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October 25 2019 INSECTA 306 urn:lsid:zoobank. A Journal of World Insect Systematics org:pub:930EAB11-37FA-41B0- UNDI M 980A-1A4736527842 0732 An annotated checklist of the Scarabaeoidea (Insecta: Coleoptera) of the Guianas Auke J. Hielkema Curitibastraat 46A, Beni’s Park Paramaribo, Suriname Meindert A. Hielkema Veenenburg 6 2804WX Gouda, Netherlands Date of issue: October 25, 2019 CENTER FOR SYSTEMATIC ENTOMOLOGY, INC., Gainesville, FL Auke J. Hielkema and Meindert A. Hielkema An annotated checklist of the Scarabaeoidea (Insecta: Coleoptera) of the Guianas Insecta Mundi 0732: 1–306 ZooBank Registered: urn:lsid:zoobank.org:pub:930EAB11-37FA-41B0-980A-1A4736527842 Published in 2019 by Center for Systematic Entomology, Inc. P.O. Box 141874 Gainesville, FL 32614-1874 USA http://centerforsystematicentomology.org/ Insecta Mundi is a journal primarily devoted to insect systematics, but articles can be published on any non- marine arthropod. Topics considered for publication include systematics, taxonomy, nomenclature, checklists, faunal works, and natural history. Insecta Mundi will not consider works in the applied sciences (i.e. medical entomology, pest control research, etc.), and no longer publishes book reviews or editorials. Insecta Mundi publishes original research or discoveries in an inexpensive and timely manner, distributing them free via open access on the internet on the date of publication. Insecta Mundi is referenced or abstracted by several sources, including the Zoological Record and CAB Abstracts. Insecta Mundi is published irregularly throughout the year, with completed manuscripts assigned an individual number. Manuscripts must be peer reviewed prior to submission, after which they are reviewed by the editorial board to ensure quality. One author of each submitted manuscript must be a current member of the Center for Systematic Entomology. Guidelines and requirements for the preparation of manuscripts are available on the Insecta Mundi website at http://centerforsystematicentomology.org/insectamundi/ Chief Editor: David Plotkin, [email protected] Assistant Editor: Paul E. Skelley, [email protected] Head Layout Editor: Robert G. Forsyth Editorial Board: J. H. Frank, M. J. Paulsen, Michael C. 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Florida Virtual Campus: http://purl.fcla.edu/fcla/insectamundi University of Nebraska-Lincoln, Digital Commons: http://digitalcommons.unl.edu/insectamundi/ Goethe-Universität, Frankfurt am Main: http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:hebis:30:3-135240 Copyright held by the author(s). This is an open access article distributed under the terms of the Creative Commons, Attribution Non-Commercial License, which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original author(s) and source are credited. http://creativecommons. org/licenses/by-nc/3.0/ Layout Editor for this article: Robert G. Forsyth 0732: 1–306 2019 An annotated checklist of the Scarabaeoidea (Insecta: Coleoptera) of the Guianas Auke J. Hielkema Curitibastraat 46A, Beni’s Park Paramaribo, Suriname [email protected] Meindert A. Hielkema Veenenburg 6 2804WX Gouda, Netherlands [email protected] Abstract. A checklist of the Scarabaeoidea (except Passalidae) of the Guianas (Guyana, Suriname and French Guiana) is presented for the first time. Both recorded and potential species are included, the latter defined as being recorded from Brazil north of the Amazon/Rio Negro main channel or from Venezuela east of the Orinoco main channel. The checklist contains a total of 930 species and subspecies, with 265 validly recorded from Guy- ana (eight new country records), 388 from Suriname (146 new country records), 531 from French Guiana (four new country records) and 177 additional species recorded from contiguous areas of neighboring Brazil and Ven- ezuela. Valid names as well as selected synonyms are provided. Multiple notes on presently recognized subspecies, nomenclatural problems, doubtful type localities, historical misidentifications, etc. are included. An extensive reference list is provided. Key words. Distribution, new country record, literature, Geotrupidae, Trogidae, Lucanidae, Hybosoridae, Scara- baeidae, Aphodiinae, Scarabaeinae, Orphninae, Melolonthinae, Rutelinae, Dynastinae, Cetoniinae. Table of Contents Table 1 follows the Literature Cited and contains condensed distributional information, including the text page location for each valid species cited in this checklist. Introduction The cosmopolitan superfamily Scarabaeoidea comprises over 31,000 described species (Jameson and Ratcliffe 2005). Members can be found in most terrestrial habitats and use a wide range of food sources. They also have widely varying and sometimes highly developed life histories, with many species, especially in the subfamily Scarabaeinae, providing parental care. Members of the family Passalidae and some of the subfamily Scarabaeinae even express clear subsocial behavior (Halffter and Edmonds 1982: 63; Boucher 2006: 264). Species range in length from less than 2 mm to over 10 cm and may be dull and inconspicuous or colorful and shiny. Some species, especially their males, have extravagant horns or mandibles. The abundance of some species suggests that they have considerable influence on their ecosystems. Due to their appearance and behavior, they have attracted the attention of people throughout history. As such, dung beetles rolling balls of dung were a symbol for the sun moving across the sky to the early Egyptians (Remler 2010: 169), and Linnaeus started his invertebrate species descrip- tions in the tenth edition of Systema Naturae (Linnaeus 1758) with scarabs. At present, scarabaeoids are considered important as, among many other things, pest species (Bedford 1980; Valois et al. 2019), biocontrol agents against dung flies (Edwards 2007), and indicators of ecosystem health (Spector 2006; Nichols et al. 2007). In 2010 we started collecting scarabaeoid species in a semi-structured way to increase the knowledge of their diversity in Suriname, and to augment the collections of the National Zoological Collection of Suriname (NZCS) and of the second author. While identifying the collected specimens, we came to the conclusion that a list of species already known to occur in the area would greatly facilitate this task. We 2 • INSECTA MUNDI 0732, October 2019 HIELKEMA AND HIELKEMA then found that no such species list existed for Suriname, and that there were hardly any checklists of (sub)families available for the surrounding countries. To fill this gap in our knowledge, we decided to make such a list. To our surprise it was rather difficult to find the correct authors and years of descrip- tion of many species, which prompted us to seek all the original generic and specific descriptions. While creating this list of generic and specific names, we started to add papers containing useful keys and comparative (re)descriptions. For each species, we also included the page number on which the species account in each publication began. As this list grew, we realized it could be beneficial to others too, and the present paper is the result. The Guianas, situated on the northern coast of South America, consist of the two independent countries of Guyana (in the west) and Suriname (in the middle) and the French department of French Guiana (in the east). Their northern borders are formed by the Atlantic Ocean, while in the south they all border Brazil, which also forms the eastern border of French Guiana and part of the western border of Guyana. The remainder of Guyana’s border is with Venezuela. Geologically, most of the land surface of the Guianas, together with parts of northern Brazil, southern and eastern Venezuela and a small part of eastern Colombia, is part of the Guiana Shield. This is a cra- ton consisting mostly of about 1.9-billion-year-old granites, gneisses, metasediments and metavolcanics (Noordam 1993). The soils of the Guiana Shield are generally very poor and acidic (Noordam 1993). The biodiversity of the Guiana Shield is related to, but notably different from, that of the Amazon Basin. Speciation on the Guiana Shield has been driven by various super- and intra-continental events (see Noonan 2018 for details) and has likely been influenced by the differences in availability of nutrients between the rich Amazonian floodplains and the poor Guianan soils. Most of the sparsely inhabited interior of the Guianas is accessible only by motorized dug-out canoes and small aircraft. Due to its isolation, the interior is still largely intact, biologically interesting, and logistically challenging for researchers. Still, various insect