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Exocelina australiae (Clark, 1863) Source of material. The three instar III larvae studied were collected in association with adults at the following locality: South Australia. Ten kilometers North of Forreston, 9.VIII.1999; C. H. S. Watts leg. Diagnosis (instar III). The third instar of Exocelina australiae can easily be distinguished from that of the other species studied in this paper by the following combination of characters: HL = 1.85–1.90 mm; L3> 2.90 mm; U> 0.50 mm; head capsule subquadrate, strongly constricted at level of occipital region (Fig. 54), scale-like sculpticels present over frontoclypeus and parietale; anterior margin of frontoclypeus broadly convex, extending mesally at level of adnasalia (Fig. 54); adnasalia margined with mostly short bluntly rounded teeth (Fig. 65); MP2/MP1> 1.50; GA/MP1> 1.50; LP2/LP1> 2.00; inner margin of stipes lacking a dorsal linear row of spinulae; profemur with more than 5 PV and 5 AV secondary setae; metafemur with more than 4 AV secondary setae; urogomphus composed of one urogomphomere; U/HW Description, instar III (Figs 54–55, 65) Body: Measurements and ratios aimed to characterize body shape are shown in Table 6. Head (Figs 54, 65): Head capsule subquadrate, strongly constricted at level of occiput; anterior margin of frontoclypeus broadly convex, extending mesally at about level of adnasalia; scale-like sculpticels present over frontoclypeus and parietale; adnasalia margined with mostly short bluntly rounded teeth (Fig. 65); HL = 1.85–1.90 mm; A/MP = 1.37–1.41; MP/LP = 2.36–2.41; MP2/MP1 = 1.56–1.67; GA/MP1 = 1.53–1.73; LP2/LP1 = 2.00–2.67; MNL/MNW = 2.72–2.96. Thorax: L3 = 3.71–3.87 mm. Abdomen (Fig. 55): LAS = 1.33–1.63 mm; LAS subcylindrical, not constricted posteriorly at point of insertion of urogomphi. Urogomphus, U = 0.56–0.58 mm, composed of one urogomphomere; U/HW = 0.25–0.26; U/LAS = 0.35–0.44. Chaetotaxy: Parietale with 9–10 temporal spines; position and number of secondary setae on legs are shown in Table 7., Published as part of Alarie, Yves, Michat, Mariano C., Watanabe, Kohei, Shaverdo, Helena, Wang, Liang-Jong & Watts, Chris H. S., 2022, An outlook on larval morphology of Copelatinae diving beetles with phylogenetic considerations (Coleoptera: Adephaga, Dytiscidae), pp. 151-205 in Zootaxa 5175 (2) on page 187, DOI: 10.11646/zootaxa.5175.2.1, http://zenodo.org/record/7003344, {"references":["Clark, H. (1863) Catalogue of the Dytiscidae and Gyrinidae of Australasia, with descriptions of new species. The Journal of Entomology Descriptive and Geographical, 2, 14 - 23."]}

Copelatus alternatus Sharp, 1882 Source of material. All instar larvae studied were associated with adults collected at the following locality: Argentina, Tucumán Province. El Cochuna Provincial Park, 3/ 9-XII-2007, roadside pond; M. C. Michat leg. The identification is firm as C. alternatus is the only Copelatus species known from that locality. Diagnosis (instar III). The third instar of Copelatus alternatus can easily be distinguished from that of the other species studied in this paper by the following combination of characters: HL = 0.90 mm; L3 1.50; GA/MP1> 1.50; 1.30 Description, instar I (Figs 1–14) Body: Measurements and ratios aimed to characterize body shape are shown in Table 2. (1) Michat & Torres (2009) (1) Michat & Torres (2009) Head (Figs 1–9): HL = 0.45–0.47 mm; Head capsule rounded, lacking a clear constriction at level of occiput; occipital suture absent; anterior margin of frontoclypeus lightly convex, extending mesally at about level of adnasalia; A/MP = 1.48–1.60; MP/LP = 1.70–1.79; MP2/MP1 = 1.96–2.17; GA/MP1 = 2.50–3.04; LP2/LP1 = 2.27–2.50; MNL/MNW = 2.75–2.88. Thorax (Figs 10–11): L3 = 0.83–0.88 mm. Abdomen (Figs 12–14): LAS = 0.30–0.32 mm; LAS subcylindrical, not constricted posteriorly at point of insertion of urogomphi. Urogomphus: U = 0.18–0.19 mm, composed of one urogomphomere; U/HW = 0.36–0.39; U/LAS = 0.56–0.65. Chaetotaxy (Figs 1–14): Position and number of additional setae on legs are shown in Table 3 and Figs 10– 11. Description, instar II. As first-instar larva except as follows: Body: Measurements and ratios aimed to characterize body shape are shown in Table 4. Head: HL = 0.65 mm; anterior margin of frontoclypeus more narrowly convex; scale-like sculpticels welldeveloped over frontoclypeus and parietale; adnasalia margined with short bluntly rounded and truncated teeth (see Fig. 63); occipital suture present; A/MP = 1.59–1.64; MP/LP = 1.64–1.76; MP2/MP1 = 2.00–2.14; GA/MP1 = 2.38–2.57; LP2/LP1 = 1.78–1.89; MNL/MNW = 2.60–2.78. Thorax: L3 = 1.32–1.34 mm. Abdomen: LAS = 0.43–0.44 mm; LAS subcylindrical to subconical, less abruptly converging posteriorly at point of insertion of urogomphi. Urogomphus: U = 0.23–0.24 mm, composed of one urogomphomere; U/HW = 0.31–0.32; U/LAS = 0.54–0.56. Chaetotaxy: Parietale with 3–5 temporal spines; position and number of secondary setae on legs are shown in Table 5. Description, instar III (Figs 15–16, 63). As second-instar larva except as follows: Body: Measurements and ratios aimed to characterize body shape are shown in Table 6. Head (Figs 15, 63): HL = 0.88–0.89 mm; A/MP = 1.52–1.56; MP/LP = 1.77–1.90; MP2/MP1 = 1.58–1.75; GA/MP1 = 1.83–2.00; LP2/LP1 = 1.42–1.58; MNL/MNW = 2.43–2.83. Thorax: L3 = 1.74–1.81 mm. Abdomen (Fig. 16): LAS = 0.68–0.73 mm; U = 0.27–0.30 mm; U/HW = 0.27–0.32; U/LAS = 0.37–0.44. Chaetotaxy: Parietale with 3–7 temporal spines. Position and number of secondary setae on legs are shown in Table 7., Published as part of Alarie, Yves, Michat, Mariano C., Watanabe, Kohei, Shaverdo, Helena, Wang, Liang-Jong & Watts, Chris H. S., 2022, An outlook on larval morphology of Copelatinae diving beetles with phylogenetic considerations (Coleoptera: Adephaga, Dytiscidae), pp. 151-205 in Zootaxa 5175 (2) on pages 157-161, DOI: 10.11646/zootaxa.5175.2.1, http://zenodo.org/record/7003344, {"references":["Sharp, D. (1882) On aquatic carnivorous Coleoptera or Dytiscidae. The Scientific Transactions of the Royal Dublin Society, Series 2, 2, 179 - 1003, pls. 7 - 18.","Michat, M. C. & Torres, P. (2009) A preliminary study on the phylogenetic relationships of Copelatus Erichson (Coleoptera: Dytiscidae: Copelatinae) based on larval chaetotaxy and morphology. Hydrobiologia, 632, 309 - 327. https: // doi. org / 10.1007 / s 10750 - 009 - 9853 - 2"]}

Copelatus nakamurai Guéorguiev, 1970 Source of material. The two instar II and five instar III larvae studied were reared ex ovo from adults collected at the following locality: Japan. Kasamatshu-cho, Mudoji, Gifu Prefecture, 4.V.2018. Y. Yoshimura, leg. Diagnosis (instar III). The third instar of Copelatus nakamurai can easily be distinguished from that of the other species studied in this paper by the following combination of characters: HL = 0.80 mm; L3 1.50; GA/MP1> 1.50; 1.30 Description, instar II Body: Measurements and ratios aimed to characterize body shape are shown in Table 4. Head: Head capsule subquadrate, strongly constricted at level of occiput; anterior margin of frontoclypeus narrowly convex, extending well beyond level of adnasalia; scale-like sculpticels lacking over frontoclypeus, welldeveloped over parietale; adnasalia margined with elongate acute teeth (see Fig. 64); HL = 0.61 mm; A/MP = 1.49; MP/LP = 2.06; MP2/MP1 = 1.60; GA/MP1 = 1.91; LP2/LP1 = 2.24; MNL/MNW = 3.11. Thorax: L3 = 1.23 mm. Abdomen: LAS = 0.45 mm; LAS subcylindrical, slightly constricted posteriorly at point of insertion of urogomphi. Urogomphus: U = 0.26 mm, composed of one urogomphomere; U/HW = 0.41; U/LAS = 0.58. Chaetotaxy: Parietale with 8 temporal spines; position and number of secondary setae on legs are shown in Table 5. Description, instar III (Figs 28–29, 64). As second-instar larva except as follows: Body: Measurements and ratios aimed to characterize body shape are shown in Table 6. Head (Figs 28, 64): HL = 0.81–0.84 mm; A/MP = 1.42–1.46; MP/LP = 2.02–2.25; MP2/MP1 = 1.51–1.74; GA/MP1 = 1.75–1.99; LP2/LP1 = 1.55–1.78; MNL/MNW = 2.85–2.98. Thorax: L3 = 1.66–1.72 mm. Abdomen (Fig 29): LAS = 0.66–0.69 mm; Urogomphus, U = 0.31–0.33 mm, composed of one urogomphomere; U/HW = 0.35–0.38; U/LAS = 0.45–0.49. Chaetotaxy: Parietale with 9–10 temporal spines; position and number of secondary setae on legs are shown in Table 7., Published as part of Alarie, Yves, Michat, Mariano C., Watanabe, Kohei, Shaverdo, Helena, Wang, Liang-Jong & Watts, Chris H. S., 2022, An outlook on larval morphology of Copelatinae diving beetles with phylogenetic considerations (Coleoptera: Adephaga, Dytiscidae), pp. 151-205 in Zootaxa 5175 (2) on page 175, DOI: 10.11646/zootaxa.5175.2.1, http://zenodo.org/record/7003344

Copelatus caelatipennis princeps Young, 1963 Source of material. The two larvae studied (one instar II and one instar III) were associated with adults collected at the following locality: USA. Florida, Orlando. Junction International Drive & Hwy 523, 26–28.IX.2016. 28°53.83’N 81°37.92’W; Y. Alarie leg. The identification is firm as C. caelatipennis princeps is the only Copelatus species known from that locality. Diagnosis (instar III). The third instar of Copelatus c. princeps can easily be distinguished from that of the other species studied in this paper by the following combination of characters: HL = 0.80 mm; L3 1.50; GA/MP1> 1.50; 1.30 Description, instar II Body: Measurements and ratios aimed to characterize body shape are shown in Table 4. Head: Head capsule rounded to subquadrate, strongly constricted at level of occiput; anterior margin of frontoclypeus narrowly convex, extending well beyond level of adnasalia; adnasalia margined with short bluntly rounded and truncated teeth (see Fig. 63); scale-like sculpticels well-developed over frontoclypeus and parietale; HL = 0.58 mm; A/MP = 1.41; MP/LP = 1.85; MP2/MP1 = 1.73; GA/MP1 = 2.00; LP2/LP1 = 1.73; MNL/MNW = 2.72. Thorax: L3 = 1.05 mm. Abdomen: LAS = 0.39 mm; LAS subconical, not constricted posteriorly at point of insertion of urogomphi. Urogomphus: U = 0.22 mm, composed of one urogomphomere; U/HW = 0.38; U/LAS = 0.58. Chaetotaxy: Parietale with 7 temporal spines; position and number of secondary setae on legs are shown in Table 5. Description, instar III (Figs 17–18, 63). As second-instar larva except as follows: Body: Measurements and ratios aimed to characterize body shape are shown in Table 6. Head (Figs 18, 63): HL = 0.81 mm; A/MP = 1.50; MP/LP = 1.98; MP2/MP1 = 1.67; GA/MP1 = 2.00; LP2/LP1 = 1.51; MNL/MNW = 2.58. Thorax: L3 = 1.58 mm. Abdomen (Fig. 18): LAS = 0.64 mm; U = 0.34 mm; U/HW = 0.43; U/LAS = 0.52. Chaetotaxy: Parietale with 10–12 temporal spines; position and number of secondary setae on legs are shown in Table 7., Published as part of Alarie, Yves, Michat, Mariano C., Watanabe, Kohei, Shaverdo, Helena, Wang, Liang-Jong & Watts, Chris H. S., 2022, An outlook on larval morphology of Copelatinae diving beetles with phylogenetic considerations (Coleoptera: Adephaga, Dytiscidae), pp. 151-205 in Zootaxa 5175 (2) on pages 161-168, DOI: 10.11646/zootaxa.5175.2.1, http://zenodo.org/record/7003344, {"references":["Young, F. N. (1963) The Nearctic species of Copelatus Erichson (Coleoptera: Dytiscidae). Quarterly Journal of the Florida Academy of Sciences, 26, 56 - 77.","Sharp, D. (1882) On aquatic carnivorous Coleoptera or Dytiscidae. The Scientific Transactions of the Royal Dublin Society, Series 2, 2, 179 - 1003, pls. 7 - 18."]}

Liopterus haemorrhoidalis (Fabricius, 1787) Source of material. The four instar III larvae were collected in association with adults at the following locality: Belarus, Vicebsk Region, area near upy Village, wetland of evinka River, Carex, moss. 12.VIII.1998;H.Shaverdo leg. Diagnosis (instar III). The third instar of Liopterus haemorrhoidalis can easily be distinguished from that of the other species studied in this paper by the following combination of characters: HL = 1.40 mm; L3> 2.90 mm; U> 0.50 mm; head capsule subquadrate, strongly constricted at level of occipital region (Fig. 58), scale-like sculpticels lacking over frontoclypeus and parietale; anterior margin of frontoclypeus broadly convex, extending mesally at level of adnasalia (Fig. 58); adnasalia margined with short bluntly rounded teeth (Fig. 65); MP2/MP1 0.50; Palaearctic. Description, instar III (Figs 58–62, 65) Body: Measurements and ratios aimed to characterize body shape are shown in Table 6. Head (Figs 58, 60, 65): Head capsule subquadrate, strongly constricted at level of occiput; anterior margin of frontoclypeus broadly convex, extending mesally at about level of adnasalia; scale-like sculpticels lacking over frontoclypeus and parietale; adnasalia margined with mostly short bluntly rounded teeth (Fig. 65); HL = 1.36–1.39 mm; A/MP = 1.16–1.27; MP/LP = 1.67–1.74; MP2/MP1 = 1.15–1.28; GA/MP1 = 1.27–1.49; LP2/LP1 = 1.54–1.71; MNL/MNW = 2.63–2.71. Thorax (Figs 61–62): L3 = 2.86–3.06 mm. Abdomen (Fig. 59): LAS = 1.09–1.17 mm; LAS subconical, constricted posteriorly at point of insertion of urogomphi. Urogomphus, U = 0.60–0.63 mm, composed of one urogomphomere; U/HW = 0.38–0.40; U/LAS = 0.54–0.57. Chaetotaxy: Parietale with 7–10 temporal spines; position and number of secondary setae on legs are shown in Figs 61–62 and Table 7., Published as part of Alarie, Yves, Michat, Mariano C., Watanabe, Kohei, Shaverdo, Helena, Wang, Liang-Jong & Watts, Chris H. S., 2022, An outlook on larval morphology of Copelatinae diving beetles with phylogenetic considerations (Coleoptera: Adephaga, Dytiscidae), pp. 151-205 in Zootaxa 5175 (2) on pages 193-197, DOI: 10.11646/zootaxa.5175.2.1, http://zenodo.org/record/7003344, {"references":["Fabricius, J. C. (1787) Mantissa Insectorum sistens species nuper detectas adiectis characteribus genericis, differentiis specificis, emendationibus, observationibus. 2 Vols. C. G. Proft, Hafniae, xx + 348 + 382 pp."]}

Copelatus oblitus Sharp, 1882 Source of material. The five instar II and five instar III larvae studied were reared ex ovo from adults collected at the following locality: Japan. Mantaburu, Yonaguni-cho, Yonaguni-jima Island, Okinawa Prefecture. 24°27’04.1”N 122°57’36.8”E, 13VI.2019; K. Watanabe leg. Diagnosis (instar III). The third instar of Copelatus oblitus can easily be distinguished from that of the other species studied in this paper by the following combination of characters: HL = 0.80 mm; L3 1.50; GA/MP1> 1.50; 1.30 Description, instar II Body: Measurements and ratios aimed to characterize body shape are shown in Table 4. Head: Head capsule ovate, not constricted at level of occiput; anterior margin of frontoclypeus narrowly convex, extending mesally below level of adnasalia; scale-like sculpticels present over frontoclypeus and parietale; adnasalia margined with short bluntly rounded teeth (see Fig. 64); HL = 0.55–0.58 mm; A/MP = 1.39–1.51; MP/LP = 1.70–1.82; MP2/MP1 = 1.67–1.95; GA/MP1 = 1.85–2.32; LP2/LP1 = 1.56–1.89; MNL/MNW = 2.41–2.61. Thorax: L3 = 1.10–1.13 mm. Abdomen: LAS = 0.37–0.39 mm; LAS subconical, not constricted posteriorly at point of insertion of urogomphi. Urogomphus: U = 0.17–0.19 mm, composed of two urogomphomeres; U/HW = 0.26–0.29; U/LAS = 0.45–0.51. Chaetotaxy: Parietale with 3–5 temporal spines; position and number of secondary setae on legs are shown in Table 5. Description, instar III (Figs 30–33, 64). As second-instar larva except as follows: Body: Measurements and ratios aimed to characterize body shape are shown in Table 6. Head (Figs 30, 64): HL = 0.74–0.80 mm; A/MP = 1.50–1.60; MP/LP = 1.65–1.82; MP2/MP1 = 1.44–1.67; GA/MP1 = 1.59–1.83; LP2/LP1 = 1.31–1.47; MNL/MNW = 2.40–2.60. Thorax (Figs 32–33): L3 = 1.57–1.64 mm. Abdomen (Fig 31): LAS = 0.54–0.60 mm; U = 0.20–0.25 mm, U/HW = 0.22–0.27; U/LAS = 0.37–0.47. Chaetotaxy: Parietale with 3–4 temporal spines; position and number of secondary setae on legs are shown in Figs 32–33 and Table 7., Published as part of Alarie, Yves, Michat, Mariano C., Watanabe, Kohei, Shaverdo, Helena, Wang, Liang-Jong & Watts, Chris H. S., 2022, An outlook on larval morphology of Copelatinae diving beetles with phylogenetic considerations (Coleoptera: Adephaga, Dytiscidae), pp. 151-205 in Zootaxa 5175 (2) on pages 175-177, DOI: 10.11646/zootaxa.5175.2.1, http://zenodo.org/record/7003344, {"references":["Sharp, D. (1882) On aquatic carnivorous Coleoptera or Dytiscidae. The Scientific Transactions of the Royal Dublin Society, Series 2, 2, 179 - 1003, pls. 7 - 18."]}

Copelatus parallelus Zimmermann, 1920 Source of material. All larvae studied were reared ex ovo from were adults collected at the following locality: Japan. Kyoto Prefecture. Uji-gawa, Fushimi-ku, Kyoto-shi, Kyoto Prefecture, 34°55’08.8”N 135°45’28.3”E, 6.XI.2016; K. Watanabe leg. Diagnosis (instar III). The third instar of Copelatus parallelus can easily be distinguished from that of the other species studied in this paper by the following combination of characters: HL = 0.80 mm; L3 1.50; GA/MP1> 1.50; 1.30 0.40; U/LAS> 0.60; Palaearctic. Description, instar I (Figs 34–47) Body: Measurements and ratios aimed to characterize body shape are shown in Table 2. Head (Figs 34–42): HL = 0.43 mm; Head capsule subquadrate, strongly constricted at level of occiput; occipital suture absent; anterior margin of frontoclypeus broadly convex extending beyond level of adnasalia mesally; scale-like sculpticels well-developed over frontoclypeus and parietale; adnasalia margined with strong acute teeth; occipital carina absent; A/MP = 1.27; MP/LP = 2.30; MP2/MP1 = 2.17; GA/MP1 = 2.60; LP2/LP1 = 2.74; MNL/ MNW = 3.66. Thorax (Figs 43–44): L3 = 0.80 mm. Abdomen (Figs 45–47): LAS = 0.32 mm; LAS subconical, slightly constricted posteriorly at point of insertion of urogomphi. Urogomphus: U = 0.17 mm, composed of one urogomphomere; U/HW = 0.36; U/LAS = 0.52. Chaetotaxy (Figs 34–47): Position and number of additional setae on legs are shown in Table 3 and Figs 43–44. Description, instar II. As for first-instar larva except as follows: Body: Measurements and ratios aimed to characterize body shape are shown in Table 4. Head: Anterior margin of frontoclypeus extending mesally at about same level of adnasalia; adnasalia margined with strong acute teeth (see Fig. 64); occipital suture present; HL = 0.56 mm; A/MP = 1.39; MP/LP = 2.24; MP2/ MP1 = 1.86; GA/MP1 = 2.44; LP2/LP1 = 2.12; MNL/MNW = 3.55. Thorax: L3 = 1.07 mm. Abdomen: LAS = 0.41 mm. Urogomphus: U = 0.18 mm, composed of one urogomphomere; U/HW = 0.30; U/LAS = 0.45. Chaetotaxy: Parietale with 7 temporal spines; position and number of secondary setae on legs are shown in Table 5. Description, instar III (Figs 48–51, 64). As second-instar larva except as follows: Body: Measurements and ratios aimed to characterize body shape are shown in Table 6. Head (Figs 48, 64): HL = 0.78 mm; A/MP = 1.29; MP/LP = 2.38; MP2/MP1 = 1.69; GA/MP1 = 1.82; LP2/LP1 = 1.67; MNL/MNW = 3.07. Thorax (Figs 50–51): L3 = 1.47 mm. Abdomen (Fig. 49): LAS = 0.61 mm; U = 0.40 mm; U/HW = 0.50; U/LAS = 0.65. Chaetotaxy: Parietale with 7 temporal spines. Position and number of secondary setae on legs are shown in Figs 50–51 and Table 7., Published as part of Alarie, Yves, Michat, Mariano C., Watanabe, Kohei, Shaverdo, Helena, Wang, Liang-Jong & Watts, Chris H. S., 2022, An outlook on larval morphology of Copelatinae diving beetles with phylogenetic considerations (Coleoptera: Adephaga, Dytiscidae), pp. 151-205 in Zootaxa 5175 (2) on pages 177-186, DOI: 10.11646/zootaxa.5175.2.1, http://zenodo.org/record/7003344, {"references":["Zimmermann, A. (1920) Bemerkenswerte Neuerwerbungen des Zoologischen Museum in Hamburg, Haliplidae, Dytiscidae et Gyrinidae. Entomologische Blatter, 16, 224 - 234.","Sharp, D. (1882) On aquatic carnivorous Coleoptera or Dytiscidae. The Scientific Transactions of the Royal Dublin Society, Series 2, 2, 179 - 1003, pls. 7 - 18."]}

Copelatus glyphicus (Say, 1823) Source of material. The five instar III larvae studied were associated with adults collected at the following locality: Canada. Québec, Trois-Rivières. Ephemeral pond in red maple forest at Chemin des Oblats, 06.VIII.1986, 08.VIII.1986, 12.VIII.1986; Y. Alarie leg. The identification is firm as C. glyphicus is the only Copelatus species known from that region. Diagnosis (instar III). The third instar of Copelatus glyphicus can easily be distinguished from that of the other species studied in this paper by the following combination of characters: HL = 0.80–0.90 mm; L3 1.50; GA/MP1> 1.50; 1.30 Description, instar III (Figs 19–20, 63) Body: Measurements and ratios aimed to characterize body shape are shown in Table 6. Head (Figs 19, 63): Head capsule rounded, strongly constricted at level of occiput; anterior margin of frontoclypeus broadly convex, extending beyond level of adnasalia; scale-like sculpticels well-developed both on frontoclypeus and parietale; adnasalia margined with short bluntly rounded and truncated teeth (Fig. 63); HL = 0.81–0.94 mm; A/MP = 1.34–1.38; MP/LP = 2.12–2.17; MP2/MP1 = 1.50–1.73; GA/MP1 = 1.79–1.88; LP2/LP1 = 1.40–1.56; MNL/MNW = 2.85–3.04. Thorax: L3 = 1.60–1.74 mm. Abdomen (Fig. 20): LAS = 0.65–0.74 mm; LAS subcylindrical, not constricted posteriorly at point of insertion of urogomphi. Urogomphus: U = 0.28–0.32 mm, composed of one urogomphomere; U/HW = 0.31–0.33; U/LAS = 0.42–0.44. Chaetotaxy: Parietale with 6–9 temporal spines; position and number of secondary setae on legs are shown in Table 7., Published as part of Alarie, Yves, Michat, Mariano C., Watanabe, Kohei, Shaverdo, Helena, Wang, Liang-Jong & Watts, Chris H. S., 2022, An outlook on larval morphology of Copelatinae diving beetles with phylogenetic considerations (Coleoptera: Adephaga, Dytiscidae), pp. 151-205 in Zootaxa 5175 (2) on page 168, DOI: 10.11646/zootaxa.5175.2.1, http://zenodo.org/record/7003344

Copelatus tenebrosus Régimbart, 1880 Source of material. The single instar III larvae studied was reared ex ovo from were adults collected at the following locality: Japan. Mantaburu, Yonaguni-cho, Yonaguni-jima Island, Okinawa Prefecture, 24°27’04.1”N 122°57’36.8”E, 13.VI.2019; E. K. Watanabe leg. Diagnosis (instar III). The third instar of Copelatus tenebrosus can easily be distinguished from that of the other species studied in this paper by the following combination of characters: HL = 0.80 mm; L3 1.50; GA/MP1> 1.50; 1.30 Description, instar III (Figs 52–53, 65) Body: Measurements and ratios aimed to characterize body shape are shown in Table 6. Head (Figs 52, 65): Head capsule rounded, slightly constricted at level of occiput; anterior margin of frontoclypeus narrowly convex, extending mesally well beyond level of adnasalia; scale-like sculpticels present over frontoclypeus and parietale; adnasalia margined with mostly short bluntly rounded teeth (Fig. 65); HL = 0.84 mm; A/MP = 1.40; MP/LP = 2.03; MP2/MP1 = 1.64; GA/MP1 = 1.85; LP2/LP1 = 1.43; MNL/MNW = 2.87. Thorax: L3 = 1.73 mm. Abdomen (Fig. 53): LAS = 0.63 mm; LAS subconical, not constricted posteriorly at point of insertion of urogomphi. Urogomphus, U = 0.30 mm, composed of two urogomphomeres; U/HW = 0.33; U/LAS = 0.47. Chaetotaxy: Parietale with 5–6 temporal spines; position and number of secondary setae on legs are shown in Table 7., Published as part of Alarie, Yves, Michat, Mariano C., Watanabe, Kohei, Shaverdo, Helena, Wang, Liang-Jong & Watts, Chris H. S., 2022, An outlook on larval morphology of Copelatinae diving beetles with phylogenetic considerations (Coleoptera: Adephaga, Dytiscidae), pp. 151-205 in Zootaxa 5175 (2) on page 187, DOI: 10.11646/zootaxa.5175.2.1, http://zenodo.org/record/7003344, {"references":["Regimbart, M. (1880) The new Dytiscidae and Gyrinidae collected during the recent scientific Sumatra-expedition. Notes from the Leyden Museum, 2, 209 - 216."]}

Exocelina ferruginea (Sharp, 1882) Source of material. The three instar II and three instar III larvae studied were collected in association with adults at the following locality: South Australia. One kilometer North of Forreston, 2.IX.2000; C. H. S. Watts leg. Diagnosis (instar III). The third instar of Exocelina ferruginea can easily be distinguished from that of the other species studied in this paper by the following combination of characters: HL = 1.10–1.20 mm; L3 = 2.20–2.50 mm; U> 0.50 mm; head capsule subquadrate, strongly constricted at level of occipital region (Fig. 56), scale-like sculpticels present over frontoclypeus and parietale; anterior margin of frontoclypeus broadly convex, extending mesally at level of adnasalia (Fig. 54); adnasalia margined with short acute teeth (Fig. 65); MP2/MP1> 1.50; GA/ MP1> 1.50; LP2/LP1> 2.00; inner margin of stipes lacking a dorsal linear row of spinulae; profemur with less than 3 PV and more than 5 AV secondary setae; metafemur with less than 3 AV secondary setae; urogomphus composed of one urogomphomere; U/HW> 0.40; U/LAS> 0.50; Australian. Description, instar II Body: Measurements and ratios aimed to characterize body shape are shown in Table 4. Head: Head capsule subquadrate, strongly constricted at level of occiput; anterior margin of frontoclypeus broadly convex, extending mesally at level of adnasalia; scale-like sculpticels present over frontoclypeus and parietale; adnasalia margined with short acute teeth (see Fig. 65); HL = 0.85–0.90 mm; A/MP = 1.33–1.38; MP/LP = 2.52–3.00; MP2/MP1 = 1.83; GA/MP1 = 2.00–2.33; LP2/LP1 = 3.17–3.80; MNL/MNW = 2.77–3.18. Thorax: L3 = 1.70–1.84 mm. Abdomen: LAS = 0.54–0.63 mm; LAS subcylindrical, not constricted posteriorly at point of insertion of urogomphi. Urogomphus: U = 0.46–0.51 mm, composed of one urogomphomere; U/HW = 0.50–0.53; U/LAS = 0.74–0.96. Chaetotaxy: Parietale with 4–6 temporal spines; position and number of secondary setae on legs are shown in Table 5. Description, instar III (Figs 56–57, 65). As second-instar larva except as follows: Body: Measurements and ratios aimed to characterize body shape are shown in Table 6. Head (Figs 56, 65): HL = 1.08–1.23 mm; A/MP = 1.27–1.33; MP/LP = 2.79–3.27; MP2/MP1 = 1.55–1.82; GA/MP1 = 1.65–2.06; LP2/LP1 = 2.22–2.50; MNL/MNW = 2.88–2.94. Thorax: L3 = 2.19–2.44 mm. Abdomen (Fig. 57): LAS = 0.79–0.85 mm; U = 0.44–0.71 mm, U/HW = 0.38–0.57; U/LAS = 0.56–0.88. Chaetotaxy: Parietale with 4–7 temporal spines; position and number of secondary setae on legs are shown in Table 7., Published as part of Alarie, Yves, Michat, Mariano C., Watanabe, Kohei, Shaverdo, Helena, Wang, Liang-Jong & Watts, Chris H. S., 2022, An outlook on larval morphology of Copelatinae diving beetles with phylogenetic considerations (Coleoptera: Adephaga, Dytiscidae), pp. 151-205 in Zootaxa 5175 (2) on page 193, DOI: 10.11646/zootaxa.5175.2.1, http://zenodo.org/record/7003344, {"references":["Sharp, D. (1882) On aquatic carnivorous Coleoptera or Dytiscidae. The Scientific Transactions of the Royal Dublin Society, Series 2, 2, 179 - 1003, pls. 7 - 18."]}

Barretthydrus Lea, 1927 (Figs. 1–19) Diagnosis. Instar III of Barretthydrus can be distinguished from those of other genera of Australian Hydroporini that have been well studied (i.e., Antiporus and Paroster) by the following combination of characters: HL = 1.24–1.36 mm; HL/HW 0.30; labial palpus composed of two palpomeres (Figs. 5–6); prementum lacking spinulae along lateral margins (Figs. 5–6); L3/HW Instar I (Figs. 1–12) Description. Body: Subcylindrical, narrowing towards abdominal apex. Measurements and ratios that characterize the body shape are shown in Table 1. Head: Head capsule (Figs. 1–2) pearshaped, tapering posteriorly, not constricted at level of occipital suture; ecdysial suture welldeveloped, coronal suture short; frontoclypeus elongate, bluntly rounded, subtriangular with welldeveloped lateral branches, anteroventral margin with 12 ventral lamellae clypeales (Bertrand 1972); dorsal surface with 2 spine-like egg bursters (ruptor ovi of Bertrand 1972) at about mid-length; ocularium present, stemmata not visible ventrally and subdivided into 2 vertical series; epicranial plates meeting ventrally; tentorial pits visible medioventrally at about mid-length. Antenna (Figs. 3–4) elongate, slightly shorter than HW; composed of 4 antennomeres, A2 and A3 longest, A1 shortest; A3’ relatively elongate, shorter than A4; A3 lacking ventroapical spinula. Mandible (Fig. 7) prominent, falciform, distal half projecting inwards and upwards, apex sharp; mandibular channel present. Labium (Figs. 5–6), prementum subrectangular, about as long as broad, lacking lateral marginal spinulae; LP elongate, distinctly shorter than MP, composed of 2 palpomeres; LP2 subfusiform, distinctly longer than LP1. Maxilla (Figs. 13–14) with short, thick stipes, incompletely sclerotized ventrally; cardo fused to stipes; galea and lacinia absent; MP elongate, slightly shorter than antenna, composed of 3 palpomeres; MP1 and MP2 longest, MP2 distinctly longer than MP1. Thorax: Thoracic terga convex, pronotum slightly shorter than meso- and metanota combined, meso- and metanota subequal; protergite subrectangular to subovate, more developed than meso- and metatergites; [We were unable to determine the presence of an anterotransverse carina owing to the bad condition of the only specimen available.]; thoracic sterna membranous; spiracles absent. Legs: Long (Figs. 8–9), composed of 6 articles (sensu Lawrence 1991); L1 shortest, L3 longest; CO robust, elongate, TR divided into 2 parts by an annulus, FE, TI, and TA slender, subcylindrical, PTwith 2 long, slender, slightly curved claws; posterior claw shorter than anterior claw on L1 and L2, posterior claw longer than anterior claw on L3; femora, tibiae, and tarsi lacking spinulae along ventral margin. Abdomen: Eight-segmented (Figs 10–11); segments I–VI sclerotized dorsally, membranous ventrally; segment VII sclerotized both dorsally and ventrally, ventral sclerite independent from dorsal one; tergites I–VII narrow, transverse, rounded laterally, lacking sagittal line; [We were unable to determine the presence of an anterotransverse carina owing to the bad condition of the only specimen available.]; segment VIII (=LAS) longest, completely sclerotized, ring-like, strongly constricted at point of insertion of urogomphus; projecting backwards into a very short, subconical siphon; spiracles absent lateroventrally on segments I–VII. Urogomphus very long, composed of 2 urogomphomeres; U1 long, much longer than segment VIII; U2 narrower, setiform, much shorter than U1. Chaetotaxy: Similar to that of generalized Hydroporinae larva (Alarie and Harper 1990; Alarie et al. 1990; Alarie 1991; Alarie and Michat 2007 a) except for the following features (Figs. 1–12): Pores PAd and ANf absent; pore ANh distal; setae MX 4 and TR2 absent; pore FEa articulated distally, close to seta FE5; seta TI7 short, spine-like; seta AB10 spine-like; setae UR2 and UR3 contiguous, seta UR4 articulated posteriorly; setae UR5, UR6, and UR7 elongate; seta UR8 inserted subapically. [Pore PAk and seta PA13 could not be located. We are reluctant to consider them as lacking due to the condition of the only instar I specimen available.] Instar II No specimens were available for study. Instar III (Figs. 13–19) Description. As instar I except as follows: Body: Measurements and ratios that characterize the body shape in Tables 1 and 2. Head: Head capsule (Figs. 15–16) constricted at level of occipital suture; egg bursters lacking. Antenna elongate, distinctly shorter than HW. LP 2 subequal to slightly shorter than LP 1. MP subequal in length to antenna, MP 1 slightly shorter than MP 2 (Figs. 13–14). Thorax: Protergum lacking anterotransverse carina; both meso- and metathoracic terga with an anterotransverse carinae; sagittal line visible; mesopleural region with a spiracular opening on each side. Legs: Position and number of secondary setae in Table 3; natatory setae present on dorsal margin of femora, tibiae, and tarsi (Figs. 17–18). Abdomen: Segment VII completely sclerotized both dorsally and ventrally, all tergites with anterotransverse carina (Fig. 19); mesopleural region of segments I–VII with a spiracular opening on each side. Chaetotaxy: Head capsule with numerous secondary setae; lateroventral margin of PA with several secondary spine-like setae (Figs. 15–16); anteroventral margin of nasale with half circle of about 60 lamellae clypeales of different lengths, directed downwards; AN, MX, and LA lacking secondary setae; MN with 1 hair-like secondary seta on basoexternal margin; thoracic and abdominal sclerites I–VIII with numerous secondary setae mainly on posterior half; natatory setae present on dorsal margin of femora, tibiae, and tarsi; secondary leg setation in Tables 3-4 and Figs. 17–18; U with secondary setae (Fig. 19). Remarks. Larvae of Barrethydrus can readily be distinguished from those of other Australian Hydroporini described in detail (i.e., Paroster and Antiporus) by the presence of natatory setae on the dorsal margins of the femur, tibia, and tarsi and metric characters presented in Table 2, and, superficially, from the less well-studied genera by the strong, dark yellow banding on the body. Distribution. Endemic to Australia.

Barretthydrus tibialis Lea , 1927 (Figs. 1–14, 16–19) Source of Material. Larvae were collected in association with adults at the following localities: Australia, New South Wales, 20 km North of Nelligan, 15-VIII.1997, coll. C. H. S. Watts; Sardine Creek, 30 km North of Orbost V, 30-XI-1998, C. H. S. Watts leg. Diagnostic Combination (Instar III). The third instar of B. tibialis can easily be distinguished from that of the closely similar B. geminatus by the presence of a broad, blackish macula on the anterior portion of the frontoclypeus (Fig. 16) in addition to a relatively shorter head capsule compared to the length of abdominal segment VIII (HL/LAS 4.60) and a larger total number of secondary setae on selected leg articles (Table 3). Instar I (Figs. 1–12) Description. Color: [Not available owing to the bad state of preservation of the only specimen available.] Body: Measurements and ratios that characterize the body shape in Table 1. Head: Head capsule as in Figs. 1–2, HL = 0.60 mm, HL/LAS = 4.61. Abdomen: As in Figs. 10–11, U 1 = 0.75 mm; U1/HW = 1.55. Instar II No specimen available for study. Instar III (Figs. 13–14, 16–19) Description. Color: Head capsule predominantly yellow; frontoclypeus with a broad, black, subapical macula (Fig. 16); head appendages dark yellow; thoracic tergites dark brown; abdominal tergites I, IV – VII dark brown, II and III dark yellow to pale brown and VIII pale yellow; urogomphi dark yellow to pale brown; legs dark brown proximally, pale yellow distally. Body: Measurements and ratios that characterize the body shape in Table 1. Head: Head capsule as in Fig. 16, HL = 1.24–1.30, HL/LAS Abdomen: U1 = 1.51–1.69 mm, U1/HW = 1.56–1.67 (Fig. 19). Chaetotaxy: ProFE with more than 52 secondary setae; mesoFE with more than 56 secondary setae; mesoTI with more than 47 secondary setae; mesoTA with more than 34 secondary setae; metaCO with more than 26 secondary setae; metaFE with more than 66 secondary setae; metaTI with more than 50 secondary setae; metaTA with more than 43 secondary setae., Published as part of Alarie, Yves, Michat, Mariano C., Hendrich, Lars & Watts, Chris H. S., 2018, Larval Description and Phylogenetic Placement of the Australian Endemic Genus Barretthydrus Lea, 1927 (Coleoptera: Dytiscidae: Hydroporinae: Hydroporini: Sternopriscina), pp. 639-661 in The Coleopterists Bulletin 72 (4) on page 645, DOI: 10.1649/0010-065X-72.4.639, http://zenodo.org/record/4789513

Barretthydrus geminatus Lea, 1927 (Fig. 15) Source of Material. Larvae were collected in association with adults at the following locality: Australia, New South Wales, 13 km NNW Dungog, Williams River, Tillegra Bridge, 109 m., 19. X.2006, 32.19.078S 151.41.250 E, L. Hendrich leg. (NSW 84). Diagnostic Combination (Instar III). The third instar of B. geminatus can easily be distinguished from that of the closely similar B. tibialis by the absence of maculae on the dorsal surface of the head capsule (Fig. 15), in addition to a relatively longer head capsule compared to the length of abdominal segment VIII (HL/LAS>4.60 compared to Instar I No specimen available for study. Instar II No specimen available for study. Instar III (Fig. 15) Description. Color: Head capsule testaceous (Fig. 15); head appendages pale yellow; thoracic tergites dark brown; abdominal tergites I – III brown mesally, broadly yellow laterally, IV – VII dark brown, VIII pale yellow; urogomphi dark yellow; legs predominantly dark yellow except coxae brownish. Body: Measurements and ratios that characterize the body shape in Table 1. Head: Head capsule as in Fig. 15, HL = 1.27–1.36, HL/LAS>4.60. Abdomen: U1 = 1.29–1.45 mm, U1/HW = 1.37–1.49. Chaetotaxy: ProFE with less than 45 secondary setae; mesoFE with less than 51 secondary setae; mesoTI with less than 45 secondary setae; mesoTA with less than 30 secondary setae; metaCO with less than 18 secondary setae; metaFE with less than 53 secondary setae; metaTI with less than 46secondary setae; metaTA with less than 38 secondary setae., Published as part of Alarie, Yves, Michat, Mariano C., Hendrich, Lars & Watts, Chris H. S., 2018, Larval Description and Phylogenetic Placement of the Australian Endemic Genus Barretthydrus Lea, 1927 (Coleoptera: Dytiscidae: Hydroporinae: Hydroporini: Sternopriscina), pp. 639-661 in The Coleopterists Bulletin 72 (4) on pages 646-647, DOI: 10.1649/0010-065X-72.4.639, http://zenodo.org/record/4789513

Scirtes serratus sp. nov. (Figs 18, 27) Type material. Holotype ♂, ���SA 1 Km S Nangwarry 22/9/07 CHS Watts/ Bred from larva/ On slide with larval exuviae���, SAMA. Description (number examined, 1). Habitus. Length approximately 3.4 mm, oval. Head. Testaceous. Antenna light testaceous, basal antennomeres lighter. Eyes rather small, width between eyes dorsally about 3.2�� dorsal width of eye. Setae moderately long. Frons with sides moderately diverging, front edge weakly concave, edges weakly beaded. Antennomere 1 of antenna barrel-shaped; antennomere 2 about two-thirds as long and not much narrower, barrel-shaped; antennomere 3 about same size as antennomere 2, slightly narrower, wider towards apex; antennomere 4 about 2�� length of antennomere 3; antennomeres 5���10 subequal, narrow, cylindrical, a little shorter than antennomere 4; antennomere 11 elongate/oval, 1.5�� length of and flatter than antennomere 10. Pronotum. Testaceous; broad, 2.2�� as wide as long. Punctures moderately large, about a puncture width apart, each puncture with a moderately long golden seta. Hind angles weakly obtuse, front edge sinuate, margins weakly beaded. Scutellar shield. Testaceous. Equilateral triangle, lateral sides weakly convex. Elytra. Testaceous. Sides weakly flanged in basal third. Strongly and evenly punctate, somewhat larger than on pronotum, most punctures less than a puncture width apart or less, each puncture with a moderately long pale seta. Epipleuron relatively wide in front quarter becoming narrower over rest of elytron, front portion flat. Ventral surface. Testaceous. Pronotal process very narrow at apex. Mesoventral notch ���U���-shaped. Mesoventral process long, narrow, reaching to metaventrite. Metacoxal plate almost square, anterior-lateral corner extending a short way along metaventrite; hind edge straight; sides beaded, posterolateral angles rounded. Metatrochanter small, relatively squat, about 1.2�� as long as wide. Metafemur greatly swollen, widest a little before middle, weakly notched on rear margin near apex. Dorsal metatibial spine relatively short, about a third length of ventral spine, ventral spine about two-thirds length of segment 1 of metatarsus; segment 1 of metatarsus a little longer than other segments combined. Ventrites with very small, shallow punctures each with a short seta; reticulation moderate, fine, more pronounced towards rear. Male. Basal piece short, round; trigonium long and thin, 1.6�� length of basal piece; tip rounded; ventral edge deeply serrated; two parameroids, right hand one a little shorter than trigonium with weak apical hook, left hand one broad, apex rounded, about a third length of right hand one. Tegmen as long as penis, lobes thin, elongate, well separated (Fig. 18). Abdominal segments not known. Female. Not known. Etymology. Name refers to the serrated ventral edge of the trigonium. Notes. Known from only one specimen reared from a larva collected from a shallow seasonal swamp. The failure to collect more over several springs suggests that the species is rare. It will run to S. orientalis in the key in Watts (2004) but can be separated by the strongly serrated ventral edge of the trigonium. The larva is known from a single exuvia in poor condition, lacking terminal abdominal segments. It will run to the S. helmsi / S. orientalis couplet in the key to Australian Scirtes larvae in Watts (2014). It is larger than S. orientalis (approximately 8 mm long vs 6 mm) and larger than S. lynnae with which it is sympatric. It cannot be separated from S. helmsi, which also occurs in the vicinity, on the limited material available., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on pages 524-525, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Watts, C. H. S. (2004) Revision of Australian Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae). Transactions of the Royal Society of South Australia, 128, 131 - 168."]}

Scirtes helmsi species group (Watts 2004) Defined here as: broadly oval; mandible incisivus relatively short, first segment of labial palpus shorter than second, caudal margin of metacoxal plates straight or nearly so, caudal margin of sternite 7 truncated rather than concave, metatrochanters elongate-oval, 2.0�� as long as wide, aedeagus asymmetrical, pala well developed, one or two parameroids, typically the right hand one longer and with an apical hook; tegmen variable, usually simple, parameres simple, usually long and thin but sometimes broader and tending to wrap around the penis but are never fused with it (Watts 2004) (Figs 18���20); female coxites and styles elongate, prehensor moderately developed, occasionally absent, basal sclerite typically pelican-bill-like but occasionally ring-shaped with internal prongs or bullet-shaped with internal spines (Figs 7���16). Larva with maxillary palpus four-segmented, five anal papillae and mandible with single-pointed incisivus. Most of the species of Australian Scirtes studied fell into this group. The sequence data identified two major subgroupings with some internal substructure within each. There is no correlation between the different female prehensors and bursal sclerites and the sequence data. The male terminalia of S. zwicki are quite unlike those of other species in the group (and from any other known Scirtes species), having elongate, symmetrical genitalia and a greatly modified sternite 9. Sternite 8 appears to be missing (Fig. 23). Genetically it is unequivocally within the S. helmsi subgroup (Fig. 1). There are 27 described species in the Scirtes helmsi group in Australia (Watts 2004 & this paper). Yoshitomi & Ruta (2010) noted the close similarity of 11 New Caledonian species with Australian species. From the illustrations of the male genitalia, S. babeldaobensis Yoshitomi from Micronesia in Yoshitomi (2009) and possibly S. zerchei Klausnitzer from Luzon Island may also belong in the S. helmsi group., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on page 520, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Watts, C. H. S. (2004) Revision of Australian Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae). Transactions of the Royal Society of South Australia, 128, 131 - 168.","Yoshitomi, H. (2009) The Scirtes (Coleoptera: Scirtidae: Scirtinae) of Micronesia. Zootaxa, 1974, 1 - 16."]}

Scirtes rutai nom. nov. = Scirtes beccus Ruta, Kiałka & Yoshitomi, 2014 Notes. Scirtes beccus Ruta, Kiałka & Yoshitomi, 2014 from Kinabalu Park, Ranu, Sabah is preoccupied by Scirtes beccus Watts, 2004. We propose the replacement name, Scirtes rutai, after Rafał Ruta of Wroclaw in recognition of his extensive and important work on the Scirtes of Southeast Asia., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on page 527, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Ruta, R., Kialka, A. & Yoshitomi, H. (2014) A supplement to the revision of the Scirtes flavoguttatus species-group (Coleoptera: Scirtidae: Scirtinae). Zootaxa, 3901 (1), 1 - 62. https: // doi. org / 10.11646 / zootaxa. 3902.1.1","Watts, C. H. S. (2004) Revision of Australian Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae). Transactions of the Royal Society of South Australia, 128, 131 - 168."]}

Ora improtecta Watts, 2004 Notes. This species remains little known and no specimens were available for our DNA study. Morphologically it is distinct from the two other Australian species in the genus, which appear to be more typical of the genus, and its position in Ora is problematic. In this context it needs to be recognized that Ora is a poorly characterized cosmopolitan genus much in need of a revision. There is a recent specimen in the South Australian Museum���s collection from Halmahera, Indonesia. It is thus probable that the species is widespread in the islands to the north of Australia., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on page 520, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Watts, C. H. S. (2004) Revision of Australian Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae). Transactions of the Royal Society of South Australia, 128, 131 - 168."]}

Scirtes zwicki sp. nov. (Figs 23, 24) Type material. Holotype ♂, ���NT Holmes Jungle/CHS Watts 10/1/14 / at light.���, NTM. Paratypes 12 ♂, 1 ♀ as for holotype, SAMA. Description (number of dissected males examined, 13). Habitus. Length 2.5���2.9 mm, elongate oval. Head. Area between eyes black, frons light testaceous, division between areas of colour distinct. Antenna light testaceous, becoming darker towards apex. Eyes moderately large, width between eyes dorsally about 3.0�� dorsal width of eye. Setae moderately long, pale. Frons with sides moderately diverging, front edge weakly concave, edges quite strongly beaded. Antennomere 1 of antenna barrel-shaped; antennomere 2 about half as long, narrower, barrel-shaped: antennomere 3 about as long as antennomere 2, about half as wide, widening towards apex; antennomere 4 about twice length of antennomere 3; antennomeres 5���10 subequal, narrow, cylindrical, a little shorter than antennomere 4; antennomere 11 elongate/oval, a little longer and flatter than antennomere 10, 1.6�� length of and flatter than antennomere 10. Pronotum. Central half dark testaceous-black, margins abruptly light testaceous. Short, 2.3�� as wide as long. Punctures moderately sized, about 1.5 puncture widths apart, each puncture with a moderately long pale seta. Hind angles weakly obtuse, front edge sinuate, margins weakly beaded. Scutellar shield. Light testaceous. Equilateral triangle, lateral sides weakly convex. Elytron. Light testaceous, area adjacent to scutellum sometimes darker. Sides weakly flanged in basal third. Moderately and evenly punctate, most punctures about 1.5 puncture widths apart, each puncture with a short pale seta. Epipleuron relatively wide in front quarter, becoming narrower over rest of elytron, front portion flat. Ventral surface. Light testaceous, prosternum often diffusely darker. Prosternal process very narrow. Receiving notch on mesoventrite well-marked, narrowly diamond-shaped, reaching past front of mesocoxae. Mesoventral process long, narrow, reaching to metaventrite. Anterior extension of metaventrite in midline small, triangular, not bounded behind by ridge; rear midline extension of metaventrite short, about twice as wide as long. Metacoxal plate square, anterior-lateral corner extending about half way along metaventrite; hind edge weakly angled towards midline; sides beaded, posterolateral angles rounded. Metatrochanter small, elongate, about 1.7�� as long as wide. Metafemur greatly swollen, widest about middle, weakly notched on rear margin near apex. Dorsal metatibial spine relatively short, about twice length of ventral spine and about half length of segment 1 of metatarsus; segment 1 of metatarsus a little longer than other segments combined. Ventrites with small, shallow punctures each with a short seta, reticulation moderate, fine. Male. Little external difference between the sexes. Basal piece of penis indistinct, trigonium very long and thin, apical half bifid, lobes weakly paddle-shaped; without parameroids. Tegmen a little more than half as long as penis, lobes thin, elongate, well separated. Apices broad with two well-developed sharply pointed lobes (Fig. 23e). Sternite 9: well developed, elongate, apical portion bullet-shaped with strongly sclerotized apical cap, apodemes about twice as long as apical portion, anterior ends fused (Fig. 23c). Tergite 9: membranous, without setae, apodemes moderately long (Fig. 23a). Tergite 8: relatively narrow, moderately sclerotized, apical portion with ruffled margin, without setae, apodemes weakly sclerotized, a little longer than apical portion, weakly diverging (Fig. 23b). Female. Coxites broad, elongate/triangular with numerous pores, styli small, narrowly oval, positioned slightly inside of apex. Bursal sclerite well developed, anterior end rounded behind which is a flared structure with two variable rows of short spines (Fig. 10). Prehensor either fused with basal sclerite or absent. Etymology. Named after Peter Zwick, a long time student of Australian Scirtidae who has helped CHSW significantly in his studies of the family. Notes. A small species easily recognised by the black areas on the head and pronotum strongly contrasting with the testaceous elytra, greatly swollen metafemora and the large, relatively close-set eyes. Collected from or near riverine rainforest in coastal Northern Territory. It will run to S. emmaae in the key in Watts (2004), but can be separated by its smaller size and extremely elongate bifid penis (Fig. 23). Larva. Not known., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on pages 523-524, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Watts, C. H. S. (2004) Revision of Australian Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae). Transactions of the Royal Society of South Australia, 128, 131 - 168."]}

Scirtes lynnae sp. nov. (Figs 20, 26) Type material. Holotype ♂, ���SA 1 Km S Nangwarry 37 33 140 46 20/10 /14 CHS Watts/ Bred from larva.���, SAMA. Paratype ♂ as for holotype, SAMA. Description (number examined, 2). Habitus. Length 2.2���2.4 mm, oval. Head. Dark testaceous; antenna testaceous, basal antennomeres lighter. Eyes rather small, width between eyes about 3.8�� dorsal width of eye. Setae moderately long, pale. Frons with sides weakly diverging, front edge straight, edges weakly beaded. Antennomere 1 of antenna narrowly barrel-shaped; antennomere 2 about three-quarters as long and not much narrower, barrel-shaped; antennomere 3 about three-quarters length of antennomere 2, slightly narrower, wider towards apex; antennomere 4 about 1.5�� length of antennomere 3; antennomeres 5���10 subequal, narrow, cylindrical, a little shorter than antennomere 4; antennomere 11 elongate/oval, 1.4�� length of and flatter than antennomere 10. Pronotum. Dark testaceous; broad, 2.7�� as wide as long. Punctures moderately large, about a puncture width apart, each puncture with a moderately long golden seta. Hind angles weakly obtuse, front edge sinuate, margins weakly beaded. Scutellar shield. Testaceous. Equilateral triangle, lateral sides weakly convex. Elytra. Light testaceous, base and humeral angles darker. Sides weakly flanged in basal third. Strongly and evenly punctate, somewhat larger than on pronotum, most punctures less than a puncture width apart or less, each puncture with a moderately long pale seta. Epipleuron relatively wide in front quarter, becoming narrower over rest of elytron, front portion flat. Ventral surface. Testaceous. Pronotal process very narrow at apex. Mesoventral notch ���U���-shaped. Mesoventral process long, narrow, reaching to metaventrite. Metacoxal plate almost square, anterior-lateral corner extending a short way along metaventrite; hind edge straight; sides beaded, posterolateral angles rounded. Metatrochanter small, relatively squat, about 1.2�� as long as wide. Metafemur greatly swollen, widest a little before middle, weakly notched on rear margin near apex. Dorsal metatibial spine, about half length of ventral spine, ventral spine about two-thirds length of segment 1 of metatarsus; segment 1 of metatarsus a little longer than other segments combined. Ventrites with very small, shallow punctures each with a short seta, reticulation moderate, fine, more pronounced towards rear. Male. Basal piece short, round; trigonium long, broad at base, evenly narrowing to rounded tip, about 2�� length of pala; ventral edge smooth; two parameroids, right hand one about half length of trigonium, thin, without apical hook, left hand parameroid very small, oval. Tegmen two-thirds as long as penis, lobes thin, finger-like, well separated (Fig. 20). Tergite 8 broad short, apodemes straight well sclerotized. Tergite 9 smaller and weaker. Sternite 8 not known. Female. Not known. Etymology. Named after Lynn Strefford of the South Australian Museum who has very competently done much of the unsung administrative work in support of our studies over many years. Notes. In small size and dark colour it greatly resembles Scirtes pinjarraensis but differs from that species in the straight right parameroid, and the absence of spines at the apex of the trigonium (Fig. 22). Sequence data (Fig. 1) place the species as sister species to S. orientalis. In the key to Australian Scirtes in Watts (2004) it will run to S. brisbanensis / S. pinjarraensis, but can be separated from S. brisbanensis by its small size, rounded rather than pointed apex of the trigonium and dark rather than mainly chestnut coloured dorsal surface and from S. pinjarraensis by the very different apex of the trigonium (Figs 20, 22). The small black larvae were abundant in a drying pool in a reed/weed choked swamp. Larvae. Scirtes lynnae runs to the S. helmsi / S. orientalis couplet in Watts (2014). Its small size separates it from S. helmsi and also S. serratus with which it is sympatric. It differs from S. orientalis in generally having fewer claw teeth (6���7 vs 7���9) and broader tergite 9. From Scirtes triangularis, which also runs to S. orientalis, it differs in its smaller size and single row of palisade setae on the clypeolabrum (Fig. 26)., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on pages 525-526, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Watts, C. H. S. (2004) Revision of Australian Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae). Transactions of the Royal Society of South Australia, 128, 131 - 168."]}

Scirtes elegans species group (Yoshitomi 2009) Yoshitomi (2009) defined the S. elegans group as ���having the tegmen connected to an asymmetrical penis���. We add: first segment of labial palpus as long as second, mandible incisivus long, metatrochanter elongate, caudal margin of sternite 7 moderately concave. The relationship between tegmen and penis perhaps could be described as having the elements of each complexly intermingled (Fig. 21). Within the group the female of only one species is known���that of the only Australian species, S. albamaculatus, from North Queensland. Assuming that the female of S. albamaculatus is representative of the group we could also add: members are dark coloured with white/cream macula or stripes on the elytra and females with elytral excitators (Fig. 31) and a ring-like bursal sclerite (Fig. 6). No larvae have been described. The Scirtes elegans group as presently defined has only three species, S. elegans from Malaysia, S. albamaculatus from Australia and S. albotaeniatus Yoshitomi from Guam. The sequence data place S. albamaculatus as unique within Australian Scirtes. However, it also strongly links S. albamaculatus to S. teruhisai Yoshitomi & Ruta from Borneo (Fig. 1) which is the only representative of the species-rich Southeast Asian S. flavoguttatus group (Yoshitomi & Ruta 2010; Ruta et al. 2014) included in our study. This placement is not unexpected as a number of species currently placed in this group have female excitators, pale colour patterns on the elytra, all have fused tegmen and penis and some, e.g. S. sarawakensis Ruta & Yoshitomi, have an aedeagus approaching the complexity seen in S. albamaculatus. The form of the mandible and labial palpus also point to membership of the S. flavoguttatus group. With the discovery of additional species in the S. flavoguttatus group in Southeast Asia (Ruta et al. 2014) the distinction between these two groups has become rather tenuous and relies on the asymmetric rather than symmetric penis. The strong linkage of S. albamaculatus and S. teruhisai by the sequence data suggests that the two species belong in the same species group. However, with only one species of each group available to us we feel it is premature to formally combine these species groups on the results of this study., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on page 519, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Yoshitomi, H. (2009) The Scirtes (Coleoptera: Scirtidae: Scirtinae) of Micronesia. Zootaxa, 1974, 1 - 16."]}

Scirtes brisbanensis Pic, 1956 / Scirtes triangularis Watts, 2004 Notes. In Cooper et al. (2014) it was noted that two closely related taxa, each represented by a single female and both identified as Scirtes brisbanensis, were present at Mt Molloy in North Queensland. With access to additional specimens, including males, the present study shows that one specimen was S. brisbanensis and the other S. triangularis (Fig. 1). Larva. A single larva of S. triangularis is now known, identified by DNA sequencing. In the key to the larvae of Australian Scirtes in Watts (2014) it runs to the S. helmsi / S. orientalis couplet. However, it is smaller than S. helmsi (head width 0.9 mm vs 1.2 mm) and with fewer setae on tergite 1 (8 vs 30���40). From the similar sized S. orientalis it differs in having a more uneven row of palisade setae on the clypeolabrum (Fig. 28) and eight rather than nine claw teeth but is otherwise very similar., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on page 526, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Watts, C. H. S. (2004) Revision of Australian Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae). Transactions of the Royal Society of South Australia, 128, 131 - 168.","Cooper, S. J. B., Watts, C. H. S., Saint, K. M. & Leijs, R. (2014) Phylogenetic relationships of Australian Scirtidae (Coleoptera) based on mitochondrial and nuclear sequences. Invertebrate Systematics, 28, 628 - 642. https: // doi. org / 10.1071 / IS 13046"]}

Scirtes japonicus species group (Nyholm 2002) Nyholm (2002) based this group primarily on the form of the penis: ���symmetric with parameroids longitudinally deeply cleft and tegmen with a lateral lobe (paramere) on each side���. We would add: first segment of labial palpus shorter than second, mandible incisivus moderately long, caudal margin of metacoxal plate concave and caudal margin of sternite 7 moderately concave; larvae with tip of mandible bifid and maxillary palpus threesegmented (Watts 2004; Yoshitomi 2005). The prehensors and bursal sclerites of all three Australian species (Figs 3���5) are closely similar to those of S. japonicus (Yoshitomi 2005). The penis of no Australian species has this morphology. The aedeagus of S. emmaae is simple and appears plesiomorphic as it conforms closely to the scirtid ground plan (Zwick 2016). Scirtes kaytae and S. tindaleensis have asymmetric aedeagi resembling those of species in the S. helmsi group. The aedeagus of S. emmaae, particularly the apical triangular lobes of the parameres, is similar to those of S. micronesianus Yoshitomi, which was placed in a separate group by Yoshitomi (2009) and named the S. micronesianus group. However, other than their aedeagi, S. emmaae and S. kaytae are morphologically and genetically close and S. tindaleensis is also close on both larval and adult morphology yet only S. emmaae could be classified in the S. micronesianus group. We think that on this evidence these three species belong in the same species group and have placed all three Australian species in the S. japonicus group based on sequence data and adult and larval morphology but recognise that future study is needed to confirm this placement, particularly in light of the conflicting morphology of the male genitalia. Scirtes nehouensis Ruta & Yoshitomi, 2010 from New Caledonia appears to us on both general morphology and male and female genitalia (Ruta & Yoshitomi 2010b) to be conspecific with S. emmaae. We herein place the species in synonymy under Scirtes emmaae Watts, 2004., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on pages 518-519, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Nyholm, T. (2002) Scirtes japonicus Kiesenwetter and its allies with descriptions of Scirtes ussuriensis n. sp. (Coleoptera Scirtidae). Entomologische Blatter, 98, 49 - 60.","Watts, C. H. S. (2004) Revision of Australian Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae). Transactions of the Royal Society of South Australia, 128, 131 - 168.","Yoshitomi, H. (2005) Systematic revision of the family Scirtidae of Japan, with phylogeny, morphology and bionomics (Insecta: Coleoptera, Scirtoidea). Japanese Journal of systematic Entomology, Monographic Series, 3, 1 - 212. [Matsuyama]","Zwick, P. (2016) Australian Marsh Beetles (Coleoptera: Scirtidae). 9. The relations of Australian Ypsiloncyphon species to their Asian congeners, additions mainly to Petrocyphon and Prionocyphon, and a key to Australian genera of Scirtinae. Zootaxa, 4085 (2), 151 - 198. https: // doi. org / 10.11646 / zootaxa. 4085.2.1","Yoshitomi, H. (2009) The Scirtes (Coleoptera: Scirtidae: Scirtinae) of Micronesia. Zootaxa, 1974, 1 - 16.","Ruta, R. & Yoshitomi, H. (2010 b) Scirtidae: The genus Scirtes Illiger (Coleoptera). In: Jach, M. A. & Balke, M. (Eds.), Water Beetles of New Caledonia. Part 1. Monographs on Coleoptera 3. Natural History Museum, Wien, pp. 403 - 438."]}

Scirtes haemisphaericus species group (Yoshitomi 2005) Yoshitomi (2005) defined this group as having the ���body oval, parameres simple, penis symmetrical with a long slender pala (basal piece)���. We add: mandibles blunt, first segment of labial palpus shorter than second segment, caudal margin of metacoxal plates weakly concave, metatrochanters elongate (about 2.5�� as long as wide), caudal margin of sternite 7 deeply concave; females with relatively short coxites and broad short styles and a moderately sized, compact, well sclerotized prehensor and bursal sclerite (Fig. 10; Yoshitomi 2005); larvae with a pronounced spindle-shape with multidentate mandibles, maxillary palpus three-segmented, anal palpi>5 (Klausnitzer 2009; Watts 2014). Both the sequence data and adult and larval morphology support a relationship of the single Australian species, S. auratus, with the European S. haemisphaericus, the type species of the genus Scirtes. The Japanese S. sobrinus Lewis clearly also belongs in this group on larval characters, prehensor and bursal sclerite although less clearly on the male genitalia (Yoshitomi 2005). The sequence data also placed two closely related species from Vietnam, each represented by a single male, in this grouping (Fig. 1). The male genitalia of both are similar and closely resemble the illustration of the male genitalia of S. sobrinus in Yoshitomi (2005)., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on page 519, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Yoshitomi, H. (2005) Systematic revision of the family Scirtidae of Japan, with phylogeny, morphology and bionomics (Insecta: Coleoptera, Scirtoidea). Japanese Journal of systematic Entomology, Monographic Series, 3, 1 - 212. [Matsuyama]","Klausnitzer, B. (2009) Insecta: Coleoptera: Scirtidae. Subwasserfauna von Mitteleuropa Bd. 20 / 7. Spektrum Akademischer, Verlag, Heidelberg, 134 pp."]}

Scirtes pinjarraensis Watts, 2004 = Scirtes pygmaeus Watts, 2004, syn. nov. Notes. Cooper et al. (2014), using sequence data, suggested that Scirtes pinjarraensis and S. pygmaeus Watts were probably conspecific. Examination of further specimens of both taxa confirms this. Watts (2004) separated the two taxa by the presence of a distinctly bent tip to the trigonium in S. pygmaeus which was absent in S. pinjarraensis. Examination of additional specimens of both taxa shows that this bent tip is actually a distinct spine projecting at right angles to the trigonium and surrounded by a number of much smaller spines (Fig. 22). Dissection of more paratypes of S. pinjarraensis showed that a number had identical structures and that the absence of a spine was caused by it being broken off, probably during dissection. We consider, on both morphological and sequence grounds, that S, pinjarraensis and S. pygmaeus are conspecific. Therefore, S. pygmaeus on page preference becomes a junior synonym of S. pinjarraensis. This terminal structure on the trigonium is unique within Australian Scirtes and readily identifies the species, which is often the commonest scirtid in swamps in southwest Australia. Other than by the male genitalia, the species is recognized from other species in southwest Australia by its small size, dark testaceous colour with pale antennal base and pale sides to the pronotum. In size, colour and male genitalia S. pinjarraensis resembles S. lynnae, but differs from this species in the hooked left parameroid, presence of a small right parameroid and spines at the apex of the trigonium. Sequence data place it close to S. helmsi and S. brisbanensis (Fig. 1)., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on page 526, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Watts, C. H. S. (2004) Revision of Australian Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae). Transactions of the Royal Society of South Australia, 128, 131 - 168.","Cooper, S. J. B., Watts, C. H. S., Saint, K. M. & Leijs, R. (2014) Phylogenetic relationships of Australian Scirtidae (Coleoptera) based on mitochondrial and nuclear sequences. Invertebrate Systematics, 28, 628 - 642. https: // doi. org / 10.1071 / IS 13046"]}

Scirtes exoletus Waterhouse, 1880 Notes. A very widespread, quite common species found in all states and mainland territories of Australia. Specimens from the Northern Territory, New South Wales and Victoria were identical in the genes sequenced; however, the specimen from Wongabel on the Atherton Tableland of North Queensland differed by 7.2% (pdistance) for COI, potentially indicative of a separate species., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on page 527, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629

Scirtes auratus Watts, 2004 (Figs 2, 17) Supplementary description. Male. Tergite 8: small, apical portion very weakly sclerotized, hind edge with dense row of extremely small fine setae or microtrichia. Apodemes relatively short, strongly sclerotized, parallel. Tergite 9: short, broad, apical portion short with apical margin with long setae, apodemes strongly sclerotized, about three times length of apical portion, weakly diverging. Sternite 9: small, weakly sclerotized, broadly bilobed, apical third with numerous pores and thin setae, without obvious apodemes. Aedeagus complex, symmetrical, basal piece large, expanding anteriorly; trigonium long, thin, with complex set of huge curved spines at base, apex with a few small stout spines; parameroids in two parts, inner lobe longer than trigonium, broad in basal half, narrow in apical half, bulbous at apex; outer lobe broad, flat, about half as long as inner lobe numerous pores and spines on outer margin and towards apex. Tegmen consists of two separate narrow rods, about as long as aedeagus, central portions covered with short strong spines, apical portion with outer spur of varying length (Fig. 17). Female. Coxites elongate, styles relatively short and broad. Tergite 9: long, with spurs; tergite 8 long; tergite 8 shorter, apex with thick row of moderately long setae. Prehensor moderately sclerotized, long, apical end with two triangular plates. Bursal sclerite elongate, strongly sclerotized, triangular apical end with two strong prongs, surface strongly pimpled (Fig. 2). Notes. The species is found in coastal swamps from the Northern Territory around the northern coast to at least Townsville and has been collected in large numbers at light. The larvae are found in shallow grassy swamps (Watts 2014). It is not known if the pupae of the species use a snorkel as described by Zwick & Zwick (2008) for S. haemisphaericus. In the laboratory without access to plant stems the larvae form pupal cells in damp sand or pupate on the undersides of leaves., Published as part of Watts, Chris H. S., Cooper, Steven J. B. & Saint, Kathleen M., 2017, Review of Australian Scirtes Illiger, Ora Clark and Exochomoscirtes Pic (Coleoptera: Scirtidae) including descriptions of new species, new groups and a multi-gene molecular phylogeny of Australian and non-Australian species, pp. 511-532 in Zootaxa 4347 (3) on page 523, DOI: 10.11646/zootaxa.4347.3.5, http://zenodo.org/record/1048629, {"references":["Watts, C. H. S. (2004) Revision of Australian Scirtes Illiger and Ora Clark (Coleoptera: Scirtidae). Transactions of the Royal Society of South Australia, 128, 131 - 168.","Zwick, P. & Zwick, H. (2008) Number of larval instars, early instar structure, and life history of Scirtes hemisphaericus (Coleoptera: Scirtidae) in central Germany. Lauterbornia¸ 63, 87 - 99."]}

Tiporus undecimmaculatus (Clark, 1862) Type material studied. Syntypes, 2 females: ��� Paralectotype ��� [printed, round and blue bordered label], ��� New Holland ��� [handwritten, round], ��� H. maculatus Clark ��� [yellow, handwritten], ��� Hypodes undecimmaculatus Clk ��� det. C. Watts 1974��� [handwritten, printed] (NHM). An additional historical specimen, most probably from Clark��s collection, was also studied. Male: ���Lectotype��� [printed, round and violet bordered label], ���67.56���, ��� H. maculatus Clark ��� [yellow, handwritten], ���Hypodes undecimmaculatus Clk ��� det. C. Watts 1974��� [handwritten and printed white label] (NHM). Remarks. According to the original description by Clark (1862: 412) the species was described after two specimens from ��� New Holland ���. Therefore the designation of third specimen as a lectotype is invalid. We do not consider the designation of a lectotype necessary, as this would not serve in any way the stability of the nomenclature. Both syntypes together constitute the name-bearing type of Hydroporus undecimmaculatus Clark, 1862., Published as part of Hendrich, Lars, Balke, Michael & Watts, Chris H. S., 2016, Description of a new Tiporus Watts, 1985 from northern Queensland, Australia (Coleoptera: Dytiscidae, Hydroporinae), pp. 174-182 in Zootaxa 4189 (1) on page 180, DOI: 10.11646/zootaxa.4189.1.10, http://zenodo.org/record/165723, {"references":["Clark, H. (1862) Catalogue of the Dytiscidae and Gyrinidae of Australasia, with descriptions of new species. The Journal of Entomology. Descriptive and Geographical, 1, 399 - 421."]}

Tiporus queenslandicus sp. n. (Figs 1–2, 5, 8, 11) Type locality. Australia, Northern Queensland, Silver Valley, 17°34'S 145°18'E. Type material. Holotype, male: “ Australia: No. Qld. Silver Valley Dec 8, 1997 G.Challet ” [17°34'S 145°18'E, 700 m], “ Holotype Tiporus queenslandicus sp. n. Hendrich, Watts & Balke des. 2016” [printed red label] (SAMA). Paratypes. 6 exs., with same data as holotype (CGC, CLH, NHM); 18 exs., “Qld. Watson [Watsonville] 14 km W Herberton 31/3/96 C. Watts ”, “ SAMA Database No 25-001341” (SAMA); 5 exs., 12k N. Laura Qld 2/8/74 C. Watts, “SAMA Database No 25-001344” (SAMA); 16 exs., “Lakeland Downs Qld. 10/83 C.Watts”, “SAMA Database No 25-001342” (SAMA); 2 exs., “Qld. Irvine Bank 17.2524S 145 12 53E 2/8/03 CHSWatts”, “SAMA Database No 25-009342”, one specimen “DNA VOUCHER”, “SAMA Database No 25-009325”, (SAMA); 1 ex., “McIlwraith Rng. Weather Stn. N Qld. 23/7/82 C. Watts”, “SAMA Database No 25-001350” (SAMA); 1 ex., Qld. Herberton 10 km W 31/3/96 C.Watts”, “SAMA Database No 25-001349” (SAMA); 2 exs., “Cairns Qld.”, “SAMA Database No 25-001347” (SAMA); 1 ex., “Qld. Petford 20 km W 28/3/96 C.Watts”, “SAMA Database No 25- 001348” (SAMA); 1 ex., “25k. N. Coen Qld. 29/9/84 ”, 1 ex., Qld. Herberton 10 km W 31/3/96 C.Watts”, “SAMA Database No 25-001346” (SAMA); 1 ex., “Helenvale Qld. 29/7/82 C.Watts”, “SAMA Database No 25-001345” (SAMA); 2 exs., “ 10 km W Herberton Qld CHS Watts 31/3/96 ”, “ DNA VOUCHER ”, “2476” and “2479”, “ SAMA Database No 25 - 013137 ” (SAMA); 8 exs., “ Ewan Road, 10 miles west of Paluma N.Q. 5.I.66 J.G. Brooks Q.243”, “ SAMA Database No 25-001351” (SAMA); 4 exs., “ AUSTRALIA Qld Luster Cr 15 km se Mt Carbine Nov 15/90 D. Larson ” (ANIC); 3 exs., “ AUSTRALIA Qld.Watsonville 10 kmW Herberton Dec 9/90 Larson ” (ANIC); 2 exs., “ AUSTRALIA Qld Luster Cr 15 km se Mt Carbine Nov 15/90 D. Larson ” (ANIC); 6 exs., “ Mt. Spec. NQ 5.1.1966 J.G.Brooks ” (ANIC); 6 exs., “ Australia QLD. 15 km W Petford pool dry stream Nov. 3/90 Larson” (ANIC); 8 exs., “ Australia QLD. Silver Vlly 15 km s Herberton Dec 17/90 Larson ” (ANIC); 36 exs., “ AUSTRALIA Qld. Emu Cr 5 km E Petford, Nov 3 1990 D. Larson (ANIC, CLH, ZSM); 4 exs., “ AUSTRALIA Qld. Catherine Cr nr Collins [Collinsville] Weir Nov. 20.1990” (ANIC, ZSM); 1 ex., “ AUSTRALIA Qld. 10 km N Ravenshoe Dec 9/90 Larson & Storey ” (ANIC); 4 exs., “1306 S 142.56 E QLD Wenlock River Xing 26 Oct.1992 T.Weir, P.Zborowski still stagnant pools in dry river bed” (ANIC); 3 exs., “ Australia: No. Qld. Emu Creek near Petford Dec 4, 1997 G.L. Challet (CGC, CLH, NHM); 2 exs., “ Australia: No. Qld. pond @ Mcleod River Dec 4, 1997 GChallet” (CGC, NHM); 2 exs., “ Australia: No. Qld. Pond on Development Rd 2km S Cookshire Dec9, 1997 GChallet” (CGC, CLH); 1 ex., “ Australia: No. Qld. Silver Valley 14.4 km from Hwy 1 Dec 7, 1997 G.Challet ” (CGC); 1 ex., “ Australia: No. Qld. pond @ Mcleod River No. of Mt. Carbine Dec 3, 1997 G.Challet ” (CGC, NHM); 1 ex., “ Australia: No. Qld. Chilagoe Creek Dec 8, 1997 G.L. Challet ” (CGC). Each paratype was provided with a printed red paratype label. Remarks: Most of the specimens deposited in SAMA and ANIC were published under T. undecimmaculatus in Watts (2000) before. Description. A large, blackish and broadly-elongate species, widest in middle, with reddish markings on elytra and pronotum (Figs 1, 2). Measurements. TL: 4.6–4.7 mm (holotype 4.7 mm); TL–H: 4.1–4.2 mm (holotype 4.2 mm); MW: 2.4–2.5 mm (holotype 2.5 mm); TL/MW: 1.88–1.91. Colour. Head black. Pronotum sides, base and portions of middle dark red-brown; with two reddish spots basally. Elytron with six vague red patches dorsally and laterally. Ventral surface dark reddish-brown. Epipleuron, prosternum and legs mainly rufo-piceous; metatibia and -tarsus dark brown. Appendages rufo-piceous, apical segments of male protarsus almost black. Sculpture. Dorsal surface reticulate, strongly and densely rugose-punctate throughout, but punctures on dorsal surface shallow. Pronotum with a distinct raised ridge parallel to and a little distant from each side, area just inside ridge depressed, ridge and depression strongest anteriorly, weak posteriorly. Elytron weakly margined, strongly convex, rounded and slightly broadened behind middle. Prothoracic process narrow, strongly convex, roundly pointed at apex, little constricted between procoxae. Metacoxal lines raised, slightly to moderately diverging anteriorly. Male. Protarsus 3-segmented. Protarsomeres broadened and proximally expanded, those on mesotarsomeres little expanded. Single claw on protarsus (there is only one claw in males of Tiporus) weakly curved, slightly thickened and with a small tooth on underside near base. Male protibia with small tooth just beyond middle (Fig. 1). Tip of last abdominal ventrite weakly to strongly tuberculate in middle and with short carina. Metatibia normal, not expanded. Median lobe of aedeagus broad in centre but narrow at tip (Figs 5 a, b); right paramere as in Fig. 5 c. Female. Protarsus 5-segmented, robust somewhat expanded on inside, with two simple claws. Protibia on posterior side without tooth (Fig. 2). Tip of last abdominal ventrite flattened. Lateral extension of elytron near tip weak but visible. Etymology. Named after the Queensland State in Australia where all the type material has been collected. The species name is an adjective in the nominative singular. Distribution. North-eastern Queensland, Australia, from the Wenlock River in the north to Collinsville in the south (Fig. 11). Habitat. A lotic species. At Watsonville T. queenslandicus sp. n. was collected in deeper rest pools of a broad, shallow and sandy creek. The associated fauna comprised Copelatus irregularis W.J. Macleay, 1871, Laccophilus clarki Sharp, 1882, Necterosoma regulare Sharp, 1882, Sternopriscus hansardii (Clark, 1862), Hydroglyphus mastersii (W.J. Macleay, 1871), Hydroglyphus daemeli (Sharp, 1882) and Limbodessus rivulus (Larson, 1994). Near Herberton it was collected in a still flowing but temporary stream, forming pools together with Laccophilus walkeri Balfour-Browne, 1939 and the hydrophilid Berosus trishae Watts, 1987. Affinities. In habitus T. queenslandicus sp. n. is similar to T. giuliani (Fig. 3) and T. undecimmaculatus (Fig. 4). From the darker T. undecimmaculatus it can be separated by its well-developed dorsal colour pattern, the spine on the male protibia which is close to the centre (Fig. 8), not toward the apex (Fig. 10), and the broader central lobe of the aedeagus (Figs 7 a, b). From T. giuliani it can be distinguished by the form of the median lobe and parameres (Figs 6 a, b, c), and the almost black legs and tarsi. Furthermore, in T. giuliani the margin in lateral view of the elytron curves forward for a short distance immediately before it meets the edge of the pronotum, whereas it is straight or almost so in T. queenslandicus sp. n. Tiporus undecimmaculatus and T. giuliani are also distinctly smaller (TL of T. undecimmaculatus: 3.6–3.7 mm and T. giuliani: 4.0– 4.1 mm) than Tiporus queenslandicus sp. n. (TL: 4.6–4.7 mm). The new species is restricted to north-eastern Queensland whereas T. giuliani and T. undecimmaculatus are only known from the Northern Territory and north-western Australia (Fig. 11). Genetically, all Tiporus species are comparably strongly differentiated from each other based on mitochondrial cox1 gene divergences. This has been shown by Hendrich et al. (2010, fig. 2 D), where clustering specimens using uncorrected p-distances even at 10% threshold will retrieve the same clusters corresponding to morphologically delineated species. Accordingly, when we compared a cox1 sequence for Tiporus queenslandicus sp. n. available from Genbank (EU617001, from R. Leys and under the name T. undecimmaculatus) with one of our true T. undecimmaculatus (Genbank FR733053) we find divergence of about 14% which is very high for Coleoptera and strong indicator for presence of different species. Comparison with T. giuliani (FR733049) revealed the same magnitude of divergence.

Tiporus Watts, 1985 Tribe Hydroporini. Small (3.3���5.0 mm), oval and black beetles. Some species with reddish spots on the elytra and the pronotum. From the related genus Sekaliporus Watts, 1997, where the lateral margins of the elytron and of the pronotum form a nearly continuous straight or slightly sinuate line in combination, it can be distinguished by the elytral margin which is curved upwards to shoulders, and the 3-segmented male protarsi (Hendrich & Balke 2015). For a detailed generic analysis see Watts (2000)., Published as part of Hendrich, Lars, Balke, Michael & Watts, Chris H. S., 2016, Description of a new Tiporus Watts, 1985 from northern Queensland, Australia (Coleoptera: Dytiscidae, Hydroporinae), pp. 174-182 in Zootaxa 4189 (1) on page 175, DOI: 10.11646/zootaxa.4189.1.10, http://zenodo.org/record/165723, {"references":["Watts, C. H. S. (1985) A faunal assessment of Australian Hydradephaga. Proceedings of the Academy of Natural Sciences of Philadelphia, 137, 22 - 28.","Watts, C. H. S. (2000) Three new species of Tiporus Watts (Coleoptera: Dytiscidae) with re-descriptions of the other species of the genus. Records of the South Australian Museum, 33 (2), 89 - 99."]}

Fikáček, Martin, Watts, Chris H. S. (2015): Notes on the Australian Anacaenini (Coleoptera: Hydrophilidae): description of male of Phelea breviceps Hansen and unravelling the identity of Crenitis neogallica Gentili. Zootaxa 3980 (3): 427-434, DOI: http://dx.doi.org/10.11646/zootaxa.3980.3.6

Phelea breviceps Hansen, 1999 (Figs. 1���20) Phelea breviceps Hansen, 1999 a: 128. Phelea breviceps: Hansen (1999 b: 118, catalogue), Watts (2002: 93, notes on morphology and biology), Komarek & Beutel (2007: 221, phylogenetic analysis confirming assignment to the Anacaenini). Type locality. Australia, Tasmania, northern side of Cradle Mt., ca. 1370 m a.s.l. [ca. 41 �� 40.8 ��S 145 ��57.0��E]. Type material examined. Holotype: female (MCZ): ���N. side of CradleMtTas / c 4500 �� Feb 57 / Darlingtons // HOLOTYPE / Phelea breviceps / M. Hansen [red label, handwritten] // MCZ TYPE / 35208 [red label, partly handwritten] // Aug���Dec 2005 / MCZ Image / Database���. Additional material examined. AUSTRALIA: Tasmania: 1 male (BMNH): ���N. W. Tasmania / Cradle Valley / 3000 feet / 27.i. 1923 // Brit. Mus. / 1925 ��� 220 [GPS coordinates probably ca. 41 �� 38.5 ��S 145 �� 57.5 ��E]���; 8 females, 1 male, 2 unsexed specimens (SAMA): western side of Cradle Mt., i. 1959, in a shallow pond/tarn beside the overland track, 41 �� 42 ��� 36 ��� S 145 �� 56 ��� 16 ���E, leg. C.H.S. Watts. Complementary description (only taxonomically/phylogenetically important characters not mentioned by Hansen (1999 a) listed here): Head subquadrate in dorsal view, clypeus arcuately excised when seen in frontodorsal or frontal views (Figs. 1, 5); labrum large, strongly sclerotized, deflexed into a vertical position and hence well visible in frontal view only (Figs. 1, 2, 5, 7��� 8), bearing a transverse row of setae dorsally along anterior margin (Fig. 8); apical maxillary palpomere with a few digitiform sensilla on dorsoposterior face (Fig. 11); trichobothria present on clypeus, frons and pronotum (not examined on elytra), basal socket large but ca. of the size of surrounding punctation and hence not distinguishable from it under binocular microscope, hairs of trichobotria moderately long and rather stout (Fig. 9, 13); lateral glabrous portion of prothoracic hypomeron very wide, defined from pubescent portion by a ridge anteriorly (Figs. 3, 12); mesoventrite without distinct median posterior bulge and anterior pits or ridges (Figs. 3, 17, 19); mesofurca with short arms, dorsal projection connecting them with dorsal body wall absent; aedeagus (Fig. 4) 0.7 mm long, weakly sclerotized, phallobase long and wide, with large and wide, but distinctly detached manubrium, parameres distinctly shorter than the main part of phallobase (i.e. not considering manubrium), weakly sinuate on outer margin, angulate on inner margin apically, bearing numerous pore-like sensilla on the whole surface, median lobe much longer than parameres, slightly longer than main part of phallobase, gradually narrowing towards apex, with apical gonopore and basal struts slightly longer than body of median lobe. Diagnosis. Phelea breviceps is easy-to-recognize from other hydrophilid genera by the uniformly dark dorsal coloration (it is frequently even darker than in the illustrated specimen), rather compressed body, quadrate head with large fully sclerotized labrum which is deflexed and hence fully seen in frontal view only, mesoventrite nearly flat, without median projection, and by the unique pattern of abdominal pubescence (pubescence covering basal 2.5 ventrites, its posterior limit forms a bisinuate pattern on ventrite 3, and ventrites 4 and 5 are bare, just with few spine-like setae posteriorly; Figs. 3, 20). Distribution and biology. The species seems to be extremely restricted in distribution, as all known specimens were collected in close vicinity to Cradle Mt. in the western part of the mountain range in central Tasmania between ca. 900 and 1300 m a. s. l. (Fig. 6). Collecting circumstances are unknown for the holotype and the additional specimen from BMNH. Last known record of the species is from 1959 when a short series of that species was collected by the second author from small tarns on the plateau at the base of Cradle Mt. This may prove that the species is aquatic. On the other hand, the partly reduced ventral hydrophobic pubescence (absent in metafemora and part of abdominal ventrites, Figs. 3, 14, 20) may indicate that the species may be hygropetric, i.e. inhabiting wet places but normally not submerged underwater (e.g. in the similar way as in anacaenine genera Pseudorygmodus and Horelophus, see Fik��ček et al. 2012 and Fik��ček & Vondr��ček 2014). No specimens were collected after 1959, despite the efforts of C. Watts and L. Hendrich in similar habitats across Tasmania (but the 1959 collecting site was never re-visited as it is not easy to access). Comments on the phylogenetic position. Phelea was assigned to the Anacaenini in its original description by Hansen (1999 a) and later confirmed as a member of the tribe by Komarek & Beutel (2007) who considered it as a member of the basal anacaenine stock together with Crenitis. For this study we examined the external morphology and genitalia, studies on the structure of the mouthparts or the use of DNA were impossible due to a limited number of old specimens available. The absence of long mesofurca definitely excludes Phelea from a position within any group of the Hydrophilinae. Within the remaining hydrophilid groups, the absence of the scutellary stria and the presence of digitiform sensilla on the apical maxillary palpomere indicates the position within the subfamily Chaetarthriinae, and the absence of an abdominal fringe of setae excludes assignment to the Chaetarthriini (see the discussion in Fik��ček & Vondr��ček 2014 for details). Phelea also agrees with most anacaenine genera in the presence of a transverse row of setae on the labrum (character 11 in Komarek & Beutel 2007, overlooked for Phelea by the authors but actually present in this genus, see Fig. 8). The assignment to the Anacaenini hence seems well justified. Within Anacaenini, Phelea strongly resembles some members of Crenitis s.str. in the general body shape, large deflexed labrum, wide mentum with nearly straight anterior margin, pronotum with bisinuate posterior margin and lateral impressions (present in some Crenitis only, Komarek & Beutel 2007), absence of the accessory ridge below posterior margin of pronotum, wide horizontal epipleura (Fig. 18), the shape of the mesoventrite (with only slightly sinuate anapleural sutures meeting together at anterior margin of mesothorax), more-or-less cylindrical metatibiae and a large manubrium on the phallobase (see e.g. Ji & Komarek 2003). Some of these characters are also shared with the New Zealand endemic Horelophus which, moreover, has a very similar morphology of the aedeagus (with wide but short parameres and very long phallobase, see Fik��ček et al. 2012). Horelophus also agrees with Phelea completely in wing venation (compare drawings in Hansen (1999 a) and Fik��ček et al. (2012)) and shares with Phelea the presence of short club-like setae arising from dorsal punctures. On the other hand, Phelea does not seem to have modified setae on the male pro- and mesotarsi (Figs. 15���16) which are present in male Horelophus. Nevertheless, it seems justified to consider Phelea tentatively as a member of the Horelophus - Crenitis clade of the Anacaenini. Thanks to its autapomorphies (quadrate head, limited pubescence of the abdomen) and the morphology of mesoventrite (flat, without transverse ridge), Phelea is easily recognizable from Crenitis. Additional effort is however needed to understand its phylogenetic position and the relationship to Crenitis., Published as part of Fik����ek, Martin & Watts, Chris H. S., 2015, Notes on the Australian Anacaenini (Coleoptera: Hydrophilidae): description of male of Phelea breviceps Hansen and unravelling the identity of Crenitis neogallica Gentili, pp. 427-434 in Zootaxa 3980 (3) on pages 428-430, DOI: 10.11646/zootaxa.3980.3.6, http://zenodo.org/record/244776, {"references":["Hansen, M. (1999 a) Fifteen new genera of Hydrophilidae (Coleoptera), with remarks on generic classification of the family. Entomologica Scandinavica 30, 121 - 172. http: // dx. doi. org / 10.1163 / 187631200 X 00228","Hansen, M. (1999 b) World Catalogue of Insects. Vol. 2. Hydrophiloidea (Coleoptera). Apollo Books, Stenstrup, 416 pp.","Watts, C. H. S. (2002) Checklists & Guides to the identification, to genus, of adult & larval Australian water beetles of the families Dytiscidae, Noteridae, Hygrobiidae, Halipilidae, Gyrinidae, Hydraenidae and the superfamily Hydrophiloidea (Insecta: Coleoptera). Identification and Ecology guide 43. Cooperative Research Centre for freshwater Ecology, Albury, 110 pp.","Komarek, A. & Beutel, R. G. (2007) Phylogenetic analysis of Anacaenini (Coleoptera: Hydrophilidae: Hydrophilinae) based on morphological characters of adults. Systematic Entomology 32, 205 - 226. http: // dx. doi. org / 10.1111 / j. 1365 - 3113.2006.00359. x","Fikacek, M., Leschen, R. A. B., Newton, A. F. & Gunter, N. (2012) Horelophus walkeri rediscovered: adult morphology and notes on biology (Coleoptera: Hydrophilidae). Acta Entomologica Musei Nationalis Pragae, 52, 129 - 146. Fikacek, M. & Vondracek, D. (2014) A review of Pseudorygmodus (Coleoptera: Hydrophilidae), with notes on the classification of the Anacaenini and on distribution of genera endemic to southern South America. Acta Entomologica Musei Nationalis Pragae, 54, 479 - 514.","Ji, L. & Komarek, A. (2003) HYDROPHILIDAE: II. The Chinese species of Crenitis Bedel, with descriptions of two new species (Hydrophilidae). In: Jach, M. A. & Ji, L. (Eds.), Water Beetles of China Vol. III. Wiener Coleopterologenverein, Wien, pp. 397 - 409."]}