Wikipedia

Agraphydrus

Agraphydrus is a genus of water scavenger beetle in the family Hydrophilidae represented by 205 described species.[1] It is distributed across the Afrotropical, Australasian, and Indomalayan realms.

Agraphydrus
Dorsal view of Agraphydrussp.
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Family: Hydrophilidae
Subfamily: Acidocerinae
Genus: Agraphydrus
Régimbart, 1903
Diversity
205 species
Synonyms
  • Gymnhelochares d’Orchymont, 1932
  • Pseudohelochares Satô, 1960
  • Horelophopsis Hansen, 1997
  • Pseudopelthydrus Jia, 1998
  • Megagraphydrus Hansen, 1999

Taxonomy

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The genus Agraphydrus was described for the first time by Régimbart in 1903.[2]

It currently contains 205 described species, partly thanks to a series revisionary work developed by Albretch Komarek.[3][4][5][6][7]

Description

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Small beetles (1.4–4.8 mm), pale/yellowish to dark brown in coloration, eyes not emarginate, labrum exposed, maxillary palps moderately long. Elytra without sutural striae, not laterally explanate; the elytral punctation ranges from very fine and subtle to coarse.

A complete diagnosis was presented by Girón and Short,[1] but also see Komarek and collaborators.[3][4][5][6][7]

Habitat

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According to Girón and Short,[1] based on annotations by Komarek and collaborators:[3][4][5][6][7]

Agraphydrus can be found in an extremely broad range of habitats, from rivers, streams and forest pools, to hygropetric environments around waterfalls or seepages over rocks; a few species have been collected in terrestrial habitats by sifting moss and leaves from near water bodies, or in the gravel along the bank of a river; in many cases specimens have been found associated with floating vegetation, mosses and algae.[1]

Species

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  1. Agraphydrus abrasus Komarek and Freitag, 2020 [6]
  2. Agraphydrus activus Komarek and Hebauer, 2018 [7]
  3. Agraphydrus acutus Komarek, 2020 [5]
  4. Agraphydrus aethiopicus Komarek, 2020 [5]
  5. Agraphydrus agilis Komarek and Hebauer, 2018 [7]
  6. Agraphydrus albescens (Régimbart, 1903) [8]
  7. Agraphydrus ampullatus Komarek and Freitag, 2020 [6]
  8. Agraphydrus anacaenoides Komarek, 2019 [4]
  9. Agraphydrus anatinus Komarek, 2018 [3]
  10. Agraphydrus andamanicus Komarek, 2018 [3]
  11. Agraphydrus andringitra Komarek, 2020 [5]
  12. Agraphydrus angulatus Komarek, 2019 [4]
  13. Agraphydrus angustatus Komarek, 2020 [5]
  14. Agraphydrus angustipenis Komarek, 2018 [3]
  15. Agraphydrus anhuianus (Hebauer, 2000) [9]
  16. Agraphydrus annapurnensis Komarek, 2018 [3]
  17. Agraphydrus arduus Komarek and Hebauer, 2018 [7]
  18. Agraphydrus ater Komarek, 2018 [3]
  19. Agraphydrus atripalpis Komarek, 2020 [5]
  20. Agraphydrus attenuatus (Hansen, 1999) [10]
  21. Agraphydrus audax Komarek and Hebauer, 2018 [7]
  22. Agraphydrus avita (Hansen, 1997) [11]
  23. Agraphydrus bacchusi Komarek, 2019 [4]
  24. Agraphydrus balkeorum Komarek, 2019 [4]
  25. Agraphydrus batak Komarek and Freitag, 2020 [6]
  26. Agraphydrus bhutanensis Komarek, 2018 [3]
  27. Agraphydrus bicoloratus Komarek, 2020 [5]
  28. Agraphydrus bilardoi Komarek, 2020 [5]
  29. Agraphydrus biltoni Komarek, 2020 [5]
  30. Agraphydrus biprojectus Minoshima, Komarek, and Ôhara, 2015 [12]
  31. Agraphydrus borneensis Komarek, 2019 [4]
  32. Agraphydrus boukali Komarek, 2018 [3]
  33. Agraphydrus brevilobatus Komarek and Freitag, 2020 [6]
  34. Agraphydrus brevipenis Komarek, 2019 [4]
  35. Agraphydrus burmensis Komarek, 2019 [4]
  36. Agraphydrus calvus Komarek and Hebauer, 2018 [7]
  37. Agraphydrus camerunensis Komarek, 2020 [5]
  38. Agraphydrus cantonensis Komarek and Hebauer, 2018 [7]
  39. Agraphydrus carinatulus Komarek, 2019 [4]
  40. Agraphydrus cervus Komarek, 2019 [4]
  41. Agraphydrus ceylonensis Komarek, 2018 [3]
  42. Agraphydrus chinensis Komarek and Hebauer, 2018 [7]
  43. Agraphydrus cinnamum Komarek, 2018 [3]
  44. Agraphydrus clarus Komarek, 2019 [4]
  45. Agraphydrus comes Komarek and Hebauer, 2018 [7]
  46. Agraphydrus communis Komarek, 2018 [3]
  47. Agraphydrus confusus Komarek and Hebauer, 2018 [7]
  48. Agraphydrus congolensis Komarek, 2020 [5]
  49. Agraphydrus conicus Komarek and Hebauer, 2018 [7]
  50. Agraphydrus connexus Komarek and Hebauer, 2018 [7]
  51. Agraphydrus constrictus Komarek, 2018 [3]
  52. Agraphydrus contractus Komarek and Hebauer, 2018 [7]
  53. Agraphydrus coomani (d'Orchymont, 1927) [13]
  54. Agraphydrus coronarius Minoshima, Komarek, and Ôhara, 2015 [12]
  55. Agraphydrus crassipenis Komarek, 2018 [3]
  56. Agraphydrus dapengensis Yang & Jia, 2021 [14]
  57. Agraphydrus decipiens Minoshima, Komarek, and Ôhara, 2015 [12]
  58. Agraphydrus delineatus Komarek, 2019 [4]
  59. Agraphydrus elongatus Ribera, Hernando, and Cieslak, 2019 [15]
  60. Agraphydrus engkari Komarek, 2019 [4]
  61. Agraphydrus excisus Komarek, 2019 [4]
  62. Agraphydrus exedis (d'Orchymont, 1937) [16]
  63. Agraphydrus exiguus Komarek, 2019 [4]
  64. Agraphydrus falcatus Komarek, 2018 [3]
  65. Agraphydrus fasciatus Komarek and Hebauer, 2018 [7]
  66. Agraphydrus fikaceki Komarek and Hebauer, 2018 [7]
  67. Agraphydrus flavescens Komarek, 2020 [5]
  68. Agraphydrus flavipes Komarek, 2020 [5]
  69. Agraphydrus flavonotus Komarek, 2018 [3]
  70. Agraphydrus floresinus Komarek, 2019 [4]
  71. Agraphydrus fontis Komarek, 2020 [5]
  72. Agraphydrus forcipatus Komarek and Hebauer, 2018 [7]
  73. Agraphydrus fortis Komarek, 2018 [3]
  74. Agraphydrus fujianensis Komarek and Hebauer, 2018 [7]
  75. Agraphydrus geminus (d'Orchymont, 1932) [17]
  76. Agraphydrus gereckei Komarek, 2020 [5]
  77. Agraphydrus gilvus Komarek, 2018 [3]
  78. Agraphydrus glaber Komarek, 2018 [3]
  79. Agraphydrus globipenis Komarek and Hebauer, 2018 [7]
  80. Agraphydrus goldschmidti Komarek, 2020 [5]
  81. Agraphydrus gracilipalpis Komarek, 2018 [3]
  82. Agraphydrus hamatus Komarek, 2019 [4]
  83. Agraphydrus hanseni (Satô and Yoshitomi, 2004) [18]
  84. Agraphydrus heinrichi Komarek, 2018 [3]
  85. Agraphydrus helicopter Komarek, 2019 [4]
  86. Agraphydrus hendrichi Komarek, 2019 [4]
  87. Agraphydrus heterochromatus Komarek, 2019 [4]
  88. Agraphydrus hortensis Komarek, 2019 [4]
  89. Agraphydrus hygropetricus Komarek, 2018 [3]
  90. Agraphydrus igneus Komarek and Hebauer, 2018 [7]
  91. Agraphydrus imitans Komarek, 2019 [4]
  92. Agraphydrus indicus (d'Orchymont, 1932) [17]
  93. Agraphydrus inflatus Komarek, 2018 [3]
  94. Agraphydrus infuscatus Komarek, 2019 [4]
  95. Agraphydrus insidiator Minoshima, Komarek, and Ôhara, 2015 [12]
  96. Agraphydrus ishiharai (Matsui, 1994) [19]
  97. Agraphydrus jaechi (Hansen, 1999) [10]
  98. Agraphydrus jankodadai Komarek, 2019 [4]
  99. Agraphydrus jilanzhui Komarek and Hebauer, 2018 [7]
  100. Agraphydrus kallar Komarek, 2018 [3]
  101. Agraphydrus kathapa Komarek, 2019 [4]
  102. Agraphydrus kempi (d'Orchymont, 1922)
  103. Agraphydrus khasiensis Komarek, 2018 [3]
  104. Agraphydrus kodaguensis Komarek, 2018 [3]
  105. Agraphydrus komareki Yang & Jia, 2021 [14]
  106. Agraphydrus laocaiensis Komarek, 2019 [4]
  107. Agraphydrus latus Komarek, 2019 [4]
  108. Agraphydrus longipalpus (Jia, 1998) [20]
  109. Agraphydrus longipenis Komarek and Hebauer, 2018 [7]
  110. Agraphydrus lunaris Komarek, 2019 [4]
  111. Agraphydrus luteilateralis (Minoshima and Fujiwara, 2009) [21]
  112. Agraphydrus madagascarensis Komarek, 2020 [5]
  113. Agraphydrus maehongsonensis Komarek, 2019 [4]
  114. Agraphydrus malayanus (Hebauer, 2000) [9]
  115. Agraphydrus malkini Komarek, 2020 [5]
  116. Agraphydrus manfredjaechi Komarek, 2019 [4]
  117. Agraphydrus masatakai Minoshima, Komarek, and Ôhara, 2015 [12]
  118. Agraphydrus matoposensis Komarek, 2020 [5]
  119. Agraphydrus mazzoldii Komarek, 2019 [4]
  120. Agraphydrus meghalayanus Komarek, 2018 [3]
  121. Agraphydrus microphthalmus Komarek, 2019 [4]
  122. Agraphydrus minutissimus (Kuwert, 1890) [22]
  123. Agraphydrus mirabilis Komarek, 2019 [4]
  124. Agraphydrus montanus Minoshima, Komarek, and Ôhara, 2015 [12]
  125. Agraphydrus muluensis Komarek, 2019 [4]
  126. Agraphydrus musculus Komarek, 2019 [4]
  127. Agraphydrus namthaensis Komarek, 2019 [4]
  128. Agraphydrus nanus Komarek, 2018 [3]
  129. Agraphydrus narusei (Satô, 1960) [23]
  130. Agraphydrus nemorosus Komarek, 2019 [4]
  131. Agraphydrus nepalensis Komarek, 2018 [3]
  132. Agraphydrus niger Komarek, 2018 [3]
  133. Agraphydrus nigroflavus Komarek, 2019 [4]
  134. Agraphydrus obesus Komarek, 2019 [4]
  135. Agraphydrus obscuratus Komarek, 2018 [3]
  136. Agraphydrus obsoletus Komarek, 2018 [3]
  137. Agraphydrus occultus Komarek and Freitag, 2020 [6]
  138. Agraphydrus ogatai Minoshima, 2016 [24]
  139. Agraphydrus orbicularis Komarek, 2019 [4]
  140. Agraphydrus orientalis (d'Orchymont, 1932) [17]
  141. Agraphydrus palawanensis Komarek and Freitag, 2020 [6]
  142. Agraphydrus pallidus Komarek, 2019 [4]
  143. Agraphydrus papuanus Komarek, 2019 [4]
  144. Agraphydrus pauculus (Knisch, 1924) [25]
  145. Agraphydrus pauper Komarek, 2020 [5]
  146. Agraphydrus pelingeni Komarek and Freitag, 2020 [6]
  147. Agraphydrus penangensis Komarek, 2019 [4]
  148. Agraphydrus piceus Komarek, 2019 [4]
  149. Agraphydrus politus (Hansen, 1999) [10]
  150. Agraphydrus praecipuus (d'Orchymont, 1937) [26]
  151. Agraphydrus protentus Komarek, 2018 [3]
  152. Agraphydrus pseudoniger Yang & Jia, 2021 [14]
  153. Agraphydrus pullus Komarek, 2018 [3]
  154. Agraphydrus punctatellus Régimbart, 1903 [2]
  155. Agraphydrus punctulatus Komarek, 2018 [3]
  156. Agraphydrus puzhelongi (Jia, 2010) [27]
  157. Agraphydrus pygmaeus (Knisch, 1924) [25]
  158. Agraphydrus raucus Komarek, 2019 [4]
  159. Agraphydrus reductus Komarek and Hebauer, 2018 [7]
  160. Agraphydrus regularis (Hansen, 1999) [10]
  161. Agraphydrus reticulatus Komarek, 2019 [4]
  162. Agraphydrus reticuliceps Komarek and Hebauer, 2018 [7]
  163. Agraphydrus rhodesiensis Komarek, 2020 [5]
  164. Agraphydrus rhomboideus Komarek, 2019 [4]
  165. Agraphydrus rivalis Komarek, 2020 [5]
  166. Agraphydrus robustus Komarek and Hebauer, 2018 [7]
  167. Agraphydrus rostratus Komarek, 2018 [3]
  168. Agraphydrus rugosus Komarek, 2018 [3]
  169. Agraphydrus sabulosus Yang & Jia, 2021 [14]
  170. Agraphydrus sarawakensis Komarek, 2019 [4]
  171. Agraphydrus schoedli Komarek, 2019 [4]
  172. Agraphydrus schoenmanni Komarek and Hebauer, 2018 [7]
  173. Agraphydrus scintillans Komarek, 2019 [4]
  174. Agraphydrus scutifer Komarek, 2020 [5]
  175. Agraphydrus setifer Komarek and Hebauer, 2018 [7]
  176. Agraphydrus shaverdoae Komarek, 2019 [4]
  177. Agraphydrus siamensis (Hansen, 1999) [10]
  178. Agraphydrus sipekorum Komarek, 2018 [3]
  179. Agraphydrus skalei Komarek, 2019 [4]
  180. Agraphydrus spadix Komarek, 2019 [4]
  181. Agraphydrus spinosus Komarek, 2019 [4]
  182. Agraphydrus splendens Komarek and Hebauer, 2018 [7]
  183. Agraphydrus stagnalis (d'Orchymont, 1937)
  184. Agraphydrus stramineus Komarek, 2019 [4]
  185. Agraphydrus sucineus Komarek, 2019 [4]
  186. Agraphydrus sundaicus Komarek, 2019 [4]
  187. Agraphydrus tamdao Komarek, 2019 [4]
  188. Agraphydrus taprobanensis Komarek, 2018 [3]
  189. Agraphydrus tenuipalpis Komarek and Freitag, 2020 [6]
  190. Agraphydrus thaiensis Minoshima, Komarek, and Ôhara, 2015 [12]
  191. Agraphydrus tristis Komarek, 2019 [4]
  192. Agraphydrus tulipa Komarek, 2019 [4]
  193. Agraphydrus tumidus Komarek, 2020 [5]
  194. Agraphydrus tumulosus Komarek, 2018 [3]
  195. Agraphydrus umbrosus Komarek and Hebauer, 2018 [7]
  196. Agraphydrus uncinatus Komarek and Hebauer, 2018 [7]
  197. Agraphydrus usambaraensis Komarek, 2020 [5]
  198. Agraphydrus uvaensis (Hebauer, 2000) [9]
  199. Agraphydrus vadoni Komarek, 2020 [5]
  200. Agraphydrus variabilis Komarek and Hebauer, 2018 [7]
  201. Agraphydrus vietnamensis Komarek, 2019 [4]
  202. Agraphydrus villiersi (Balfour-Browne, 1958) [28]
  203. Agraphydrus wangmiaoi Komarek and Hebauer, 2018 [7]
  204. Agraphydrus yunnanensis Komarek and Hebauer, 2018 [7]
  205. Agraphydrus zetteli Komarek and Freitag, 2020 [6]

References

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  1. ^ a b c d Girón, Jennifer C.; Short, Andrew Edward Z. (2021-06-18). "The Acidocerinae (Coleoptera, Hydrophilidae): taxonomy, classification, and catalog of species". ZooKeys (1045): 1–236. Bibcode:2021ZooK.1045....1G. doi:10.3897/zookeys.1045.63810. ISSN 1313-2970. PMC 8233300. PMID 34228772.
  2. ^ a b Régimbart, MA (1903). "Coléoptères aquatiques (Haliplidae, Dytiscidae, Gyrinidae et Hydrophilidae) recueillis dans le Sud de Madagascar par M. Ch. Alluaud (Juillet 1900 – Mai 1901)". Annales de la Société Entomologique de France. 72: 1–51.
  3. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao Komarek, A. (2018). "Taxonomic revision of AgraphydrusRégimbart, 1903 II. The Indian Subcontinent" (PDF). Koleopterologische Rundschau. 88: 103–204. Archived from the original (PDF) on 2021-06-24. Retrieved 2022-01-07.
  4. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw ax ay az ba bb bc bd be bf bg bh bi bj bk Komarek, A. (2019). "Taxonomic revision of AgraphydrusRégimbart, 1903 III. Southeast Asia (except Philippines) and Australian Region (Coleoptera: Hydrophilidae: Acidocerinae)" (PDF). Koleopterologische Rundschau. 89: 151–316. Archived from the original (PDF) on 2021-06-24. Retrieved 2022-01-07.
  5. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab Komarek, A. (2020). "Taxonomic revision of AgraphydrusRégimbart, IV. Africa, Western Asia, and redescription of the genus (Coleoptera: Hydrophilidae: Acidocerinae)" (PDF). Koleopterologische Rundschau. 90: 127–200. Archived from the original (PDF) on 2021-02-14. Retrieved 2022-01-07.
  6. ^ a b c d e f g h i j k l Komarek, A.; Freitag, H. (2020). "Taxonomic revision of AgraphydrusRégimbart, 1903 V. Philippine species and their first DNA barcodes" (PDF). Koleopterologische Rundschau. 90: 201–242. Archived from the original (PDF) on 2021-02-14. Retrieved 2022-01-07.
  7. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah Komarek, A.; Hebauer, F. (2018-07-30). "Taxonomic revision of Agraphydrus Régimbart, 1903 I. China and Taiwan (Coleoptera: Hydrophilidae: Acidocerinae)". Zootaxa. 4452 (1): 1–101. doi:10.11646/zootaxa.4452.1.1. ISSN 1175-5334. PMID 30314249. S2CID 52975164.
  8. ^ Régimbart, M. (1903). "Coléoptères aquatiques (Haliplidae, Dytiscidae, Gyrinidae et Hydrophi-lidae) recueillis dans le Sud de Madagascar par M. Ch. Alluaud (Juillet 1900 – Mai 1901)". Annales de la Société Entomologique de France. 72: 1–51.
  9. ^ a b c Hebauer, F. (2000). "The genus Megagraphydrus Hansen, 1999, with description of new species (Coleoptera: Hydrophilidae)". Acta Coleopterologica. 16: 14–22.
  10. ^ a b c d e Hansen, Michael (1999-01-01). "Fifteen new genera of Hydrophilidae (Coleoptera), with remarks on the generic classification of the family". Insect Systematics & Evolution. 30 (2): 121–172. doi:10.1163/187631200X00228. ISSN 1876-312X.
  11. ^ Hansen, M. (1997). "A new subfamily for a remarkable new genus and species of Hydrophilidae from New Guinea (Coleoptera: Hydrophilidae)". Annales Zoologici. 47: 107–110.
  12. ^ a b c d e f g Minoshima, Y. N.; Komarek, A.; Ôhara, M. (2015). "A revision of Megagraphydrus Hansen (Coleoptera, Hydrophilidae): synonymization with Agraphydrus Régimbart and description of seven new species". Zootaxa. 3930: 1–63. PMID 25781810.
  13. ^ d’Orchymont, A. (1927). "Papers on Malayan aquatic biology. V. Notes on the Hydrophilidae in the Federated Malay States Museums". Journal of the Federated Malay States Museums. 13 (4): 246–252.
  14. ^ a b c d Yang, Zhen-ming; Jia, Fenglong; Jiang, Lu; Guo, Qiang (2021-05-18). "Four new species of Agraphydrus Régimbart, 1903 with additional faunistic record from China (Coleoptera, Hydrophilidae, Acidocerinae)". Deutsche Entomologische Zeitschrift. 68 (1): 189–205. doi:10.3897/dez.68.66200. ISSN 1860-1324. S2CID 236330170.
  15. ^ Ribera, I.; Hernando, C.; Cieslak, A. (2019). "Aquatic Coleoptera of North Oman, with description of new species of Hydraenidae and Hydrophilidae" (PDF). Acta Entomologica Musei Nationalis Pragae. 59: 253–272. doi:10.2478/aemnp-2019-0021. hdl:10230/44839. S2CID 198381025.
  16. ^ d’Orchymont, A. (1937). "Descriptions of three new Hydrophilidae from India". Records of the Indian Museum. 39: 29–31.
  17. ^ a b c d’Orchymont, A. (1932). "Zur Kenntnis der Kolbenwasserkäfer (Palpicornia) von Sumatra, Java und Bali". Archiv für Hydrobiologie. Supplement Band IX: 623–714. [pl. XIV–XVII.]
  18. ^ Satô, M.; Yoshitomi, H. (2004). "Discovery of a second representative of the genus Horelophopsis (Coleoptera, Hydrophilidae) from the Ryukyu archipelago, Japan". Elytra. 32: 41–49.
  19. ^ Matsui, E. (1994). "Three new species of the genus Enochrus from Japan and Taiwan (Coleoptera: Hydrophilidae)". Transactions of the Shikoku Entomological Society. 20: 215–220.
  20. ^ Jia, F. (1998). "A new genus Pseudopelthydrus gen. n. from Hainan Island, China (Coleoptera: Hydrophilidae: Hydrophilinae)". Chinese Journal of Entomology. 18: 225–230.
  21. ^ Minoshima, Y. N.; Fujiwara, J. (2009). "Discovery of water scavenger beetle genus Megagraphydrus Hansen (Coleoptera, Hydrophilidae) from Japan, with description of a new species" (PDF). Elytra. 37: 53–64.
  22. ^ Kuwert, A. (1890). "Bestimmungs-Tabellen der europäischen Coleopteren. XX. Heft. Hydrophilidae. II. Abteilung: Sphaeridiini und Helophorini". Verhandlungen des naturforschenden Vereins in Brünn. 28: 159–328.
  23. ^ Satô, M. (1960). "One new genus and two new species of the subtribe Helocharae from Japan (Coleoptera: Hydrophilidae)". Transactions of the Shikoku Entomological Society. 6: 76–80.
  24. ^ Minoshima, Yûsuke N. (2016). "Taxonomic review of Agraphydrus from Japan (Coleoptera: Hydrophilidae: Acidocerinae)". Entomological Science. 19 (4): 351–366. doi:10.1111/ens.12213. ISSN 1479-8298. S2CID 89394202.
  25. ^ a b Knisch, A. (1924). "Neue neotropische Palpicornier. (Col. Hydrophilidae. - Op. 16.)". Wiener Entomologische Zeitung. 41: 114–140.
  26. ^ d’Orchymont, A. (1937). "Contribution à l'étude des Palpicornia IX". Bulletin et Annales de la Société Entomologique de Belgique. 77: 213–255.
  27. ^ Jia, Fenglong (2010-06-08). "Megagraphydrus puzhelongi, sp. n., a new water scavenger beetle from China (Coleoptera: Hydrophilidae: Hydrophilinae)". Zootaxa. 2498 (1): 65. doi:10.11646/zootaxa.2498.1.6. ISSN 1175-5334.
  28. ^ Balfour-Browne, J. (1958). "La Reserve naturelle integrale du Mont Nimba, Guinea. VI. Coléoptères Hydrophiloides". Mémoires de l'Institut français d'Afrique noire. 53: 169–190.
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