Bol. Mus. Nac. Hist. Nat. Parag. Vol. 17, nº 1 (Ago. 2013): 100-10076-82

THE OCCURRENCE OF (COLEOPTERA: ) IN PARAGUAY AND A WORLD CHECKLIST OF SPECIES

William D. Shepard1, 2, Cheryl B. Barr1, 3 & Carlos A. Aguilar Julio4

1Essig Museum of Entomology, 1107 Valley Life Sciences Bldg., University of California, Berkeley, California 94720 USA. E-mail: 2william.shepard @csus.edu and [email protected] 4Museo Nacional de Historia Natural del Paraguay, Sucursal 1 Campus, Central XI, San Lorenzo, Paraguay. E-mail: [email protected]

Abstract.- The occurrence of the family Torridincolidae is reported from Paraguay. Adults, pupae and larvae were collected within the Atlantic Forest ecosystem from the vicinity of waterfalls in the Yvytyruzú Cordillera, De- partment Guairá. A checklist is given for all Torridincolidae species described to date. Key words: Torridincollidae, Myxophaga, Paraguay, new records, World checklist.

Resumen.- La ocurrencia de la familia Torridincolidae se divulga de Paraguay. Se colectaron adultos, pupas y larvas en la ecosistema del Bosque Atlántico cerca de saltos de Cordillera del Yvytyruzú, departamento Guairá. Se da una checklist de todas las especies de Torridincolidae actualmente descrito. Palabras clave: Torridincollidae, Myxophaga, Paraguay, nuevos reportes, checklist mundial.

Torridincolidae, commonly called torrent beet- cycle are aquatic, and most species occur in les, is a small, little-known family of aquatic thin layers of water, especially in fast-flowing in the Suborder Myxophaga with 37 habitats such as waterfalls. formally described species in seven genera The literature on Torridincolidae is largely (Braule-Pinto et al., 2011; Hajek et al., 2011; systematic and was well reviewed by Vanin Vanin, 2011). They are largely restricted to the (2011). Aquatic respiration in adults of most Southern Hemisphere although some species torridincolid species is via a plastron and lar- occur in Japan and China. All stages of the life vae breathe via segmented trachael gills (Hin- 1 2

Figure 1-2. new species. 1) Adult. 2) Larva. Scale bar for each = 1 mm.

Nota Boletín del Museo nacional de Historia Natural del Paraguay Vol. 17, Nº 1 (Agosto 2013) 77

3 4

Figures 3-4. Salto Paí. 3) upper cascade. 4) Bedrock ledge inhabited by Ytu new species. ton, 1967; Endrödy-Younga, 1997). The possi- water. The specimens from Salto Suizo were ble occurrence of the family in Paraguay was collected from the surface of a boulder, below cited previously by Aguilar (2010) and Jäch & the plunge pool (Fig. 5), over which flowed a Balke (2008), but the specimens were thought thin layer of fast water. All specimens are cu- to be lost. The location of those specimens is rrently housed at Essig Museum of Entomolo- now known and their data are reported below. gy, University of California, Berkeley, where Shepard and Aguilar have been surveying they are being studied and described. the aquatic byrrhoid beetles of Paraguay for These two collections represent the far- several years and in 2010 published a prelimi- thest west that members of the Ytu zeus spe- nary report which included taxonomic keys, cies group have been collected in the Atlantic geographic distribution and habitat notes for Forest ecosystem. Ytu brutus Spangler, another the known taxa. This paper was the first to deal member of the Ytu zeus species group, occurs with the aquatic beetle fauna of Paraguay. The farther west, but in the Amazon basin near the torridincolid specimens discussed here were Brazilian/Bolivian border outside of the Atlan- found in February 2011 during their continuing tic Forest ecosystem (Spangler, 1980). Because survey work. The collections, including larvae, both Paraguayan locations are in remnants of pupae and adults (Figs. 1–2), represent an un- the Interior Atlantic Forest, we expect to find described member of the Ytu zeus Reichardt additional populations, and perhaps additio- species group. The species was collected at nal species, when more of this ecosystem is two localities by Shepard and Barr, both in the surveyed in Paraguay. The presence of these Yvytyruzú Cordillera in Departamento Guai- two populations further bolsters the case for rá: Salto Paí, Arroyo Tacuara, 3.7 km SSW of the conservation and protection of this unique, Melgarejo, S25º44.916’ W56º15.231’, eleva- endangered ecosystem, as advocated in Ga- tion 221 m; and Salto Suizo, Arroyo Libertad, lindo-Leal & Câmara (2003). Over time more 6 km SSE of Melgarejo, S25º46.299’ W56º and more river systems are threatened by the 13.575’, elevation 243 m. construction of hydroelectric dams (Fahey and The specimens from Salto Paí were collec- Langhammer, 2003) and by the loss of forests ted in a series of cascades separated by deep in river basins due to soybean farming. plunge pools (Figs. 3–4). The specimens were The four specimens that were thought to on massive sandstone outcroppings and were be lost are now known to be in the Naturhisto- covered by a relatively thin layer of swift rische Museum Wien, in Vienna, Austria. The

The occurrence of Torridincolidae in Paraguay and a world checklist of species 78 Boletín del Museo nacional de Historia Natural del Paraguay Vol. 17, Nº 1 (Agosto 2013) collection data is: PARAGUAY: Paraguari, 1. Delevea bertrandi Reichardt, 1976: 209 (TYPE Sapucay, IV 1994, Ulf Drechsel. They are also SPECIES)...... South Africa. members of Ytu (Manfred Jäch, pers. comm.) 2. Delevea namibensis Endrödy-Younga, 1997: but they have not yet been examined to deter- 320...... South Africa. mine if they are the same species as the one Genus: Satonius Endrödy-Younga, 1997: 317 reported above. Revision of the genus: Hájek & Fikáček (2008). An updated world checklist of the Torridin- 1. Satonius kurosawai (Satô, 1982): 279 (TYPE colidae is provided as follows: SPECIES)...... Japan. = Delevea kurosawai Satô, 1982: 279 (origi- CHECKLIST OF TORRIDINCOLIDAE nal description). OF THE WORLD = Satonius kurosawai (Satô, 1982); (En- drödy-Younga (1997: 318) (new combina- Family: Torridincolidae Steffan, 1964: 199 tion, redescription); Subfamily: Deleveinae Endrödy-Younga, Beutel (1998: 54) (description of larva); Ha- 1997:3171 yashi (2007: 77) (faunistics, photographs of imago and larva); Hayashi & Kadowaki (2007: Genus: Delevea Reichardt, 1976: 209 149) (faunistics, photograph of imago); Hayas- hi (2008: 61) (faunistics, photographs of pupa); *A phylogenetic analysis by Beutel (1998/99) questions the va- Hájek & Fikáček (2008: 662) (adult diagnosis lidity of the subfamily Deleveinae, but no taxonomic alternative was proposed. and bionomic and description of the larvae);

Figure 5. Salto Suizo, waterfall and arroyo upstream from boulder where Ytu new species was collected.

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Hájek et al.(2011: 56) (faunistics, description of (TYPE SPECIES)...... Madagascar. wing polymorphy). Genus: Torridincola Steffan, 1964: 193 2. Satonius fui Hájek, Yoshitomi, Fikáček, Hayas- 1. Torridincola rhodesica Steffan, 1964: 194 hi & Jia, 2011: 52...... China. (TYPE SPECIES)...... Zimbabwe. 3. Satonius jaechi Hájek, Yoshitomi, Fikáček, 2. Torridincola congolesica Steffan, 1973: Hayashi & Jia, 2011: 53...... China. 639...... Congo-Kinshasa. 4. Satonius schoenmanni Hájek & Fikáček, 2008: 3. Torridincola natalesica Steffan, 1973: 642...... 663...... China...... South Africa. 5. Satonius stysi Hájek & Fikáček, Genus: Ytu Reichardt, 1973: 131 2008: 663...... China. Revision of the genus: Reichardt & Vanin (1977). 6. Satonius wangi Hájek & Fikáček, 2008: 1. Ytu zeus Reichardt, 1973: 137 (TYPE SPE- 665...... China. CIES)...... Brazil.

Subfamily: Torridincolinae Steffan, 1964 2. Ytu angra Reichardt & Vanin, 1977: 131...... (Endrödy-Younga 1997) ...... Brazil. = Ptyopterinae Abdullah, 1974 3. Ytu artemis Reichardt, 1973: 143...... Brazil. (syn. Endrödy-Younga 1997) 4. Ytu athena Reichardt, 1973: 139...... Brazil. Genus: Claudiella Reichardt & Vanin, 1976: 212 5. Ytu brutus Spangler, 1980: 145...... Brazil. 1. Claudiella ingens Reichardt & Vanin, 1976: 214 (TYPE SPECIES)...... Brazil. 6. Ytu cleideae Vanin, 1991: 573...... Brazil. Genus: Iapir Py-Daniel, Fonseca & Barbosa, 7. Ytu cupidus Reichardt, 1973: 146...... Brazil. 1993: 671 8. Ytu cuyaba Reichardt & Vanin, 1977: 127...... = Hintonia Reichardt, 1973:125 (nomen ...... Brazil. nova; praeocc. Fraser Brunner 1949 in Pis- ces), (= Ptyopteryx Reichardt & Costa, 1967) 9. Ytu demeter Reichardt, 1973: 142...... Brazil. (praeocc. Kolenati, 1848) 10. Ytu godoyi Reichardt & Vanin, 1977: 129...... 1. Iapir borgmeieri (Reichardt & Vanin, 1976): ...... Brazil. 216...... Brazil. = Ptyopteryx borgmeieri Reichardt & Vanin, 11. Ytu hephaestus Reichardt, 1973: 138...... Brazil. 1976. 12. Ytu itati Reichardt & Vanin, 1977: 132.. Brazil. 2. Iapir britskii (Reichardt & Costa, 1967): 14 13. Ytu mirandus Reichardt & Vanin, 1977: 126.... (TYPE SPECIES)...... Brazil...... Brazil. = Ptyopteryx britskii Reichardt & Costa, 1967. 14. Ytu mirim Reichardt & Vanin, 1977: 125...... Brazil. 3. Iapir castalia Reichardt, 1973: 131...... Brazil. 15. Ytu morpheus Reichardt, 1973: 145...... Brazil. 4. Iapir quadridentatus Braule-Pinto, Fonseca & Hamada, 2011: 48...... Brazil. 16. Ytu phebo Reichardt, 1973: 140...... Brazil. 5. Iapir trombetensis (Fonseca, Py-Daniel & Bar- 17. Ytu reichardti Vanin, 1978: 1...... Brazil. bosa, 1991): 293...... Brazil. 18. Ytu yaguar Reichardt & Vanin, 1977: 130...... = Hintonia trombetensis Fonseca, Py-Daniel ...... Brazil. & Barbosa, 1991. 19. Ytu ysypo Reichardt & Vanin, 1977: 126...... Genus: Incoltorrida Steffan, 1973: 634 ...... Brazil. 1. Incoltorrida madagassica Steffan, 1973: 635 20. Ytu n. sp. of zeus species group...... Paraguay.

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ACKNOWLEDGMENTS (Coleoptera: Myxophaga). Annals of the We thank John A. Kochalka of the Museo Na- Transvaal Museum, 36(24): 313-332. cional de Historia Natural de Paraguay who Fahey, C. & P. F. Langhammer. 2003. The provided official support for our field work, effects of dams on biodiversity in the collecting permits, and access to the museum Atlantic Forest. Pp. 413–425, in: Galin- collections. Bolivar Garcete Barrett is ack- da-Leal, C. and I. G. Câmara (eds.). The nowledged for his valuable comments and Atlantic Forest of South America: Bio- language revision of the manuscript. Special diversity Status, Threats, and Outlook. thanks go to two anonymous reviewers whose Island Press, Washington. 488 pp. comments improved the manuscript. Finally Fonseca, C. R. V. da, V. Py-Daniel & U. C. our gratitude to Sergio Rios Diaz for facilita- Barbosa. 1991. A new species of Hin- ting the publication of this paper in the Boletín tonia Reichardt 1973 from Amazon del Museo Nacional de Historia Natural del Basin, Brazil (Coleoptera: Torridincoli- Paraguay. dae). Revista Brasileira de Entomologia, 35(2): 293-300. LITERATURE Fraser-Brunner, A. 1949. A classification of the Aguilar Julio, C. A. 2010. Methods for Cat- fishes of the family Myctophidae. Pro- ching Beetles. Naturalia Scientific -Co ceedings of the Zoological Society of llection. Montevideo, Uruguay. 303 pp. London, 118(pt. 4): 1019-1106. Abdullah, M. 1974. Notes on Sphaeriformia Galinda-Leal, C. & I. G. Câmara. 2003. The Abdullah, particularly Torridincolidae Atlantic Forest of South America: Bio- Steffan (Col. Myxophaga). Annales de diversity Status, Threats, and Outlook. la Société Entomologique de France, Island Press. Washington. 488 pp. 10(4): 959-962. Hájek, J. & M. Fikáček. 2008. A review of the Beutel, R. G. 1998. Torridincolidae: II. Des- genus Satonius (Coleoptera: Myxopha- cription of the larva of Satonius kuro- ga: Torridincolidae): taxonomic revi- sawai (Satô, 1982) (Coleoptera). Pp. 53- sion, larval morphology, notes on wing 59, in: Jäch M. A. & L. Ji (eds.). Water polymorphism, and phylogenetic impli- Beetles of China, Vol. II. Zoologisch- cations. Acta Entomologica Musei Na- Botanische Gesellschaft in Österreich tionalis Pragae, 48(2): 655-676. and Wiener, Coleopterologenverein, Hájek, J., H. Yoshitomi, M. Fikáček, M. Ha- Wien. 371 pp. yashi & F.-L. Jia. 2011. Two new spe- Beutel, R. G. 1998/99. Phylogentic analysis of cies of Satonius Endrödy-Younga from Myxophaga (Coleoptera) with a redes- China and notes on the wing polymor- cription of horni (Lepiceri- phism of S. kurosawai Satô (Coleoptera: dae). Zoologischer Anzeiger, 237: 291- Myxophaga: Torridincolidae). Zootaxa, 308. 3016:51-62. Braule-Pinto, V. A., C. R.V. da Fonseca & N. Hayashi, M. 2007. Distributional records and Hamada. 2011. Iapir quadridentatus sp. ecological notes on aquatic Coleoptera nov., a new torridincolid beetle from of Shimane Prefecture. Bulletin of the Brazil (Coleoptera: Myxophaga: Torri- Hoshizaki Green Foundation, 10: 77- dincolidae). Zootaxa, 3031: 47-53. 113. Endrödy-Younga, S. 1977. Active extraction Hayashi, M. & H. Kadowaki. 2007. Records of water-dissolved oxygen and descrip- on aquatic Coleoptera from rivers in and tions of new taxa of Torridincolidae around Mt. Daisen, Tottori Prefecture,

W. D. Shepard, C. B. Barr & C. A. Aguilar Julio Boletín del Museo nacional de Historia Natural del Paraguay Vol. 17, Nº 1 (Agosto 2013) 81

Japan. Bulletin of the Hoshizaki Green Reichardt, H. & S. A. Vanin. 1976. Two new Foundation, 10: 149-168. Torridincolidae from Serra do Cipo, Hayashi, M. 2008. Distributional records and Minas Gerais, Brazil (Coleoptera, ecological notes on aquatic Coleoptera Myxophaga). Studia Entomologica, of Shimane Prefecture, part II. Bulletin 19(1-4): 211-218. of the Hoshizaki Green Foundation, 11: Reichardt, H. & S. A. Vanin. 1977. The to- 61-91. rridincolid genus Ytu (Coleoptera, Hinton, H. E. 1967. On the spiracles of larvae Myxophaga). Papeis Avulsos Zoologia, of the Suborder Myxophaga (Coleopte- São Paulo, 31(7): 119-140. ra). Australian Journal of Zoology, 15: Satô, M. 1982. Discovery of Torridincolidae 955-959. (Coleoptera) in Japan. Annotationes Jäch, M. A. & M. Balke. 2008. Global diversi- Zoologicae Japonenses, 55(4): 276-283. ty of water beetles (Coleoptera) in fres- Shepard, W. D. & C. A. Aguilar Julio. 2010. hwater. Hydrobiologia, 595: 419-442. Estudio preliminary de las familias de Kolenati, F. A. 1848. Genera et species Tri- escarabajos acuáticos , El- chopterorum. 1. Heteropalpoidea, Pra- midae, Lutrochidae, y Psephenidae co- gae. 141-296 pp. nocidos de Paraguay (Coleoptera: By- Mesa, A. & C. S. Fontanetti. 1985. The chro- rrhoidea). Boletín del Museo Nacional mosomes of a primitive species of beet- de Historia Natural del Paraguay, 16(1): le: Ytu zeus (Coleoptera, Myxophaga, 30-42. Torridincolidae). Proceedings of the Spangler, P. J. 1980. A new species of Ytu from Academy of Natural Sciences of Phila- Brazil (Coleoptera: Torridincolidae). delphia, 137(2): 102-105. The Coleopterists Bulletin, 34(2): 145- Pinto, V. A. B., C. R. V. da Fonseca & N. Ha- 158. mada. 2011. Iapir quadridentatus sp. Steffan, A.W. 1964. Torridincolidae, coleopte- nov., a new torridincolid beetle from rorum nova familia e regione aethiopi- Brazil (Coleoptera: Myxophaga: Torri- ca. Entomologische Zeitschrift, 74 (17): dincolidae). Zootaxa, 3031: 47-53. 193-200. Py-Daniel, V., C. R. V. da Fonseca & U. C. Bar- Vanin, S. A. 1978. Ytu reichardti, a new torri- bosa. 1993. Iapir nom. n. para Hintonia colid from Serra do Cipó, Minas Gerais, Reichardt, 1973 (Coleoptera, Torridin- Brazil (Coleoptera, Myxophaga, Torri- colidae). Revista Brasileira de Entomo- dincolidae). Revista Brasileira de Ento- logia, 37(4): 671. mologia, 22(1): 1-4. Reichardt, H. & C. Costa. 1967. Ptyopteryx Vanin, S. A. 1991. Ytu cleideae, a new species britskii, a new Neotropical genus and of torridincolid from Mato Grosso, Bra- species of the hitherto Ethiopian Torri- zil (Coleoptera, Torridincolidae). Re- dincolidae (Coleoptera, Myxophaga). vista Brasileira de Entomologia, 35(3): Papéis Avulsos de Zoologia, São Paulo, 573-576. 21: 13-19. Vanin, S. A. & C. Costa. 2001. Description of Reichardt, H. 1973. A critical study of the su- immature stages of Claudiella ingens border Myxophaga, with a taxonomic Reichardt & Vanin, 1976 and compa- revision of the Brazilian Torridincolidae rative notes on other Torridincolidae and Hydroscaphidae (Coleoptera). Ar- (Coleoptera, Torridincolidae). Aquatic quivos de Zoologia, São Paulo, 24 (2): , 23(1): 1-10. 73-162. Vanin, S. A. 2011. Torridincolidae. Torrent

The occurrence of Torridincolidae in Paraguay and a world checklist of species 82 Boletín del Museo nacional de Historia Natural del Paraguay Vol. 17, Nº 1 (Agosto 2013)

Beetles. Version 27 February 2011. Tree of Life Web Project. http://tolweb. http://tolweb.org/Torridincolidae/ in The org/.

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