François Génier

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.

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.

Reclassification of Homocopris Burmeister as a Valid Genus to Accommodate Three Species Formerly in Dichotomius Hope (Scarabaeidae: Scarabaeinae: Coprini)

Burmeister (1846) created Homocopris as a subgenus of Copris Geoffroy for the Chilean species Copris torulosus Eschscholtz, 1822. Harold (1869) latermovedC. torulosus to the genusPinotusErichson, 1847, thereby placingHomocopris in synonymywith Pinotus. The species Pinotus torulosus remained in this combination until Martínez (1951) determined Pinotus to be a junior synonym of Dichotomius Hope, 1838. Dichotomius is a very large genus centered in South America, but widely distributed in the New World. The purpose of this note is to reassess the status of Homocopris in order to facilitate use of the name in upcoming publications. Our ongoing studies ofDichotomius have led us to reaffirm Burmeister’s original conclusions about the taxonomic uniqueness of C. torulosus. It, along with two other species, Dichotomius achamas (Harold, 1880) and Dichotomius buckleyi (Waterhouse, 1891), share a combination of characters that distinguish them from other Dichotomius and related genera of Coprini. These characters are: a) a transverse, keel-like, ventral clypeal process (conical and usually bifurcate in Dichotomius); b) presence of a supplemental, lateral, pronotal carina above the carina separating the pronotum and hypomeron (supplemental carina lacking in Dichotomius); c) metatibial spur flattened and truncated apically (bifurcate or subapically dentate inDichotomius); and d) the absence of a large medial lamella-copulatrix-like structure in the internal sac (present inDichotomius). Based on this evidence, Homocopris Burmeister, new status, is hereby considered a valid genus of Coprini (Scarabaeidae: Scarabaeinae). The type species of the genus is Copris torulosus Eschscholtz by monotypy. The species here included in the genus are: Homocopris torulosus (Eschscholtz), new combination, recorded from Chile, Argentina, and Brazil; Homocopris achamas (Harold), new combination, recorded from Colombia; and Homocopris buckleyi (Waterhouse), new combination, recorded from Ecuador. Several new species are also known to us and will be described in an upcoming revision of the genus. We thank W. D. Edmonds for his constructive review of the preliminary manuscript. FZVMwas sponsored by Fundação de Amparo à Pesquisa do Estado deMato Grosso (570847/2008 and 447441/2009) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq PDJ 151603/2007-3). Part of this work was funded by Fundação de Amparo à Pesquisa do Estado de Minas Gerais/Biota Minas (CRA-APQ 03554-0).

A New Species And Notes On The Subgenus Deltochilum (Deltochilum) Eschscholtz, 1822 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini)

Deltochilum eurymedon new species is described and illustrated. Two new synonymies are proposed, Deltochilum (Telhyboma) Kolbe, 1893 = Deltochilum (Deltochilum) Eschscholtz, 1822 and D. (D.) scabriusculum montanum Howden, 1966 = D. (D.) scabriusculum Bates, 1887. Diagnostic characters are given and aedeagi are illustrated for each species currently included in the subgenus Deltochilum. Finally, distributional data and literature records are also provided.

A New North American ATEUCHUS Weber (Coleoptera: Scarabaeidae, Scarabaeinae)

Abstract Ateuchus floridensis, new species, from Florida is described and illustrated. Its relationship with A. histeroides Weber and A. lecontei (Harold) is discussed. A new key to identify species of Ateuchus occurring north of Mexico is provided.

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.

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.

Onthophagus mirabilis Bates, Description of the Newly Discovered Female (Coleoptera: Scarabaeidae, Scarabaeinae)

Abstract The previously unknown female of Onthophagus mirabilis Bates is described. Its affinities to other species of the group are discussed and observations on its biology and collecting methods are given.

Dendropaemon Perty, 1830 Nomenclature Revisited: On the Unavailability of “Onthoecus Lacordaire, 1856” (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini)

Phanaeini are by far the dung beetle tribe with the most understood systematics in the NewWorld. For the past three decades, a series of papers has been published addressing the taxonomy of most of its now recognized 11 genera (see list in Cupello and Vaz-de-Mello 2013), and the last of those was Génier and Arnaud’s (2016) revision of the highly enigmatic genus Dendropaemon Perty, 1830. Although rich in species (41 recognized by Génier and Arnaud 2016, 18 of them as new), representatives of the genus are very rare in collections compared to other phanaeine groups. This fact is likely related to the putative unusual biology of at least some Dendropaemon species, which are thought to be ant-nest inquilines (Vaz-de-Mello and Génier 2009; Génier and Arnaud 2016). As a direct consequence of the rarity of the genus in collections,Dendropaemon taxonomy was largely neglected for the most part of the 20 century after Blut’s (1939) revision of the genus. One exception was Edmonds’ (1972) review of the phanaeine supraspecific classification, where Dendropaemon was divided into three subgenera: Dendropaemon s. str., Paradendropaemon Edmonds, 1972, and Coprophanaeoides Edmonds, 1972, the former including three species-groups. Forty-four years later, that number would quadruple. In addition to those three subgenera, Génier and Arnaud (2016) recognized nine others: their new taxa Glaphyropaemon Génier and Arnaud,NigropaemonGénier and Arnaud, Rutilopaemon Génier and Arnaud, Streblopaemon Génier and Arnaud, Sulcopaemon Génier and Arnaud, and Titthopaemon Génier and Arnaud, as well as Eurypodea Klages, 1906 (until then considered a valid monospecific genus under the name Tetramereia Klages, 1907, which is now considered a junior synonym of Eurypodea), Enicotarsus Laporte, 1831, and Onthoecus Lacordaire, 1856, the latter two names previously regarded as available junior synonyms of Dendropaemon. As reported in their work, Génier and Arnaud (2016) came across a significant number of nomenclatural problems involving those genus-group names and, for most of them, their treatment of those issues has proven adequate. However, while working on a catalogue of the dung beetles of the Brazilian state of Rio de Janeiro, the first author (MC) encountered a nomenclatural issue related to the availability of the subgenus name Onthoecus, which is addressed below. The nameOnthoecus first appeared in the second edition of Dejean’s catalogue of his beetle collection. On page 140, Dejean (1833) listed “Onthoecus Dejean” to include two unpublished species names, “Amyntas Dej.” and “Depressus Dupont,” both from unspecified localities in Brazil (“Brasilia”). On page 156 in the catalogue’s third edition, Dejean (1836) included a third unpublished species name, “Æneipennis Buquet”, from “Brasilia”, and listed “Telephus Lacordaire” from Cayenne as a synonym of “Depressus”. As previously noted by Bousquet and Bouchard (2013), no available species names

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.