Coleoptera: Elmidae and Protelmidae World Catalogue of Insects VOLUME 14 The titles published in this series are listed at brill.com/wci Ancyronyx acaroides (left) and A. malickyi (middle and right) on submerged wood in a Sumatran lowland river. Painting by W. Zelenka (†), ca. 2000. Coleoptera: Elmidae and Protelmidae By Manfred A. Jäch Ján Kodada Michaela Brojer William D. Shepard Fedor Čiampor, Jr. LEIDEN | BOSTON Cover illustrations: Habitus illustrations (from above): Cuspidevia, Macronychus, Potamophilus, Stenelmis, Troglelmis. Paintings by W. Zelenka (†). Library of Congress Control Number: 2006356329 Want or need Open Access? Brill Open offers you the choice to make your research freely accessible online in exchange for a publication charge. Review your various options on brill.com/brill-open. Typeface for the Latin, Greek, and Cyrillic scripts: “Brill”. See and download: brill.com/brill-typeface. issn 1398-8700 isbn 978-90-04-29176-8 (hardback) isbn 978-90-04-29177-5 (e-book) Copyright 2016 by Koninklijke Brill nv, Leiden, The Netherlands. Koninklijke Brill nv incorporates the imprints Brill, Brill Hes & De Graaf, Brill Nijhoff, Brill Rodopi and Hotei Publishing. All rights reserved. No part of this publication may be reproduced, translated, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior written permission from the publisher. Authorization to photocopy items for internal or personal use is granted by Koninklijke Brill nv provided that the appropriate fees are paid directly to The Copyright Clearance Center, 222 Rosewood Drive, Suite 910, Danvers, ma 01923, usa. Fees are subject to change. This book is printed on acid-free paper and produced in a sustainable manner. Contents Acknowledgements ix List of Authors x Introduction xi New nomenclatorial and taxonomic acts xviii Family Group Names 1 Genus and Species Group Names 3 Elmidae 5 Extant taxa 5 Species incertae sedis 242 Fossil taxa 243 Taxa removed from Elmidae 244 Unavailable names 245 Nomina exclusa 249 Protelmidae 251 References 253 Index 297 Acknowledgements The following persons are sincerely thanked for various advice, for providing lit erature and/or distributional data, or for sending specimens: V.I. Alekseev (Kaliningrad, Russia), M.A. Alonso-Zarazaga (Madrid, Spain), R.B. Angus (London, U.K.), M. Barcley (London, U.K.), E. Bauernfeind (Vienna, Austria), R. Booth (London, U.K.), D.S. Boukal (České Budějovice, Czech Rep.), S. Bouzid (Annaba, Algeria), W. Brunnbauer (Vienna, Austria), B. Bruvo Mađarić (Zagreb, Croatia), P. Buzinsky (Lublin, Poland), E. de Coninck (Tervuren, Belgium), G.N. Foster (Ayr, U.K.), N. Ferreira (Rio de Janeiro, Brazil), M. Fikáček (Prague, Czech Rep.), H. Freitag (Manila, Philippines), J. Garrido (Vigo, Spain), M. Gonzáles-Córdoba (Cali, Colombia), M. Hess (Munich, Germany), H. Huijbregts (Leiden, Netherlands), L. Ji (Shenyang, China), S.-W. Jung (Seoul, South Korea), Y. Kamite (Nagoya, Japan), A. Komarek (Vienna, Austria), A. Kovalev (St. Petersburg, Russia), J. Krikken (Leiden, Netherlands), M. Madl (Frauenkirchen, Austria), A. Mantilleri (Paris, France), M. Maruyama (Fukuoka, Japan), O. Merkl (Budapest, Hungary), V. Mičetić Stanković (Zagreb, Croatia), A. Millán (Murcia, Spain), H. Nasserzadeh (Tehran, Iran), N.B. Nikitskyi (Moscow, Russia), A.N. Nilsson (Umeå, Sweden), A. Prokin (Voronezh, Russia), M. Przewocźny (Poznań, Poland), I. Ribera (Barcelona, Spain), W. Sondermann (Bogotá, Colombia), M. Tavano (Genova, Italy), C. Taylor (London, U.K.), D. Telnov (Riga, Latvia) and H. Yoshitomi (Matsuyama, Japan). P.J. Spangler (†) and H.P. Brown (†) are thanked for sending an unpublished checklist to W.D. Shepard many years ago. All habitus illustrations (book cover and frontispiece) were kindly provided by W. Zelenka (†). This study was partly supported by the Slovak Research and Development Agency: Project No. APVV-0213-10 and VEGA: Project No. 1/0395/14. World Catalogue of Insects. Vol. 14. Coleoptera: Elmidae, Protelmidae M. A. Jäch & M. Brojer Naturhistorisches Museum Wien, Burgring 7, A – 1010 Wien, Austria ([email protected] (corresponding author), michaela.brojer @nhm-wien.ac.at) J. Kodada Faculty of Natural Science, Comenius University in Bratislava, Mlynská dolina Ilkovičova 6, 842 15 Bratislava 4, Slovakia ([email protected]) W.D. Shepard Essig Museum of Entomology, 1101 Valley Life Sciences Bldg., #4780, University of California, Berkeley, California 94720 USA ([email protected]) F. Čiampor, Jr. Zoology Lab, Institute of Botany, Slovak Academy of Sciences, Dúbravská cesta 9, SK – 845 23 Bratislava, Slovakia ([email protected]) Introduction Elmidae or Riffle Beetles are a moderately large cosmopolitan family of True Water Beetles (see Jäch & Balke 2008). Adults and larvae are usually considered to be aquatic, however, adults of several genera, e.g. Potamophilus Germar, 1811 are often encountered a little above the water line or in spray zones of water- falls and cascades. Members of this family are generally living in lotic habitats, few species may be found in lakes or ponds and a few species are recorded from subterranean waters. Strictly terrestrial representatives, which do exist in other typical water beetle families (e.g. Dytiscidae, Hydrophilidae, Hydraenidae, Dryopidae) are completely unknown among elmids. Pupation takes place above the water line. Elmidae are often used in ecological running water monitoring programs (Moog & Jäch 2003). In the past, elmids have even been part of the diet of man. Austrelmis condimentarius was used as seasoning for food in South America. This species was reported to have considerable commercial value (Philippi 1864). The morphology of Elmidae was described in detail by Kodada & Jäch (2005) (second edition in press). Elmidae belong to Polyphaga and are currently placed in Elateriformia. Within Elateriformia, they are usually attributed to Byrrhoidea (Kodada & Jäch 2005, Bouchard et al. 2011, Kundrata et al. 2013, Bocak et al. 2014), or Dryopoidea (Crowson 1955, 1981, Lawrence & Newton 1982, Hunt et al. 2007). Protelmidae, hitherto regarded as a tribe of Elminae, are here elevated to family rank, because of strongly deviating external and genitalic features. Adults of Protelmidae are aquatic (three genera live in riffles of running water, one genus is troglobiontic). Protelmid larvae are still unknown. Among the “great six” water beetle families (i.e. families with more than thousand species, see Jäch & Balke 2008), only the three largest ones (Dytiscidae, Hydraenidae, Hydrophilidae) have been covered by world catalogues recently. The last world catalogue of Elmidae was published more than one hundred years ago, by Zaitzev (1910) and included 297 species. Elmidae were still regarded as a subfamily of Dryopidae by Zaitzev (1910), although various authors have already treated Elmidae as a distinct family earlier (e.g. Grouvelle 1889b, 1892a, 1896a, d, 1900b, 1906b, c, 1908, Reitter 1886, 1899). The North American Elmidae were catalogued by Brown (1983), the Palearctic elmids were published by Jäch et al. (2006) (second edition in press), and the Neotropical taxa were compiled by Segura et al. (2013). Catalogues for the Australian, Afrotropical, and the Oriental fauna have never been published, but a check list of the Australian species was provided by Glaister (1999). xii Introduction A preliminary version of this catalogue was published by Joel K. Hallan in the internet (“Synopsis of the described Coleoptera of the world”: http://insects.tamu.edu/research/collection/hallan/test/Arthropoda/Insects/ Coleoptera/Family/Coleoptera1.htm), based on a card catalogue by H.P. Brown (1921–2008), authored by M.A. Jäch and J. Kodada. This preliminary version was updated until June 2010. The reason for placing this preliminary version in the internet was simply to provide a widely accessible source of useful informa- tion, although it was still very incomplete, especially with regard to distribu- tional data. Particular Problems In the course of the compilation of this catalogue we faced numerous nomen- clatorial problems. These concerned for instance the correct spelling of taxa names (e.g. decisions between correct and incorrect original spellings (sensu ICZN 1999: Art. 32), or treatment of epithets that are nouns in apposition), treatment of “subsequent spellings” (sensu ICZN 1999: Art. 33), correct identity and spelling of author names, correct publication dates, correct type localities, the availability and correct authority of some very poorly described species published in Zhang & Yang (1995), and finally the unsatisfactory quality of vari- ous global online lists. We were able to check all references listed in this catalogue, except for one: Crotch (1872), who published a nomen nudum (Limnius sinaiticus). Taxa Names Many of the species group epithets erected by Hinton are definitely nouns in the nominative singular standing in apposition to the generic name (ICZN 1999: Art. 11.9.1.2). They are independent of the grammatical gender of the generic name (ICZN 1999: Art. 31.2.1). Subsequent authors, who did not realize, that these epithet names were in fact nouns, unwarrantedly used their adjecti- val forms or changed their gender when transferring the species to genera of different sex. Some examples: Macrelmis aeolus (Hinton, 1946b) [not aeolis], M. aleus (Hinton, 1946b) [not alea], M. aristaeus (Hinton, 1946b) [not aristea], M. celsus (Hinton, 1946b) [not celsa], M.