The impact of dung beetles on the free-living stages of ruminant parasites in faeces and their role as biological control agents in grazing livestock.

Dung beetles provide a variety of ecosystem services in both natural and farmed landscapes. Amongst these services, reductions in the abundance of the free-living stages of pests and parasites that develop in faeces is considered to be of great importance. There is evidence from Australia that enhanced dung beetle populations can reduce populations of pest fly species, particularly the bush fly, however, there is little empirical evidence for reductions in the incidence and impact of nematode parasitism in grazing ruminants. There are two main pathways whereby beetles can disrupt worm life-cycles: predaceous species that feed on eggs or larvae can directly reduce populations in dung whereas coprophagous species can affect parasite development, survival and translocation by altering the location, microclimate and infrastructure of dung deposits. In addition, predaceous mites that are phoretic on dung beetles, can also prey on larval stages in the faeces. To date, reductions in both larval survival and the acquisition of gastrointestinal nematode burdens in ruminants on pasture has been reported only in association with the activity of large tunnelers that bury dung 15 cm or more below ground. The activity of dwellers, rollers and shallow tunnelers can either limit or enhance larval development and translocation, depending on the influence of other factors, notably rainfall. Currently, the scientific evidence for dung beetles playing a major role in the control of gastrointestinal nematodes in domestic ruminants is very limited and may have been overestimated in assessments of their ecosystem services.

Dung removal increases under higher dung beetle functional diversity regardless of grazing intensification.

Dung removal by macrofauna such as dung beetles is an important process for nutrient cycling in pasturelands. Intensification of farming practices generally reduces species and functional diversity of terrestrial invertebrates, which may negatively affect ecosystem services. Here, we investigate the effects of cattle-grazing intensification on dung removal by dung beetles in field experiments replicated in 38 pastures around the world. Within each study site, we measured dung removal in pastures managed with low- and high-intensity regimes to assess between-regime differences in dung beetle diversity and dung removal, whilst also considering climate and regional variations. The impacts of intensification were heterogeneous, either diminishing or increasing dung beetle species richness, functional diversity, and dung removal rates. The effects of beetle diversity on dung removal were more variable across sites than within sites. Dung removal increased with species richness across sites, while functional diversity consistently enhanced dung removal within sites, independently of cattle grazing intensity or climate. Our findings indicate that, despite intensified cattle stocking rates, ecosystem services related to decomposition and nutrient cycling can be maintained when a functionally diverse dung beetle community inhabits the human-modified landscape.

Upsa centennial, a new genus and species (Coleoptera: Scarabaeidae: Scarabaeinae: Endroedyolini) from the Southern Mistbelt forest in South Africa: celebrating 100 years of entomology at the University of Pretoria.

Upsa centennial Deschodt, Sole Scholtz, new genus and species is described, celebrating a century of entomology at the University of Pretoria. This new genus in the dung beetle tribe Endroedyolini (Scarabaeidae: Scarabaeinae) occurs in the Albany Coastal Mistbelt forest in the Eastern Cape Province in South Africa. A distribution map and key for all genera and species in the tribe are provided.[Genus Zoobank url: urn:lsid:zoobank.org:act:92A64613-A8CE-48DF-84D9-7450EA1D40AA].

A new Phoberus MacLeay species (Coleoptera: Scarabaeoidea: Trogidae) from the Maloti-Drakensberg Mountains of southern Africa and the Phoberus caffer species group reviewed.

Phoberus ntlenyanae Strümpher, new species is described from Lesotho and South Africa and is placed in the southern African Phoberus caffer group of species. The P. caffer species group is redefined to include the new species. One taxonomic status change is proposed: Phoberus lilianae (Scholtz, 1980) from the Eastern Highlands of Zimbabwe and the eastern escarpment of South Africa is elevated to species level. Phoberus caffer (Harold, 1872) is recorded for the first time from Eastern Cape and North West Province of South Africa. A key is provided for the identification of the members of the group. Photographs illustrate the species, and a map shows their geographical distributions. [Zoobank URL: urn:lsid:zoobank.org:act:0CB1BAD0-F9C2-4303-B746-487BCADF0D07].

Defining new dung beetle tribes to resolve discrepancies between phylogeny and tribal classification in the subfamily Scarabaeinae (Coleoptera: Scarabaeidae).

The monophyletic, dung beetle subfamily, Scarabaeinae, has a long history of subdivision into tribes and subtribes represented by groups of genera unified by designated characters. Various tribal names have been proposed although only 11 or 12 were considered valid in recent reviews. Three of these 12 tribes have now been invalidated due to extensive polyphyly indicated by both morphological and molecular phylogenies. Although partial revision of tribal classification was proposed in the most recent global molecular phylogeny of the subfamily, this was primarily limited to redefinition of membership or provision of support for pre-existing tribes. Complete revision was not attempted as the phylogeny examined interrelationships between only ~52% of the valid described genera. It was also hampered by inconsistencies in topology using different analytical methods, weakened by limited bootstrap support for many basal nodes, as well predicting several phylogenetic relationships that seemed anomalous with regards to biogeographical hypotheses. Nevertheless, using this phylogeny, we propose further revision of tribal classification in a clade of primarily African genera included within a group defined as "basal Scarabaeinae". Using three different methods, the sister lineages of this clade were consistently recovered, mostly with strong bootstrap support for their molecular relatedness. Therefore, we provide a number of defining morphological characters to support their proposed designation as three new tribes: Byrrhidiini, Endroedyolini and Odontolomini. We also discuss approaches to a full tribal revision as well as the implications for historical biogeography of the Scarabaeinae.

Redescription of the holotype of Bostrichoclerus bicornus Van Dyke, 1938, a rare species of the New World Tillinae Fauna (Coleoptera: Cleridae), with some taxonomic notes.

Bostrichoclerus bicornus Van Dyke is known from southwest United States and northwest Mexico. To date, only two specimens have been captured: the holotype, collected on Isla Angel de la Guarda, in the Gulf of California, Mexico, and a second individual collected in San Bernardino County, California, United States. The original description of B. bicornus is brief and lacks any images. Considering its rarity, we present the redescription of this species based on the examination of the holotype and compare this taxon to similar genera of New World Tillinae. Images of the holotype and the Bostrichoclerus specimen collected in southern California are given. We conclude that B. bicornus is undoubtedly a member of the subfamily Tillinae with unclear intergeneric relations in the group.

Grebennikovius, a new genus to accommodate Epactoides basilewskyi (Balthasar, 1960) (Coleoptera: Scarabaeidae: Deltochilini).

Phacosoma basilewskyi Balthasar, 1960 was described for a peculiar East African deltochiline dung beetle species. Years later, Paulian (1975) erected the deltochiline genus Madaphacosoma Paulian, 1975 for two new Malagasy species, namely Madaphacosoma humberti Paulian, 1975, as the type species, and M. betschi Paulian, 1975. Phacosoma basilewskyi was transferred to Madaphacosoma in that same publication. Later, Paulian (1991) added Madaphacosoma major Paulian, 1991 to the genus for a total of three Madagascan and one mainland African species. Recently, Madaphacosoma together with three other genera occurring in Madagascar, namely Aleiantus Olsoufieff, 1947, Phacosomoides Martinez Pereira, 1959 and Sikorantus Paulian, 1976, were synonymized with Epactoides Olsoufieff, 1947 by Wirta Montreuil (2008). Although Madaphacosoma basilewskyi (Balthasar, 1960) is not specifically mentioned as a new combination with three other species, M. major, M. humberti and M. betschi, catalogued on pages 661-662 in Wirta Montreuil (2008), and the genus is treated as valid by Tarasov Dimitrov (2016) and Tarasov (2017), it is currently lumped with the other Malagasy species in the genus Epactoides. The current accepted name for the species is thus Epactoides basilewskyi (Balthasar, 1960).

An update to the taxonomy of the genus Macroderes Westwood 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) with descriptions of new species from South Africa.

The genus Macroderes Westwood, 1842 (Scarabaeidae: Scarabaeinae, tribe incertae sedis) is practically endemic to South Africa with only one species that is found little north in southern Namibia. The ranges of the species are limited to the winter-fall and bimodal biomes of South Africa comprising the regions of Richtersveld as extreme points of its northerly distribution, and extending to the south through Namaqualand to Cape Agulhas. The taxonomy of the genus is revised. Fourteen valid species are recognised and three others are of doubtful validity. Seven new species are described, these are: M. cederbergensis Abdalla Deschodt, new species, M. tortuosus Abdalla Scholtz, new species, M. gifboomi Abdalla Scholtz, new species, M. leipoldti Abdalla Deschodt, new species, M. oreatus Abdalla Deschodt, new species, M. porselinus Abdalla, new species, and M. soleiana Abdalla Deschodt, new species. Macroderes nitidus Harold, 1877 is redescribed and its lectotype is designated. The diagnostic characters as well as an updated geographic distribution of each species are summarised. An updated key of the genus, photographs of habitus, sclerites of internal sac, pronotum, pronotal punctures and elytra also are provided.

Multi-locus phylogeny of southern African Acontias aurantiacus (Peters) subspecies (Scincidae: Acontinae) confirms the presence of three genetically, geographically and morphologically discrete taxa.

Acontias aurantiacus (Peters) presently consists of five subspecies that are largely restricted to the coastal regions and offshore islands of Mozambique and northern South Africa, with subspecies being defined on the basis of morphological differences and/or parapatric distributions. In this study one nuclear (RAG1) and two mitochondrial (cyt b and 16S) genes of 30 individuals sampled from four Mozambican and five South African localities were used to assess the taxonomic relationships of three mainland A. aurantiacus subspecies. Our analyses suggest that A. aurantiacus is a species complex that is sister to A. bicolor (Hewitt), A. cregoi (Boulenger) and A. rieppeli (Sternfeld). Our molecular results support other lines of evidence (consistent morphological differences and parapatric distributions between the subspecies) in suggesting that each of these subspecies are taxonomically distinct. In light of this, and supported by the genetic distances between these subspecies which exceed those of other recognised Acontias species, we propose elevating A. fitzsimonsi (Broadley) and A. parietalis (Broadley) to specific status. Our results further suggest that the melanistic and typically-coloured forms of each taxon are nested within the same clades, supporting previous contentions that melanistic individuals are not cryptic taxa.

Five new species of Cymatodera Gray, 1832 (Coleoptera: Cleridae: Tillinae) from Mexico and Central America.

Five new species of Cymatodera from Mexico, Guatemala, Honduras, Costa Rica and Panama are described here based on adult male and female specimens: Cymatodera humeralis sp. n; Cymatodera lorenae sp. n.; Cymatodera crassa sp. n.; Cymatodera depressa sp. n; and Cymatodera nigrofasciata sp. n. Male genitalia and other characters of taxonomic value are used for diagnostic purposes.

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.

Correction: The Gut Microbiomes of Two Pachysoma MacLeay Desert Dung Beetle Species (Coleoptera: Scarabaeidae: Scarabaeinae) Feeding on Different Diets.

[This corrects the article DOI: 10.1371/journal.pone.0161118.].

The Gut Microbiomes of Two Pachysoma MacLeay Desert Dung Beetle Species (Coleoptera: Scarabaeidae: Scarabaeinae) Feeding on Different Diets.

Micro-organisms inhabiting animal guts benefit from a protected and nutrient-rich environment while assisting the host with digestion and nutrition. In this study we compare, for the first time, the bacterial and fungal gut communities of two species of the small desert dung beetle genus Pachysoma feeding on different diets: the detritivorous P. endroedyi and the dry-dung-feeding P. striatum. Whole-gut microbial communities from 5 individuals of each species were assessed using 454 pyrosequencing of the bacterial 16S rRNA gene and fungal ITS gene regions. The two bacterial communities were significantly different, with only 3.7% of operational taxonomic units shared, and displayed intra-specific variation. The number of bacterial phyla present within the guts of P. endroedyi and P. striatum individuals ranged from 6-11 and 4-7, respectively. Fungal phylotypes could only be detected within the gut of P. striatum. Although the role of host phylogeny in Pachysoma microbiome assembly remains unknown, evidence presented in this study suggests that host diet may be a deterministic factor.

Three new Sisyphus species (Coleoptera: Scarabaeidae: Scarabaeinae) from shaded vegetation in southern Africa.

Descriptions are provided for three new shade-associated species in the dung beetle genus, Sisyphus Latreille, 1807: Sisyphus umbraphilus sp. nov., Sisyphus oralensis sp. nov., and Sisyphus neobornemisszanus sp. nov. All three species are illustrated by photographs of habitus, diagnostic features, and male aedeagi. A map is provided showing their geographical distributions.

A new monotypic genus, a species synonymy and nomenclatural corrections in the arid-adapted Canthonini (Scarabaeidae, Scarabaeinae) from the Succulent Karoo Biome of south-western Africa.

Although it has been demonstrated that the tribe Canthonini (Scarabaeidae: Scarabaeinae) is polyphyletic, those canthonine genera occurring in arid south-western Africa apparently form a monophyletic group. In this paper we add one new monotypic genus, Drogo stalsi gen. et. sp. n. to this group, and synonymize another member, Byrrhidium namaquense Scholtz and Howden, 1987 syn. n. with Byrrhidium ovale Harold, 1869. We correct the spelling of two species names that were not in agreement with the gender rule. A phylogenetic analysis based on morphology supports the erection of Drogo gen. n. An updated key to the genera and species of the Byrrhidium group is presented.

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.

A new synonymy in the fidius group of Copris Müller 1764 (Coleoptera: Scarabaeidae: Scarabaeinae) and a new species from the highland grasslands of South Africa.

The fidius group constitutes the basal clade in a phylogeny of the Afrotropical members of the genus, Copris Müller, 1764. In this paper we describe Copris crassus Deschodt and Davis as a new species in this group. We also demonstrate that the distributional function between two body dimensions (lateral horn length and mid-line elytron length) differs between Copris fidius (Olivier 1789) and C. crassus. Copris bihamatus Balthasar, 1965 is also a member of the fidius group. Nguyen-Phung (1988) suspected that it was a synonym of C. fidius (Olivier) so we compare specimens of similar body size and use the distributional function between body dimensions to demonstrate that C. bihamatus is indeed a synonym of C. fidius. We provide a key for all known valid species in the fidius group, except C. serius Nguyen-Phung, 1987.

A review on the effect of macrocyclic lactones on dung-dwelling insects: Toxicity of macrocyclic lactones to dung beetles.

Avermectins and milbemycins are commonly used in agro-ecosystems for the control of parasites in domestic livestock. As integral members of agro-ecosystems with importance in maintaining pasture health through dung burial behaviour, dung beetles are an excellent nontarget bio-indicator taxon for examining potential detrimental effects of pesticide application. The current review focuses on the relative toxicity of four different anthelmintics (ivermectin, eprinomectin, doramectin and moxidectin) in dung residues using dung beetles as a bioindicator species. One of the implications of this review is that there could be an effect that extends to the entire natural assemblage of insects inhabiting and feeding on the dung of cattle treated with avermectin or milbemycin products. Over time, reduced reproductive rate would result in decreased dung beetle populations and ultimately, a decrease in the rate of dung degradation and dung burial.

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

Distribution of the subgenus Scarabaeus (Scarabaeolus) Balthasar 1965 (tribe Scarabaeini) is centred on southern and south central Africa with only three out of 27 species recorded from West and northeast Africa. After taxonomic corrections and descriptions of seven new southern African species this somewhat controversial subgenus now comprises 33 valid species of which one is flightless. In this paper, Scarabaeus (Scarabaeolus) vansoni Ferreira 1958 syn. nov. is synonymized with Scarabaeus (Scarabaeolus) lucidulus (Boheman 1860) and Scarabaeus (Scarabaeolus) xavieri Ferreira 1968 syn. nov. is synonymized with Scarabaeus (Scarabaeolus) andreaei zur Strassen 1963. Scarabaeus (Scarabaeolus) reichei Waterhouse 1890 stat. rev. is removed from synonymy with Scarabaeus (Scarabaeolus) canaliculatus Fairmaire, 1888 and reinstated as a valid species. Distribution maps for S. (S.) reichei, S. (S.) canaliculatus and a third close relative, Scarabaeus (Scarabaeolus) fritschi Harold 1868 are provided. The seven new species comprise: Scarabaeus (Scarabaeolus) soutpansbergensis Deschodt and Davis spec. nov., Scarabaeus (Scarabaeolus) megaparvulus Davis and Deschodt spec. nov., Scarabaeus (Scarabaeolus) niemandi Deschodt and Davis spec. nov., Scarabaeus (Scarabaeolus) carniphilus Davis and Deschodt spec. nov., Scarabaeus (Scarabaeolus) ermienae Deschodt and Davis spec. nov., Scarabaeus (Scarabaeolus) planipennis Davis and Deschodt spec. nov. and Scarabaeus (Scarabaeolus) nitidus Davis and Deschodt spec. nov. A key is provided for all the known winged species together with notes on some of the previously described species.

A new galloping gait in an insect.

An estimated three million insect species all walk using variations of the alternating tripod gait [1]. At any one time, these animals hold one stable triangle of legs steady while swinging the opposite triangle forward. Here, we report the discovery that three different flightless desert dung beetles use an additional gallop-like gait, which has never been described in any insect before. Like a bounding hare, the beetles propel their body forward by synchronously stepping with both middle legs and then both front legs. Surprisingly, this peculiar galloping gait delivers lower speeds than the alternating tripod gait. Why these beetles have shifted so radically away from the most widely used walking style on our planet is as yet unknown.