Catalogue of the Incini with the description of the first Archedinus species from Honduras (Coleoptera: Scarabaeidae: Cetoniinae)

ACTA ENTOMOLOGICA MUSEI NATIONALIS PRAGAE www.aemnp.euISSN 1804-6487 (online) – 0374-1036 (print) RESEARCH PAPER Catalogue of the Incini with the description of the fi rst Archedinus species from Honduras (Coleoptera: Scarabaeidae: Cetoniinae) Matthias SEIDEL1,2,4), Emmanuel ARRIAGA-VARELA1,2) & Rafael SOUSA3) 1) Department of Zoology, Faculty of Science, Charles University, Prague, Viničná 7, CZ-12843, Prague, Czech Republic; e-mails: [email protected], [email protected] 2) Department of Entomology, National Museum, Cirkusová 1740, CZ-19300 Prague, Czech Republic; 3) Museu de Zoologia da Universidade de São Paulo, Avenida Nazaré 481, Ipiranga, 04263–000, São Paulo, São Paulo, Brazil; e-mail: [email protected] 4) Corresponding author Abstract. We present an annotated catalogue for the tribe Incini (Coleoptera: Scarabaeidae: Cetoniinae) including references to all taxonomic and nomenclatural acts, clarifying the spelling of names, providing type depositories and occurrence records for the species. The spelling of Golinca davisii (Waterhouse, 1877) is fi xed, and the incorrect subsequent spelling Pantodinus klugi Burmeister, 1847 is preserved. A comprehensive list of all valid names in Incini is provided. Furthermore, we describe the third species in the genus Archedinus Morón & Krikken, 1990, and fi rst one known from outside of Mexico, A. antoshkai Seidel & Arriaga-Varela sp. nov. from Cerro las Minas, the highest mountain in Honduras. We provide an updated determination key for the species of Archedinus. The new species is compared with Archedinus howdeni Morón & Vaz-de-Mello, 2007, the most similar species in terms of genital and habitus morphology. An updated key to identifi cation of males of Archedinus is provided. Key words. Coleoptera, Scarabaeidae, Cetoniinae, Coelocratus, Inca, Golinca, Pantodinus, fl ower chafer, faunistics, gender, nomenclature, spelling, types, taxonomy, year of description, Neotropical Region Zoobank: http://zoobank.org/urn:lsid:zoobank.org:pub:0610B52C-FE2C-49F1-A16F-0B14881AF4F2 © 2018 The Authors. This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Licence. Accepted: 1st August 2018 Published online: 10th September 2018 Introduction Incini is a Neotropical tribe of fl ower chafers (Coleoptera: Scarabaeidae: Cetoniinae). Previously, this group of beetles has been treated as a subtribe Incaina in the subfamily Trichiinae (e.g., MORÓN & KRIKKEN 1990; MORÓN & VAZ-DE-MELLO 2007; SMITH 2006). SMITH et al. (2006) indicated the need for elevation of the subtribe to tribal level, since the historical concept of the tribe Trichiini was paraphyletic based on their preliminary molecular data. Agreeing with that, more recent works treat this group as a tribe in the subfamily Cetoniinae based on morphological and molecular phylogenetic analyses (e.g., MICÓ et al. 2008, ŠÍPEK et al. 2016). BOUCHARD et al. (2011) corrected the stem and spelling of the family group name to Incand Incini. The taxonomic history of the species and genera in the tribe is complex. The spelling of species names, authors and year of publication of species have been cited differently by cataloguers, taxonomists, and ecologists. The lack of a complete and comprehensive work dealing with the nomenclature in this tribe is an obstacle for taxonomists aiming to continue research on Incini. Five Incini genera have been reported from Mexico to Argentina (Fig. 6): Archedinus Morón & Krikken, 1990 (Figs 1–3, 5A–B), Coelocratus Burmeister, 1842 (Figs 5C–D), Inca LePeletier & Serville, 1828 (Figs 5E–G), Golinca Thomson, 1878 (Figs 5H–J) and Pantodinus Burmeister, 1847 (Figs 5K–M). The genus Archedinus was described by MORÓN & KRIKKEN (1990) in order to include a single species, A. relictus 2018 58(2): 389-405 doi: 10.2478/aemnp-2018-0031 Seidel.indd 389 10.9.2018 15:36:27 SEIDEL et al.: Catalogue of Incini and description of a new Archedinus (Coleoptera: Scarabaeidae) 390 Morón & Krikken, 1 990 from Chiapas, Mexico. Archedinus is currently classifi ed within the tribe Incini, close to Pantodinus as a “relictual” representative with reduced sexual dimosphism. Males and females of A. relictus were described based on nine specimens collected in the cloud forest of El Triunfo in the Sierra Madre de Chiapas mountain range. Later, MORÓN & VAZ-DE-MELLO (2007) described a second species, A. howdeni from Chimalapas region in Oaxaca, Mexico based on four male specimens and expanded the generic defi nition of Archedinus. The position of Archedinus within the other Incini was tested based on a morphological phylogenetic analysis, where it was sister taxon of Pantodinus and, along this one, sister to the remaining Incini. Not much is known about the life history of the genus. Archedinus relictus was reported having “diurnal activity, living under forest litter and rotten logs in montane humid forests located at elevations going from 1000 to 1850m” (MORÓN & KRIKKEN 1990). Archedinus howdeni was found in “tropical subdeciduous forest at 1600 m” (MORÓN & VAZ-DE-MELLO 2007). Besides these two type localities in Mexico, no other faunistic records have been published. Therefore, the distribution range of Archedinus is very poorly known. In the present contribution we describe a third species recently collected in Honduras, showing that the genus is not solely endemic to Mexico and has a much larger distribution range than previously known. We update the generic diagnosis and the determination key based on the work of MORÓN & VAZ-DE-MELLO (2007). In addition, we present an annotated catalogue of the Incini aiming to resolve all the nomenclatural problems in the tribe. Material and methods Collections. The material cited is deposited in the following collections: ADPC Alain Drumont Personal Collection, Brussels, Belgium; AKPC Anton Kozlov Personal Collection, Moscow, Russia; ARPC Andreas Reichenbach Personal Collection, Leipzig, Germany; BCRC Brett Ratcliffe Personal Collection, Lincoln, Nebraska, USA; BMNH Natural History Museum, London, United Kingdom (Max Barclay, Michael Geiser); CEMT Setor de Entomologia da Coleção Zoológica, Universidade Federal do Mato Grosso, Cuiabá, Brazil (Fernando Vaz-deMello); CMNC Canadian Museum of Nature Collection, Ottawa, Canada (Robert Anderson, François Génier); DBPC Denis Bouchard Personal Collection, Autouillet, France; DZUP Coleção Entomológica Padre Jesus Santiago Moure, Universidade Federal do Paraná, Curitiba, Brazil (Lúcia Massutti de Almeida); EAPZ Escuela Agrícola Panamericana, Tegucigalpa, Honduras (Ron Cave, Jesús Orozco); FZBRS Fundação Zoobotânica do Rio Grande do Sul, Porto Alegre, Brazil (Luciano de Azevedo Moura); IEXA Colección Entomológica, Instituto de Ecología, A.C., Xalapa, México (Leonardo Delgado); KSPC Kaoru Sakai Personal Collection, Tokyo, Japan; LGPC Leccinum García Personal Collection, Ciudad Victoria, Tamaulipas, Mexico; MACN Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina (Arturo Roig Alsina); MLUH Zentralmagazin Naturwissenschaftlicher Sammlungen, Martin- -Luther-Universität Halle-Wittenberg, Halle, Germany (Karla Schneider, Joachim Händel); MNCN Museo Nacional de Ciencias Naturales, Madrid, Spain (Mercedes París); MNCR Museo Nacional de Costa Rica, San José, Costa Rica (Ángel Solís); MNHN Muséum national dʼHistoire naturelle, Paris, France (Antoine Mantilleri, Olivier Montreuil); MNHG Museé dʼHistoire Naturelle, Geneva, Switzerland (Giulio Cuccodoro); MNHY Museum of Natural History, Yokohama, Japan (Kyohei Watanabe); MSPC Matthias Seidel Personal Collection, Prague, Czech Republic; MXAL Miguel Angel Morón Personal Collection, Xalapa, Mexico; MZSP Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (Sonia A. Casari); MZUEFS Museu de Zoologia da Universidade Estadual de Feira de Santana, Feira de Santana, Brazil (Freddy Bravo); NHRS Naturhistoriska riksmuseet, Stockholm, Sweden (Johannes Bergsten); NMPC Department of Entomology, National Museum, Prague, Czech Republic (Lukáš Sekerka, Jiří Hájek); PSPC Petr Šípek Personal Collection, Prague, Czech Republic; RBINS Royal Belgian Institute of Natural Sciences, Brussels, Belgium (Alain Drumont, Pol Limbourg); RMNH Naturalis Biodiversity Centre, Leiden, Netherlands; SLCC Stephane LeTirant Collection, Montreal, Canada; SMF Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main, Germany (Damir Kovac); TMPC Tetsuo Mizunuma Personal Collection, Osaka, Japan; UFRPE Coleção Entomológica, Universidade Rural de Pernambuco, Recife, Brazil (Paschoal Grossi); UNSM University of Nebraska State Museum, Lincoln, Nebraska, USA (Brett Ratcliffe, M. J. Paulsen); USNM United States National Museum, Washington, District of Columbia, USA (currently housed at the University of Nebraska State Museum for off-site enhancement) (Brett Ratcliffe); UUZM Evolutionsmuseet, Uppsala University, Uppsala, Sweden (Hans Mejlon); ZMHB Museum für Naturkunde Leibniz-Institut für Evolutionsund Biodiversitätsforschung, Berlin, Germany (Johannes Frisch, Joachim Willers); ZMUH Zoologisches Museum, Centrum für Naturkunde, Hamburg, Germany (Martin Husemann). Measurements. Total body length and width were measured using an electronic caliper, and measurements of other structures were obtained by using an ocular micrometer attached to an Olympus (SZX 16) dissecting microscope. Illustrations. Habitus photographs of the new species were taken using a Canon EOS 550D digital camera with a Canon MP-E65mm f/2.8 1–5× macro lens. Photographs of all other species were provided by curators of different institutions (see Acknowledgements). Line drawings were made by hand. All morphological illustrations were subsequently edited in Adobe Photoshop CS5. Pictures of the habitat in the type locality of A. antoshkai were provided by Viktor Sinyaev. Annotated catalogue. The style for the entries for species in the catalog follows MOORE et al. (2017) and provide: 1) the valid species name, author and date, 2) original spelling and combination, misspellings, new combinations, and invalid names, 3) synonyms and the reference in which the synonym was designated, 4) general distribution data including the country (in capital letters) and states/provinces/departments when they are known. Distributions are based on the literature and on specimens that we examined. Seidel.indd 390 10.9.2018 15:36:28 Acta Entomologica Musei Nationalis Pragae, volume 58, number 2, 2018 391 Results Archedinus Morón & Krikken, 1990 Type species. Archedinus relictus Morón & Krikken, 1990: 76. Diagnosis. This genus is distinguished from other members of the tribe Incini by the following combination of characters: body length 17–27 mm, dorsal integument shiny, dark-brown to black; clypeal anterior margin sinuated, male with short clypeal horn or elevated clypeal margin lacking lateral or medial clypeal horns (present in Golinca, Inca and Pantodinus); eyes with strong supraocular ridges; male with antennal club longer than pedicel and funicle together, mentum with anterior margin sinuate or rounded; pronotum coarsely punctate, without posterior impression; anterior prosternal process projected, longer than procoxal length, elytra deeply striate without waxy surface; pterothoracic setae sparse; profemur with a preapical projection; protibia with 3 external teeth; mesoand metatibiae expanded, tridentate; aedeagus with parameres simple or bifurcate. Archedinus is most similar to Pantodinus due to the sinuate antero-ventral clypeal margin and the well marked elytral striae. However, Archedinus can be distiguished by the strong supraocular ridges, the prosternal process being longer than the procoxal anterior face, and by the sparse setae on the pterosterna (MORÓN & VAZ DE MELLO 2007). The third stage larva of A. relictus was diagnosed from other Incini (MORÓN 1995) by having mandibles without a stridulatory area; premolar area on left mandible with 2 teeth; dorsoepicranial setae absent; maxillary stridulatory area with 12 irregular, non-pointed teeth; last antennomere with 7–13 dorsal sensory spots; tarsal claws with 2 internal preapical setae; spiracle respiratory plates with irregular, ameboid “holes”; dorsal and ventral abdominal sclerites with long setae. Archedinus antoshkai Seidel & Arriaga-Varela, sp. nov. (Figs 1A; 2A,C,E; 3A–B,D,F) Type material. HOLOTYPE:  (NMPC), label data: “HONDURAS: Lempira / Department, P.N. Celaque Cerro / Las Minas / 14°32´50´´N, 88°40´11´´W / 01-04.vii.2014, 2600m / V. Sinyaev & M. Márquez lgt. // ex. Matthias Seidel / Collection // Archedinus n. sp. / det. M. Seidel 2016 // HOLOTYPE / Archedinus antoshkai / sp. nov. / M. Seidel & E. Arriaga-Varela / des. 2017”. Description. Male (holotype). Total length: 17.5 mm, humeral width: 7.5 mm, maximum width: 8.5 mm (Figs 1A–B). Color. Ventrally and dorsally shiny black, surface in punctures dull. Head. Surface coarsely and irregularly Fig. 1. Habitus views of Archedinus spp. A–B – A. antoshkai sp. nov., C – A. howdeni Morón & Vaz-de-Mello, 2007. A, C – dorsal habitus. B – lateral habitus. Image 1C by courtesy of François Génier. Seidel.indd 391 10.9.2018 15:36:28 SEIDEL et al.: Catalogue of Incini and description of a new Archedinus (Coleoptera: Scarabaeidae) 392 punctate (Fig. 2A), with punctures ranging in diameter from 27.5–87.5 μm. With a deep, long, triangular excavation, of irregular triangular shape narrowing down from frons to clypeus to a third of its width. Clypeus covered by short, tawny setae, abruptly constricted at base to 0.7 times of apical width, anterior clypeal margins sinuate with anterolateral corners refl exed. Maximum width of head capsule 3.1 mm; maximum width of clypeus 1.5 mm. Interocular distance equals 3.1 maximum transverse eye diameters. Labium elongate, narrowing anteriorly with anterior margin slightly rounded (Fig. 3D), deeply punctate, setigerous; setae long, tawny. Distal maxillary palpomere 0.8 mm in length, subcylindrical, weakly curved, apex truncate. Third palpomere subfusiform, rounded, not connate with second, enlarged, 0.6 mm long. Antennae with 10 antennomeres, antennal club composed of three enlarged, lamellate antennomeres, club 2.6 mm long, 1.7 times longer than preceding six antennomeres combined; antennomere 7 wider than long, with narrowed, short anterior projection; antennomere 6 longer than wide, with apex obliquely truncate, without anterior projection; antennomere 3 as long as antennomere 4 or 5, antennomere 3 as long as wide, pedicel nearly globular, as wide as long; scape elongate-pyriform. Fig. 2. Details of Archedinus spp. A, C, E – A. antoshkai sp. nov. B, D, F – A. howdeni Morón & Vaz-de-Mello 2007. A‒B – head dorsal view. C‒D – head antero-lateral view. E‒F – pronotum dorsal view. Images 2B,D,F courtesy of François Génier. Seidel.indd 392 10.9.2018 15:36:28 Acta Entomologica Musei Nationalis Pragae, volume 58, number 2, 2018 393 Prothorax. Pronotum about 1.3 times wider than long; 1.5 times wider at posterior angles than at anterior angles, margins rounded, slightly more strongly convergent to posterior angles (Figs 2C,E). Dorsal surface coarsely and irregularly punctate, punctures more dense in anterior half, punctures deep, ranging from 37–325 μm in diameter. Disc with a longitudinal depression on midline, and 2 depressions on each side, 1 large, elongate at anterior half, vanishing towards antero-lateral corners, and a small, shallower one at posterior 3/4, depressions with punctures wider and confl uent. Prosternum smooth on sides, with long prosternal process, acute, with briefl y rounded apex, and many long, tawny setae projected anteriad. Elytra with margins curved in anterior fi fth, then subparallel to rounded apical fourth; with 10 wide, deep striae, each with irregular row of rounded to elongate ellipsoid, foveate punctures; striae I–V extend from anterior border to apical callus; striae VI–IX extend from humerus to apical umbone; stria X extends along lateral margin but does not reach apex; interstriae shiny, with fi ne punctures and sparser, larger, foveate punctures; epipleura continuously narrowing posteriorly to just before apex. Mesothorax. Mesoventrite coarsely punctate, depressed at center, abruptly upturned near anterior borders of mesocoxae. Wings well developed. Metaventrite convex at sides, moderately depressed around discrimen; coarsely punctate towards anterolateral corners Fig. 3. Details of Archedinus spp. A, B, D, F – A. antoshkai sp. nov. C, E, G – A. howdeni Morón & Vaz-de-Mello 2007. A, C – aedeagus dorsal view. B – aedeagus lateral view. D–E – outline of mentum. F–G – antennae with arrow indicating third antennomere. Images 3C,G by courtesy of François Génier. Seidel.indd 393 10.9.2018 15:36:29 SEIDEL et al.: Catalogue of Incini and description of a new Archedinus (Coleoptera: Scarabaeidae) 394 with many slender, decumbent to erect setae. Legs. Profemur with a preapical, blade-like, dorsal projection, with rounded apex. Protibia with a wide inner basal notch. All tarsi robust, with large, sickle-shaped claws. Abdomen. Ventrites convex, almost glabrous, with sparse, small decumbent setae, fi nely punctate, punctures narrowly transverse; last ventrite with deeper, more rounded punctures on posterior margin which is slightly raised and briefl y emarginate at middle. Pygidium glabrous, shiny, with dense, fi ne micropunctures mixed with sparse, large, sometimes confl uent, round punctures, fi nely rugose toward basal angles; without waxy spots; posterior margin nearly rounded. Genitalia. Aedeagus with large basal piece, slightly convex; tectum shallowly concave; parameres elongate, sinuous in lateral view, with apical third bifurcate in apical third in dorsal view, apex rounded (Figs 3A–B). Female. Unknown. Differential diagnosis. Archedinus antoshkai is most similar to A. howdeni based on the emarginate clypeal projection, anterior prosternal process with rounded apex, and the apices of the parameres not being bifurcate. However, it can be distinguished from it by its smaller size (17.5 mm in A. antoshkai versus 25–27 mm in A. howdeni), the clypeal projection with anterior margin more strongly emarginate medially (Fig. 2A in A. antoshkai versus Fig. 2B,D in A. howdeni), in A. howdeni the frons is with an elongate, triangular depression (Figs 2A,C in A. antoshkai versus Fig. 2B in A. howdeni), the third antennomere is small, being as long as the fourth (Fig. 3F in A. antoshkai versus Fig. 3G in A. howdeni), the mentum is elongate with a rounded anterior margin (Fig. 3D in A. antoshkai versus Fig. 3E in A. howdeni), pronotum with coarser lateral margins (Figs 2C,E in A. antoshkai versus Figs 2D,F in A. howdeni) and the aedeagus with apices of the parameres simple, without a small lateral projection (Fig. 3A in A. antoshkai versus Fig. 3C in A. howdeni). Etymology. The new species is named after Anton Olegovich Kozlov (Moscow, Russia), who made the specimen available for study and generously donated the holotype to us. The name refers to the diminutive form of his fi rst name, “Antoshka”, used by friends; noun in apposition. Natural history. The species occurs in the highest mountain in Honduras in a tropical cloud forest at 2600 m elevation. Female specimens, immature stages, and life cycle are unknown. Distribution. Only known from the type locality: Cerro Las Minas in Celaque National Park, Lempira Department, Honduras. Key to the males of Archedinus (modifi ed from MORÓN & VAZ-DE-MELLO 2007) 1 Apex of prosternal process acute. Profemur with an acute preapical tooth. Elytral striae with waxy gray secretion. Male clypeus with short, bifurcate horn. Apex of parameres bifurcate. ..................................... ........................... A. relictus Morón & Krikken, 1990 ‒ Apex of prosternal process rounded. Profemur with a rounded preapical tooth. Striae without waxy secretion. Male clypeus with elevated anterior margin briefl y sinuate. Apex of parameres acute. ............... 2 2 Antennomere 3 as long as antennomeres 4 and 5 combined. Mentum short and anteriorly sinuate. Apex of parameres with small, lateral projection. Total length 25–27 mm. ....................................................... ................ A. howdeni Morón & Vaz-de-Mello, 2007 ‒ Antennomere 3 half as long as antennomeres 4 and 5 combined. Mentum elongate and anteriorly rounded. Apex of parameres simply rounded. Total length ca. 17.5 mm. ..................................................................... ......... A. antoshkai Seidel & Arriaga-Varela sp. nov. Fig. 4. Distribution and habitat Archedinus species. A – distribution map: black square – A. relictus Morón & Krikken, 1990; white circle – A. howdeni Morón & Vaz-de-Mello, 2007; black circle – A. antoshkai sp. nov. B–C – type locality of A. antoshkai sp. nov. in Celaque National Park at 2600 m. Images 4B–C by courtesy of Viktor Sinyaev. Seidel.indd 394 10.9.2018 15:36:30 Acta Entomologica Musei Nationalis Pragae, volume 58, number 2, 2018 395 Catalogue of the tribe Incini Tribe Incini Burmeister, 1842 Incadae Burmeister, 1842: 704 [original spelling] Type genus. Inca LePeletier & Serville, 1828. Remarks. BOUCHARD et al. (2011) corrected the stem of the family group name from Incato Inc-, lacking an explanation for their action. As discussed below under the section Inca, Inca is a masculine genus name derived from Spanish. While creating family-group names BURMEISTER (e.g., 1842, 1844) removed a few fi nal letters from every genus name independently from their gender and added suffi xes such as -idae, -didae or -adae (e.g., Lomapteridae Burmeister, 1842 [from Lomaptera Gory & Percheron, 1833]; Macraspididae Burmeister, 1844 [from Macraspis MacLeay, 1819]; Lichniadae Burmeister, 1844 [from Lichnia Erichson, 1835]). Therefore, it can be securely assumed that the correct stem of Incadae Burmeister, 1842 is Incand Article 29.3.3. (ICZN 1999) is followed: “If a generic name is or ends in a word not Greek or Latin, or is an arbitrary combination of letters, the stem for the purposes of the Code is that adopted by the author who establishes the new family-group taxon, either the entire generic name (see Article 29.6), or the entire generic name with the ending elided, or the entire generic name with one or more appropriate linking letters incorporated in order to form a more euphonious family-group name”. Genus Archedinus Morón & Krikken, 1990 Archedinus Morón & Krikken, 1990: 74 [original description] Type species. Archedinus relictus Morón & Krikken, 1990 [by original designation]. Gender. Masculine. Species included. 3 species. Distribution. The genus is found in Mexico and Honduras, likely occurring in Guatemala as well. Archedinus antoshkai Seidel & Arriaga-Varela sp. nov. (Figs 1A; 2A,C,E; 3A–B,D,F) Archedinus antoshkai Seidel & Arriaga-Varela sp. nov. (this paper). Locality records. HONDURAS: LEMPIRA (this paper). Types. Holotype  in NMPC (Figs 1A; 2A,C,E; 3A‒B,D,F). Arch edinus howdeni Morón & Vaz-de-Mello, 2007 (Figs 1B; 2B,D,F; 3C,G) Archedinus howdeni Morón & Vaz-de-Mello, 2007: 111 [original combination] Locality records from literature. MEXICO: OAXACA (MORÓN & VAZ-DE-MELLO 2007). Types. Holotype  and 1 paratype  in MXAL/IEXA, 1 paratype  in CMNC (Figs 1B; 2B,D,F; 3C,G), 1 paratype  in CEMT (MORÓN & VAZ-DE-MELLO 2007). Archedinus relictus Morón & Krikken, 1990 (Figs 5A‒B) Archedinus relictus Morón & Krikken, 1990: 74 [original combination] Locality records from literature. MEXICO: CHIAPAS (MORÓN & KRIKKEN 1990). Types. Holotype , allotype  and 2 paratypes (1  1 ) in MXAL, 3 paratypes (2  1 ) in SMF (Figs 5A‒B), 2 paratypes (1  1 ) in RMNH (MORÓN & KRIKKEN 1990). Genus Coelocratus Burmeister, 1842 Coelocratus Burmeister, 1842: 767 [original description] Coleocratus: MORÓN & VAZ-DE-MELLO (2007): 118 [incorrect subsequent spelling] Type species. Inca rufi pennis Gory & Percheron, 1833 [by monotypy]. Gender. Masculine. Species included. 1 species. Remarks. BLACKWELDER (1944) and KRAJCIK (1999, 2012) erroneously listed 1841 instead of 1842 as the year of description of the genus. RICCHIARDI (2002) transferred the genus from Trichiini to Incini. Distribution. According to GORY & PERCHERON (1833), Coelocratus rufi pennis is from Brazil, but they did not provide a more detailed locality, and BURMEISTER (1842) restricted it to northern Brazil. Actual specimens in the Burmeister collection (MLUH) are labelled as being from Cayenne (French Guiana) (Fig. 6). Coelocratus rufi pennis (Gory & Percheron, 1833) (Figs 5 C‒D) Inca rufi pennis Gory & Percheron, 1833: 109 [original combination] Ynca rufi pennis: LAPORTE (1840): 159 [combination with misspelled genus name] Coelocratus rufi pennis: BURMEISTER (1842): 768 [new combination] Types. Lectotype  in MNHG (Figs 5C‒D) (designated by RICCHIARDI 2002: 4), 1 paralectotype  in MACN (invalid lectotype designation by DI IORIO 2013: 82). Locality records from literature. BRAZIL (GORY & PERCHERON 1833, BURMEISTER 1842). Additional material examined. FRENCH GUIANA: CAYENNE: 1  1  (MLUH): Cayenne. Remarks. As noted by MACLEAY (1838) the author of the species is GORY & PERCHERON (1833) and not Dupont as stated in GORY & PERCHERON (1833). DI IORIO (2013) overlooked the lectotype designation by RICCHIARDI (2002) and designated a lectotype in the Burmeister collection housed in the MACN. RICCHIARDI (2002) validly designated the lectotype based on one syntype of Inca rufi pennis housed in the MNHG collection. The syntype sent by Gory & Percheron to Burmeister in Argentina therefore has to be regarded as a paralectotype. Genus Golinca Thomson, 1878 Golinca Thomson, 1878: 31 [original description] Type species. Cetonia bifrons Olivier, 1789: 82 [by original designation]. Gender. Masculine. Species included. 4 species. Remarks. When describing Golinca, THOMSON (1878) did not state the gender of the genus name and as such it has not been subsequently established with the combination of specifi c epithets. We assume that Thomson derived Golinca from the generic names Goliathus and Inca, two genera of which Incini species were previously combined with. Therefore, the gender of the genus should be regarded as masculine, like in Inca (see above). Seidel.indd 395 10.9.2018 15:36:30 SEIDEL et al.: Catalogue of Incini and description of a new Archedinus (Coleoptera: Scarabaeidae) 396 Distribution. The genus is found in Bolivia, Brazil, Colombia, Costa Rica (SOLÍS 2007), Ecuador, French Guiana, and Peru (Fig. 6). Examined material not identifi ed to species. BOLIVIA: SANTA CRUZ: 1  (PDPC): Bolivia, Sara province, J. Steinbach leg. BRAZIL: AMAZONAS: 3  (ZMUH): Brazil, Manaus, Rio Negro; 3  (ZMUH): Brazil, Uypiranga, Amazone; 1  (MZSP) Brazil, Amazonas, Manaus, Rio Negro, vi.1921, J. F. Zikán leg., 1  (MZSP): Brazil, Amazonas, Manaus, Reserva Ducke, viii.1965. V. C. Araujo leg. MATO GROSSO: 1  (USNM): Brazil, Mato Grosso, 1886. COLOMBIA: HUILA: 1  (DBPC): Colombia, Huila, Gigante, viii.1992; 1  (RBINS): Colombia, Huila, Gigante, P. Bleuzen leg. NARIÑO: 1  1  (DBPC): Colombia, Nariño, v.2002. COSTA RICA: PUNTARENAS: 1 spec. (MNCR): Costa Rica, Osa, Sierpe, Rancho Quemado, 8.679096, -83.566714, 30.xi.1991, Freddy A. Quesada leg., Número de catálogo: 346020; 1 spec. (MNCR): Costa Rica, Area de Conservación Osa, Golfi to, Parque Nacional Corcovado, Estación Sirena, Playa Sirena, 8.480171, -83.591289, 30.xi.1993, Gilberto Fonseca leg., Número de catálogo: 1957850. ECUADOR: PASTAZA: 1  1  (DBPC): Ecuador, Pastaza, Puyo, iv.1994; 1  (MSPC): Ecuador, Pastaza, Puyo, ca. 2011. PERU: 2  (ZMUH): Peru, Nypiranga, coll. Le Moult; 1  (ADPC): Peru, Rio Huallaga, xi.–xii.1980. A. Bonc Haas leg. LORETO: 1  1  (MSPC): Peru, Loreto, Iquitos, 10.i.2007; 1  1  (ZMUH): Peru, Iquitos, San-Roque. SAN MARTIN: 3  6  (PDPC): Peru, San Martin, Altomayo, Naranjos , iii.2017. UCAYALI: 2  (PDPC): Peru, Ucayali, Atalaya, i.2017. Golinca bifrons (Olivier, 1789) (Figs 5H‒J) Cetonia bifrons Olivier, 1789: 82 [original combination] Inca bifrons: LEPELETIER & SERVILLE (1828): 381 [new combination] Ynca bifrons: LAPORTE (1840): 158 [combination with misspelled genus name] Golinca bifrons: THOMSON (1878): 31 [new combination] Locality records from literature. COLOMBIA: AMAZONAS (OTAVO et al. 2013). ECUADOR: PASTAZA (CAMPOS 1921). FRENCH GUIANA: CAYENNE (OLIVIER 1789). Additional material examined. BRAZIL: AMAZONAS: 1  (USNM): Brazil, Amazonas, Maues, 1934; 1  (USNM): Brazil, Amazonas, Maues, v.1939; 1  1  (USNM): Brazil, Amazonas, Ipiranga, iv.1936. COLOMBIA: HUILA: 1  1  (SLPC): Huila prov., Gigante, 15.ii.2007. FRENCH GUIANA: CAYENNE: 1  (NMPC): Cayenne; 7  7  (DBPC): French Guiana, piste Belizon, ii.2005; 2  (DBPC): French Guiana, near Cacao, ii.2009; 2  (DBPC): French Guiana, piste Regina, xii.2009; 1  1  (ARPC): French Guyana, Matiti ZA Wayabo, i.2015, Jean Louis Giuglaris leg.; 1  (PDPC): French Guiana, Montagne de Kaw, PK29 +1(D6), ii.2006, Bondil F. leg.; 1  (PDPC): French Guiana, Crique Blanche, PK 59,5 +1 (N2), ii.2009; 1  1  (PDPC): French Guiana, piste de Kaw, 26.i.2017; 1  (BCRC): French Guiana: Montagne des Chevaux, RN2, PK 22, xii.2014, S. Brule leg.; 1  (BCRC): French Guiana: Montagne des Chevaux, RN2, PK 22, ii.2012, P.-H Dalens leg.; 1  (BCRC): French Guiana: Montagne des Chevaux, RN2, PK 22, xii.2011; 3  3  (CMNC): French Guiana, Piste de Saint Elie, PK32, i.2000, banana trap, Y. Ponchel leg.; 1  1  (ZMUH): Guyana, Cayenne; 1  (ZMUH): Cayene, coll. Madon. SAINT-LAURENT-DUMARONI: 10  5  (ZMUH): French Guiana, Nouveau Chantier, collection Le Moult; 3  8  (ZMUH): French Guiana, St-Jean du Maroni, collection Le Moult. Remarks. OTAVO et al. (2013) erroneously listed the year of publication for this species as Olivier, 1758. Golinca davisii (Waterhouse, 1877) Inca davisii Waterhouse, 1877: 228 [original combination] Golinca davisi: SCHENKLING (1922): 5 [new combination and incorrect subsequent spelling] Locality records from literature. PERU (WATERHOUSE 1877). Additional material examined. BOLIVIA: SANTA CRUZ: 1  1  (CMNC): Bolivia, Santa Cruz, Ichilo, Steinbach leg.; 1  (CMNC): Bolivia, Santa Cruz, Ichilo, Buenavista, ii.1950, Martínez leg.; 1  (CMNC): Bolivia, Santa Cruz, Ichilo, Buenavista, iii.1955, Martínez leg. ECUADOR: EL ORO: 1  (MNCN): St. Rosa (Ecuador), 24.v.–5.vi.1865, F. Martinez, Expedicion al Pacifi co (1862–1865). MORONA SANTIAGO: 1  (NMPC): Ecuador, Macas, Normandie, 1500 m. Types. Female holotype in BMNH. Remarks. SCHENKLING (1922), while transferring Inca davisii Waterhouse, 1877 to the genus Golinca, misspelled the specifi c epithet as “davisi” lacking one “i”. This incorrect subsequent spelling has since been used by several authors (BERGÉ 1884, BLACKWELDER 1944, KRAJCIK 1999, MORÓN & VAZ-DE-MELLO 2007, VALOIS & SILVA 2015). Nevertheless, the correct spelling, G. davisii, remained in use (NAGAI 1994, RATCLIFFE et al. 2015, DE SOUZA SILVA et al. 2017). We follow Art. 33.3 (ICZN 1999) which states that “any subsequent spelling of a name different from the correct original spelling, other than a mandatory change or an emendation, is an “incorrect subsequent spelling”; it is not an available name”. Since both the correct and incorrect subsequent spellings were recently in use, the incorrect subsequent spelling is not in prevailing usage and cannot be preserved under Art. 33.3. The name to be used is Golinca davisii (Waterhouse, 1877). Golinca ishiharai Nagai, 1994 Golinca ishiharai Nagai, 1994: 243 [original combination] Types. Holotype  in MNHY, 1 paratype  in TMPC, 1 paratype  in KSPC (NAGAI 1994). Locality records from literature. PERU: SATIPO (NAGAI 1994). Remarks. The holotype of G. ishiharai was not found in the MNHY and possibly could have been destroyed by a fi re that consumed Shinji Nagai’s personal collection (pers. comm. by Kyohei Watanabe, 2018). Golinca trevisani Valois & Silva, 2015 Golinca trevisani Valois & Silva, 2015: 193 [original combination] Types. Holotype  in MPEG, 1 paratype  in BCRC (VALOIS & SILVA 2015). Locality records from literature. BRAZIL: RONDÔNIA (VALOIS & SILVA 2015), AMAZONAS (VALOIS & SILVA 2015). Genus Inca LePeletier & Serville, 1828 Inca LePeletier & Serville, 1828: 380 [original description] Ynca Chevrolat, 1833: unpaginated [unjustifi ed emendation] Type species. Cetonia ynca Weber, 1801 [subsequent designation by HOWDEN (1968: 12)] Gender. Masculine. Remarks. The authorship and date of publication of the name Inca has been the subject of confusion. BLACKWELDER (1944), MORÓN et al. (1997), KRAJCIK (1999) and RESTREPO-GIRALDO et al. (2003) erroneously attributed the authorship of Inca to “Serville (1825)”. BOUSQUET (2016) showed that the second livraison of the “Encyclopédie méthodique” containing pages 345 to 832 was published in December 1828. The authors of Inca are LEPELETIER & SERVILLE (1828) as stated on page 346 in the second livraison. KRAJCIK (1999) erroneously listed Ynca as a synonym to Inca and attributed authorship to GORY & PERCHERON Seidel.indd 396 10.9.2018 15:36:30 Acta Entomologica Musei Nationalis Pragae, volume 58, number 2, 2018 397 (1833). However, the name Ynca was introduced by CHEVROLAT (1833) as an unjustifi ed emendation and later used by LAPORTE (1840). LEPELETIER & SERVILLE (1828) did not explain the origin and gender of the genus name. Furthermore, the specifi c epithets associated with the genus in the original description are either gender neutral or can be either masculine or feminine. Since, LEPELETIER & SERVILLE (1828) changed the specifi c epithet “barbicorne” (neutral) to “barbicornis” (masculine and feminine), it is clear that they deemed the gender of the genus not to be neutral. The genus name Inca is derived from the Spanish noun “Inca” (masculine). According to the Diccionario de la Lengua Española (REAL ACADEMIA ESPAÑOLA 2014) the masculine noun “Inca” has three meanings: 1) sovereign who ruled the Inca Empire; 2) descendant of the Inca; 3) old gold coin of Peru. We follow Art. 30.2.1. (ICZN 1999), which states: “If a name reproduces exactly a noun having a gender in a modern European language (without having to be transliterated from a non-Latin alphabet into the Latin alphabet) it takes the gender of that noun.” Since these conditions are met in this case, the gender of the genus Inca is masculine. Furthermore, BURMEISTER (1842) considered the genus name to be masculine and therefore adapted previously feminine species epithets into the masculine form. The type species of the genus was not selected until HOWDEN (1968) designated Cetonia ynca Weber, 1801 as type species. HOWDEN (1968) thought that the original specifi c epithet spelling was “inca” not “ynca” and therefore designated the type species by tautonomy following Art. 68.4 (ICZN). Even though in this case the absolute tautonymy was not met, we follow Art. 69.1.1 (ICZN 1999), which states that: “in the absence of a prior type fi xation for a nominal genus or subgenus, an author is deemed to have designated one of the originally included nominal species as type species, if he or she states (for whatever reason, right or wrong) that it is the type or type species, or uses an equivalent term, and if it is clear that that author accepts it as the type species.” Therefore, the subsequent designation of Cetonia ynca as type species of Inca by HOWDEN (1968) is valid. Distribution. Known from Tamaulipas in Mexico through Central and South America to Paraguay and northern Argentina (Fig. 6). There are no records known to us from Venezuela, Guyana, and Suriname, but specimens collected in Trinidad and close to the Venezuela and Suriname borders indicate the genus might well be distributed in those countries. Examined material not identifi ed to species (I. irroratus/burmeisteri group). BRAZIL: RIO DE JANEIRO: 1  (BCRC): Brazil, Rio de Janeiro, Rio Friburgo, x.1983, C. Behduin leg.; 1  (USNM): Brazil, Serra da Carioca. SANTA CATARINA: 1  1  (MZSP): Brasil, Santa Catarina, Joinville, Dirings leg.; 1  (MZSP): Brasil, Santa Catarina, Rio Vermelho, ix.1957; 1  (MZSP): Brasil, Rio Vermelho, Santa Catarina, xi.1960, Dirings leg.; 1  (MZSP): Brasil, Santa Catarina, Rio Vermelho, i.1952; 1  (MZSP): Brasil, Rio Vermelho, Santa Catarina, iii.1949, Dirings leg.; 1  (MZSP): Brasil: Santa Catarina, Timbó; 2  8  (MZSP): Brasil: Santa Catarina, Timbó, iii.1960, Dirings leg.; 3  (MZSP): Brasil: Santa Catarina, Timbó, vi.1969; 1  (MZSP): Brasil: Santa Catarina, Timbó, v.1956. SÃO PAULO: 1  (MZSP): Brasil, São Paulo, (Capital), i.1960, Dirings leg.; 1  (MZSP): Brasil, São Paulo, Pindamonhagaba, Eugênio Lefevre, iii.1963, Exp. Dep. Zoologia leg.; 1  (MZSP): Brasil, São Paulo, Salesópolis, Est. Biol. Boraceia, 1.‒4.ii.1973, Vanin leg. Inca besckii Burmeister & Schaum, 1840 Inca besckii Burmeister & Schaum, 1840: 380 [original combination] Inca beskii: BURMEISTER (1842): 711 [incorrect subsequent spelling, unavailable name] Inca beskei: OHAUS (1900): 219 [incorrect subsequent spelling, unavailable name] Inca besckei: SCHENKLING (1922): 3 [incorrect subsequent spelling, unavailable name] Types. Six specimens (possible syntypes) of Inca besckii in MLUH. No types were found in ZMHB. Locality records from literature. BRAZIL (BURMEISTER & SCHAUM 1840), RIO DE JANEIRO (OHAUS 1900). Additional material examined. BRAZIL: MINAS GERAIS: 1  (CEMT): Brazil, Minas Gerais, Mata do Paraíso, 20°47ʹS; 42° 51ʹW, F. Vaz de Mello leg.; 1  (MZSP): Minas Gerais, Passa Quatro, 915 m, 24.i.1923, J. Zikán leg., Banana Koder; 1  (MZSP): Brazil, Minas Gerais, Viçosa, 648 m, xii.1957, col. E. Amante; 1  (MZSP): Minas (Brazil), Passa Quatro, Faz. Dos Campos, 31.i.1916, J. F. Zikán leg.; 1  (MZSP): Minas Gerais, Virginia, Faz. Dos Campos 1500 m, 9.ii.1916, J. F. Zikán leg. PARANÁ: 1  (NMPC): Brazil, Parana; 1  (BCRC): Brazil, Parana, Roladia, III; 1  (BCRC): Brazil, Parana, Caviuna, XII; 1  1  (AKPC): Brazil, Parana, i.2007; 1  (CMNC): Brazil, Campina Grande nr. Curitiba, 15.ii.1966; H. & M. Townes leg.; 1  (ZMUH): Parana, Curitiba; 2  1  (ZMUH): Brazil, Parana, Lucena. RIO DE JANEIRO: 1  (MZSP): Brazil, Rio de Janeiro, Itatiaya, 1100 m, Maromba, 18.ii.1939, J. F. Zikán leg.; 2  (BCRC): Brazil, Rio de Janeiro, Itatiaya, ii.1969; 1  (NHRS): Rio de Janeiro, Petropolis, 19.xi.1904, F. Ohaus; 1  (NHRS): Rio de Janeiro, Petropolis, 2.ii.1899, F. Ohaus; 1  (NHRS): Rio de Janeiro, Petropolis, 12.i.1899, F. Ohaus. RIO GRANDE DO SUL: 1  (MZUEFS): Brazil, Rio Grande do Sul, Vila Olivia, 14.ii.1950, J. Becker leg. (#30591); 1  (MZUEFS): Brazil, Rio Grande do Sul, Vila Olivia, 14.ii.1950, J. Becker leg. (#30599). SANTA CATARINA: 1  1  (ARPC): Brazil, Santa Catarina, Corupá, iii.1966; 1  (MSPC): Brazil, Santa Catarina, iii.2004; 1  (ARPC): Brazil Santa Catarina, Rio Natal, xii.1975; 1  (MZSP): Brazil, Santa Catarina, Rio Vermelho, vi.1961, Dirings leg.; 3  (BCRC): Brazil, Santa Catarina, Rio Vermelho, III; 2  (BCRC): Brazil, Santa Catarina, 1980; 1  (BCRC): Brazil, Sao Bento do Sul; 1  (MZSP): Brasilien, Santa Catarina, Nova Teutonia, ii.1935, B. Pohl leg.; 1  1  (USNM): Brazil, Santa Catarina, Nova Teutonia, ii.1965; 1  1  (USNM): Brazil, Santa Catarina, Nova Teutona, iii.1974; 1  1  (USNM): Brazil, Santa Catarina; 1  (CMNC): Brazil, Santa Catarina, Corupá, xi.1977; 2  1  (CMNC): Brazil, Santa Catarina, Jaraguá, iii.1963; Maller leg.; 1  (CMNC): Brazil, Santa Catarina, Mafi a, iv.1974, W. Johnson leg.; 1  (ZMUH): Santa Catharina. Remarks. SCHENKLING (1922), BLACKWELDER (1944), KRAJCIK (1999, 2012) and MORÓN & VAZ-DE-MELLO (2007) erroneously listed Schaum as the only author. Inca bonplandi (Gyllenhal, 1817) Trichius bonplandi Gyllenhal, 1817: 196 [original combination] Inca serricollis (partim): GORY & PERCHERON (1833): 108 [I. bonplandi synonymized with I. serricollis ignoring priority] Inca bonplandi: BURMEISTER & SCHAUM (1840): 380 [new combination and resurrection of the senior synonym] Ynca bonplandi: LAPORTE (1840): 159 [combination with misspelled genus name] Inca bomplandi: BRUCH (1911): 216 [incorrect subsequent spelling, unavailable name] Inca tapujo Perty, 1830. Junior subjective synonym. Inca tapujo Perty, 1830: 51 [original combination]. Synonymized with I. serricollis by GORY & PERCHERON (1833: 108). Ynca tapago: LAPORTE (1840): 159 [combination with misspelled genus name and incorrect subsequent spelling of specifi c epithet, unavailable name] Inca tapayo: GORY & PERCHERON (1833): 108, 401 [incorrect subsequent spelling, unavailable name] Inca bonplandi var. tapujo: SCHENKLING (1922): 4 [downgraded to variety, unjustifi ed act] Seidel.indd 397 10.9.2018 15:36:30 SEIDEL et al.: Catalogue of Incini and description of a new Archedinus (Coleoptera: Scarabaeidae) 404 der (MLUH), Joachim Händel (MLUH), Johannes Bergsten (NHRS), Martin Husemann (ZMUH), and Hans Mejlon (UUZM) for hosting the fi rst author in their collections. Max Barclay (BMNH) is thanked for loaning the Incini types under his care. We thank Viktor Sinyaev (Moscow, Russia) for providing the habitat photos and Aleš Bezděk (Biology Centre of the Czech Academy of Sciences CAS, Institute of Entomology, České Budějovice, Czech Republic), Brett Ratcliffe (UNSM), and Lukáš Sekerka (NMPC) for helping us with literature. 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