Research Article
Print
Research Article
New species, new synonymies and a new record of the genus Cryptogonus Mulsant, 1850 (Coleoptera, Coccinellidae) from China
expand article infoLizhi Huo, Wenjing Li, Xiaosheng Chen, Shunxiang Ren, Xingmin Wang
‡ South China Agricultural University, Guangzhou, China
Open Access

Abstract

Three new species of the genus Cryptogonus Mulsant, 1850 from China are described and illustrated: C. dulongjiangensis, C. fusiformis and C. reniformis Huo and Ren. Cryptogonus octoguttatus Mader, 1954 and C. kurosawai Sasaji, 1968 are recognized as synonymous with C. schraiki Mader, 1933. Cryptogonus hingstoni Kapur, 1948 is newly recorded from China. A species list of the genus Cryptogonus is presented.

Key Words

Cryptogonus, new species, new synonymy, new record, China

Introduction

The genus Cryptogonus belongs to Aspidimerini (Coleoptera, Coccinellidae). The Aspidimerini species are widely distributed in South and Southeast Asia. They are natural enemies of coccidae, aphidae, aleyrodidae and have important application value in the control of insect pests.

The genus Cryptogonus was erected by Mulsant (1850) with C. orbiculus (Gyllenhal, 1808) as the type species by monotypy. Weise (1900) separated the genera Cryptogonus and Aspidimerus Mulsant, 1850 from Scymnini based on the structure of male genitalia and erected the tribe Aspidimerini with Aspidimerus Mulsant, 1850 as the type genus. Kapur (1948) revised the tribe Aspidimerini and proposed two new genera: Pseudaspidimerus Kapur and Acarinus Kapur, and subdivided the genus Cryptogonus into six species groups based on the shape of prosternal carinae. In Kapur’s revision, 19 Cryptogonus species were included, and then during the past several decades 30 new species were attributed to this genus (Mader 1954, Sasaji 1968, Bielawski 1972, 1979, Ghorpade 1974, Pang and Mao 1979, Hoàng 1982, 1985, Cao and Xiao 1984, Canepari 1986, Kuznotsov and Pang Hong 1991, Xiao and Li 1992, Yu 1995, Pang 1998). Among those 30 new species, C. nigritus Pang & Mao, 1979 and C. montanus Hoàng, 1985 were transferred to Aspidimerus Mulsant, 1850 and Trigonocarinatus Huo & Ren, 2015, respectively (Kovář 2007, Huo et al. 2015).

During studies on Aspidimerini from the Oriental Region, the genus Aspidimerus from China and Pseudaspidimerus from Laos have been reviewed (Huo et al. 2013, 2014). In the present paper, three new species of the genus Cryptogonus Kapur, 1948 from China are described and illustrated. Cryptogonus octoguttatus Mader, 1954 and C. kurosawai Sasaji, 1968 are recognized as synonymous with C. schraiki Mader, 1933. Cryptogonus hingstoni Kapur, 1948 is newly recorded from China. A species list of the genus Cryptogonus is also presented.

Material and methods

All studied materials were deposited in the Department of Entomology, South China Agricultural University (SCAU). Type specimens designated in the current article were deposited in SCAU and the Institute of Zoology (IOZ), Chinese Academy of Science, Beijing.

Measurements were made using an ocular micrometer attached to a stereomicroscope (SteREO Discovery V20, Zeiss) as follows: (TL) total length, from apical margin of clypeus to apex of elytra; (TW) total width, across both elytra at widest part; (TH) total height, through the highest point of elytra to elytral outer margins; (HW) head width, including eyes; (PL) pronotal length, from middle of anterior margin to base of pronotum; (PW) pronotal width at widest part; (EL) elytral length, along suture, from apex to base including scutellum; (EW) elytral width, across both elytra at widest part; (ID) interocular distance, nearest distance between two eyes. Morphological terms follow Ślipiński (2007) and Ślipiński and Tomaszewska (2010) and are applied as in our previous studies on Chinese species of former Scymninae (e.g. Chen et al. 2013, Chen et al. 2014).

External morphology was observed with a stereomicroscope (SteREO Discovery V20, Zeiss). Male and female genitalia were dissected, cleared in 10% NaOH solution by boiling for several minutes and observed under a compound microscope, Olympus BX51. Images were photographed with digital cameras (AxioCam HRc and Coolsnap-Procf & CRI Micro*Color). The software AXIOVISION REL. 4.8 and IMAGE-PRO PLUS 5.1 were used to capture images from both cameras. Images were cleaned up and arranged in plates with ADOBE PHOTOSHOP CS5.

Taxonomy

Description of new species

Cryptogonus dulongjiangensis Huo & Ren, sp. n.

Figure 1

Diagnosis

This species can be distinguished from other Cryptogonus species by its long penis, regularly narrowing to pointed apex, penis capsule with both branches very short (Fig. 1e).

Figure 1.

Cryptogonus dulongjiangensis Huo et Ren, sp. n. (a) dorsal view; (b) frontal view; (c) lateral view; (d) abdomen, male; (e) penis; (f) apex of penis; (g) ventral view of tegmen; (h) lateral view of tegmen; (i) coxites. Scale bars: a–d, 0.5 mm; e–h, 0.1 mm.

Description

TL: 2.37–2.60 mm, TW: 1.86–2.11 mm, TH: 1.21–1.32 mm, TL/TW: 1.23–1.27; PL/PW: 0.51–0.53; EL/EW: 0.99–1.00, HW/PW: 0.61–0.63; PW/EW: 0.71–0.73. ID/HW: 0.49–0.53.

Body rounded, densely covered with short pubescence, golden on head and pronotum and silver white on elytra (Fig. 1a–c). Head yellow in male and black in female, clypeus dark brown. Pronotum black except a triangular yellow spot at anterior corner in male. Scutellum and elytra black. Ventral side black except legs and abdomen partially yellow to dark brown.

Punctures on frons coarse and dense, 0.3–0.5 diameters apart. Punctures on pronotum and elytra fine and sparse, 1–3 diameters apart. Punctures on metaventrite fine and sparse at middle, 5 diameters apart, coarse and dense on both sides, 0.2 diameters apart.

Male genitalia. Penis long, regularly narrowing to pointed apex (Fig. 1f), penis capsule with both branches very short (Fig. 1e). Tegminal strut as long as tegmen (Fig. 1h). Parameres 2 times length of phallobase with apices sparsely setose. Penis guide, in lateral view, gradually narrowing to pointed apex. In ventral view, subparallel at basal 2/3, narrowing to pointed apex (Fig. 1g).

Female genitalia. Coxites subtriangular (Fig. 1i), with dense, long terminal setae. Spermatheca missing.

Types

Holotype: 1 male, CHINA: Yunnan Prov.: Qinglandan, Maku Village, Dulongjiang, Gongshan County, 27°41.12′N, 98°16.35′E, ca 1260 m, 4–7.VIII.2010, Wang XM et al. leg. (SCAU). Paratypes (2): CHINA: Yunnan Prov.: 1 male, Maku Village, Dulongjiang, Gongshan County, 27°40.57′N, 98°18.15′E, ca 1600 m, 1.VIII.2010, Wang XM et al. leg; 1 female, Bapo Village, Dulongjiang, Gongshan County, 27°44.08′N, 98°20.59′E, ca 1400 m, 28.VII.2010, Wang XM et al. leg. (SCAU).

Distribution

China (Yunnan).

Etymology

The specific epithet refers to the type locality, Dulongjiang, Yunnan.

Cryptogonus fusiformis Huo & Ren, sp. n.

Figure 2

Diagnosis

This species can be distinguished from other Cryptogonus species by its fusiform penis guide in ventral view (Fig. 2f).

Figure 2.

Cryptogonus fusiformis Huo et Ren, sp. n. (a) dorsal view; (b) frontal view; (c) lateral view; (d) abdomen, male; (e) penis; (f) apex of penis; (g) ventral view of tegmen; (h) lateral view of tegmen. Scale bars: a–d, 0.5 mm; e–g, 0.1 mm.

Description

TL: 2.68 mm, TW: 2.14 mm, TH: 1.36 mm, TL/TW: 1.25; PL/PW: 0.51; EL/EW: 1.00, HW/PW: 0.58; PW/EW: 0.72. ID/HW: 0.50.

Body rounded, densely covered with short, silver white pubescence (Fig. 2a–b). Base of head yellow, anterior part black and clypeus dark brown. Dorsum entirely black. Ventral side black except legs and abdomen partially reddish brown.

Punctures on frons dense and coarse, 0.5–1.0 diameters apart. Punctures on pronotum and elytra fine and sparse, 2–4 diameters apart. Punctures on metaventrite fine and sparse at middle, 5 diameters apart, coarse and dense on both sides, 0.3 diameters apart.

Male genitalia. Penis long with apex pointed. Penis capsule with longer outer branch and short inner one. Tegminal strut slightly longer than tegmen. Parameres 2 times length of phallobase with apex densely setose (Fig. 2g). Penis guide, in lateral view strongly curved as S-shape, equivalent to parameres, in ventral view gradually broadening toward midlength, narrowing to pointed apex (Fig. 2f).

Female genitalia. Unknown.

Types

Holotype: 1 male, CHINA: Yunnan Prov.: Mengdui Town, Zhenkang County, 23°54.16′N, 98°54.02′E, ca 1400 m, 18.V.2008, Wang XM et al. leg. (SCAU).

Distribution

China (Yunnan).

Etymology

The specific epithet is a Latin adjective referring to its fusiform penis guide in ventral view.

Cryptogonus reniformis Huo & Ren, sp. n.

Figure 3

Diagnosis

This species can be easily identified by its reniform spot on elytra and trifurcate penis apex (Fig. 3a, h).

Figure 3.

Cryptogonus reniformis Huo & Ren, sp. n. (a) dorsal view; (b) frontal view; (c) lateral view; (d) abdomen, male; (e) coxites; (f) spermatheca; (g) penis; (h) apex of penis; (i) ventral view of tegmen; (j) lateral view of tegmen. Scale bars: a–d, 0.5 mm; e, g–j, 0.1 mm; f, 0.05 mm.

Description

TL: 1.90–2.40 mm, TW: 1.52–1.94 mm, TH: 1.00–1.27 mm, TL/TW: 1.24–1.25; PL/PW: 0.53–0.55; EL/EW: 0.96–1.01, HW/PW: 0.61–0.64; PW/EW: 0.71–0.72. ID/HW: 0.49–0.52.

Body rounded, densely covered with short, silver white pubescence (Fig. 3a–c). Head yellowish in male and black in female, clypeus dark brown. Pronotum black except a triangular yellowish spot at anterior corner in male. Scutellum black. Elytra black with a reniform spot at middle (Fig. 3a). Underside black except legs partially yellowish.

Punctures on frons coarse and dense, 0.5–1.5 diameters apart. Punctures on pronotum and elytra fine and sparse, 2–4 diameters apart. Punctures on metaventrite fine and sparse at middle, 8 diameters apart, coarse and dense on both sides, 0.5 diameters apart.

Male genitalia. Penis long with apex trifurcate (Fig. 3h). Penis capsule with outer branch bigger than inner one, anterior margin deeply concave (Fig. 3g). Tegminal strut as long as tegmen. Parameres 2 times length of phallobase with apex sparsely setose (Fig. 3j). Penis guide, in lateral view gradually narrowing to pointed apex, a little longer than parameres, in ventral view 3 times as long as wide, slightly broadening to apical 1/3, gradually narrowing to rounded apex (Fig. 3i).

Female genitalia. Coxites reniform with a small projection on the apical end (Fig. 3e). Spermatheca short and strongly arcuate without ramus (Fig. 3f).

Types

Holotype: 1 male, CHINA: Yunnan Prov.: Qinglandan, Maku Village, Dulongjiang, Gongshan County, 27°41.12′N, 98°16.35′E, ca 1260 m, 4–7.VIII.2010, Wang XM et al. leg. (SCAU). Paratypes (38): CHINA: Yunnan Prov.: 6 males, 14 females (2 males and 2 females in IOZ, 4 males and 12 females in SCAU), with the same data as holotype; 1 male, Kongdang Village, Dulongjiang, Gongshan County, 27°52.18′N, 98°20.24′E, ca 1600 m, 27.VII.2010, Wang XM et al. leg. (SCAU); 4 males, Bapo Village, Dulongjiang, Gongshan County, 27°44.09′N, 98°21.02′E, ca 1400 m, 28.VII.2010, Wang XM et al. leg. (SCAU); 3 males, 3 females, Bapo Village–Maku Village, Dulongjiang, Gongshan County, 27°42.49′N, 98°20.18′E, ca 1450 m, 29.VII.2010, Wang XM et al. leg. (SCAU); 4 males, 1 female, Maku Village, Dulongjiang, Gongshan County, 27°40.57′N, 98°18.15′E, ca 1600 m, 1.VIII.2010, Wang XM et al. leg. (SCAU); Tibet: 2 females, Beibeng Village, Motuo County, ca 850 m, 4.X.2011, Huo LZ et al. leg. (SCAU).

Distribution

China (Tibet, Yunnan).

Etymology

The specific epithet is a Latin adjective referring to its reniform spot on elytra.

New synonymies and new record

Cryptogonus schraiki Mader, 1933

Figures 4, 5

Cryptogonus schraiki Mader, 1933: 80; Kapur 1948: 101; Pang and Mao 1979: 69; Kovář 2007: 576; Ren et al. 2009: 120.

Cryptogonus octoguttatus Mader, 1954: 129; Pang and Mao 1979: 68; Kovář 2007: 576; Ren et al. 2009: 118. Syn. n.

Cryptogonus kurosawai Sasaji, 1968: 11; Yu 1995: 140, 2011: 161; Kovář 2007: 576. Syn. n.

Remarks

Leopold Mader described C. schraiki Mader, 1933 from Sichuan, China with a brief description of elytral coloration (Fig. 4a–i). Kapur (1948) reviewed this species and illustrated its appearance. Later, Mader described C. octoguttatus Mader, 1954 also from Sichuan, China, only with description of elytral coloration. Sasaji (1968) described C. kurosawai Sasaji, 1968 from Taiwan, China. Photographs of the holotype were available on the website of The Digital Museum of Natural & Science. Pang and Mao (1979) reviewed C. schraiki and C. octoguttatus and illustrated their appearance and male genitalia, but didn’t notice the similarities of these two species. We examined these three species from China and found that they are just the same species with different elytral coloration. The elytral coloration is variable, from entirely black to entirely yellowish (Fig. 5a–l). Besides, we found the male genitalia of specimens are slightly different, even in the same coloration. Sometimes penis guide equal to, slightly longer or shorter than parameres. They are considered as individual differences.

Figure 4.

Cryptogonus schraiki Mader, 1933. (a) dorsal view; (b) frontal view; (c) lateral view; (d) abdomen, male; (e) lateral view of tegmen; (f) ventral view of tegmen; (g) penis; (h) coxites; (i) spermatheca. Scale bars: a–d, 0.5 mm; e–h, 0.2 mm; i, 0.1 mm.

Figure 5.

Different elytral coloration of Cryptogonus schraiki Mader, 1933. Scale bars: a–l, 0.5 mm.

Material examined

205 specimens from China were examined (see the details in supplementary material).

Distribution

China (Anhui, Fujian, Gansu, Guangdong, Guizhou, Hubei, Hunan, Sichuan, Taiwan, Yunnan).

Cryptogonus hingstoni Kapur, 1948

Figure 6

Cryptogonus hingstoni Kapur, 1948: 103; Kapur 1963: 24; Kovář 2007: 575.

Remarks

Kapur (1948) described this species from Sikkim, India. Two specimens from Tibet, China match the description except coloration variation on elytra. In the original description, the spots on sutural and middle part of the elytron distinctly larger than that on humeral callus, the spot on pronotum is oval. However, in the present specimens we examined, the spots on sutural and middle part are as large as that on humeral callus, the spot on pronotum is triangular (Fig. 6a–c).

Figure 6.

Cryptogonus hingstoni Kapur, 1948. (a) dorsal view; (b) frontal view; (c) lateral view; (d) abdomen, male; (e) ventral view of tegmen; (f) lateral view of tegmen; (g) penis; (h) coxites. Scale bars: a–d, 0.5 mm; e–h, 0.1 mm.

Material examined

CHINA: Tibet: 1 male, Bangxin Village, Motuo County, 29°34.58′N, 95°23.60′E, ca 1840 m, 12.X.2011, Huo LZ et al. leg; 1 female, Beibeng Village–Hanmi Village, Motuo County, 29°14.31′N, 95°10.58′E, ca 800–2100 m, 5-8.X.2011, Huo LZ et al. leg.

Distribution

China (Tibet) new distribution, India (Sikkim).

List of the genus Cryptogonus Mulsant

  1. Cryptogonus angusticarinatus Sasaji, 1968

  2. Cryptogonus ariasi (Mulsant, 1853)

  3. Cryptogonus bilineatus Kapur, 1948

  4. Cryptogonus bimaculatus Kapur, 1948

  5. Cryptogonus brachylobius Pang, 1998

  6. Cryptogonus bryanti Kapur, 1948

  7. Cryptogonus complexus Kapur, 1948

  8. Cryptogonus deltodirus Kapur, 1948

  9. Cryptogonus deltoides Kapur, 1948

  10. Cryptogonus downingi Kapur, 1948

  11. Cryptogonus dulongjiangensis Huo & Ren, sp. n.

  12. Cryptogonus forficulae Cao & Xiao, 1984

  13. Cryptogonus fractemaculatus Pang, 1998

  14. Cryptogonus fulvoterminatus Boheman, 1858

  15. Cryptogonus fusiformis Huo & Ren, sp. n.

  16. Cryptogonus guangdongiensis Pang & Mao, 1979

  17. Cryptogonus hainanensis Pang & Mao, 1979

  18. Cryptogonus hanoiensi Hoàng, 1982

  19. Cryptogonus himalayensis Kapur, 1948

  20. Cryptogonus hingstoni Kapur, 1948

  21. Cryptogonus horishanus (Ohta, 1929)

  22. Cryptogonus kapuri Ghorpade, 1974: 55

  23. Cryptogonus laetus (Weise, 1885)

  24. Cryptogonus langchanhensis Hoàng, 1982

  25. Cryptogonus lepidus (Weise, 1885)

  26. Cryptogonus lijiangiensis Pang & Mao, 1979

  27. Cryptogonus linguilatus Pang, 1998

  28. Cryptogonus lobulus Xiao, 1992

  29. Cryptogonus loebli Canepari, 1986: 27

  30. Cryptogonus nepalensis Bielawski, 1972

    Cryptogonus nepalensis bhutanensis Bielawski, 1979

  31. Cryptogonus nitidus Kapur, 1948

  32. Cryptogonus ocellatus Hoàng, 1985

  33. Cryptogonus ohtai Sasaji, 1968

  34. Cryptogonus orbiculus (Gyllenhal, 1808)

  35. Cryptogonus parorbiculus Kuznotsov & Pang Hong, 1991

  36. Cryptogonus polytrichus Kuznotsov & Pang Hong, 1991

  37. Cryptogonus postmedialis Kapur, 1948: 95

  38. Cryptogonus qianjiangensis Xiao, 1992

  39. Cryptogonus quadriguttatus (Weise, 1895)

  40. Cryptogonus reniformis Huo & Ren, sp. n.

  41. Cryptogonus robustus Yu, 1995

  42. Cryptogonus sagittiformis Pang & Mao, 1979

  43. Cryptogonus schraiki Mader, 1933

  44. Cryptogonus trifurcatus Pang & Mao, 1979

  45. Cryptogonus trioblitus (Gorham, 1895)

  46. Cryptogonus tristis (Weise, 1910)

  47. Cryptogonus wuzhishanus Pang & Mao, 1979

  48. Cryptogonus xiushanensis Xiao, 1993

  49. Cryptogonus yunnanensis Cao & Xiao, 1984

Acknowledgements

The authors sincerely thank Rashid Azad (SCAU) for improving this manuscript. The research was supported by the Science and Technology Partnership Program, Ministry of Science and Technology of China (KY201402014) and Science and Technology Program of Guangzhou, China (151800033).

References

  • Bielawski R (1979) Ergebnisse der Bhutan-Expedition 1972 des Naturhistorischen Museums in Basal, Coleoptera: Fam. Coccinellidae. Entomologica Basiliensia 4: 83–125.
  • Boheman CH (1859) Coleoptera. Species novas descripsit. In: Virgin C (Ed.) Kongliga Svenska Fregatten Eugenies Resa omkring jorden under befäl af C. A. Virgin Ahren 1851–1853. Vetenskapliga iakttagelser på H.M. Konung Oscar Den Förstes befallning utgifna af K. Svenska Vetenskaps Akademien, Zoologi III, Inseckter. P.A. Norstedt & söner, Stockholm, Sweden, 113–218.
  • Canepari C (1986) On some coccinellids of northern India and Nepal in the Geneva Museum of Natural History (Coleoptera Coccinellidae). Revue Suisse de Zoologie 93(1): 21–36.
  • Cao CY, Xiao NN (1984) New species of Coccinellidae from Yunnan, China. Entomotaxonomia 6(2–3): 109–125.
  • Chen XS, Li WJ, Wang XM, Ren SX (2014) A review of the subgenus Neopullus of Scymnus (Coleoptera: Coccinellidae) from China. Annales Zoologici 64(2): 299–326. doi: 10.3161/000345413X672483
  • Chen XS, Wang XM, Ren SX (2013) A review of the subgenus Scymnus of Scymnus from China (Coleoptera: Coccinellidae). Annales Zoologici 63(3): 417–499. doi: 10.3161/000345413X672483
  • Ghorpade KD (1974) Description of a new Cryptogonus Mulsant from Bangalore, Southern India. Oriental Insects 8(1): 55–60. doi: 10.1080/00305316.1974.10434442
  • Gorham HS (1895) On the Coccinellidae collected by Mr. L. Fea in Burma. Annali del Museo Civicodi Storia Naturale, Genova (2)34: 683–695.
  • Gyllenhal L (1808) Synonymia Insectorum, oder: versuch einer synonymie aller bisher bekannten Insecten, nach Fabricii Systema Eleutheratorum geordnet von C. J. Schönherr. Erster Band, Eleutherata oder Käfer. Zweiter Theil. C.F. Marquard, Stockholm, 424 pp.
  • Hoàng DN (1982) Coccinellidae of Vietnam Part I. Nha xuat ban khoa hoc va ky thuat, Hanoi, 211 pp.
  • Huo LZ, Chen XS, Li WJ, Wang XM, Ren SX (2015) A new genus of the tribe Aspidimerini (Coleoptera Coccinellidae) from the Oriental Region. Annales Zoologici 65(2): 171–185. doi: 10.3161/00034541ANZ2015.65.2.005
  • Huo LZ, Wang XM, Chen XS, Ren SX (2013) The genus Aspidimerus Mulsant, 1850 (Coleoptera, Coccinellidae) from China, with descriptions of two new species. ZooKeys 348: 47–75. doi: 10.3897/zookeys.348.5746
  • Huo LZ, Wang XM, Chen XS, Ren SX (2014) The genus Pseudaspidimerus Kapur, 1948 (Coleoptera: Coccinellidae) from Laos, with descriptions of two new species. The Pan-Pacific Entomologist 90(4): 182–190. doi: 10.3956/2014-90.4.182
  • Kapur AP (1948) A revision of the tribe Aspidimerini Weise (Coleoptera, Coccinellidae). Transactions of the Royal Entomological Society of London 99: 77–128. doi: 10.1111/j.1365-2311.1948.tb01233.x
  • Kapur AP (1963) The Coccinellidae of the Third Mount Everest Expedition, 1924 (Coleoptera). Bulletin of the British Museum (Natural History) Entomology 14: 1–48.
  • Kovář I (2007) Coccinellidae. In: Löbl I, Smetana A (Eds) Catalogue of Palaearctic Coleoptera, Volume 4. Apollo Books, Stenstrup, 568–631.
  • Kuznetsov V, Pang H (1991) Description of Four New Aspidimerini (Coleoptera, Coccinellidae) from Vietnam. Elytra, Tokyo 19(2): 185–190.
  • Mader L (1954) Weiteres über Coccinelliden aus der Sammlung des Naturhistorischen Museums in Wien. Koleopterologische Rundschau, 32(1951–1954): 123–131.
  • Mulsant E (1850) Species des Coléoptères trimères sécuripalpes. Annales des Sciences Physiques et Naturelles, d’Agriculture et d’Industrie, Lyon (2) 2: 1–1104. doi: 10.5962/bhl.title.8953
  • Ohta Y (1929) Scymninen Japans. Insecta Matsumurana 4(1–2): 1–16.
  • Pang H (1998) The new record and new species of Aspidimerinae (Coleoptera: Coccinellidae) from China. Entomotaxonomia 20(3): 185–193.
  • Pang XF, Mao JL (1979) Economic Entomology of China 14: Coleoptera Coccinellidae II. Science Press, Beijing, 170 pp.
  • Poorani J (2002) An annotated checklist of the Coccinellidae (Coleoptera) (excluding Epilachninae) of the Indian subregion. Oriental Insects 36: 307–383. doi: 10.1080/00305316.2002.10417335
  • Ren SX, Wang XM, Pang H, Peng ZQ, Zeng T (2009) Colored pictorial handbook of ladybird beetles in China. Science Press, Beijing, 336 pp.
  • Sasaji H (1968) A revision of the Formosan Coccinellidae (II): Tribe Stethorini, Aspidimerini and Chilocorini (Coleoptera). Etizenia, Fukui (32): 1–24.
  • Ślipiński A (2007) Australian ladybird beetles (Coleoptera: Coccinellidae) their biology and classification. ABRS, Canberra, 286 pp.
  • Ślipiński A, Tomaszewska W (2010) Coccinellidae Latreille, 1802. In: Leschen RAB, Beutel RG, Lawrence JF (Eds) Handbook of Zoology, 2, Coleoptera. Walter de Gruyter GmbH & Co KG, Berlin, New York, 454–472.
  • Weise J (1895) Deutsche Entomologische Zeitschrift, 1895: 326.
  • Weise J (1900) Coccinelliden aus Ceylon gesammelt von Dr. Horn. Deutsche Entomologische Zeitschrift 44: 417–448. doi: 10.1002/mmnd.48019000237
  • Weise J (1910) Uber Chrysomeliden und Coccinelliden der Philippinen. Philippine Journal of Science 5D(4): 229–232.
  • Xiao NN, Li HX (1992) Coleoptera: Coccinellidae. In: Huang FS (Ed.) Insects of Wuling Mountains area, southwestern China. Science Press, Beijing, 368–390.
  • Yu GY (2011) The Coccinellidae of Taiwan. Chemical Industry Press, Beijing, 198 pp.