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.

Phylogeny and biogeography of southern African spoon-winged lacewings (Neuroptera: Nemopteridae: Nemopterinae).

Nemopteridae are a charismatic family of lacewings characterised by uniquely extended hind wings. They are an ancient widespread group in the drier regions of the world. The family comprises two subfamilies, Crocinae (thread-wings) and Nemopterinae (spoon- and ribbon-wings). The present distribution of the family has been largely influenced by the vicariant events of plate tectonics, resulting in relict populations in some parts of the world and extensive evolutionary radiations in others, particularly southern Africa where the vast majority of the species are endemic to the Western and Northern Cape Provinces of South Africa. This study aimed to establish the validity of the 11 currently recognised genera and infer their biogeographic history using molecular sequence data from four gene regions. The hypothesis that the Cape nemopterines co-evolved with certain taxa in the Cape Floristic Region was also tested. Phylogenetic analysis supports seven of the 11 currently recognised genera. The crown age of the Nemopterinae is estimated to be at ca. 145.6 Mya, indicating that the group has been present since the late Jurassic. Most of the genera appear to have diversified during the middle Eocene and into the middle Miocene (ca. 44-11 Mya) with recent rapid radiation of several of the genera occurring during the late Miocene (ca. 6-4.5 Mya). While these data support an initial radiation with the Rushioideae (Aizoaceae) it is recommended that further study including observations and gut content be carried out.

Morphology and terminology of dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) male genitalia.

The external and internal male genitalia of 327 species of 11 tribes of the subfamily Scarabaeinae, including species of Deltochilini, Scarabaeini, Gymnopleurini, Ateuchini, and Coprini, among others, were examined. Descriptions of the variations in the genital segment, the aedeagus, the internal sac, and its sclerites and raspules are presented. An exhaustive comparison of structures, names, and terminology used in literature for Scarabaeinae male genitalia are discussed. The internal sac of the aedeagus is divided in areas for an easer comparison of its internal structures; basal, submedial, medial, and apical areas are described in detail and compared. The variation of apical and medial sclerites, as well as the raspules of the submedial area, are described and compared in detail among all the taxa studied.

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.

Did dung beetles arise in Africa? A phylogenetic hypothesis based on five gene regions.

Scarabaeinae dung beetle phylogenetics are poorly understood, although recent phylogenetic reconstruction based on morphology and molecular analyses are congruent on the hypothesis that the oldest Gondwana tribes are polyphyletic and that representatives of the ancestral groups are found in Africa. We present a molecular phylogeny of the African representatives of the two oldest tribes, Canthonini (the putative ancestor of all "rollers") and Dichotomiini (thought to represent the ancestral "tunnellers"), based on partial sequence data from two mitochondrial and three nuclear genes, extracted from eight of the nine dichotomiine genera and 17 of the 23 genera of Canthonini. Three well-supported lineages were consistently obtained. Divergence times estimated the origin of the two tribes at around 56 million years ago (MYA) with the splits of the three dung beetle lineages being estimated to have taken place between 40 and 34MYA. The ages of these splits and subsequent radiation of the modern dung beetle groups concur with those predicted by the fossil record and coincide with the proposed age of radiation of the large mammal groups with whose dung most African dung beetles are still associated. Dispersal of dung beetle groups from Africa is proposed as a biogeographic model, and evidence is presented that dung beetles disperse quickly and widely across continents, and even oceans.

Bearing selection in ball-rolling dung beetles: is it constant?

Ball rolling in dung beetles is thought to have evolved as a means to escape intense inter- and intra-specific competition at the dung pile. Accordingly, dung beetles typically roll along a straight-line path away from the pile, this being the most effective escape strategy for transporting dung to a suitable burial site. In this study, we investigate how individual diurnal dung beetles, Scarabaeus (Kheper) nigroaeneus, select the compass bearing of their straight-line rolls. In particular, we examine whether roll bearings are constant with respect to geographic cues, celestial cues, or other environmental cues (such as wind direction). Our results reveal that the roll bearings taken by individual beetles are not constant with respect to geographic or celestial references. Environmental cues appear to have some influence over bearing selection, although the relationship is not strong. Furthermore, the variance in roll bearing that we observe is not affected by the presence or absence of other beetles. Thus, rather than being constant for individual beetles, bearing selection varies each time a beetle makes a ball and rolls it away from the dung pile. This strategy allows beetles to make an efficient escape from the dung pile while minimizing the chance of encountering competition.

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.

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).

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.

Intraspecific patterns of mitochondrial variation in natural population fragments of a localized desert dung beetle species, Pachysoma gariepinum (Coleoptera: Scarabaeidae).

Phylogenetic, population, and coalescent methods were used to examine the genetic structuring of Pachysoma gariepinum, a flightless dung beetle species endemic to the arid west coast of southern Africa that exhibits interrupted south to north morphological clinal variation along a distributional gradient. Mitochondrial cytochrome oxidase I sequence data of 67 individuals from 5 localities revealed the presence of 3 geographically distinct evolutionary lineages (with an overall nucleotide divergence of 5.7% and a per-locality divergence of 1.9-3.8%) which display significant levels of genetic structuring. The separation of the lineages was estimated to have occurred between 2.2 and 5.7 million years ago--which is the late Miocene, early Plio-Pleistocene era--possibly in response to the ebb and flow of the Orange and Holgat River systems as well as the interactions between the moving and stable sand dune systems. Moreover the species' current range appears to have been influenced by the formation of advective fog resulting in a constant source of water in an area with low precipitation thereby allowing for the beetles to radiate to areas that were previously inhospitable. Fu's F-statistics and population parameters based on recent mutations indicated that little to no recent population growth has occurred. This together with changing anthropogenic factors and the recovery of 3 geographically discrete management units, points to a need for census data in order to monitor and conserve the genetic diversity of this species.

Trophic associations of a dung beetle assemblage (Scarabaeidae: Scarabaeinae) in a woodland savanna of Botswana.

Species richness and abundance of dung beetles were assessed across a range of bait types that acted as surrogates for the food resources available in Chobe National Park, Botswana. These bait types were comprised of the dung of pig (omnivore), cattle (ruminant herbivore dropping fine-fiberd pads), sheep (pellet-dropping ruminant herbivore), and elephant (monogastric, nonruminant herbivore producing coarse-fibered droppings), and chicken livers (carrion). Species richness was similar between traps baited with pig, cattle, and elephant dung but was relatively lower in those baited with sheep dung and carrion. In traps baited with pig dung, abundance was relatively greater than in all other bait types. A cluster analysis of species abundance distributions for the 30 most abundant species identified four different patterns of bait type association at a 60% level of similarity. All but 1 of the 15 species in cluster A were attracted primarily to the dung of omnivores and pad-dropping ruminant herbivores (pig and cattle). All seven species of cluster B were attracted primarily to coarse-fibered, nonruminant herbivore dung (elephant). All four species of cluster C were primarily carrion and pig dung associated, whereas all four species of cluster D were carrion specialists. In conclusion, the most abundant species were attracted to all bait types, but most species were largely specialized to different dung types or carrion, with dung attracting the majority of the fauna in terms of both species richness and abundance.

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.

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.

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.

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.

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.

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.].