Nisotra gemella (Erichson)

Lee, Chi-Feng, Chiang, Ming-Yao, Geiser, Michael F. & Chuang, Kuo-Hung, 2024, The genus Nisotra Baly, 1864 (Coleoptera, Chrysomeliae, Galerucinae, Alticini) in Taiwan, with redescriptions of four Asian species and notes on the immature stages of N. gemella (Erichson, 1834), ZooKeys 1205, pp. 299-331 : 299-331

Nisotra gemella (Erichson)

Figs 3 G – I , 6 A – N , 7 , 8 , 9 , 10 , 11 , 12

Haltica gemella Erichson, 1834: 275 ( Philippines: Luzon).

Nisotra gemella : Jacoby 1885: 34 ( Indonesia: Java); Scherer 1969: 148 (illustration of male aedeagus); Kimoto and Takizawa 1973: 177 ( Nepal); Scherer 1979: 138 ( India, Bhutan); Gruev 1985: 40 ( Nepal); Kimoto 1986: 60 (additional records in Taiwan); Kimoto 1987: 191 (additional records in Taiwan); Kimoto 1989: 263 (additional records in Taiwan); Kimoto 1991: 16 (additional records in Taiwan); Medvedev 1993: 17 ( Philippines: Luzon, Palawan (misidentification! )); Kimoto 2000: 233 ( Laos, Thailand); Medvedev 2000: 21 ( Nepal); Kimoto 2001: 201 (catalogue); Aston 2009: 5 ( China: Hong Kong).

Nisotra gemellata [sic!]: Duvivier 1885: 49 ( Indonesia: Sumatra).

Sphaeroderma javana Motschulsky, 1866: 421 ( Indonesia: Java). Synonymized with N. gemella by Medvedev (2006). Synonym confirmed.

Podagrica javana : von Harold 1876: 3481.

Nisotra javana : Weise 1922: 126; Ogloblin 1930: 106.

Sphaeroderma orbiculata Motschulsky, 1866: 421 ( India); Harold 1876: 3548 (catalogue). Synonymized with N. gemella by Scherer (1969). Synonym confirmed.

Nisotra orbiculata : Ogloblin 1930: 106; Chen 1933: 55 ( China: Guandong); Chen 1934 a: 181 (Taiwan); Chen 1934 b: 278 (South China, Vietnam, India); Chûjô 1935: 474 (catalogue of Taiwan fauna); Gressitt and Kimoto 1963: 793 ( China: Sichuan, Jiangxi, Guanxi, Fujian, Hainan Island); Kimoto 1966: 35 (additional records in Taiwan); Scherer 1969: 148 ( India); Kimoto 1970: 215 (New Taipei City: Pinglin (坪林 )); Takizawa et al. 1995: 14 (additional records in Taiwan).

Nisotra bowringi Baly, 1876: 584 ( China: Hong Kong); Jacoby 1889: 196 ( Myanmar). Synonymized with N. orbiculata by Ogloblin (1930). Synonym confirmed.

Podagrica bowringi : Maulik 1926: 278 ( India).

Podagrica hibisci Bryant, 1941: 286 ( Malaysia; host plant: Hibiscus rosa-sinensis L.). Synonymized with N. gemella by Scherer (1969). Synonym confirmed.

Types.

Haltica gemella . Lectotype ♂ ( MNHUB, here designated to preserve stability of nomenclature), labeled (Fig. 6 A – C ): “ Manila / Meyer [h, y] // Paratypus [p, r] // HOLOTYPUS [p, red letters] / Nisotra ♂ / gemella / (Erichson) [h, black letters] / det Dr. G. Scherer 19 [p, red letters] 67 [h, w, black letters] ”. Paratypes ( MNHUB): three specimens bear same labels as lectotype except “ PARATYPUS [p, red letters] / Nisotra ♂ or ♀ / gemella / (Erichson) [h, black letters] / Dr. G. Scherer 19 [p, red letters] 67 [h, w, black letters] ”. One specimen labeled (Fig. 6 D ): “ 45025 [p, w] // Haltica / gemella / Eri / Manila / Meyer [h, y] // Typus [p, r] // PARATYPUS [p, red letters] / Nisotra ♂ / gemella / (Erichson) [h, black letters] / Dr. G. Scherer 19 [p, red letters] 67 [h, w, black letters] ”. Although labels for holotype and paratypes were written by Scherer, but it was not officially published.

Sphaeroderma javana . Lectotype ♂ ( ZIN, designated by Medvedev 2006) (Fig. 6 E – G ): “ Sphaeroderma / javana / Motsch. / Java [h, y] // LECTOTYPUS / Nisotra / javana Motsch. / L. Medvedev design. [p, r] ”. Medvedev (2006) indicated that there are three paralectotypes female bearing same labels.

Sphaeroderma orbiculata . Lectotype ♂ ( ZIN, designated by Medvedev 2006) (Fig. 6 H – J ): “ Sphaeroderma / orbiculata / Motsch / Ind. or. [h, y] // Nisotra / orbiculata Mts / (= borwing Baly / 1927 [h] D. Ogloblin det. [p. w] // LECTOTYPUS / Nisotra / orbiculata Motsch. / L. Medvedev design. [p, r] ”.

Nisotra bowringi . Lectotype ♀ ( NHMUK, here designated to preserve stability of nomenclature) (Fig. 6 K, L ), labeled: “ Hong / Kong [h, w, circle label] // Nisotra / Bowringi / Baly / Hong Kong [h, b] // Type / H. T. [p, w, circle label with red border] ” . Paralectotype: 1 ♀ ( NHMUK), shared with the same pin as lectotype.

Podagrica hibisci . Lectotype ♀ ( NHMUK, here designated to preserve stability of nomenclature) (Fig. 6 M, N ), labeled: “ 01880 [h] / Malaya [p] / Kula Lumpur / 16.12. [h] 19 [p] 37 [h] / Entom. Div. [p] / Hibiscus / Rosa-sinensis [h, w] // Type [p, r, circle label with red border] // Pres. by / Imp. Inst. Ent. / B. M. 1941-61. [p, w] // Podagrica / hibisci / Bryant [h] / Det. G. E. Bryant [p, w] // NHMUK 015024484 [p, w, with QR-Code] ” . Paralectotype. 1 ♀ ( NHMUK), bearing same labels as lectotype but without type and QR-Code labels.

Additional material examined.

China. Fujian: 1 ♀ ( TARI), Chishui (赤水), 20. VI. 2014, leg. Y. - T. Chung ; 1 ♂ ( KMNH), Chungan: Bohea Hills , 11. I. 1940, leg. T. C. Maa ; 1 ♀ ( TARI), Zhaizhong (宅中), 24. VIII. 2014, leg. Y. - T. Wang ; Guandong: 1 ♀ ( KMNH), Fei-ha to Fei-loi , 1. VII. 1950, leg. J. L. Gressitt ; 1 ♀ ( NHMUK), nr. Canton (= Guanzhou , 廣州), 27. IX. 1980, leg. P. M. Hammond, B. M. 1980-491 ; Guanxi: 1 ♂ ( NHMUK), Huanjiang Xian, Maonan , leg. M. Häckel ; Hainan: 1 ♀ ( TARI), Nanhsitsum (南喜村), 15. XI. 2018, leg. Y. - T. Wang ; Hong Kong: 1 ♂, 1 ♀ ( NHMUK), coll. Walker, 1893—58 ; Hunnan: 1 ♀ ( NHMUK), mts. Daiongshan, Xinhua , VII. 2004, leg. Jing ; Yunnan: 1 ♂ ( TARI), Bakaxiaozhai (巴卡小寨), 1. VIII. 2017, leg. Y. - T. Wang ; 1 ♂ ( TARI), Chinoshan (基諾山), 31. V. 2018, leg. Y. - T. Wang ; 4 ♂ ( TARI), Manfen (曼粉), 20. IX. 2017, leg. Y. - T. Wang ; 1 ♀ ( TARI), Menglun (勐侖), 2. VIII. 2017, leg. Y. - T. Wang ; 1 ♀ ( TARI), same but with “ 20. IV. 2018 ” ; 2 ♀ ( TARI), same but with “ 2. V. 2019 ” ; 1 ♀ ( TARI), Mohan (磨憨), 14. V. 2016, leg. Y. - T. Wang ; 1 ♀ ( TARI), Nabang (那邦), 25. IX. 2018, leg. Y. - T. Wang ; 2 ♀ ( TARI), Ruili (瑞麗), 15. IX. 2014, leg. Y. - T. Wang ; Zhejiang: 3 ♂ ( NHMUK), Da-Laen-Saen, near Ning-Po , col. Walker, 1893-18 ; 2 ♂ ( NHMUK), same locality, leg. J. J. Waliker, G. C. Champion Coll., B. M. 1927-409 ; India. Andaman Islands : 3 ♂ ( NHMUK), Capt. Wimberley, Fry Coll. 1905 100 ; 2 ♂, 1 ♀ ( NHMUK), same locality, leg. F. A. de Roepstorff, 1884-15 ; 1 ♂, 1 ♀ ( NHMUK), Port Blair , 10. X. 1989, leg. B. S. Bhumannava, feeding on Urena lobata ; Assam: 1 ♀ ( KMNH), Kaziranga nödl. Mikir-Hills, Brahmaputra , V. 1961, leg. G. Scherer ; Sikkim: 1 ♀ ( NHMUK), Dikchu , 23. IV. 1924, leg. R. W. G. Hingaton ; Tamil Nadu: 5 ♂ ( SEHU), Nilgiri, Mettupalayam , 30. II. 1978 ; Uttarakhand: 1 ♀ ( NHMUK), Ranikhet, Kumaon , coll. H. G. Champion, 1953-156 ; 1 ♂ ( SEHU), Mohand Forest nr Dehar Dun , 7–9. XI. 1978 ; 3 ♂, 1 ♀ ( SEHU): FRI, Dehra Dun , 10–13. XI. 1978 ; Indonesia. Java: 1 ♀ ( SEHU), Bremi, Mt. Argopuro , 31. V. 2005, leg. H. Takizawa ; 9 ♂, 12 ♀ ( MCSN), Buitenzorg (= Bogor), 1875, leg. G. B. Ferrari ; 1 ♀ ( MCSN), same but with “ X. 1872 ” ; 1 ♀ ( MCSN), same locality, 1878, leg. D. Lansberge ; 1 ♂, 1 ♀ ( SDEI); Sulawesi: 1 ♀ ( SEHU), Kendari: Amoito , 1. XII. 1974, leg. K. Kusigemati ; 1 ♂ ( SEHU), Kendari: Andonuhu , 3. XII. 1974, K. Kusigemati ; 3 ♂, 3 ♀ ( MCSN), Kandari , III. (18) 74, leg. O. Beccari ; Sumatra: 2 ♂ ( SEHU), Aceh, Kota Cane , 26–28. IV. 1998, leg. H. Takizawa ; 3 ♂, 4 ♀ ( MCSN), Ayer Manicior (= Ajer Mantjoer), VIII. 1878, leg. O. Beccari ; 2 ♂, 1 ♀ ( SDEI), Bangkei Island , 1885, leg. H. Kühn ; 1 ♀ ( MCSN), Benculen , IV. 1891, leg. E. Modigliani ; 1 ♂, 1 ♀ ( SEHU), Bukit Gompong, Sukarami , NE 20 km from Padang, 19. VIII. 1998, leg. M. Ohara ; 1 ♂ ( SEHU), Jambi, Lapak Aur , 6. II. 2006, leg. H. Takizawa ; 1 ♂ ( MCSN), Padang , 1890, leg. E. Modigliani ; 1 ♂ ( MCSN), Sing Hara , X. 1878, leg. O. Beccari ; 1 ♂, 2 ♀ ( SDEI), Tebing Tinggi , leg. Schultheiss ; Laos. 3 ♂, 4 ♀ ( NHMUK), Betw. Vientiane & Luang Prabang , end of 1919, leg. R. V. de Salvaza ; Attapu: 1 ♂ ( NHMUK), Bolaven Plateau, 15 km SE of Ban Huang Kong, Nong Lom (lake), 18–30. IV. 1999, leg. E. Jendek & O. Šauša ; Bolikhamsai: 1 ♂, 1 ♀ ( NHMUK), Lak Sao , 18. VIII. 2004, leg. M. Geiser; Borikhan : 1 ♂ ( NHMUK), Borikhan env., 20 km N of Muang Pakxan , 18. V. 2003, leg. O. Šafránek ; Champasak: 1 ♀ ( NHMUK), Bolaven Paleau, Rout (No. 23), Pakse-Paksong , Ban Itou env (km 35), 10–18. IV. 1998, leg. E. Jendek & O. Šauša; Hua Pan : 1 ♀ ( NHMUK), Ban Saleui, Phou Pan Mt. , 11. IV. - 15. V. 2012, leg. C. Holzschuh ; 1 ♂ ( NHMUK), same but with “ 27. IV. – 1. VI. 2011 ” ; Phongsaly: 1 ♂ ( NHMUK), Phongsaly, Phu Fa , 26. VII. 2006, leg. M. Geiser ; Savannakhét: 1 ♂ ( SEHU), 1. V. 2006, leg. K. Maruyama ; Vientiane: 1 ♀ ( SEHU), Vang vieng , 28–29. V. 2004, leg. T. Tsuru ; 1 ♀ ( NHMUK), same locality, 20. VII. 2006, leg. M. Geiser ; 2 ♂, 1 ♀ ( NHMUK), same but with “ 21. VII. 2006 ” ; Malaysia. Kuala Lumpur: 1 ♂ ( SEHU), Labu , 31. III. 2007, leg. H. Takizawa ; Perak: 4 ♂, 2 ♀ ( MCSN), 25 km NE of Ipoh, Banjaran Titi Wangsa mts., Korbu mt. , 27. I. – 2. II. 1999, leg. P. Čechovský ; 3 ♂, 2 ♀ ( MCSN), same but with “ 4–13. III. 1998 ” ; Sabah: 1 ♂, 2 ♀ ( SDEI), Kinabalu , S. W. 11, leg. H. Rolle ; Myanmar. 1 ♂, 1 ♀ ( SDEI), Carin Cheba , V. – VII. - (18) 88, leg. L. Fea ; 1 ♂, 1 ♀ ( NHMUK), same locality, coll. Fry, 1905.100 ; 1 ♀ ( MCSN), Carin, Asciuii Cheba , I. (18) 88, leg. L. Fea ; 4 ♂, 4 ♀ ( SDEI), Palon , Pegu, VIII. – IX. (18) 87, leg. L. Fea ; 1 ♀ ( NHMUK), Momeit , leg. Doherty, Fry Coll., 1905.100 ; 1 ♀ ( NHMUK), Paungde , coll. Andrewes, 1922-221 ; 1 ♂, 2 ♀ ( NHMUK), Prome , coll. Andrewes, 1922–221 ; 2 ♂, 2 ♀ ( SDEI), Rangoon (= Yangon), 1887, leg. Fea ; 1 ♀ ( NHMUK), same but with “ Atkinson Coll., 1892–3 ” ; 2 ♀ ( NHMUK), same but with “ coll. Fry, 1905.100 ” ; 1 ♂ ( NHMUK), same locality, 1933–34, leg. F. J. Meggitt ; 4 ♂, 4 ♀ (4 ♂, 2 ♀: SDEI; 1 ♀: MCSN; 1 ♀: NHMUK), Teinzo , V. 1886, leg. Fea ; 1 ♂, 2 ♀ ( NHMUK), Tharrawaddy , leg. H. Swale, 1913-117 ; 4 ♀ ( NHMUK), same locality, leg. H. E. Andrewes, 1922-221 ; 2 ♂, 3 ♀ ( NHMUK), same locality, 26. XII. 1953, leg. H. G. Champion ; 1 ♀ ( NHMUK), Toungoo , 26. XII. 1953, leg. H. G. Champion ; Kachin State: 1 ♂ ( MCSN), VIII. 1885, leg. Fea ; 1 ♀ ( NHMUK), Bhamo , VIII. 1885, leg. Fea ; 2 ♂, 1 ♀ ( NHMUK), Nam Tamai , 2. VIII. 1938, leg. R. Kaulback ; Kayin Sate: 1 ♀ ( TARI), Than Daung Gyi , 18. V. 2017, leg. Y. - T. Wang ; Yangon: 2 ♂, 2 ♀ ( NHMUK), Shwentha, 23. VII. 1988, leg. T. T. Nwe, collected from Urena lobata ; Nepal. 3 ♂, 2 ♀ ( SEHU), Balaju Kathmandu Valley , 11. IX. 1987, leg. H. Takizawa ; Papua New Guinea. 1 ♀ ( SDEI), Haveri , VII. – XI. (18) 93, leg. Loia ; 1 ♀ ( SDEI), Ighibirei , VII. - VIII. (18) 90, leg. Loria ; 1 ♂ ( TARI), West Highland, Begesin Missions station 43 km 237 ° von Madan, Kulturlandschan , 6. V. 1996, leg. H. Deumer ; Philippines. Luzon. Kalinga: 1 ♂ ( MCSN), Tulgao , 23. VI. 1988, coll. Medvedev ; Singapore. 3 ♂, 1 ♀ ( NHMUK), leg. C. J. Saunders, B. M. 1933-227 ; Taiwan. Hsinchu: 3 ♂, 3 ♀ ( TARI), Hsinpu (新埔), 2. I. 2010, leg. K. - H. Chuang ; 18 ♂, 23 ♀ ( TARI), same but with “ 5. X. 2021 ” ; 12 ♂, 14 ♀ ( TARI), same but with “ 18. XII. 2021 ” ; Kinmen: 2 ♂, 7 ♀ ( TARI), Kinmen Island (金門島), 18. VI. 2014, leg. Y. - T. Chung ; 4 ♂, 8 ♀ ( TARI), same island, Jhongshanlin (中山林), 28. VI. 2023, leg. C. - F. Lee ; 5 ♀ ( TARI), 太武苗圃 (= Kinmen Botanical Gardens , 金門植物園), 24. IV. 2002, leg. H. T. Shih ; 1 ♀ ( TARI), same but with “ Taiwushan (太武山) ” ; Matsu: 1 ♀ ( TARI), Beigan Island (北竿島), 14. V. 2018, leg. H. - T. Fang ; New Taipei City: 1 ♀ ( KMNH), Pinglin (坪林), 23. VI. 1965, leg. Y. Kurosawa ; 4 ♂ ( TARI), Watzuwei (挖仔尾), 28. VI. 2008, leg. H. Lee ; 1 ♂ ( KMNH), Urai (烏來), 31. V. 1976, leg. H. Makihara ; Nantou: 1 ♀ ( SDEI), Fuhosho (= Wucheng , 五城), IX. 1909, leg. Sauter ; Pingtung: 1 ♂, 1 ♀ ( TARI), Shantimen (三地門), 1–5. III. 1982, leg. K. C. Chou & C. C. Pan ; 1 ♀ ( TARI), Tahan trail (大漢林道), 10. IV. 2023, leg. J. - C. Chen ; Taipei: 1 ♀ ( SDEI), Hokuto (= Peitou, 北投), III. 1912, leg. Sauter ; Taitung: 2 ♀ ( SDEI), Paroe (= Tawu, 大武), IX. 1912, leg. H. Sauter ; Taoyuan: 1 ♀ ( TARI), Tayuan (大園), 15. VIII. 2011, leg. L. - F. Chu ; Thailand. Chiang Mai: 1 ♀ ( NHMUK), Fah Luan Univ. Campus , 20–22. VII. 2009, leg. D. Quicke & B. et R. Butcher ; 1 ♀ ( NHMUK), Queen Sirikit Botanic Gardens , VII. 2006, leg. H. Mendel & M. V. L. Barclay ; 1 ♂ ( NHMUK), Tha Ton Env. , 20. IV. 2003, leg. O. Šafránek ; Kanchanaburi: 1 ♂, 1 ♀ ( NHMUK), Thongpapoom , 11–12. VII. 2009, leg. D. Quicke & B. et R. Butcher ; Mae Hong Son: 1 ♂ ( NHMUK), Mae Hong Son , 17–21. VI. 1993, leg. Schneider ; Nakhon Nayok: 2 ♂, 1 ♀ ( SEHU), Ban Na , 27. VII. 1997, leg. S. Ohmomo ; Nan: 1 ♂, 2 ♀ ( NHMUK), Doi Phuka N. P. , V. 2000, leg. local collector ; Songkhla: 1 ♀ ( NHMUK), Chon Thong , 24–27. IV. 1991, leg. L. Dembický ; Yala: 1 ♂ ( NHMUK), Betong, Gunung Cang Dun vill., 25. III. – 22. IV. 1993, leg. J. Horák ; Vietnam. Băc Giang: 2 ♂, 2 ♀ ( TARI), Tây Yên Tů , 2. VI. 2014, leg. Y. - T. Wang ; Hà Nôi: 1 ♂, 1 ♀ ( TARI), Huyên Mê Linh , 30. V. 2014, leg. Y. - T. Wang ; Hòa Binh: 3 ♀ ( NHMUK), VIII. 1918, leg. R. V. de Salvaza ; 1 ♂ ( MCSN), Ha Son-Binh , 10 km SW Hoa Binh, 17. X. 1976, leg. L. Medvedev ; Lao Cai: 2 ♂ ( NHMUK), Bao Ha , 12. IV. 1962, leg. A. Warchalowski ; 1 ♂ ( NHMUK), same but with “ 4. IV. 1962 ” ; 2 ♂, 5 ♀ ( NHMUK), Cha Pa (= Sa Pa), 14. IV. 1962, leg. A. Warchalowski ; Nam Dinh: 1 ♀ ( NHMUK), Van Diem , 19. III. 1962, leg. A. Warchalowski ; 1 ♂ ( NHMUK), same but with “ 7. II. 1962 ” ; Ninh Binh: 1 ♂ ( NHMUK), Cuc Phuong , 8. VI. 1966, leg. R. Bizlawski & B. Pisarski ; Quang Ninh: 1 ♂ ( TARI), 10 km SE Tien Yen , 1–14. IV. 2004, leg. H. Mühle ; Vinh Phuc: 1 ♂, 2 ♀ ( NHMUK), Tam Dao , 25. IV. 1962, leg. A. Warchalowski ; 1 ♂, 1 ♀ ( NHMUK), same locality, 17. X. 1966, leg. J. Kania .

Redescription.

Adults. Length 3.1–3.9 mm, width 2.0– 2.5 mm (n = 376). General color yellowish brown (Fig. 3 G – I ); elytra, meso- and metathoracic, and abdominal ventrites metallic purple; four basal antennomeres I – IV yellowish brown, V dark brown, VI – XI black. Antennae (Fig. 7 A ) filiform in males, ratios of lengths of antennomeres I to XI 1.0: 0.5: 0.6: 0.5: 0.5: 0.5: 0.5: 0.6: 0.6: 0.7: 1.0; ratios of length to width from antennomeres I to XI 3.2: 2.4: 2.9: 2.6: 2.2: 2.2: 2.2: 2.6: 2.6: 2.6: 4.2; similar in females, ratios of lengths of antennomeres I to XI (Fig. 7 B ) 1.0: 0.5: 0.5: 0.5: 0.5: 0.5: 0.6: 0.6: 0.6: 0.6: 1.0; ratios of length to width from antennomeres I to XI 3.6: 2.3: 2.9: 2.5: 2.4: 1.9: 2.0: 2.2: 2.6: 2.2: 3.8. Pronotum 1.8–1.9 × wider than long; disc shining, moderately convex, with sparse, fine punctures; longitudinal groove on each side of apical margin reduced; longitudinal groove on basal margin short and shallow; lateral margins rounded; apical margins slightly concave; basal margin medially convex. Elytra 1.2 × longer than wide; disc with coarse punctures arranged into longitudinal lines, with fine punctures between coarse punctures; lateral margins rounded, narrowed behind middle. Aedeagus (Fig. 7 C, D ) wide, ~ 4.8 × longer than wide; parallel sided, but slightly wider at apical 1 / 7, and then moderately and apically narrowed; apex truncate, but with one median rounded process; strongly curved in lateral view, apex directed upward; tectum membranous, with several stout setae at apex of internal sac. Endophallic spiculae reduced. Gonocoxae (Fig. 7 G ) longer than wide, and basally connected; apex of each gonocoxa widely rounded, with eight long setae along apical margin and outer margin. Ventrite VIII (Fig. 7 E ) with apex weakly sclerotized, with one semicircular membranous area at middle of apical margin, several long setae arranged into transverse line near apical margin, and several shorter setae along apical margin, both types of setae absent medially. Spiculum extremely long. Spermathecal receptaculum (Fig. 6 F ) strongly swollen; pump long and curved, with long apical process; sclerotized spermathecal duct moderately long after base of spermathecal gland.

Variations. Adults in some populations have more ovate body shapes (elytra 1.1 × as long as wide), including those on the Andaman Islands ( India).

Remarks. Adults of N. gemella are characterized by more convex pronotum and reduced longitudinal grooves on the sides of the apical margins (less convex pronotum and long, deep longitudinal grooves on sides of apical margins in others). In males of N. gemella , the truncate apex of the aedeagus bears a small process (Fig. 7 C ) that differs from the acute apex in N. chrysomeloides (Fig. 4 C ) and the widely rounded apex and small medial process in N. nigripes (Fig. 14 C ). The moderately curved aedeagus in lateral view (Fig. 7 D ) differs from the strongly curved aedeagus in N. dohertyi (Fig. 5 E ) and slightly curved aedeagus in N. nigripes (Fig. 14 D ). The membranous tectum (Fig. 7 C ) differs from the sclerotized tectum in N. chrysomeloides (Fig. 4 C ). In females of N. gemella , the straight gonocoxae (Fig. 7 G ) differ from the dorsally directed gonocoxal apices in N. chrysomeloides (Fig. 4 H ), the inwardly directed apices in N. dohertyi (Fig. 5 I ), and laterally directed apices in N. nigripes (Fig. 14 G ). Abdominal ventrite VIII, with one transverse line of long setae inside the apical margin and dense short setae along the apical margin (Fig. 7 E ), differs from ventrite VIII of N. dohertyi (Fig. 5 F, G ), with several pairs of long setae along the apical margin.

Third-instar larvae. Length 4.8 mm, width 1.0 mm, cylindrical, cream colored, with tubercles well-developed, head dark brown (Fig. 8 A, B ). Head (Fig. 9 A ) hypognathous, rounded, strongly sclerotized; surface generally smooth. Endocarina (ec) visible, extending into clypeus. Frontal suture (fs) V-shaped, starting from apical 1 / 3 of endocarina; apically abbreviated halfway between endocarina and lateral margin. Vertex with four pairs of setae (v 1 – v 4). Frons with three pairs of setae (f 1 – f 3). Sides with five pairs of setae (s 1 – s 5). Clypeus transverse, apical margin straight, with one pair of small setae at sides. Labrum transverse, apical margin irregular, with two pairs of long setae; epipharynx (Fig. 9 E ) with several setae at sides, and flattened, apically pointed setae. Mandible (Fig. 9 F ) robust; five-toothed, with tiny serrations on mesal margins of teeth; two setae on opposite sides; mola reduced; prostheca composed of several flattened, slender setae. Maxillary palpus (mp) (Fig. 9 B ) three-segmented, palpomere II with one long seta at one side and one short seta at opposite side; palpifer (pf) with one long seta at opposite sides; mala (ml) with two-segmented conical sensorium and surrounding by three pairs of short setae, mesal side margined with one row of elongate, flattened setae with rounded apices; stipes (st) with one long seta and one short seta. Labial palp (lp) two-segmented, palpomere II cylindrical, palpomere I much wider than II; ligula (lg) rounded, with one pair of short setae; prementum (prm) with one pair of long setae; postmentum (pom) with two pair of long setae. Stemmata reduced and invisible. Antenna (Fig. 9 C, D ) three-segmented, one tubercle with one prominent and several tiny conical sensoria on antennomere II, one large conical sensorium on antennomere III.

Prothorax (Fig. 10 ) with dorsal, dorso-lateral, and anterior epipleural tubercles fused, D- DL - EPa type; basal and apical margins with two pairs of setae, lateral margins with four pairs of setae; epipleural region with one small tubercle EPp, with one seta; trochantin (ti) with one seta; sternal regions with two tubercles: ES and SS, ES not divided medially, with one pair of setae, SS separated, with one seta on each tubercle; one additional seta between ES and SS. Meso- and metathoraces with dorsal tubercle divided into Dai, Dae, Dpi, and Dpe; with one pair of setae on each tubercle, except Dae; DL tubercle not divided, with two setae; epipleuron with one tubercle, EP, surrounding spiracle, with one seta; trochantin (ti) without setae; sterna with two tubercles, ES and SS, separated medially, with one seta on each tubercle. Legs (Fig. 8 C ) five-segmented, coxa (co) with four long setae apically and two short setae basally; trochanter (tr) with four long mesal setae, one long seta apically, and one short seta at lateral area; femur (fe) with three pairs of apical setae on opposite; tibia (tb) with one short mesal seta near margin, two pairs of setae near outer margin, and one pair of setae apically; tarsungulus (ta) slightly curved, with broad base, with one mesal seta near margin of broad base.

Abdominal segment I – VIII (Fig. 10 ) with dorsal tubercles divided into three tubercles: Da, Dpi, and Dpe, Da and Dpi not divided medially, Da with two pairs of setae, Dpi with one pair of setae, Dpe with one seta; dorso-lateral region with two tubercles: DLa and DLp, with one seta on each tubercle; epipleural tubercle with one seta separated from spiracle, spiracle rounded, tubercle P with one seta; sternal region with three tubercles: PS, ES, and SS; ES and SS not divided medially, ES with one pair of setae but SS with two pairs of setae; PS large and with two setae. Abdominal segment IX covered by pygopod dorsally (Fig. 7 E ); pygopod oblong, slightly and apically widened, with three pairs of setae margined apically; ventral region (Fig. 8 D ) sclerotized and not divided, with two pairs of setae.

Pupa. Length 3.8 mm; width 1.4 mm, yellowish white. Head (Fig. 11 B ) with three pairs of setae: one pair on vertex, one pair on mesal margins of eyes, the other pair between antennae. Prothorax (Fig. 11 A ) with five pairs of setae on outer margin; one pair on baso-lateral angles; two pairs near basal margin and close to each other, outer pair shorter; one pair at central part near middle line. Meso- and metathoraces with two pairs of setae. Abdominal segment I – VIII (Fig. 11 A ) with four pairs of setae; setae on abdominal segment I – VI near posterior margin; VII (Fig. 11 C ) expanded posteriorly, setae along posterior margin; VIII similar to VII but narrower; IX with apical processes sclerotized at apex and strongly pointed, two pairs of setae at sides, two pairs on base of apical processes, one on each, dorsal and ventral surfaces, respectively. Bases of femora of front and middle legs with three pairs of setae, but only two pairs of setae on bases of femora of hind legs.

Host plants.

Malvaceae : Hibiscus rosa-sinensis ( Bryant 1941 and current study) (Fig. 11 F ), H. tiliaceus (current study), Urena lobata ( Kimoto 2003) , and Lamiaceae : Mesona chinensis (current study). Takizawa (1979) recorded two species of Urticaceae ( Boehmeria nivea var. nivea and Gonostegia hirta ) as host plants for Taiwanese populations of Nisotra orbiculata (= N. gemella ) but this may be incorrect because flea beetles collected from both plants belong to the genus Euphitrea Baly. Both plants were also regarded as host plants for Neorthaea (= Euphitrea ) nisotroides Chen and Neorthaea flavicornis Chen in the same paper ( Takizawa 1979). We assessed five plants for adults of Nisotra gemella , which fed originally on Mesona chinensis in the laboratory. These five were Hibiscus rosa-sinensis , H. tiliaceus , Urena lobata ( Malvaceae ), Boehmeria nivea var. nivea ( Urticaceae ), and Mentha canadensis ( Lamiaceae ). They did not feed on leaves of Boehmeria nivea var. nivea or Mentha canadensis .

Biology.

Females deposited eggs on the soil or leaf litter (Fig. 12 A ). Each egg (Fig. 12 B ) was oblong and pale yellow, 1.0 mm in length and 0.5 mm in width. Larvae hatched after 10–14 days, and have three instars Larvae were pale yellow with black heads, pronota, and pygopods (Fig. 12 C ). They fed on roots. Larval durations varied from 25–46 days. Mature larvae (Fig. 12 D ) crawled into the soil and constructed underground chambers for pupation. Pupal stage (Fig. 12 E ) durations were 7–8 days. Adults fed by cutting the leaf lamina. They jumped promptly when disturbed. Adults remained on the adaxial side of leaves.

Remarks.

The specimens collected from Palawan, Philippines ( Medvedev 1993) and Vietnam, were misidentified and represent Nisotra chrysomeloides . The same error has been made by other taxonomists.

Distribution.

Cambodia, China, India (including Andaman and Nicobar islands), Indonesia (Sumatra, Java, Sulawesi), Laos, Malaysia, Myanmar, Nepal, Papua New Guinea, Philippines (Luzon), Singapore, Thailand, Taiwan, and Vietnam.

Figures

Figure 3. Habitus of Nisotra species A N. chrysomeloides Jacoby, female, dorsal view B ditto, ventral view C ditto, lateral view D N. dohertyi (Maulik), female, dorsal view E ditto, ventral view F ditto, lateral view G N. gemella (Erichson), female, dorsal view H ditto, ventral view I ditto, lateral view.

Figure 4. Nisotra chrysomeloides Jacoby, adult A antenna, male B antenna, female C apex of aedeagus, front view D base of aedeagus, dorsal view E aedeagus, lateral view F abdominal ventrite VIII, female G spermatheca H gonocoxae.

Figure 5. Nisotra dohertyi (Maulik), adult A antenna, male B antenna, female C apex of aedeagus, front view D base of aedeagus, dorsal view E aedeagus, lateral view F abdominal ventrite VIII, female, from China (Yunnan) G abdominal ventrite VIII, female, from China (Laos) H spermatheca I gonocoxae.

Figure 6. Type specimens and labels A Haltica gemella Erichson, 1834, lectotype B aedeagus dissected from lectotype, left: dorsal view; right: lateral view C labels pinned with lectotype D labels pinned with a paralectotype E Sphaeroderma javana Motschulsky, 1866, lectotype F aedeagus dissected from lectotype, left: dorsal view; right: lateral view G labels pinned with lectotype H Sphaeroderma orbiculata Motschulsky, 1866, lectotype I aedeagus dissected from lectotype, left: dorsal view; right: lateral view J labels pinned with lectotype K Nisotra bowringi Baly, 1876, lectotype L labels pinned with lectotype M Podagrica hibisci Bryant, 1941 N labels pinned with lectotype O Nisotra nigripes Jacoby, 1894, holotype P labels pinned with holotype.

Figure 7. Nisotra gemella (Erichson), adult A antenna, male B antenna, female C aedeagus, dorsal view D aedeagus, lateral view E abdominal ventrite VIII, female F spermatheca G gonocoxae.

Figure 8. Nisotra gemella (Erichson), third-instar larva A larva in dorsal view B larva in lateral view C leg D abdominal ventrite IX E pygopod. Abbreviations: co- coxa; fe- femur; ta- tarsungulus; tb- tibia; tr- trochanter.

Figure 9. Nisotra gemella (Erichson), third-instar larva A head B maxilla and labium C antenna, in lateral view D antenna, in dorso-lateral view E epipharynx F mandible. Abbreviations: ec- endocarina; f 1 – f 3 - setae on frontal area; fs- frontal suture; lg- ligula; lp- labial palp; ml- mala; mp- maxillary palp; pf- palpifer; pom- postmentum; prm- prementum; s 1 – s 5 - setae on sides of head; st- stripes; v 1 – v 4 - setae on vertex.

Figure 10. Nisotra gemella (Erichson), third-instar larva: location of tubercles and body chaetotaxy. Abbreviations: Pr- prothorax; Ms- mesothorax; Mt- metathorax; A 1 – A 8 - abdominal segments I – VIII.

Figure 11. Nisotra gemella (Erichson), pupa A dorsal view B head C abdominal segments VII – IX.

Figure 12. Nisotra gemella (Erichson) A female depositing eggs on soil B eggs C larvae feeding on roots D mature larvae crawling into soil and constructing underground chamber for pupation E pupa F adults attacking Hibiscus rosa-sinensis.

Figure 14. Nisotra nigripes Jacoby, adult A antenna, male B antenna, female C aedeagus, dorsal view D aedeagus, lateral view E abdominal ventrite VIII, female F spermatheca G gonocoxae.