A new Peruvian species of Scybalocanthon Martínez, 1948 (Coleoptera, Scarabaeidae, Scarabaeinae, Deltochilini) and some remarkable intrapopulational variation in the endophallus of S. pinopterus (Kirsch, 1873).

Scybalocanthon ashei sp. nov. from Madre de Dios, Peru, is described based on differences in external and male genital morphology. Its diagnostic characters and an updated identification key to the species of the genus are provided as well as new distributional data for the following species: S. acrianus Silva & Valois, 2019, S. aereus (Schmidt, 1922), S. kaestneri (Balthasar, 1939) and S. pinopterus (Kirsch, 1873).

A new species of Onitis Fabricius, 1798 from south-eastern Africa (Coleoptera, Scarabaeidae, Scarabaeinae, Onitini).

Onitis albertcollarti sp. nov. is described and illustrated. The new species is closely related to and occurs sympatrically with O. lycophron Klug, 1855.

Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae).

The taxonomy and systematics of the genus Digitonthophagus Balthasar (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini) is revised. A detailed study of the male genitalia combined with external morphology suggests that the variability, previously recognized, for D. gazella is hiding a species complex within the Afrotropical region and the Arabian Peninsula. The current study recognizes 16 species; 13 from the Afrotropical region and Arabian Peninsula and three from the eastern portion of the Saharo-Arabian region and the continental Indomalayan region. Species are organized into six species groups based on the results of the morphology-based phylogenetic analysis. The following 12 species are described as new: D. aksumensis Génier new species; D. biflagellatus Génier new species; D. dilatatus Génier new species; D. eucatta Génier new species; D. falciger Génier new species; D. fimator Génier new species; D. namaquensis Génier new species; D. petilus Génier new species; D. sahelicus Moretto new species; D. uks Génier new species; D. ulcerosus Génier new species; and D. viridicollis Génier new species. In order to stabilize nomenclature, lectotypes are designated for Scarabaeus bonasus Fabricius, 1775; Scarabaeus catta Fabricius, 1787, and Onthophagus gazella lusinganus d'Orbigny. A neotype is designated for Scarabaeus dorcas Olivier, 1789 whose status and synonymy need to be altered in order to clarify the status of Scarabaeus gazella auctorum, the widely introduced species with economic importance. A naming scheme is presented for the sclerites of the internal sac. External and male genitalia are illustrated and distribution maps are provided for each species.

Digitonthophagus gazella auctorum: an unfortunate case of mistaken identity for a widely introduced species (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini).

At risk of committing entomological heresy, we question the identity of a dung-burying beetle species that originates from Africa and has been introduced first into Hawaii and subsequently to Australasia, North America, and South America (Fincher 1986; Edwards 2007; Noriega et al. 2010) for pasture improvement and biological control of dung-breeding flies (Waterhouse 1974; Bornemissza 1979).  Under the name Onthophagus gazella (Fabricius 1787), it was the first species selected for introduction into Australia by the CSIRO Dung Beetle Project (Bornemissza 1976; Edwards 2007). Firstly, in 1968, a "tropical strain" was introduced from Hawaii where it had become established after introduction from Zimbabwe in 1957 (Markin & Yoshioka 1998). Later, after establishment of the CSIRO Dung Beetle Research Unit in Pretoria in 1970, a "cold" or "even rainfall strain" was introduced into Australia directly from South Africa (Bornemissza 1976) (even rainfall region = south coast of Eastern Cape). The species was subsequently introduced into the southern continental United States of America (Victoria County, Texas) from Hawaii (Montes de Oca & Halffter 1998) then elsewhere into southeastern and southwestern states from Hawaii and breeding colonies from Australia (Anderson & Loomis 1978). It has since expanded its range through Mexico, Central America, and the Caribbean to coastal Colombia (Kohlmann 1994; Noriega 2002; Noriega et al. 2006, 2011). Expansion of its range within central southern South America (Noriega et al. 2010) has been assisted by introductions into Brazil from the United States of America since the 1980s (Bianchin et al. 1998), and others into Venezuela and Chile (Vidaurre et al. 2008). More recently, it has been introduced into quarantine and field trials in New Zealand (Forgie et al. 2013) using individuals originating from the south coast of the Eastern Cape and Northwest Province of South Africa (S. Forgie, personal communication).

Dendropaemon Perty, 1830: taxonomy, systematics and phylogeny of the morphologically most derived phanaeine genus (Coleoptera: Scarabaeidae, Scarabaeinae, Phanaeini).

The taxonomy and systematics of the Neotropical genus Dendropaemon Perty is revised. The current study recognize 41 species organized into 12 subgenera. The establishment of the subgenera is reflecting the presented phylogenetic analysis. Six subgenera are established from previously available genus group names: Coprophanaeoides Edmonds, 1972; Dendropaemon Perty, 1830; Enicotarsus Laporte, 1831; Eurypodea Klages, 1906; Onthoecus Lacordaire, 1856; Paradendropaemon Edmonds, 1972 and Tetramereia Klages, 1907. Six additional subgenera are described as new: Glaphyropaemon n. subg.; Nigropaemon n. subg.; Rutilopaemon n. subg.; Streblopaemon n. subg.; Sulcopaemon n. subg. and Titthopaemon n. subg. The following 18 species are described as new: Dendropaemon (Coprophanaeoides) bluti n. sp.; Dendropaemon (Coprophanaeoides) carinifer n. sp.; Dendropaemon (Coprophanaeoides) compressipennis n. sp.; Dendropaemon (Coprophanaeoides) cribrosus n. sp.; Dendropaemon (Coprophanaeoides) furtadoi n. sp.; Dendropaemon (Coprophanaeoides) inflatus n. sp.; Dendropaemon (Coprophanaeoides) pilosissimus n. sp.; Dendropaemon (Dendropaemon) aenigmaticus n. sp.; Dendropaemon (Dendropaemon) amazonicus n. sp.; Dendropaemon (Dendropaemon) angustulus n. sp.; Dendropaemon (Dendropaemon) flechtmanni n. sp.; Dendropaemon (Dendropaemon) larseni n. sp.; Dendropaemon (Glaphyropaemon) inemarginatus n. sp.; Dendropaemon (Nigropaemon) nigritulus n. sp.; Dendropaemon (Onthoecus) lydiae n. sp.; Dendropaemon (Onthoecus) morettoi n. sp.; Dendropaemon (Paradendropaemon) vazdemelloi n. sp.; Dendropaemon (Sulcopaemon) latistriatus n. sp.. The following nomen novum: Dendropaemon (Onthoecus) attalus nom. nov. is created to replace the primary junior homonym Dendropaemon amyntas Harold, 1868. Except for Dendropaemon montei Pessôa & Lane, 1936, type material of all the species have been examined and lectotypes designated for the following two species: Dendropaemon fasces Blut, 1939 and Dendropaemon lobatus Waterhouse, 1891. In order to stabilize nomenclature, neotypes were also designated for the following species: Dendropaemon convexus Harold, 1869; Enicotarsus ater Laporte, 1832; Enicotarsus quadratus Laporte, 1932 and; Enicotarsus viridipennis Laporte, 1831. Color habitus are presented for each of the valid species.

Notes on Afrotropical Catharsius species described by Edgar von Harold (Coleoptera: Scarabaeidae, Scarabaeinae).

The type material of the Afrotropical Catharsius species described by E. von Harold is studied. Lectotypes are designated for C. camillus, C. dux, C. harpagus, and C. pandion. Habiti, aedeagi and original labels of the primary types are illustrated. Catharsius mossambicanus Ferreira, 1960 is here considered a junior synonym of C. pandion Harold, 1877. An alloreferent male is designated and illustrated for C. dux Harold, 1878 in order to illustrate males of this species in which females are undistinguishable from C. duciformis Ferreira, 1959.

Ateuchus cujuchi n. sp., a new inquiline species of Scarabaeinae (Coleoptera: Scarabaeidae) from tuco-tuco burrows in Bolivia.

In 2012, P. Skelley and J. Wappes were investigating the insect fauna of Ctenomys (Blainville, 1826) (Rodentia: Ctenomyidae) burrows at low elevation in Santa Cruz de la Sierra province, Bolivia. A number of beetles were extracted from this microhabitat and among them, 50 specimens belonging to the New World genus Ateuchus Weber from the subfamily Scarabaeinae (Coleoptera: Scarabaeidae). The specimens were submitted to the author for identification and did not match any currently described species. Although South American species of the genus Ateuchus are critically in need of a modern revision, it is considered important to describe this particular species as it is the first one recorded from mammal burrows in South America and it is easily distinguishable from all other known Ateuchus.

Innovative Bayesian and parsimony phylogeny of dung beetles (coleoptera, scarabaeidae, scarabaeinae) enhanced by ontology-based partitioning of morphological characters.

Scarabaeine dung beetles are the dominant dung feeding group of insects and are widely used as model organisms in conservation, ecology and developmental biology. Due to the conflicts among 13 recently published phylogenies dealing with the higher-level relationships of dung beetles, the phylogeny of this lineage remains largely unresolved. In this study, we conduct rigorous phylogenetic analyses of dung beetles, based on an unprecedented taxon sample (110 taxa) and detailed investigation of morphology (205 characters). We provide the description of morphology and thoroughly illustrate the used characters. Along with parsimony, traditionally used in the analysis of morphological data, we also apply the Bayesian method with a novel approach that uses anatomy ontology for matrix partitioning. This approach allows for heterogeneity in evolutionary rates among characters from different anatomical regions. Anatomy ontology generates a number of parameter-partition schemes which we compare using Bayes factor. We also test the effect of inclusion of autapomorphies in the morphological analysis, which hitherto has not been examined. Generally, schemes with more parameters were favored in the Bayesian comparison suggesting that characters located on different body regions evolve at different rates and that partitioning of the data matrix using anatomy ontology is reasonable; however, trees from the parsimony and all the Bayesian analyses were quite consistent. The hypothesized phylogeny reveals many novel clades and provides additional support for some clades recovered in previous analyses. Our results provide a solid basis for a new classification of dung beetles, in which the taxonomic limits of the tribes Dichotomiini, Deltochilini and Coprini are restricted and many new tribes must be described. Based on the consistency of the phylogeny with biogeography, we speculate that dung beetles may have originated in the Mesozoic contrary to the traditional view pointing to a Cenozoic origin.

Onthophagus fragosus n.sp. a second endemic species of Onthophagus Latreille from Cuba (Coleoptera : Scarabaeidae, Scarabaeinae).

Onthophagus fragosus, a new species of Onthophagus, is described from Cienfuegos Province on the island of Cuba. Specimens were collected in leaf litter samples at elevations of 650-935 m. It appears to be most closely related to O. marginatus Castelnau, the only other Cuban endemic Onthophagus species. Images of habitus and of male genitalia are provided. 

From coprophagy to predation: a dung beetle that kills millipedes.

The dung beetle subfamily Scarabaeinae is a cosmopolitan group of insects that feed primarily on dung. We describe the first case of an obligate predatory dung beetle and contrast its behaviour and morphology with those of its coprophagous sympatric congeners. Deltochilum valgum Burmeister killed and consumed millipedes in lowland rainforest in Peru. Ancestral ball-rolling behaviour shared by other canthonine species is abandoned, and the head, hind tibiae and pygidium of D. valgum are modified for novel functions during millipede predation. Millipedes were killed by disarticulation, often through decapitation, using the clypeus as a lever. Beetles killed millipedes much larger than themselves. In pitfall traps, D. valgum was attracted exclusively to millipedes, and preferred injured over uninjured millipedes. Morphological similarities placing D. valgum in the same subgenus with non-predatory dung-feeding species suggest a major and potentially rapid behavioural shift from coprophagy to predation. Ecological transitions enabling the exploitation of dramatically atypical niches, which may be more likely to occur when competition is intense, may help explain the evolution of novel ecological guilds and the diversification of exceptionally species-rich groups such as insects.