictipennis, basal piece is nearly as long as dorsal plate of each paramere in lateral view (Fig. 7F), dorsal plate wide (Fig. 7E), laterophyse obviously bent outwards in ventral view (Fig. 7D).

Lycocerus flavipennis
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.
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.
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. 8D-F); ventral process of each paramere thin and short, feebly bent inwards and approaching to each other in ventral view (Fig. 8D), nearly straight in lateral view (Fig. 8F); dorsal plates of parameres obviously longer than ventral processes (Fig. 8D, F), with inner margins parallel, outer margins abruptly converging apically in the middle, apical margins rounded (Fig. 8E); laterophyse feebly shorter than ventral process, bent dorsally, with apices acute and directing dorso-inwards in ventral view (Fig. 8D, F).
Female (Fig. 18D).Similar to the males, but eyes less protruding, antennae thinner and shorter, extending to basal one-third length of elytra when reclined, middle antennomeres without impressions, pronotum nearly as long as wide.
Abdominal sternite VIII (Fig. 16E): triangular emargination in middle and rounded emarginations on both sides of posterior margin, lateral emarginations wider and feebly deeper than the middle one, the portions between lateral and middle emarginations narrow and acute at apices, obviously extending over apices of latero-apical angles, which are widely triangular.
Etymology.The specific name is derived from the Latin flavus (golden-yellow) and pinna (wing), referring to its yellow elytra.Diagnosis.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. 17C) with the portion between lateral and middle emarginations rounded at apices, which obviously extending over latero-apical angles.In comparison, L. zdeneki has yellow scutellum; female abdominal sternite VIII (Fig. 16F) with the portion between lateral and middle emarginations feebly extending over latero-apical angles.

Lycocerus maoershanensis
Description.Male (Fig. 19A).Head, prothorax and legs yellowish orange, vertex with a small triangular dark brown marking, mandibles dark brown at apices, antennomeres III-XI black, pronotum with two dark brown irregular markings near middle of anterior and posterior margins, scutellum black, elytra pale yellow and almost transparent, legs more or less darkened at tarsi, apices of tibiae and femora, meso-and metasterna and abdomen black.Body densely covered with yellow recumbent pubescence.
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.
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. 9A-C); ventral process of each paramere slender, feebly bent inwards and approaching to each other in ventral view (Fig. 9A), inclining ventrally in lateral view (Fig. 9C); dorsal plates of parameres slightly longer than ventral processes (Fig. 9A, C), with inner margins emarginate at apical parts, outer margins slightly converging apically, apical margins slightly arcuate and descending inwards, inner apical angle acute angled and outer angle rounded in dorsal view (Fig. 9B); laterophyse feebly longer than ventral process, with apices acute and directing dorso-inwards in ventral view (Fig. 9A).
Female (Fig. 19B).Similar to the males, but eyes less protruding, antennae shorter and extending to basal third length of elytra when reclined, middle antennomeres without impressions, pronotum nearly as long as wide, fore and middle legs with a digitiform tooth on each anterior and posterior claw.
Internal organ of reproductive system (Fig. 13C): spermatheca nearly as long as diverticulum; accessory gland nearly as long as spermatheca.
Abdominal sternite VIII (Fig. 17C): subrounded emarginations in middle and on both sides of posterior margin, lateral emarginations deeper than and as wide as the middle one, the portions between lateral and middle emarginations moderately wide and rounded at apices, obviously extending over apices of latero-apical angles, which are narrowly triangular.
Etymology.The specific name is derived from the name of the type locality, Maoershan, Guangxi, China.Diagnosis.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. 9E) and moderately longer than ventral process (Fig. 9F), while narrower (Fig. 7B) and distinctly long (Fig. 7C) in L. putzi; laterophyse feebly furcate at apices (Fig. 9F), while acute in the latter (Fig. 7C).

Lycocerus putzimimus
Description.Male (Fig. 19C).Head, prothorax and legs orange, mandibles dark brown at apices, antennae black, except for bases of antennomeres I yellow, pronotum with a small and round black marking in center of disc, scutellum yellow, elytra pale yellow and almost transparent, each with a black longitudinal band along apical two-thirds part, leaving a long triangular area pale yellow along suture, legs darkened at tarsi, meso-and metasterna and abdomen black, terminal two abdominal ventrites and sternites pale yellow.Body densely covered with yellow recumbent pubescence.
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.
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. 9D-F); ventral process of each paramere slender, slightly expanded at apices, slightly bent inwards and approaching to each other in ventral view (Fig. 9D), inclining ventrally in lateral view, with apices slightly curved dorsally (Fig. 9F); dorsal plate obviously longer than ventral process of each paramere, with apical margins rounded (Fig. 9E); laterophyse feebly longer than ventral process, with apices bifurcate and directing dorso-outwards (Fig. 9D, F).
Etymology.The specific name is derived from the Latin mimus (imitator), referring to its similarity to L. putzi Švihla, 2011.Diagnosis.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. 10B); abdominal sternite VIII (Fig. 17E) moderately narrowed posteriorly, the portion between lateral and middle emarginations rounded at apices, which slightly extending over apices of latero-apical angles.In comparison, L. centrochinensis has larger body; aedeagus: dorsal plates of parameres with inner margins abruptly diverging near middle (Fig. 3E); abdominal sternite VIII (Fig. 16B) strongly narrowed posteriorly, the portion between lateral and middle emarginations triangular at apices, which distinctly extending over apices of latero-apical angles.
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.
Legs slender, all claws simple.Aedeagus: basal piece nearly as long as dorsal plate of each paramere (Fig. 10A-C); ventral process of each paramere slender and feebly bent inwards and approaching to each other in ventral view (Fig. 10A), inclining ventrally in lateral view (Fig. 10C); dorsal plates of parameres obviously longer than ventral processes (Fig. 10A, C), with inner margins nearly parallel, feebly triangularly protuberant near base, outer margins slightly converging apically, apical margins slightly descending outwards, inner and outer apical angles rounded in dorsal view (Fig. 10B); laterophyse nearly as long as ventral process, with apices acute and directing dorso-outwards (Fig. 10A, C).
Female (Fig. 20B).Similar to the males, but eyes less protruding, antennae shorter, not reaching elytral mid-length when reclined, middle antennomeres without impressions, pronotum nearly as long as wide, fore and middle legs with a digitiform tooth on each anterior and posterior claw.
Abdominal sternite VIII (Fig. 17E): hardly emarginate in middle and subrounded emarginations on both sides of posterior margin, lateral emarginations obviously deeper than the middle one, the portion between lateral and middle emarginations wide and rounded at apices, slightly extending over apices of latero-apical angles, which are truncated.Lycocerus centrochinensis (Švihla, 2004) Diagnosis.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. 11E), while truncated in the latter (Fig. 4E); spermatheca with two spiral tubes (Fig. 15B), while only one in the latter.
Female (Fig. 20D).Similar to the males, but eyes less protruding, antennae shorter and extending to elytral mid-length when reclined, middle antennomeres without impressions, pronotum nearly as long as wide, fore and middle legs with a digitiform tooth on each anterior and posterior claw.Internal organ of reproductive system (Fig. 15B): spermathecal duct stout, spermatheca with two spiral tubes, which are subequal in length, both of them shorter than diverticulum; accessory gland ~ 2.5× longer than spermatheca.
Abdominal sternite VIII (Fig. 17F): triangular emarginations in middle and on both sides of posterior margin, lateral emarginations feebly deeper than the middle one, the portion between lateral and middle emarginations narrow and acute at apices, obviously extending over apices of latero-apical angles, which are widely triangular.
Etymology.The specific name is derived from the Latin spermatike (sperm-carrying), referring to its distinctive spermatheca, which has two spiral tubes.

Discussion
The characters of tarsal claws, which was emphasized by Wittmer (1995) to define the subgenera of former Athemus Lewis, 1895 (now a junior synonym of Lycocerus), is proven again to be variable even within a species group (Table 1), as noted by Okushima and Hsiao (2017).Since no character was considered valuable enough to define the subgenera, Okushima (2005) proposed to define species groups to subdivide the large genus Lycocerus sensu lato.
At the beginning, Okushima (2005) defined the species groups of Lycocerus based on the genitalia of both sexes.Especially, he highlighted the characters of the female internal organ of reproductive system, including the length of spermathecal duct, and shape, length, and number of tubes of spermatheca.However, in the present study, we discovered that the number of tubes of the spermatheca could be variable within the species group.In the L. pallidulus group, the spermatheca of L. bispermathecus sp.nov.has two spiral tubes, while all others have only one tube.Also, it is related to L. centrochinensis and L. chongqingensis sp.nov. on basis of the shape of aedeagus and tarsal claws, as well as the body size and coloration, so they probably belong to a natural species group.In this case, we suggest integrating the characters of both appearance and genitalia to define the species groups of Lycocerus, also we should take the distribution range into account.
All species of L. pallidulus group are distributed in the southern China, located between 21. .China is located in East Asia and lies in the transitional zone between Palaearctic and Oriental Regions (Zhang 1999).It is a region where some cantharid lineages occur only there and adjacent areas, like other insects (e.g., Bocak and Bocakova 2008).Many lineages with the highest diversity in the Chinese fauna would expand their ranges southwards to the Oriental Region, and L. pallidulus group is this case.
Originally, L. pallidulus group was regarded as a subgroup and placed in L. maculicollis group (Okushima 2005).Based on our studies and Wang et al. (2023), we found that L. pallidulus group definitely differs from L. maculicollis group in the shape of aedeagus, which is unique in the latter group noted previously by Kazantsev (1999).In addition, the former L. maculicollis group has already been suggested as a non-monophyletic (Hsiao 2021;Xi et al. 2022;Wang et al. 2022).Thus, we suggest L. pallidulus group be treated as an independent species group.
Within the L. pallidulus group, the species can be distinguished by their claws (Table 1) and the following key is designed around this feature combined with other morphological characters.

Figure 1 .
Figure 1.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.