A Taxonomic Review of the Genus Telsimia Casey (Coleoptera, Coccinellidae) from China, with Descriptions of Eight New Species

Simple Summary The Telsimia species are known as important predators of diaspidine scale insects that infest citrus, coconut, camphor, and bamboo plants, among others. A clear understanding of Telsimia species diversity and their geographical distribution forms the basis for the introduction and utilization of these natural enemies. At present, 48 species (subspecies) are known around the world, and studies on Chinese Telsimia species are relatively scarce. In this paper, 20 species of the genus Telsimia from China are reviewed, including eight new species and two new records. This taxonomic study can provide a theoretical basis for the development and utilization of Telsimia species for biological control of diaspidine scale insects. Abstract Twenty species of the genus Telsimia from China are reviewed herein. Among them, eight species are described as new to science: Telsimia chayuensis, T. forcipata, T. latus, T. lobatus, T. lunata, T. menglaensis, T. parascymnoides, and T. parvus spp. nov.; two species are reported from China for the first time: T. darjeelingensis Kapur, 1969 and T. elongate Hoàng, 1985. All species are provided with nomenclatural history, diagnoses, detailed descriptions (except for the 10 previously described species), colored illustrations, and distributions. The female genitalia of five described species are provided for the first time. A distribution map and a key to all the Chinese species are also provided.


Introduction
The genus Telsimia Casey, 1899 [1] belongs to the tribe Telsimiini of the subfamily Coccinellinae [2]. It is known in Africa and many parts of Asia and is widespread in the Pacific Islands, New Guinea, and Australia [3]. The Telsimia species are known as important predators of diaspidine scale insects that infest citrus, coconut, camphor, and bamboo plants, among others [4][5][6][7][8]. For instance, certain Telsimia species were introduced from their original places into Micronesia, Hawaii, and California, and efficient biological control of coconut scale Aspidiotus destructor Signoret and citrus scale Aonidiella aurantii (Maskell) was achieved [9][10][11][12]. A clear understanding of Telsimia species diversity and their geographical distribution forms the basis for the introduction and utilization of these natural enemies. This taxonomic study can provide a theoretical basis for the development and utilization of Telsimia species for the biological control of diaspidine scale insects.
At present, 48 species (subspecies) are known around the world, while 10 species (subspecies) are known in China (Table S1) [1,3,. No new species have been described from China since 1979, when Pang and Mao published four new species (subspecies) [33]. With the accumulation of ladybug specimens in recent years, it is necessary to carry out more taxonomic studies on Chinese Telsimia species.   Telsimia parascymnoides Wang sp. nov. 拟小毛寡节瓢虫

(Figures 2 and 3)
Etymology. The specific epithet refers to its general appearance and penis guide closely resembling T. scymnoides Miyatake. Diagnosis. In male genital structure, this species is most closely related to T. scymnoides but can be easily distinguished from the latter by detailed characteristics of penis guide: in ventral view, penis guide slightly convergent to apical one-sixth, then relatively smoothly narrowing to the apex, which is protruding and arcuate, with a tiny notch at middle (Figure 2g), while in T. scymnoides, penis guide slightly convergent to apical onefifth, then suddenly narrowing to a bifid apex (Figure 1j Body oblong oval, convex, broadest at or very slightly behind middle; elytra entirely black except apical part brownish yellow; pronotum, head and clypeus brownish yellow (Figure 3a-c); mesoventrite and metaventrite dark brown; elytral epipleuron brownish yellow; mouthpart, prothoracic hypomeron, prosternum, and legs brownish yellow. Head between eyes 0.43 times the width of head; pubescence dense, appressed, and silver-  white. Pronotum approximately twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.65 times that between posterior angles, lateral borders arcuately convergent from obtuse posterior to briefly rounded anterior angles; pubescence sparse, suberect, and silver-white. Elytra slightly longer than greatest combined breadth; external margins not explanate to borders; pubescence sparse, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 slightly recurved, punctures between coxae large and dense; ventrite 5 in midline as long as ventrites 2-4 together, apical border evenly arcuate in male (Figure 3d). Male genitalia: Penis robust (Figure 3f), in lateral view, as long as tegmen, basal onethird gradually narrowing from the middle to distal end, which is thread-like; apical onefifth gradually narrowing to a rounded apex. In lateral view of penis guide (Figure 3e), tegminal strut short, nearly one-fourth the length of tegmen; penis guide slender, gradu- are also closely related to T. shirozui but can be distinguished from the latter by the following characteristics: in ventral view of tegmen, base of phallobase half the width of proximal end, apical emargination of penis guide larger, inverted U shape; in T. shirozui, base of phallobase slightly narrower than proximal end, apical emargination smaller, inverted V shape. In lateral view of tegmen, penis guide thick, nearly parallel for the basal two-thirds, apical part sharper; in T. shirozui, penis guide relatively thinner, gradually narrowing to apical one-fourth, apical part blunt. Body oval, convex, broadest at middle; elytra black to crater brown, except apical part brownish yellow; pronotum, head and clypeus brownish yellow; mesoventrite and metaventrite dark brown; mouthparts, prothoracic hypomeron, prosternum, elytral epipleuron, and legs brownish yellow. Head between eyes nearly 0.4 times the width of head; pubescence dense, appressed, and silver-white. Pronotum approximately twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.6 times that between posterior angles, lateral borders arcuately convergent from obtuse posterior to briefly rounded anterior angles; pubescence sparse, suberect, and silver-white. Elytra slightly shorter than greatest combined breadth; external margins not explanate to borders; pubescence sparse, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 not recurved, punctures between coxae large and dense; ventrite 5 in midline as long as ventrites 2-4 together, apical border slightly notched medially in male and arcuate in female (Figure 4d,f).
Male genitalia: Penis, in lateral view (Figure 4g), as long as the tegmen, basal half distinctly narrower, apical one-fifth gradually narrowing to rounded apex. In lateral view of tegmen (Figure 4h), tegminal strut short, nearly one-sixth the length of tegmen; penis guide thick, thickness almost equals the width of penis guide in ventral view, nearly parallel for the basal two-thirds, apical part sharper; parameres distinctly shorter than penis guide. In ventral view of tegmen (Figure 4i), the base of phallobase half as wide as the proximal end, penis guide nearly parallel from base to apical one-fifth, then gradually narrowing to a bifid apex, the apical emargination larger, inverted U shape.
Female genitalia: As shown in Figure 4e. rounded anterior angles; pubescence dense, suberect, and silver-white. Elytra slightly shorter than greatest combined breadth; external margins not or very slightly explanate to borders; pubescence dense, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 slightly recurved, punctures between coxae small and sparse; ventrite 5 in midline as long as ventrites 2-4 together, apical border briefly notched medially in male (Figure 5d). Male genitalia: Penis thick (Figure 5e), curved at basal 1/3, as long as the tegmen. Tegminal strut extremely short (Figure 5f). Phallobase inflated. Parameres 1.5 times of phallobase length and 0.7 times of penis guide length, apical half with several long setae. Penis guide in lateral view gradually narrowing to a pointed apex. Penis guide tubular with a long fusiform opening on ventral side. Penis guide in ventral view 2.5 times as long as width of basal part, slightly narrowing to middle part, then expanded laterally, then convergent to a small blunt apex (Figure 5g)  Etymology. The specific epithet is the Latin adjective "latus", meaning broad, referring to its broad penis guide in ventral view.
Diagnosis. Male genitalia of this species are similar to T. daclacensis Hoàng, 1983 with proximal end of penis guide in ventral view distinctly stretched out laterally. The difference is, in Telsimia latus, the outline of the protruding part on each side is similar to an arc (Figure 5g), while, in T. daclacensis, the protruding part on each side forms a downward projection (Hoàng 1983, p. 7, Figure 2 Body oval, convex, broadest at middle; above black except anterior corners of pronotum and head brownish yellow (Figure 5a-c); underside dark brown to black, except elytral epipleuron brownish yellow, mouthparts, prothoracic hypomeron, and legs brownish yellow. Head between eyes nearly 0.4 times the width of head; pubescence dense, appressed, and silver-white. Pronotum slightly more than twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.6 times that between posterior angles, lateral borders arcuately convergent from obtuse posterior to briefly rounded anterior angles; pubescence dense, suberect, and silver-white. Elytra slightly shorter than greatest combined breadth; external margins not or very slightly explanate to borders; pubescence dense, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 slightly recurved, punctures between coxae small and sparse; ventrite 5 in midline as long as ventrites 2-4 together, apical border briefly notched medially in male (Figure 5d).
Male genitalia: Penis thick (Figure 5e), curved at basal 1/3, as long as the tegmen. Tegminal strut extremely short (Figure 5f). Phallobase inflated. Parameres 1.5 times of phallobase length and 0.7 times of penis guide length, apical half with several long setae. Penis guide in lateral view gradually narrowing to a pointed apex. Penis guide tubular with a long fusiform opening on ventral side. Penis guide in ventral view 2.5 times as long as width of basal part, slightly narrowing to middle part, then expanded laterally, then convergent to a small blunt apex ( Figure 5g).  [31].
Diagnosis. This species can be easily distinguished from other Telsimia species by its male genitalia: in lateral view (Figure 6h), tegmen bent nearly 90 • at the middle part; in ventral view (Figure 6i), tegminal strut nearly triangular, broad at base, gradually narrowing to terminal, penis guide nearly parallel for the basal two-thirds, then gradually narrowing to a pointed apex.
Materials examined (30 Remarks. This species is recorded in China for the first time. The female genitalia of this species are also provided for the first time.

(Figures 7 and 8)
Etymology. The specific epithet is the Latin adjective "lunata", meaning crescentshaped, referring to the crescent stripe on elytra.
Diagnosis. This species can be easily distinguished from all known Telsimia species by its unique elytra markings, as shown in Figure 7. The male genitalia are similar to T. postocula but can be distinguished by detailed characteristics of tegmen in ventral view: penis guide slightly longer (1.2×) than phallobase and apical notch inverted V-shaped in T. lunata ( Figure 7j); in T. postocula, penis guide distinctly longer (1.8×) than phallobase and apical notch nearly fusiform (Kapur 1967, p. 169, Figure 7b). Hoàng, 1985: 41 [31]. Diagnosis. This species can be easily distinguished from other Telsimia species by its male genitalia: in lateral view (Figure 6h), tegmen bent nearly 90° at the middle part; in ventral view (Figure 6i), tegminal strut nearly triangular, broad at base, gradually narrowing to terminal, penis guide nearly parallel for the basal two-thirds, then gradually narrowing to a pointed apex.  Body oval, convex, broadest shortly before middle; elytra black to crater brown, each elytron with two markings as showed in Figure 7a-c; pronotum black to dark brown with anterior corners brownish yellow, anterior and lateral margin dark brown; head dark brown, clypeus golden yellow; underside dark brown, except mouthparts, prothoracic hypomeron, elytral epipleuron, and legs brownish yellow. Head between eyes nearly 0.35 times the width of head; pubescence dense, appressed, and silver-white. Pronotum approximately twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.6 times that between posterior angles, lateral borders arcuate, posterior and anterior angles briefly rounded; pubescence dense, suberect, and silverwhite. Elytra generally shorter than greatest combined breadth; external margins slightly explanate to borders; pubescence dense, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 not recurved, punctures between coxae small and sparse; ventrite 5 in midline distinctly shorter than ventrites 2-4 together, apical border briefly notched medially in male and evenly arcuate in female (Figure 7d,f). Remarks. This species is recorded in China for the first time. The female genitalia of this species are also provided for the first time.

(Figures 7 and 8)
Etymology. The specific epithet is the Latin adjective "lunata", meaning crescentshaped, referring to the crescent stripe on elytra.  Male genitalia: Penis distinctly stout except basal one-third relatively slender, slightly curved at middle part, apex rounded in lateral view (Figure 7g). In ventral view, penis broadest at middle part, apex bifurcated ( Figure 7h). Penis guide in lateral view thick, almost parallel at basal two-thirds, then inner side directly narrowing to outside, forming a blunt apex; tegminal strut extremely short (Figure 7i). Penis guide slightly longer than phallobase. Parameres distinctly shorter than penis guide, apex with several long setae. Penis guide in ventral view, 1.60 times as long as wide, nearly parallel for basal two-thirds, apex bifid with a large, inverted V-shaped notch (Figure 7j).
Female genitalia: As shown in Figure 7e.   Diagnosis. This species can be easily distinguished from all known Telsimia species by its unique elytra markings, as shown in Figure 7. The male genitalia are similar to T. postocula but can be distinguished by detailed characteristics of tegmen in ventral view: penis guide slightly longer (1.2×) than phallobase and apical notch inverted V-shaped in T. lunata ( Figure 7j); in T. postocula, penis guide distinctly longer (1.8×) than phallobase and apical notch nearly fusiform (Kapur 1967 Body oval, convex, broadest shortly before middle; elytra black to crater brown, each elytron with two markings as showed in Figure 7a-c; pronotum black to dark brown with anterior corners brownish yellow, anterior and lateral margin dark brown; head dark brown, clypeus golden yellow; underside dark brown, except mouthparts, prothoracic hypomeron, elytral epipleuron, and legs brownish yellow. Head between eyes nearly 0.35 times the width of head; pubescence dense, appressed, and silver-white. Pronotum approximately twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.6 times that between posterior angles, lateral borders arcuate, posterior and anterior angles briefly rounded; pubescence dense, suberect, and silverwhite. Elytra generally shorter than greatest combined breadth; external margins slightly explanate to borders; pubescence dense, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 not recurved, punctures between coxae small and sparse; ventrite 5 in midline distinctly shorter than ventrites 2-4 together, apical border briefly notched medially in male and evenly arcuate in female (Figure 7d,f). Etymology. The specific epithet is the Latin adjective "parvus", meaning small, referring to its small penis guide in ventral view.
Diagnosis. In external appearance, it is similar to T. forcipata sp. nov., for the short oval body, narrower interocular distance (less than one-third the head width), and dense, gold pubescence on head (Figure 9a-c), but their male genitalia vary greatly (Figures 9e-h and 10g-j). In male genital structure, this species is most closely related to T. jinyangiensis, but can be easily distinguished from the latter by some detailed characteristics: in T. parvus, basal part of penis slender, without any subsidiary structure (Figure 9e); tegminal strut very short, nearly one-tenth the length of tegmen ( Figure 9g); phallobase 1.2 times the length of penis guide; apical emargination of penis guide is small and deep, the depth is five times the width of the opening ( Figure 9h); in T. jinyangiensis, basal part of penis with a broad handle-like structure; tegminal strut short, nearly one-fifth the length of tegmen; phallobase nearly the same length as penis guide; apical emargination of penis guide is short triangular, the depth is three times the width of the opening. Body short oval, convex, broadest at middle; above entirely black; underside dark brown to black, except elytral epipleuron orange-brown, mouthparts and legs brownish yellow. Head between eyes narrow, nearly 0.3 times the width of head; pubescence dense, appressed and gold. Pronotum twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.6 times that between posterior angles, lateral borders arcuately convergent from obtuse posterior to briefly rounded anterior angles; pubescence dense, suberect and gold. Elytra slightly shorter than greatest combined breadth; external margins not or slightly explanate to borders; pubescence dense, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 recurved, punctures between coxae small and sparse; ventrite 5 in midline as long as ventrites 2-4 together, apical border distinctly notched medially in male (Figure 9d).
Penis guide in ventral view, 1.60 times as long as wide, nearly parallel for basal two-thirds, apex bifid with a large, inverted V-shaped notch (Figure 7j).
Female genitalia: As shown in Figure 7e. Etymology. The specific epithet is the Latin adjective "parvus", meaning small, referring to its small penis guide in ventral view.  Male genitalia: Penis robust, in lateral view, basal half gradually narrowing from the middle to distal end, as long as the tegmen; apical half slightly narrowing to a rounded apex ( Figure 9e). In lateral view of tegmen (Figure 9g), tegminal strut extremely short, nearly one-tenth the length of tegmen; phallobase 1.2 times the length of penis guide; penis guide thick, thickness almost three-fourth the width of penis guide in ventral view, gradually narrowing from base to apical one-third, then inner side suddenly convergent to a blunt apex; parameres robust at basal two-fifth and slender at apical three-fifth, with several long setae at apex, as long as penis guide. In ventral view of tegmen, the phallobase nearly rectangle, the ratio of width to length is about 3:4; base of penis guide nearly 0.65 times the width of phallobase, penis guide slightly convergent to apical one-third, then gradually narrowing to a bifid apex, the apical emargination small and deep, the depth five times the width of the opening (Figure 9h).  Body short oval, strongly convex, broadest at middle; entirely black above; underside dark brown to black, except elytral epipleuron orange-brown, mouthparts and legs brownish yellow. Head between eyes narrow, less than one-third width of head; pubescence dense, appressed, and gold. Pronotum approximately twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.6 times that between posterior angles, lateral borders arcuately convergent from obtuse posterior to briefly rounded anterior angles; pubescence dense, suberect, and silver-white. Elytra slightly shorter than greatest combined breadth; external margins not or very slightly explanate to borders; pubescence dense, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 recurved, punctures between coxae large and sparse; ventrite 5 in midline as long as ventrites 2-4 together, apical border flattened medially in male and evenly arcuate in female (Figure 10d,f). Etymology. The specific epithet is the Latin adjective "forcipata", meaning forcipate, referring to its forcipate penis guide in ventral view.
Diagnosis. Penis guide of this species in ventral view is deeply bifid (Figure 10j), which is similar to T. ismayi (Chzeau 1984, p. 4, pl. II-4) and T. nigra (Weise). However, in the last two species, there is a distinct triangular protrusion on each lateral side at apical one-third of penis guide, which can be easily distinguished from T. forcipata. Body short oval, strongly convex, broadest at middle; entirely black above; underside dark brown to black, except elytral epipleuron orange-brown, mouthparts and legs brownish yellow. Head between eyes narrow, less than one-third width of head; pubescence dense, appressed, and gold. Pronotum approximately twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.6 times that between posterior angles, lateral borders arcuately convergent from obtuse posterior to briefly rounded anterior angles; pubescence dense, suberect, and silver-white. Elytra slightly shorter than greatest combined breadth; external margins not or very slightly explanate to borders; pubescence dense, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 recurved, punctures between coxae large and sparse; ventrite 5 in midline as long as ventrites 2-4 together, apical border flattened medially in male and evenly arcuate in female (Figure 10d,f).
Male genitalia: Penis thick, curved at middle part, apex rounded, as long as the tegmen (Figure 10h). Tegminal strut stout, broad at basal part, as long as phallobase (Figure 10j). Parameres distinctly longer than phallobase and slightly shorter than penis guide, apex with several long setae. Penis guide in lateral view thick, nearly parallel at basal two-thirds, then narrowing to blunt apex (Figure 10g). Penis guide in ventral view forcipate, splitting upward from apex to basal one-fourth (Figure 10j).
Diagnosis. This species can be easily distinguished from other Telsimia species by its inflated phallobase (Figure 11h). In ventral view, penis guide nearly half the width of phallobase, nearly 3.5 times of length than width, apex distinctly bifid (Figure 11j). Penis tubular of uniform thickness (Figure 11g) Remarks. The female genitalia of this species are provided for the first time.
Biology. Aonidiella aurantii Maskell, Aulacaspis citri Chen, Chrysomphalus aonidum L., Parlatoria ziziphi (Lucas) and Unaspis yanonensis (Kuwana) are reported as its prey [12,33,[41][42][43]. The related studies on the biological characteristics and predation efficiency of this species have shown that this species has a significant natural inhibitory effect on a variety of shield scales and has broad development and application prospects [44][45][46].   Telsimia nigra (Weise, 1879) 黑背寡节瓢虫 (Figures 8 and 12 Diagnosis. This species can be easily distinguished from other Telsimia species by its unique male genitalia: penis well developed, apex with a long filiform appendage ( Figure 12h); penis guide divided into two lobes at median line in entire length, lateral side of each lobe with a distinct angulation at apical 1/3 (Figure 12i  Remarks. Pang and Mao [33] described a new subspecies according to one specimen from Sichuan Province and explained its difference from that of Japan by the angulation on the lateral side of each lobe and the depth of the split. We compared 11 male specimens from five provinces (including Sichuan Province, the type locality of this subspecies) ( Figure S1), and we did not find any significant differences from Sasaji's illustrations [26] (p.214, Figure 88).
Biology. This species is reported to prey on Aulacaspis rosarum Borchsenius, Unaspis yanonensis (Kuwana), Aphis citricidus (Kirkaldy), and Lepidosaphes tubulorum Ferris [7,[50][51][52].   Telsimia chayuensis Wang sp. nov. 察隅寡节瓢虫 (Figures 14 and 15) Etymology. The specific name refers to its type locality. Diagnosis. In male genital structure, this species is closely related to T. emarginata and T. abdita Ślipiński, Pang and Pope, 2005, but can be distinguished from the latter by some detailed characteristics: basal one-fourth of the penis thread-like, apical one-fourth gradually narrowing to a blunt apex ( Figure 15i); in the latter two species, penis tubular, thickness almost consistent in the whole length, except the apical part slightly narrowing to a rounded apex (Figure 11g). Tegmen is mostly similar to T. abdita, but can be distinguished from the latter by detailed characters of penis guide: in lateral view, the apex bent inward (Figure 15h), while, in T. abdita, the apex bent outward; in ventral view, the apex is truncate, with a median line (Figure 15i), while, in T. abdita, the apex is rounded, without a median line. Body oblong oval, convex, broadest at middle; elytra entirely black to dark brown; pronotum black except lateral margin and anterior corners orange-brown, head and clypeus orange-brown; underside dark brown except mouthparts, prothoracic hypomeron, elytral epipleuron, and legs brownish yellow. Head between eyes nearly 0.45 times the width of head; pubescence dense, suberect, and silver-white. Pronotum approximately twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.60 times that between posterior angles, lateral borders arcuately convergent from obtuse posterior to briefly rounded anterior angles; pubescence dense, suberect, and silver-white. Elytra slightly longer than greatest combined breadth; external margins not explanate to borders; pubescence dense, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 recurved, punctures between coxae large and dense; ventrite 5 in Telsimia darjeelingensis Kapur, 1969: 50 [25]. Diagnosis. Male genitalia of this species resemble T. ceylonica Kapur 1969, but can be easily distinguished from the latter by the following characteristics: in T. darjeelingensis, tegminal strut slightly longer than penis guide; penis guide gradually narrowing to a bifid apex ( Figure 13g); basal part of penis with membranous appendages, preapical part of penis slightly enlarged but the apex curved and acute ( Figure 13e); in T. ceylonica, tegminal strut distinctly shorter than penis guide; penis guide gradually narrowing to a blunt apex; basal part of penis without membranous appendages, apex of penis bulbous, nearly dumbbell-shaped (Kapur 1969, p. 47, Figure 1).  Figures 14 and 15) one-fourth thread-like, apical one-fourth gradually narrowing to a blunt apex. In lateral view of tegmen (Figure 15h), tegminal strut short, nearly one-sixth the length of tegmen; phallobase inflated; penis guide slender, gradually narrowing to a blunt apex, which bent inward; parameres slender, nearly the same length as penis guide. In ventral view of tegmen (Figure 15i), the phallobase nearly rectangle, the ratio of width to length is about 3:2; penis guide nearly parallel from base to apical one-third, then gradually narrowing to a truncate and protruding apex, apical one-fourth of penis guide has a midline. Female genitalia: As shown in Figure 15e.  Etymology. The specific name refers to its type locality. Diagnosis. In male genital structure, this species is closely related to T. emarginata and T. abditaŚlipiński, Pang and Pope, 2005, but can be distinguished from the latter by some detailed characteristics: basal one-fourth of the penis thread-like, apical one-fourth gradually narrowing to a blunt apex (Figure 15i); in the latter two species, penis tubular, thickness almost consistent in the whole length, except the apical part slightly narrowing to a rounded apex (Figure 11g). Tegmen is mostly similar to T. abdita, but can be distinguished from the latter by detailed characters of penis guide: in lateral view, the apex bent inward (Figure 15h), while, in T. abdita, the apex bent outward; in ventral view, the apex is truncate, with a median line (Figure 15i), while, in T. abdita, the apex is rounded, without a median line. Body oblong oval, convex, broadest at middle; elytra entirely black to dark brown; pronotum black except lateral margin and anterior corners orange-brown, head and clypeus orange-brown; underside dark brown except mouthparts, prothoracic hypomeron, elytral epipleuron, and legs brownish yellow. Head between eyes nearly 0.45 times the width of head; pubescence dense, suberect, and silver-white. Pronotum approximately twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.60 times that between posterior angles, lateral borders arcuately convergent from obtuse posterior to briefly rounded anterior angles; pubescence dense, suberect, and silver-white. Elytra slightly longer than greatest combined breadth; external margins not explanate to borders; pubescence dense, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 recurved, punctures between coxae large and dense; ventrite 5 in midline distinctly shorter than ventrites 2-4 together, apical border briefly notched medially in male and evenly arcuate in female (Figure 15d,f).

Telsimia darjeelingensis
Male genitalia: Penis, in lateral view (Figure 15g), slightly shorter than tegmen, basal one-fourth thread-like, apical one-fourth gradually narrowing to a blunt apex. In lateral view of tegmen (Figure 15h), tegminal strut short, nearly one-sixth the length of tegmen; phallobase inflated; penis guide slender, gradually narrowing to a blunt apex, which bent inward; parameres slender, nearly the same length as penis guide. In ventral view of tegmen (Figure 15i), the phallobase nearly rectangle, the ratio of width to length is about 3:2; penis guide nearly parallel from base to apical one-third, then gradually narrowing to a truncate and protruding apex, apical one-fourth of penis guide has a midline.
Female genitalia: As shown in Figure 15e.  Figures 14 and 16) Etymology. The specific epithet is the Latin adjective "lobatus", meaning lobed, referring to the bifid apex of penis guide in ventral view.
Diagnosis. This species can be easily distinguished from all known Telsimia species by its male genitalia: penis distinctly stout (Figure 16f), penis guide in ventral view broad, 2.5 times of length than width, parallel at basal two-thirds, then gradually narrowing to a very small bifid apex (Figure 16h Body short oval, convex, broadest at middle; above black except apical part narrowly brownish; underside dark brown to black, except mouthpart and legs orange-brown. Head between eyes nearly 0.4 times the width of head; pubescence dense, appressed, and silver-white. Pronotum approximately twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.6 times that between posterior angles, lateral borders arcuate, posterior and anterior angles briefly rounded; pubescence dense, suberect, and silver-white. Elytra generally shorter than greatest combined breadth; external margins not explanate to borders; pubescence dense, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 distinctly recurved, punctures between coxae large and dense; ventrite 5 in midline as long as or slightly shorter than ventrites 2-4 together, apical border nearly flattened medially in male (Figure 16d).  Etymology. The specific epithet is the Latin adjective "lobatus", meaning lobed, referring to the bifid apex of penis guide in ventral view.
Diagnosis. This species can be easily distinguished from all known Telsimia species by its male genitalia: penis distinctly stout (Figure 16f), penis guide in ventral view broad, 2.5 times of length than width, parallel at basal two-thirds, then gradually narrowing to a very small bifid apex (Figure 16h Body short oval, convex, broadest at middle; above black except apical part narrowly brownish; underside dark brown to black, except mouthpart and legs orange-brown. Head between eyes nearly 0.4 times the width of head; pubescence dense, appressed, and silver-white. Pronotum approximately twice as broad as median length; anterior border arcuate medially, distance between anterior angles 0.6 times that between posterior angles, lateral borders arcuate, posterior and anterior angles briefly rounded; pubescence dense, suberect, and silver-white. Elytra generally shorter than greatest combined breadth; external margins not explanate to borders; pubescence dense, suberect, and silver-white. Abdominal postcoxal lines of ventrite 1 distinctly recurved, punctures between coxae large and dense; ventrite 5 in midline as long as or slightly shorter than ventrites 2-4 together, apical border nearly flattened medially in male (Figure 16d).
Male genitalia: Penis, in lateral view (Figure 16f), distinctly stout, curved at middle part, apex rounded; in ventral view of tegmen (Figure 16h), apical half gradually narrow to apex. Tegminal strut nearly one-third the length of main part of the tegmen. Parameres slightly shorter than penis guide, apical part with several long setae. Penis guide thick in lateral view (Figure 16g), gradually narrowing to a pointed apex; in ventral view broad, 2.5 times of length than width, parallel at basal two-thirds, and then gradually narrowing to a small bifid apex. Telsimia humidiphila Kapur, 1969: 52 [25]. Diagnosis. Male genitalia of this species are similar to T. postocula but can be easily distinguished from the latter by its slenderer penis guide and shorter parameres. In T. humidiphila, penis guide 2.5-3 times the length than width, parameres half the length of penis guide (Figure 17j); in T. postocula, penis guide nearly 2 times the length than width, parameres slightly shorter than penis guide (Kapur 1967 , p. 169, Figure 7b).
Materials examined (15    Remarks. The female genitalia of this species are provided for the first time ( Figure  18e).
Biology. Detailed larval description and additional adult diagnosis have been reported by Park and Yoon [53]. Some biological notes, including habitats, prey, seasonal observation, and the distributional tendency of the species, are also discussed.  Distribution. China: Hong Kong, Taiwan; Japan; Korea. Remarks. The female genitalia of this species are provided for the first time (Figure 18e). Biology. Detailed larval description and additional adult diagnosis have been reported by Park and Yoon [53]. Some biological notes, including habitats, prey, seasonal observation, and the distributional tendency of the species, are also discussed.

Telsimia chujoi Miyatake, 1959 朱氏寡节瓢虫
( Figures 19 and 20) the length of the penis guide, the penis guide gradually narrowing to a blunt apex, apex with a narrow, short median emargination (Figure 18i).
Diagnosis. Male genitalia of this species are similar to T. nagasakiensis Miyatake, 1978, but can be distinguished by the combined characteristics as follows: in T. chujoi, tegminal strut slightly shorter than penis guide, penis guide gradually narrowing to a pointed apex, apex with a short median line ( Figure 19g); in T. nagasakiensis, tegminal strut nearly half the length of the penis guide, the penis guide gradually narrowing to a blunt apex, apex with a narrow, short median emargination (Figure 18i  Diagnosis. This species can be easily distinguished from other Telsimia species by its unique male genitalia: in lateral view of tegmen (Figure 21g), tegminal strut very short, nearly one-sixth the length of the tegmen, main tegmen slender, gradually narrowing from base to apex, penis guide exceptionally long, more than twice the length of parameres; in ventral view (Figure 21i), penis guide six times the length as width, nearly parallel from base to apical one-sixth, then gradually narrowing to a blunt apex. Penis threadlike, distinctly longer than the penis guide (Figure 21h). Biology. This species is reported to prey on Hemiberlesia pitysophila Takagi in Guangxi, China [57]. Diagnosis. This species can be easily distinguished from other Telsimia species by its unique male genitalia: in lateral view of tegmen (Figure 21g), tegminal strut very short, nearly one-sixth the length of the tegmen, main tegmen slender, gradually narrowing from base to apex, penis guide exceptionally long, more than twice the length of parameres; in ventral view (Figure 21i), penis guide six times the length as width, nearly parallel from base to apical one-sixth, then gradually narrowing to a blunt apex. Penis thread-like, distinctly longer than the penis guide (Figure 21h). Biology. This species is reported to prey on Hemiberlesia pitysophila Takagi in Guangxi, China [57].   [23]. Diagnosis. Male genitalia of this species are closely related to T. nitida Chapin 1926, but can be easily distinguished from the latter by the ventral view of tegmen: penis guide slightly narrower than phallobase, apical emargination small, width of notch end nearly seven times of width of penis guide at middle part ( Figure 22i); in T. nitida, penis guide distinctly narrower than phallobase, apical emargination larger, width of notch end nearly twice the width of penis guide at middle part (Chapin 1965, p. 242, Figure 39 Telsimia shirozui Miyatake, 1965: 55 [22]; Miyatake, 1978: 18 [23]. Diagnosis. Male genitalia of this species are closely related to T. nitida Chapin 1926, but can be easily distinguished from the latter by the ventral view of tegmen: penis guide slightly narrower than phallobase, apical emargination small, width of notch end nearly seven times of width of penis guide at middle part ( Figure 22i); in T. nitida, penis guide distinctly narrower than phallobase, apical emargination larger, width of notch end nearly twice the width of penis guide at middle part (Chapin 1965, p. 242, Figure 39   Diagnosis. Male genitalia of this species are similar to T. elainae Chazeau 1984, especially the lateral view of penis and tegmen, but can be easily distinguished from latter by the ventral view of tegmen: in T. jinyangiensis, penis guide as long as phallobase, and distinctly narrower than phallobase, apical emargination small and inverted V shape ( Figure 23i); in T. elainae, penis guide distinctly longer than phallobase, and as wide as phallobase, apical emargination larger and inverted U shape (Chazeau 1984, p. 2, pl. I-4).

Telsimia shirozui
( Figures 20 and 23) Telsimia jinyangiensis Pang and Mao, 1979: 102 [33]; Ren et al., 2009: 150 [39]. Diagnosis. Male genitalia of this species are similar to T. elainae Chazeau 1984, especially the lateral view of penis and tegmen, but can be easily distinguished from latter by the ventral view of tegmen: in T. jinyangiensis, penis guide as long as phallobase, and distinctly narrower than phallobase, apical emargination small and inverted V shape ( Figure  23i); in T. elainae, penis guide distinctly longer than phallobase, and as wide as phallobase, apical emargination larger and inverted U shape (Chazeau 1984, p. 2, pl. I-4).        Kapur 1967, while most the other species are all black or with a little yellow spot at the end of the elytra, which hinders identification by elytral markings. In addition, color variation is common in many groups of Coccinellidae, although it has not been found in this group; its potential impact on species identification should be fully considered. Moreover, body size, shape, and punctures are occasionally used for species identification. However, as the number of species and specimens increases, the differences between species become smaller and individual differences become greater, making species identification using these characteristics very difficult. In the future, with the extensive use of 3D scanning electron microscopy and micro-CT, combined with new techniques such as geometric morphology, computer vision, and deep learning [58,59], it may be possible to make new breakthroughs in the direction of external morphology-based species identification.

Key to Chinese Species of Telsimia
Some scholars have also used female genitalia as a species identification feature, such as with T. tamdaoensis Hoàng 1981 and T. bangalorensis Kapur 1969. However, female genitalia are considered more applicable to distinguishing different species rather than identifying new species. The reason is that if female genitalia were to be used for species identification, strictly speaking, a comparative study of the female genitalia of all previous species would be necessary. The reality is that the female genitalia of many species have not been previously studied, and many species have been described on the basis of males only. In addition, male-female matching, a major problem in taxonomic studies, is also present in this taxon. In the present study, we matched males and females on the basis of a large number of external morphological characteristics and the same label information. However, this requires special caution, as multiple species could be collected at the same time and place. Through our study, we found that female genitalia differed little between most species in this taxon. Therefore, the use of female genitalia as a species identification feature in this taxon is not recommended.
In conclusion, male genitalia are more recommended for species identification of this taxon, and other characteristics can be used as an auxiliary. It is also possible to try to use DNA barcoding for species identification and male-female matching; this is one of the directions of future research.
Phylogeny. So far, we have not seen any phylogenetic studies within the genus Telsimia. Through this study, we found that there were differences in the ratio of male genitalia to body size among different groups. For example, in T. sichuanensis, T. chujoi, T. nagasakiensis, T. elongate, and T. darjeelingensis, the penis is obviously longer than the abdomen; in T. nigra, the main part of the penis is about equal to the abdomen; in T. scymnoides and T. parascymnoides sp. nov., the penis is slightly shorter than the abdomen; in the other groups, the penis is equal to or slightly longer than half the length of the abdomen. This attribute may be sourced from reproductive competition, and the changing trend of the ratio may indicate the direction of the species evolution of this taxon, forming an important feature of higher-level classification. Therefore, in the illustrations of ladybug taxonomic articles, we strongly recommend the use of scale bars. When the difference in genital size between species is not particularly large, comparable images with a particular scale bar will make the ratio difference more intuitive. In addition, antennae, mouthparts, feet, and prothorax are also important classification features for phylogenetic studies, and further research will be carried out in this area in the future.
Biology and biological control. There are relatively few studies on the biology of species in this taxon [7,[43][44][45]53], and the biology of most species is unknown. Research on biological control applications is also limited to a few species, such as T. emarginata, T. nigra, and T. nitida [4,5,9,12]. In addition to the widespread distribution of these species, the reason for this situation is largely due to the difficulty of species identification. Therefore, it is hoped that our study may provide some help.
Supplementary Materials: The following supporting information can be downloaded at: https: //www.mdpi.com/article/10.3390/insects13100869/s1, Table S1: List of Telsimia species around the world before this study; Figure S1: The tegmen of Telsimia nigra from different places in China.

Institutional Review Board Statement: Not applicable.
Data Availability Statement: All data are available in this paper.