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.

The enigmatic dung beetle genus Stiptopodius Harold, 1871 (Coleoptera: Scarabaeidae: Scarabaeinae) revisited: new species descriptions from southern Africa.

It is proposed that the genus Stiptopodius Harold, 1871 should be subdivided into two species groups based on the shape of the lateral edge of the pronotum. Three new species are described from southern Africa: Stiptopodius peringueyi Daniel Deschodt, new species, Stiptopodius savuti Daniel Deschodt, new species, and Stiptopodius muellerae Daniel Deschodt, new species. Stiptopodius savuti new species is the first record of the genus from Botswana. A distribution map for the new taxa is provided. The identification key to the known species of Stiptopodius is updated.

New southern African species and a revalidation in the dung beetle genus Gyronotus van Lansberge, 1874 (Scarabaeidae: Scarabaeinae) with an updated key.

Three new species are described in the genus Gyronotus van Lansberge, 1874 (Scarabaeidae: Scarabaeinae), from upland grasslands of South Africa. They are Gyronotus dracomontanus Deschodt Davis, new species, Gyronotus ovalis Deschodt Davis, new species and Gyronotus kearneyorum Deschodt Davis, new species. The South African coastal forest species, Gyronotus marginatus Péringuey, 1888, status revised, is removed from synonymy with Gyronotus pumilus (Boheman, 1857) and revalidated at species level. An updated key to all South African and eSwatini species is provided.

A beautiful new species of Hyalonthophagus Palestrini amp; Giacone, 1988 (Coleoptera: Scarabaeidae: Scarabaeinae), from the Northern Cape Province, South Africa.

Hyalonthophagus Palestrini Giacone, 1988, from Afrotropical savanna, is raised to generic status having been originally described as a subgenus within the genus Onthophagus Latreille, 1802. Hyalonthophagus pulcher Deschodt Davis, new species is described from the Northern Cape, South Africa. Maps are provided to show (1) the locally-restricted distribution pattern of the new species isolated from adjoining savanna in warm, dry Karoo climate to the south of the Orange River, and, (2) the disjunct distribution pattern in relation to those of three other southern African species centred in savanna to the northeast beyond areas with unsuitable climatic or ecological conditions. An assessment of the conservation status for Hyalonthophagus pulcher Deschodt Davis, new species is also provided.

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.

The endemic Southern African genus Epirinus Dejean, 1833 (Coleoptera: Scarabaeidae: Scarabaeinae) revisited: five new species and two new synonyms.

Five new species of Epirinus Dejean, 1833 were discovered during databasing of dung beetle collections. These are described here, as Epirinus inparrugosus Deschodt Davis, new species, Epirinus jacobsae Deschodt Davis, new species, Epirinus muellerae Deschodt Davis, new species, Epirinus pseudorelictus Deschodt Davis, new species and Epirinus schoolmeestersi Deschodt Davis, new species. Using comparisons based on morphology, we also studied the taxonomic status of five morphologically-allied, allopatric populations of Epirinus from isolated forest patches in northeast KwaZulu-Natal, South Africa, and eastern eSwatini (formerly Swaziland). On the basis of these comparisons we synonymize Epirinus hluhluwensis Medina Scholtz, 2005, and Epirinus ngomae Medina Scholtz, 2005, with Epirinus davisi Scholtz Howden, 1987. A distribution map is provided for all of these species. Furthermore, the first record for the genus in Namibia is reported and an updated key is provided for all known species of the genus.

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.

Two new species of Garreta Janssens, 1940 (Coleoptera: Scarabaeidae: Scarabaeinae) from Southern Africa.

Two new, putative, closely-related species of dung beetles are described in the genus Garreta Janssens, 1940 (tribe Gymnopleurini). Garreta australugens new species, is known from various dung types in the southeast lowlands of Africa (validated for South Africa, Mozambique and Zimbabwe with a probable record from Botswana). It is, here, separated at species level from its putative closest relative, Garreta lugens (Fairmaire, 1891), recorded from the lowlands of northeast Africa (validated for Kenya with additional records from Ethiopia, Somalia and Tanzania). These two taxa were formerly considered to comprise a single species. All specimens in the type series of Garreta namalugens new species, were recorded at low altitude in arid, rocky mountains from west central to northwest Namibia, mostly on dung in communal middens of the Rock Hyrax (Procavia capensis (Pallas, 1766)).

New deltochiline (Coleoptera: Scarabaeidae: Scarabaeinae) taxa associated with hyrax dung in arid south western Namibia.

A survey of rock hyrax dung middens along the arid escarpment of southwest Namibia has filled a gap in the range shown by members of the Byrrhidium group of flightless dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) belonging to the tribe Deltochilini. The survey discovered a new genus and species, Ausmontins jacobsi new genus and species, and a further four new species in two previously described genera, Namakwanus kamfferi new species, Namakwanus minutus new species, Namaphilus nubibmontanus new species and Namaphilus tirasmontanus new species. They are described and illustrated in the present paper. This addition of new taxa raises the total complement of the Byrrhidium group to seven genera and 19 species. An updated key to all the known species is also provided. It is probable that further taxa await discovery.

Digitonthophagus gazella auctorum: an unfortunate case of mistaken identity for a widely introduced species (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini).

At risk of committing entomological heresy, we question the identity of a dung-burying beetle species that originates from Africa and has been introduced first into Hawaii and subsequently to Australasia, North America, and South America (Fincher 1986; Edwards 2007; Noriega et al. 2010) for pasture improvement and biological control of dung-breeding flies (Waterhouse 1974; Bornemissza 1979).  Under the name Onthophagus gazella (Fabricius 1787), it was the first species selected for introduction into Australia by the CSIRO Dung Beetle Project (Bornemissza 1976; Edwards 2007). Firstly, in 1968, a "tropical strain" was introduced from Hawaii where it had become established after introduction from Zimbabwe in 1957 (Markin & Yoshioka 1998). Later, after establishment of the CSIRO Dung Beetle Research Unit in Pretoria in 1970, a "cold" or "even rainfall strain" was introduced into Australia directly from South Africa (Bornemissza 1976) (even rainfall region = south coast of Eastern Cape). The species was subsequently introduced into the southern continental United States of America (Victoria County, Texas) from Hawaii (Montes de Oca & Halffter 1998) then elsewhere into southeastern and southwestern states from Hawaii and breeding colonies from Australia (Anderson & Loomis 1978). It has since expanded its range through Mexico, Central America, and the Caribbean to coastal Colombia (Kohlmann 1994; Noriega 2002; Noriega et al. 2006, 2011). Expansion of its range within central southern South America (Noriega et al. 2010) has been assisted by introductions into Brazil from the United States of America since the 1980s (Bianchin et al. 1998), and others into Venezuela and Chile (Vidaurre et al. 2008). More recently, it has been introduced into quarantine and field trials in New Zealand (Forgie et al. 2013) using individuals originating from the south coast of the Eastern Cape and Northwest Province of South Africa (S. Forgie, personal communication).

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.

New replacement name for the species Scarabaeus (Scarabaeolus) nitidus Davis & Deschodt, 2015 (Coleoptera: Scarabaeidae: Scarabaeinae)--not an American pest.

In their review of the subgenus Scarabaeus (Scarabaeolus) Balthasar, 1965 (Coleoptera: Scarabaeidae: Scarabaeinae), Deschodt et al. (2015) described seven new species from Africa. One of these was Scarabaeus (Scarabaeolus) nitidus Davis & Deschodt, known by only twelve specimens from Botswana. This new name is preoccupied by Scarabaeus nitidus Linnaeus, 1758, a fact that was overlooked. The latter species has been known as Cotinis nitida (Linnaeus) since Burmeister (1842) placed the species in his new genus Cotinis. It belongs to the subfamily Cetoniinae of Scarabaeidae, is called the Green June Beetle, and is a common pest species throughout most of the eastern United States (Goodrich 1966; Woodruff 2008).

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.

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.

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

Tswalu Kalahari Reserve is a private game reserve covering 1,020 km(2) 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.

Discontinuous gas exchange and the significance of respiratory water loss in Scarabaeine beetles.

Respiratory water loss in insects is a controversial topic. Whilst earlier studies considered respiratory transpiration a significant component of overall water loss, to the extent that it was thought to be responsible not only for the evolution of discontinuous gas exchange cycles (DGCs) but also for variation in DGC patterns, later work repeatedly questioned its importance. In particular, investigations of the proportional contribution of respiratory transpiration to total water loss in species showing DGCs suggested that respiratory transpiration was unlikely to be important in these species. In turn, these studies have been criticized on analytical grounds. In this study we investigated variation in cuticular and respiratory water loss rates in five Scarabaeus dung beetle species, all of which show discontinuous gas exchange cycles, to ascertain the significance of respiratory water loss using modern analytical techniques. In particular, we determined whether there is variation in water loss rates amongst these beetles, whether both respiratory and cuticular water loss rates contribute significantly to variation in the former, and whether metabolic rate variation and variation in the duration of the DGC periods contribute significantly to variation in respiratory water loss rate. Total water loss rate varied such that species from arid areas had the lowest rates of water loss, and both cuticular and spiracular transpiration contributed significantly to variation in overall water loss rate. Moreover, variation in metabolic rate and in the duration of the DGC periods contributed significantly to variation in respiratory water loss rate. By contrast, examination of proportional water loss revealed little other than that it varies between 6.5% and 21%, depending on the species and the temperature at which it was examined. Cuticular water loss scaled as mass(0.721), but did not differ from that expected from geometric considerations alone. By contrast, respiratory water loss scaled as mass(0.531), suggesting that gas exchange takes place by diffusion and convection. Our results provide direct evidence that respiratory water loss forms a significant component of water balance, and that changes in both metabolic rate and DGC characteristics contribute to modulation of respiratory water loss.