Werner P. Strümpher

Dung beetle assemblage structure in Tswalu Kalahari Reserve : responses to a mosaic of landscape types, vegetation communities, and dung types

ABSTRACT Tswalu Kalahari Reserve is a private game reserve covering 1,020 km2 in the Northern Cape, South Africa. It has been created from a number of reclaimed farms and restocked with large indigenous mammals. Two surveys were conducted to inventory the dung beetle fauna (Coleoptera: Scarabaeidae: Scarabaeinae) and determine their spatial patterns and food type associations. The spatial survey used pig dung—baited pitfall traps to examine dung beetle distribution across three main landscape types (plains, dunes, hills) comprising six principal vegetation communities. The food study examined their relative associations with carrion and four different dung types within a single vegetation community. A total of 70 species was recorded. Because the food association study was spatially restricted and conducted under drought conditions, abundance and species richness (47 species) were much lower than in the spatial study (64 species), which was conducted after substantial rainfall. Principal spatial differences in species abundance structure of assemblages were between the sandy southwest plains and dunes; the sandy northern dune fields and plains; and the rocky hills. Forty species analyzed in the food association study showed clear distributional biases to carrion or the dung of elephant (monogastric herbivore), pig (omnivore), cattle and sheep (ruminant herbivores), or pig and cattle. The results (1) show how dung beetle assemblage structure is locally diversified across the heterogeneous landscape of the reserve and (2) indicate how the different dung types dropped by a diverse assemblage of indigenous mammals may variously favor different species of dung beetles.

Phylogeny of the family Trogidae (Coleoptera : Scarabaeoidea) inferred from mitochondrial and nuclear ribosomal DNA sequence data

Trogidae constitute a monophyletic and biologically unique family within Scarabaeoidea, being the only keratinophagous group in the superfamily. Traditionally, the family has been divided into three distinctive genera, Polynoncus Burmeister, Omorgus Erichson and Trox Fabricius. Although the taxonomy of the group is relatively well studied, changes to the existing classification have recently been proposed and the family as currently constituted has not been subjected to phylogenetic analyses. Here we present a molecular phylogeny for this cosmopolitan family based on three partially sequenced gene regions: 16S rRNA, 18S rRNA and 28S rRNA (domain 2). Included in the analyses are representatives belonging to four of the five extant genera (and three of the four subgenera) from all major zoogeographic regions, representing about 20% of the known trogid species diversity in the family. Phylogenetic analyses performed included parsimony and Bayesian inference. We deduce their historical biogeography by using trogid fossils as calibration points for divergence estimates. Our analyses resolved relationships between and within genera and subgenera that are largely congruent with existing phylogeny hypotheses based on morphological data. We recovered four well‐supported radiations: Polynoncus, Omorgus, Holarctic Trox and African Phoberus MacLeay. On the basis of this study, it is proposed that taxonomic changes to the generic classification of the family be made. The subgenera Trox and Phoberus should be elevated to genera to include the Holarctic and all the Afrotropical species, respectively, and Afromorgus returned to subgeneric rank. Estimates of divergence time are consistent with a Pangaean origin of the family in the Early Jurassic. The subsequent diversification of the major lineages is largely attributed to the break‐up of Pangaea and Gondwana in the Middle Jurassic and early Late Cretaceous, respectively.

Description of a new, possibly extinct, species of flightless Trox F. (Coleoptera: Trogidae) from South Africa

A new species, which may already be extinct, of flightless Trox, T. youngai sp.n., is described based on specimens collected 30 years ago from a now severely transformed area in South Africa. It is one of seven members of the natalensis-group of Trox species. All have very localised distributions in small fragments of once considerably larger habitats along the greater eastern escarpment and the coastal edge of southern Africa. is description brings the total number of endemic flightless Trox in the region to 14. A key to the species in the natalensis-group is provided. Furthermore, pronotal and elytral features of diagnostic value are tabulated to aid in the identification of members of this group. All species are illustrated by photographs of habitus and male aedeagi. A map is provided showing the distributions for each of them.

New species and status changes of small flightless relictual Trox Fabricius from southern Africa (Coleoptera: Trogidae)

A new species of Trox, T. mozalae, from coastal sand forest in Mozambique, is described. T. natalensis quadricostatus Scholtz is elevated to species-level and T. natalensis schaborti Scholtz and T. elizabethae van der Merwe & Scholtz are synonymised with it. A key to the subgenus Phoberus MacLeay, to which most of the flightless species in Africa belong, is provided. All species incorporated into the key are illustrated by photographs of habitus and aedeagi. A map is provided of the distribution of each of them.

Overview and revision of the extant genera and subgenera of Trogidae (Coleoptera : Scarabaeoidea)

Extant genera and subgenera of the Trogidae (Coleoptera: Scarabaeoidea) are reviewed. Contemporary classifications of this family have been based exclusively on morphological characters. The first molecular phylogeny for the family recently provided strong support for the relationships between morphologically defined genera and subgenera. On the basis of morphological, molecular and biogeographical evidence, certain taxonomic changes to the genus-level classification of the family are now proposed. The family is confirmed as consisting of two subfamilies, Omorginae Nikolajev and Troginae MacLeay, the former with two genera, Omorgus Erichson and Polynoncus Burmeister, and the latter with two genera, Trox Fabricius and Phoberus MacLeay stat. rev. Phoberus is restored to generic rank to include all Afrotropical (including Madagascan endemic) species; Afromorgus is confirmed at subgeneric rank within the genus Omorgus ; and the monotypic Madagascan genus Madagatrox syn. n. is synonymised with Phoberus. The current synonymies of Pseudotrox Robinson (with Trox ), Chesas Burmeister, Lagopelus Burmeister and Megalotrox Preudhomme de Borre (all with Omorgus ) are all accepted to avoid creating speculative synonyms before definitive phylogenetic evidence is available. New combinations resulting from restoring Phoberus to a monophyletic genus are listed in Appendix A.

Taxonomy of Colophon Gray (Coleoptera: Lucanidae): new species and a status change.

Three new species of the Cape high-mountain stag beetle genus, Colophon Gray (Coleoptera: Lucanidae), from South Africa are described. They are C. deschodti new species, C. switalae new species, and C. struempheri new species. The new taxa fall within a species complex of geographically disjunct entities related to Colophon stokoei Barnard. Furthermore, the mitochondrial COI gene shows a high degree of sequence divergence, with pairwise genetic distances between the species ranging between 7.4-10.7%. The new species are illustrated by photographs. Colophon eastmani nagaii Mizukami is raised to species level on the basis of geographic range and molecular differences between it and the nominate subspecies. This brings the total number of described species in the genus to 21. An updated checklist of the South African species of Colophon is also provided.

Trogidae (Coleoptera: Scarabaeoidea) in forensic entomology: Occurrence of known and new species in Queensland, Australia

During studies of necrophagous insect succession in pig carcases that were used as surrogates for human corpses, Trogidae were found to be fairly common at the carcases at a study site in south‐eastern Queensland. During identification of the species, one of the regular visitors to the carcases was found to be undescribed. While comparing this new species, Omorgus (Omorgus) bachorum sp. nov., with material in the Queensland Museum, another new species, Omorgus (Omorgus) undaraensis sp. nov., was discovered, which we also describe. One new synonym, Omorgus incognitus Strümpher & Scholtz, 2011 syn. nov., is also proposed, bringing the total number of Australian species of Omorgus Erichson, 1847 to 57.

A new brachypterous species of Phoberus MacLeay (Coleoptera: Trogidae) and status changes of species in the East African P. nigrociliatus group

A new brachypterous species, Phoberus nyikanus Strümpher (Coleoptera: Trogidae), is described from Malawi. In addition, three status changes are proposed: Phoberus nigrociliatus (Kolbe) from the Ethiopian highlands, P. nyansanus (Haaf) from central East Africa, and P. nyassicus (Haaf) from southern Tanzanian mountains are elevated to species level. A key is provided to aid in the identification of members of the P. nigrociliatus group, and the species in the group are illustrated by photographs. This brings the total number of species in the genus to 44.

Allopatric speciation in the flightless Phoberus capensis (Coleoptera: Trogidae) group, with description of two new species

This project was supported by funding to Clarke H. Scholtz from the National Research Foundation of South Africa (NRF).