Evolution of the Scarabaeini (Scarabaeidae: Scarabaeinae)

Abstract

Abstract. A phylogenetic analysis of the Scarabaeini, based on 244 morphological characters, including 154 multistate and three biological characters, is presented. Tree topologies generated from unweighted data and some weighted algorithms are similar and support only two clades in the tribe representing the genera Scarabaeus L. and Pachylomerus Bertoloni. The only supported subordinate groups treated in this paper as subgenera are Kheper Kirby stat. nov., Pachysoma MacLeay, Scarabaeolus Balthasar and Sceliages Westwood stat. nov.Drepanopodus Janssens syn. nov. is synonymised with Scarabaeus and six additional names, Madateuchus Paulian, Mnematidium Ritsema, Mnematium MacLeay, Neateuchus Gillet, Neomnematium Janssens and Neopachysoma Ferreira, remain synonyms. A single origin of flightlessness is supported with the subgenus Pachysoma, the most derived lineage in this clade. Rolling dung balls backwards is the ancestral behaviour and predominant mode of food relocation in Scarabaeini, although tunnelling, forward pushing, and carrying also are utilized by some lineages. Pushing food has evolved independently in Sceliages species and S. galenus (Westwood) and a novel mode of forward food relocation by dragging evolved in the subgenus Pachysoma. Feeding on wet dung is the plesiomorphic condition and maintained by the majority of species in the tribe. The most unusual feeding behaviours in the tribe are represented by the obligate millipede‐feeding species of Sceliages and the dry dung pellet and/or detritus feeders of Pachysoma.

Los Escarabeidos y Geotrúpidos de la Comunidad de Madrid: lista de especies, distribución geográfica y patrones de diversidad (Coleoptera, Scarabaeoidea, Scarabaeidae y Geotrupidae)

“…Los trabajos filogenéticos mas recientes, basados tanto en datos morfológicos (Philips et al, 2004a;Forgie et al, 2005) como moleculares (Villalba et al, 2002), confirman que la división clásica de esta familia en Scarabaeinae y Coprinae, basada principalmente en el tipo de nidificación y en el manejo del excremen-Tabla 2.-Principales datos de las 42 especies de la familia Scarabaeidae presentes en la Comunidad de Madrid. Número de registros (R), número de individuos (N), número de cuadrículas UTM de 100 km 2 de la región de estudio (C) y de 2500 km 2 la Península Ibérica (CIb) en las que aparece cada especie.…”

Status changes, new synonymies, key and descriptions of seven new species in the subgenus Scarabaeus (Scarabaeolus) Balthasar 1965 (Coleoptera: Scarabaeidae: Scarabaeinae)

“…Forgie et al (2005) proposed to move it to Scarabaeus (Scarabaeus) as it consistently falls outside their "Scarabaeolus clades".…”

“…Recently Pachysoma was reduced to subgeneric status within Scarabaeus by Harrison et al (2003), listed under Scarabaeus by Zidek & Pokorny (2004), but subsequently revalidated (Forgie et al 2006). Other recent reductions of long-established Afrotropical genera (Kheper, Sceliages) to subgeneric status within Scarabaeus, (Forgie et al 2005) may, like Pachysoma, also deserve review and revalidation as full genera on morphological, molecular and behavioral grounds. This applies particularly to Sceliages, which shows modification of the clypeal margin associated with the specialized use of millipede gut contents for breeding (Forgie et al 2002).…”

“…Furthermore, one observation of a Scarabaeus (Scarabaeolus) bohemani (Harold 1868) female following a male rolling a ball (Tribe 1976) suggests that members of the subgenus may also show behavioral differences to Scarabaeus (Scarabaeus) species in which the female clings to the side of a ball being rolled by the male (Sato 1998). In addition, the preferred morphological phylogenetic analysis of Afrotropical Scarabaeini (Forgie et al 2005) divides the tribal members into a number of clades, one of which includes Scarabaeolus species. However, in view of the controversy, comprehensive molecular analysis would be desirable to support the higher classification of the tribe.…”

Origin of Madagascan Scarabaeini dung beetles (Coleoptera: Scarabaeidae): dispersal from Africa

“…There is little doubt that the tribe is monophyletic (Philips et al 2004;Monaghan et al 2007) and phylogenetic analysis of the tribe provided strong support for monophyly of the genera (Forgie et al 2005(Forgie et al , 2006. Scarabaeus has been divided into four morphologically, behaviourally and phylogenetically distinct subgenera, the nominate subgenera, Scarabaeus s.…”