Streblopus - Van Lansberge 1874

Genus Streblopus Van Lansberge, 1874 Colonychus Harold, 1868: 10. Streblopus Van Lansberge, 1874a: 9���10. Streblopoides Balthasar, 1938: 215���216. Colonychus ��� Van Lansberge 1874a: 10 (as possible synonym of Streblopus). ��� Kolbe 1905: 552; 1907: 27. ��� Gillet 1911: 42 (as synonym of Streblopus). ��� Lucas 1920: 197 (as synonym of Streblopus). ��� Paulian1939:26 (as synonym of Streblopus).��� Blackwelder 1944:203 (as synonym of Streblopus).��� Pereira & Mart��nez 1956: 99 (as a supposed preoccupied synonym of Streblopus). ��� Halffter 1961: 226, 229 (as synonym of Streblopus). ��� Vulcano & Pereira 1964: 580 (as a supposed nomen nudum referring to Streblopus). ��� Halffter & Mart��nez 1966: 152 (as a supposed nomen nudum referring to Streblopus). ��� Krajcik 2006: 163 (as synonym of Streblopus). ��� Chamorro et al. 2019: 234 (as a supposed nomen nudum referring to Streblopus). ��� Cupello & Vaz-de-Mello 2019: 168���171 (as synonym of Streblopus). Streblopus ��� Van Lansberge1874b:187,189.��� Karsch1887:1.��� Ritsema1888:209.��� Gillet1911:42.��� Lucas 1920:617.��� Olsoufieff 1935:34.��� Paulian1938: 234; 1939: 26. ��� Balthasar 1941:345; 1951: 330. ��� Blackwelder 1944: 203. ��� Pereira & Mart��nez 1956: 94, 99, 182. ��� Halffter 1961: 229���230, 253; 1974: 257; 2003: 22. ��� Vulcano & Pereira 1964: 580; 1967: 548���549. ��� Halffter & Matthews 1966: 260. ��� Halffter & Mart��nez 1966: 103, 152���163, figs 13���21; 1967: 79; 1968: 210; 1977: 34, 43. ��� Matthews 1971: 49. ��� Mart��nez & Halffter 1972: 33. ��� Halffter & Edmonds 1982: 201. ��� Hanski & Cambefort 1991: 472. ��� Vaz-de-Mello 2000: 195. ��� Medina & Scholtz 2005: 154. ��� Krajcik 2006: 163; 2012: 249. ��� Scholtz et al. 2009: 566. ��� Carvajal-L��pez et al. 2011: 122, 316. ��� Vaz-de-Mello et al. 2011: 6, 11, 18, 25, 33, 40, 45. ��� Carvajal-L��pez 2012: 195. ��� Chamorro et al. 2018: 98; 2019: 10, 234. ��� Cupello & Vaz-de-Mello 2019: 168���171. Streblopoides ��� Balthasar 1941: 346; 1951: 331. ��� Blackwelder 1944: 203. ��� Pereira & Mart��nez 1956: 99. ��� Halffter 1961: 229; 1974: 257 (as synonym of Streblopus). ��� Vulcano & Pereira 1964: 580. ��� Halffter & Matthews 1966: 260. ��� Halffter & Mart��nez 1966: 103, 153���154, 158 (as synonym of Streblopus); 1967: 79 (as synonym of Streblopus); 1977: 34, 43 (as synonym of Streblopus). ��� Krajcik 2006: 163 (as of dubious validity). ��� Chamorro et al. 2019: 234 (as synonym of Streblopus). ��� Cupello & Vaz-de-Mello 2019: 168���169 (as synonym of Streblopus). Type species Colonychus: Never fixed (see discussion below). ��� Streblopus: Streblopus opatroides Van Lansberge, 1874, by original monotypy (Van Lansberge 1874a: 9���10). ��� Streblopoides: Streblopoides punctatus Balthasar, 1938, by original monotypy (Balthasar 1938: 215���216). Etymology Colonychus: Dubious. Possibly from the Latin prefix ��� col- ���, meaning ���with���, and the Greek word ��� onychus ���, meaning ���claw��� (Brown 1956). Masculine. ��� Streblopus: Dubious. Possibly derived from the Greek word ��� Streblos ��� for ���twisted, crooked, wrinkled��� (Brown 1956). Masculine. ��� Streblopoides: Derived from Streblopus in reference to the close relationship between Balthasar���s new genus and Van Lansberge���s. Differential diagnosis Streblopus is possibly the most readily differentiable genus of the New World dung beetle fauna. As all identification keys that included Streblopus have noted(Paulian 1938; Pereira& Mart��nez1956; Vulcano& Pereira 1967; Halffter & Mart��nez 1977; Vaz-de-Mello et al. 2011), the posteromedian dentiform process of the pronotum of both species is unique among the Scarabaeinae (Figs 1 A���C, 3A, C���D, 12A���B) and can be confidently used to separate the genus from the other groups in the subfamily. The general aspect of the body, which is very elongate, flattened and shows neither cephalic nor pronotal horns, also adds to the peculiar appearance of the genus (Figs 1 A���C, 3A, C���D), as well as the shape of its labial palpi, whose basal palpomere is unusually expanded (Fig. 11D). Indeed, Streblopus is so distinctive that it is difficult to point out any genera with which it could be confused. From the Deltochilini, apart from the characters listed above, Streblopus differs by its mesotibiae being abruptly expanded at the apex in males (Fig. 18A, C); from Deltochilum, in particular, a genus that was once said to be similar to Streblopus (Van Lansberge 1874a; Balthasar 1938) and with which it shares a wide pseudepipleura, Streblopus is different, among many other features, by the absence of tubercles at the apex of the elytra. Its slender metatibiae (Fig. 20 A���D), in turn, will separate Streblopus from Dichotomiini and Coprini, while the presence of tarsal claws (among many other characters) differentiates the genus from the Eucraniini and the absence of three pairs of sharp clypeal teeth will separate it from Scarabaeini. Finally, Streblopus is different from the Eurysternini in not having a visible scutellum, its mesocoxae are obliquely positioned (Fig. 13 C���F) and the labial palpi are 3-articulated (Fig. 11D), whereas Eurysternus, the only genus of its tribe, has a visible scutellum, parallel mesocoxae and 2-articulated labial palpi (see G��nier 2009 for more details). Although certainly far from being exclusive to Streblopus, the labrum having the anterior margin deeply emarginated (Fig. 8 A���D), the protarsi being absent in both sexes (Figs 15 A���H, 16C), the metacoxae having a lateral spur covering the epipleura (Fig. 19A), the hind wing having a deep notch at the area of the anal fold (Fig. 21F), the asymmetrical parameres (Fig. 23 A���H) and the reduced sclerites of the internal sac (Figs 24 A���F, 25A���E) are also remarkable features of this genus. Redescription HEAD. Transverse, with apex ranging from clearly emarginated (males of S. opatroides) to truncate (females of S. punctatus); emargination, when present, flanked by two small teeth; remainder of outer edge simply rounded, with no accessory teeth or notch between clypeus and paraocular areas (��genae��) (Fig. 5 A���D). Suture between clypeus and paraocular areas present and well-marked, with a tenuous tumescence (more evident in males) on clypeus adjacent to suture (Fig. 5 A���D); fronto-clypeal suture vestigial, present very shortly and only at laterals adjacent to eyes (Fig. 5 A���D). Eyes with dorsal surface wide, largest width as wide as one-fifth to one-fourth of interocular space; with a short (S. punctatus) or pronounced (S. opatroides) canthus (Fig. 5 E���F). Entire dorsal surface of head covered by small umbilicate punctation, progressively denser from apex of clypeus towards frons and paraocular area; dorsal surface almost entirely flat, without any traces of horns, tubercles or elevations; a very tenuous depression present only behind apical emargination of clypeus and in paraocular areas in front of eyes. Clypeal process transverse, with apex either rounded (Fig. 16A) or acuminate (Fig. 16B). Antennae 8- or 9-articulated (funicle with 3 or 4 articles) (Fig. 7 A���B). Labrum with lateral projections well developed and with anterior labral margin deeply U-emarginated; epipharynx with well-developed median brush and well-delimited lateral files (Fig. 8 A���D). Mandibles (Fig. 9 A���C), maxillae (Fig. 10 A���C) and labium (Fig. 11 A���C) typical of dung beetles, without any notable variation, except for shape of 3-articulated labial palpi, with basal palpomere largely expanded and almost twice as long as second and third palpomeres combined (Fig. 11D). THORAX. Pronotum transverse, convex, with no elevations, grooves or depressions (Fig. 12 A���B); lateral foveae tiny, but clearly impressed; posterior foveae absent; lateral and anterior edges marginate, anterior margin with a membranous area covering base of frons; lateral edges sinuous in both dorsal and lateral views;postero-lateral angles strongly projecting in an acuminate(or slightly rounded) projection; posterior edge highly sinuous, with a strong flange projecting backwards at centre covering base of elytral suture; surface of pronotum covered by umbilicate punctation, varying in density between species. Hypomeron with anterior region covered by small umbilicate punctures similar to and bearing setae slightly longer than those of head and pronotum, and posterior region with punctures with setae much longer than on rest of tegument (Fig. 13A); hypomeral carina completely absent; anterior region of hypomeron not excavated, posterior region declivous (Fig. 13A). Prosternum, mesoventrite, mesanepisterna and metanepisterna entirely covered by umbilicate punctation, varying in density between species (Fig. 13 B���F); tegument between punctures smooth or with a very diffuse rivulose microsculpture. Mesoventrite wide, with a broad transverse groove at centre (Fig. 13B). Meso-metaventral suture arched (Fig. 13B). Metaventrite covered by umbilicate punctation, denser towards anterior lobe and sides; centre almost devoid of any punctation; posterior region clearly depressed, varying in degree between sexes (Fig. 13 C���F). LEGS. With a highly marked sexual dimorphism and intrasexual allometric variation.All femora and tibiae with ventral surface covered by umbilicate punctation (Figs 14 A���B, 17A���F, 18A���D, 19B���E, 20A���D). Protarsi lacking in both sexes. Protibiae with three acute teeth on apical half of exterior edge; basal half marked by a row of smaller denticles (Fig. 15 A���H). Mesocoxae obliquely orientated to one another, with surface covered by umbilicate punctation, ranging in density between species (Fig. 13 C���F). Meso- and metafemora linear and lacking both anterior and posterior margins (Figs 17 A���F, 19B���E). Mesotibiae slender, but abruptly expanded near apex (Fig. 18 A���D). Mesotarsi long and slender, at least as long as mesotibiae; basal and apical mesotarsomeres subequal in length and as long as mesotarsomeres II���IV combined. Metacoxae with external spur covering epipleura (Fig. 19A, red arrow). Metatrochanters with a sexually-dimorphic brush of long setae at posterior edge (Fig. 19 B���E). Metatibiae slender, longer than metafemora (Fig. 20 A���D). Metatarsi long, at least as long as half-length of metatibiae; metatarsomeres I���IV more or less equivalent in length (tarsomeres slightly shorter towards apex); metatarsomere V very long, almost as long as combined length of metatarsomeres II���IV. Meso- and metatarsal claws curved, well developed. ELYTRA. With a broad pseudepipleura delimited by a short carina adjacent to stria VII; epipleura very narrow (Fig. 21A). With nine fine and well-delimited striae, seven on disc and two on pseudepipleura; striae interrupted by umbilicate punctures along their entire length. Striae I and IX, II and VIII, and III and IV or V connected to one another at apex of elytra; stria VIII absent at base of pseudepipleura. Elytral tegument with umbilicate punctation and with well-delimited alveolar microsculpture between umbilicate punctures. Elytral umbilicate punctures of subequal width or with variable width. Scutellary impression absent; scutellum not visible from above, entirely covered by base of elytra. ABDOMEN. With six ventrites covered by umbilicate punctation. Ventrites I���IV with punctation more clearly marked at their anterior half; ventrite V evenly punctate; ventrite VI evenly punctate at sides and smooth or with punctures very sparse at centre. Abdomen very short at centre, with length subequal to separation between meso- and metacoxae. Ventrite VI as long as ventrites III���V combined. Pygidium very short and deflected towards venter, surface covered by umbilicate punctures; completely marginate, with basal margin slightly raised (Fig. 22C); prepygidium grooved at centre, surface lacking umbilicate punctation (Fig. 22C). TERMINALIA. Aedeagus: Parameres largely asymmetrical, right paramere with clear apical projection (Fig. 23 A���H); in dorsal view, with lateral edges straight or curved; in lateral view, ventral region straight or inclined towards apex, with either well-delimited or tenuous pair of ventral keels. Internal sac with only three sclerites: axial, subaxial and superior right peripheral sclerites (Figs 24 A���F, 25A���E); superior right peripheral sclerite with well-delimited superior ring and broad, long inferior lobe (��cable��) with triangular area more sclerotized near edge (Figs 24E, 25E); axial and subapical sclerites only loosely connected, easily separable. With well-developed raspule covered by tiny scales (Figs 24 A���C, 25A���C). ��� Genital segment: With well-developed medial sclerotized plate and lateral sclerotized plates (Fig. 26 A���B). ��� Spermatheca: C-shaped, both externally and internally smooth, with or without pair of apical hooks (Fig. 27 A���C). SEXUAL DIMORPHISM. Male: Metaventrite strongly excavated posteriorly; concavity marked anteriorly by strong spur (Fig. 13C, E). Profemora robust, strongly enlarged at centre (Fig.14A,C); with anterior margin raised at centre into one or two strong spurs (Fig. 14A, C); trochantofemoral articulation with raised edge forming a short spine or not. Protibiae robust, curved inwards and downwards, with ventral carina interrupted by a longitudinal row of strong teeth; when fully closed, protibiae completely cover anterior edge of profemora; protibial spur present or not (Figs 15A, C, E, G, 16C); apical region of protibiae truncate and expanded inwards into long spur, with two transverse rows of setae (one covering anterior edge of apical lateral teeth, the other covering long spur) (Fig. 16C). Mesotibiae with internal angle of apical edge strongly projecting (Fig. 18A, C). Metatrochanter with wide brush of long setae, covering almost entire posterior edge (Fig. 19B, D). Metatibiae with apical edge strongly projecting inwards (Figs 19E, 20C, E); metatibial spur very short, rudimentary, much shorter than basal metatarsomere (Fig. 20E). Abdomen, in lateral view, contracted, concave (Fig. 22A). ��� Female: Metaventrite only shallowly excavated posteriorly, without indentation marking anterior margin of concavity (Fig. 13D, F). Profemora more linear, anterior margin simple, not raised (Fig. 14B, D); trochantofemoral articulation not developed into spur. Protibiae gracile, only slightly bent inward, not bent downwards (Fig. 15B, D, F, H); internal angle of apex projected into short spur (Fig. 15B, D, F, H); ventral carina simple, not interrupted by row of teeth; at apex, ventral carina marked by tuft of long setae. Mesotibiae with apical expansion, a result of a slight expansion of both internal and external edges, not only internal one (Fig. 18B, D). Metatrochanter with thin brush of long setae on posterior edge (Fig. 19C, E). Metatibiae with apex not expanded inwards, but with sharp spine at internal angle (Fig. 20D, F); metatibial spur well developed, very long, longer than basal metatarsomere (Fig. 20F). Abdomen, in lateral view, flat (Fig. 22B). Distribution As discussed in detail below, the distribution of Streblopus is clearly relict. While one species, S. punctatus, is known from some few sub-Andean localities in eastern Amazonia in Peru and Ecuador, the other, S. opatroides, is known from a larger number of places, but which cover a more limited range in the Atlantic Forest along the Brazilian states of Bahia and Esp��rito Santo. The genus, therefore, is present in ��� and limited to ��� the two major tropical forest ecosystems of South America east of the Andes, and its two species are separated by more than 2000 km of open and drier landscapes composing the Cerrado, in the central area of the South American Dry Diagonal (Fig. 4). Ecology Almost nothing is known about the biology of Streblopus. The specimens gathered over the past years at CEMT and CMNC were mostly collected using pitfall traps baited with human faeces; no specimens are known to have been collected using other types of bait (except literature records for pig dung) or traps. Judging from its morphology, especially the shape of its metatibiae, which are slender and long in both species, we assume the genus should be a roller (something Halffter & Mart��nez (1966: 157) had already hypothesized) and possibly belongs to the functional Pattern IV of Halffter & Edmonds��� classification of dung beetle nesting behaviour (Halffter & Edmonds 1982). Also based on the morphology, the genus is almost certainly nocturnal, as indicated by the large size of the dorsal surface of its eyes (Raine et al. 2019; Tocco et al. 2019) and the dark dorsal colouration, without any or with only few traces of metallic sheen (Hern��ndez 2002; Feer & Pincebourde 2005). Remarks As discussed in the Introduction, the first name by which the genus was referred to in the literature was Colonychus by Harold (1868). Although it was only six years later that Van Lansberge (1874) established the name Streblopus, almost all authors except Kolbe (1905, 1907) have been using Van Lansberge���s name instead of Colonychus, mostly treating the latter as either a nomen nudum (Vulcano & Pereira 1964; Halffter & Mart��nez 1966) or an available preoccupied name (Pereira & Mart��nez 1956). However, since Harold (1868) did not intend to establish a new nominal genus nor included a nominal species in it, one is led to ask whether Colonychus is available or not and, if so, whether it is indeed a synonym of Streblopus. If both questions have a positive answer, then the validity of Streblopus would be threatened in favour of Colonychus, something that would disturb a nomenclatural stability that has reigned for almost 150 years. As for the availability of Colonychus, Harold (1868: 10) cited the name in his discussion on the affinities of some Old World genera of Scarabaeinae with the New World genus Canthon, the group he was revising in that paper. There, Harold mentioned that a specimen bearing Klug���s then-unpublished name Colonychus in the Berlin museum could be a link between the African ���Epirhinen��� and Canthon, and discussed the characteristics that differentiate the two genera, stating the following: ���In between these [���Epirhinen���] and Canthon, there may be, however, placed a yet undescribed genus, which was indicated as Colonychus by Klug at the Berlin museum, and which is characterized by absent protarsi, elytra surpassing the abdomen, very slightly la
Published as part of Cupello, Mario, Ribeiro-Costa, Cibele S. & Vaz-De, Fernando Z., 2020, Systematics of the enigmatic South American Streblopus Van Lansberge, 1874 dung beetles and their transatlantic origin: a case study on the role of dispersal events in the biogeographical history of the Scarabaeinae (Coleoptera: Scarabaeidae), pp. 1-85 in European Journal of Taxonomy 603 on pages 8-17, DOI: 10.5852/ejt.2020.603, http://zenodo.org/record/3666847
{"references":["Van Lansberge J. W. 1874 a. Description de quelques Lamellicornes Coprophages. Coleopterologische Hefte 12: 4 - 12. Available from https: // biodiversitylibrary. org / page / 25118460 [accessed 7 Jan. 2020].","Harold E. 1868. Monographie der Gattung Canthon. Berliner entomologische Zeitschrift 12: 1 - 144. https: // doi. org / 10.1002 / mmnd. 18680120104","Balthasar V. 1938. Neue Gattungen und Arten der sudamerikanischen Coprophagen. (4. Beitrag zur Kenntnis der Scarabaeiden der neotropischen Region). Entomologische Blatter 34 (4): 210 - 223.","Kolbe H. J. 1905. Uber die Lebensweise und die geographische Verbreitung der coprophagen Lamellicornier. Zoologischen Jahrbucher, Supplement 8: 475 - 594.","Kolbe H. [J.] 1907. Hamburger Magalhaensische Sammelreise. Coleopteren. L. Friederichsen & Co., Hamburg.","Gillet J. J. E. 1911. Coleopterorum Catalogus. Pars 38: Scarabaeidae: Coprinae I. W. Junk, Berlin.","Lucas R. 1920. Catalogus Alphabeticus Generum et Subgenerum Coleopterorum Orbis Terrarum Totius (Famil., Trib., Subtr., Sect. incl.). Pars I. Nicolaische Verlags-Buchhandlung, Berlin.","Paulian R. 1939. Contribution a l'etude des canthonides americains [Coleopt. Lamellic.] (suite et fin). Annales de la Societe entomologique de France 108: 1 - 40.","Blackwelder R. E. 1944. Checklist of the coleopterous insects of Mexico, Central America, West Indies and South America. Part 2. United States National Museum Bulletin 185: 189 - 341. https: // doi. org / 10.5479 / si. 03629236.185.2","Pereira F. S. & Martinez A. 1956. Os generos de Canthonini americanos (Col. Scarabaeidae). Revista brasileira de Entomologia 6: 91 - 192.","Halffter G. 1961. Monografia de las especies norteamericanas del genero Canthon Hoffsg. (Coleopt., Scarab.). Ciencia, Revista Hispano-Americana de Ciencias Puras y Aplicadas 20 (9 - 12): 225 - 320.","Vulcano M. A. & Pereira F. S. 1964. Catalogue of the Canthonini (Col. Scarab.) inhabiting the Western Hemisphere. Entomologische Arbeiten aus dem Museum G. Frey 15 (2): 570 - 685. Available from https: // biodiversitylibrary. org / page / 45986646 [accessed 7 Jan. 2020].","Halffter G. & Martinez A. 1966. Revision monografica de los Canthonina americanos (Coleoptera, Scarabaeidae) (Iª Parte). Revista de la Sociedad Mexicana de Historia Natural 27: 89 - 177.","Krajcik M. 2006. Checklist of Scarabaeoidea of the World. 1. Scarabaeinae (Coleoptera: Scarabaeidae: Scarabaeinae). Animma 3 (Suppl. X): 1 - 189.","Chamorro W., Marin-Armijos D., Asenjo A. & Vaz-de-Mello F. Z. 2019. Scarabaeinae dung beetles from Ecuador: a catalog, nomenclatural acts, and distribution records. Zookeys 826: 1 - 343. https: // doi. org / 10.3897 / zookeys. 826.26488","Cupello M. & Vaz-de-Mello F. Z. 2019. Case 3788 - Streblopus Lansberge, 1874 (Insecta, Coleoptera, Scarabaeidae): proposed conservation by suppression of Colonychus Harold, 1868. Bulletin of Zoological Nomenclature 76: 167 - 174.","Van Lansberge J. W. 1874 b. Observations sur la classification des Lamellicornes Coprophages. Annales de la Societe entomologique de Belgique 17: 177 - 193. Available from https: // biodiversitylibrary. org / page / 12728027 [accessed 7 Jan. 2020].","Karsch F. 1887. Altes und neues uber Koleopteren. Berliner entomologische Zeitschrift 31: 1 - 8.","Ritsema C. 1888. Lijst der entomologische geschriften van Mr. J. W. van Lansberge, gevolgd door eene opgave der daarin beschreven nieuwe geslachten, ondergeslachten en soorten. Tijdschrift voor Entomologie 31: 201 - 234. Available from https: // biodiversitylibrary. org / page / 10853215 [accessed 7 Jan. 2020].","Olsoufieff G. 1935. Note critique sur l'essai d'une phylogenie des lamellicornes coprophages. Entomologisches Nachrichtenblatt 9: 32 - 35.","Paulian R. 1938. Contribution a l'etude des canthonides americans [Coleopt. Lamellic.]. Annales de la Societe entomologique de France 107: 213 - 296.","Balthasar V. 1941. Scarabaeidae Laparostictae (Col.). Beitrage zur Fauna Perus 1: 337 - 358.","Balthasar V. 1951. Scarabaeidae Laparostictae (Col.). Beitrage zur Fauna Perus 2: 322 - 344.","Halffter G. 2003. Tribu Scarabaeini, subtribu Canthonina. In: Moron M. A. (ed.) Atlas de los Escarabajos de Mexico. Coleoptera: Lamellicornia. Vol. II. Familias Scarabaeidae, Trogidae, Passalidae y Lucanidae: 21 - 43. Argania Editio, Barcelona.","Vulcano M. A. & Pereira F. S. 1967. Sinopse dos Passalidae e Scarabaeidae s. str. da regiao amazonica (Insecta, Coleoptera). Atas do Simposio sobre a Biota Amazonica 5 (Zoologia): 533 - 603.","Halffter G. & Matthews E. G. 1966. The natural history of dung beetles of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae). Folia Entomologica Mexicana 12 - 14: 1 - 312.","Matthews E. G. 1971. [Book review of \" Revision Monografica de los Canthonina, Parts I, II, and III. Gonzalo Halffter and Antonio Martinez \"]. The Coleopterists Bulletin 25 (2): 47 - 50.","Martinez A. & Halffter G. 1972. New taxa of American Canthonina (Coleoptera, Scarabaeinae). Entomologische Arbeiten aus dem Museum G. Frey 23: 33 - 66. Available from http: // biodiversitylibrary. org / page / 45978703 [accessed 22 Jan. 2020].","Halffter G. & Edmonds W. D. 1982. The Nesting Behavior of Dung Beetles (Scarabaeinae). An Ecological and Evolutive Approach. Instituto de Ecologia, Mexico DF.","Hanski I. & Cambefort Y. 1991. Dung Beetle Ecology. Princeton University Press, Princeton, NJ.","Vaz-de-Mello F. Z. 2000. Estado atual de conhecimentos dos Scarabaeidae s. str. (Coleoptera: Scarabaeoidea) do Brasil. In: Martin-Piera F., Morrone J. J. & Melic A. (eds) Hacia un Proyecto CYTED para el Inventario y Estimacion de la Diversidad entomologica en Iberoamerica: PRIBES- 2000. m 3 m: Monografias Tercer Milenio, Vol. 1: 183 - 195. Sociedad Entomologica Aragonesa (SEA), Zaragoza, Spain.","Medina C. A. & Scholtz C. H. 2005. Systematics of the southern African genus Epirinus Reiche (Coleoptera: Scarabaeinae: Canthonini): descriptions of a new species and phylogeny. Insect Systematics and Evolution 36: 145 - 160. https: // doi. org / 10.1163 / 187631205788838500","Krajcik M. 2012. Checklist of the World Scarabaeoidea. Animma. X, Supplement 5: 1 - 278.","Scholtz C. H., Davis A. L. V. & Kryger U. 2009. Evolutionary Biology and Conservation of Dung Beetles. Pensoft Publishers, Sofia.","Carvajal-Lopez V., Villamarin S. & Ortega A. M. 2011. Escarabajos del Ecuador. Principales Generos. Instituto de Ciencias Biologicas, Escuela Politecnica Nacional, Quito.","Vaz-de-Mello F. Z., Edmonds W. D., Ocampo F. C. & Schoolmeesters P. 2011. A multilingual key to the genera and subgenera of the subfamily Scarabaeinae of the New World (Coleoptera: Scarabaeidae). Zootaxa 2854: 1 - 73.","Carvajal-Lopez V. 2012. Primer registro de Streblopus punctatus Balthasar, 1938 y Dichotomius fonsecae (Luederwald [sic], 1925) (Coleoptera: Scarabaeidae) en las cordilleras del sur-oriente del Ecuador. Revista Politecnica 30 (3): 195 - 200.","Chamorro W., Marin-Armijos D., Granda V. & Vaz-de-Mello F. Z. 2018. Listado de especies y clave de generos y subgeneros de escarabajos estercoleros (Coleoptera: Scarabaeidae: Scarabaeinae) presentes y presuntos para Ecuador. Revista Colombiana de Entomologia 44 (1): 72 - 100. https: // doi. org / 10.25100 / socolen. v 44 i 1.6545","Halffter G. 1974. Elements anciens de l'entomofaune neotropicale: ses implications biogeographiques. Quaestiones Entomologicae 10: 223 - 262.","Brown R. W. 1956. Composition of Scientific Words. A Manual of Methods and a Lexicon of Materials for the Practice of Logotechnics. Smithsonian Books, Washington DC.","Halffter G. & Martinez A. 1977. Revision monografica de los Canthonina americanos, IV parte. Clave para generos y subgeneros. Folia Entomologica Mexicana 38: 29 - 107.","Genier F. 2009. Le genre Eurysternus Dalman, 1824 (Scarabaeidae: Scarabaeinae: Oniticellini), Revision taxonomique et Cles de Determination illustrees. Pensoft, Sofia and Moscow.","Raine E. H., Gray C. L., Mann D. J. & Slade E. M. 2019. Tropical dung beetle morphological traits predict functional traits and show intraspecific differences across land uses. Ecology and Evolution 8 (17): 8686 - 8696. https: // doi. org / 10.1002 / ece 3.4218","Tocco C., Dacke M. & Byrne M. 2019. Eye and wing structure closely reflects the visual ecology of dung beetles. Journal of Comparative Physiology A 205 (2): 211 - 221. https: // doi. org / 10.1007 / s 00359 - 019 - 01324 - 6","Hernandez M. I. M. 2002. The night and day of dung beetles (Coleoptera, Scarabaeidae) in the Serra do Japi, Brazil: elytra colour related to daily activity. Revista brasileira de Entomologia 46 (4): 597 - 600.","Feer F. & Pincebourde S. 2005. Diel flight activity and ecological segregation within an assemblage of tropical forest dung and carrion beetles. Journal of Tropical Ecology 21 (1): 21 - 30. https: // doi. org / 10.1017 / S 0266467404002056","Harold E. 1869. Catalogus Coleopterorum. Hucusque Descriptorum, Synonymicus et Systematicus. Vol. IV. Scarabaeidae: 979 - 1346. E. H. Gummi, Munich. Available from http: // biodiversitylibrary. org / page / 9641549 [accessed 7 Jan. 2020].","ICZN [International Commission on Zoological Nomenclature] 1999. International Code of Zoological Nomenclature. Fourth edition. International Trust for Zoological Nomenclature, London. Available from http: // www. nhm. ac. uk / hosted-sites / iczn / code / [accessed 7 Jan. 2020].","Horn W., Kahle I., Friese G. & Gaedike R. 1990 b. Collectiones Entomologicae. Ein Kompendium uber den Verbleib entomologischer Sammlungen der Welt bis 1960. Teil II: L bis Z. Akademie der Landwirtschaftswissenschaften der Deutschen Demokratischen Republik, Berlin.","Anonymous. 2004. Nomenclator Zoologicus. Volumes 1 - 10. A List of the Names of the Genera and Subgenera in Zoology from the Tenth Edition of Linnaeus 1758 to the End of 2004. Neave S. A. (ed.) Ver. 0.86. Available from http: // ubio. org / NomenclatorZoologicus / [accessed 8 Sep. 2018]."]}