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The subfamily Scarabaeinae has been traditionally divided into tribes on the basis of morphological similarity between groups of genera or, even, dissimilarity shown by a single genus. Although various tribal units have been described over the past 220 years, they had been recently reduced to a maximum of only 12 through synonymies within some tribes. However, following the advent of morphological and molecular cladistic systematics, it is clear that there are many discrepancies between phylogeny and phenetic tribal classification. As a result the number of tribes has now been expanded to 20 by revalidating some tribes or describing new groupings although this revision is incomplete. Thus, this study provides a comprehensive review of the current status of dung beetle tribal classification with regards to validity in terms of monophyly versus polyphyly or paraphyly. We discuss the systematic position of tribes among Scarabaeinae and provide a summary of the generic and species structure for each tribe. Given the enduring polyphyly or paraphyly in several tribes, it is clear that further tribal units will need to be described or revalidated to resolve the discrepancies. Therefore, we discuss how to use multiple lines of evidence (molecular, morphological, biogeographical, and paleontological) to build the tree of life for dung beetles and consequently provide stability in the tribal classification of the lineage. [ABSTRACT FROM AUTHOR]

Dung beetles feed mainly on scarce and ephemeral mammalian faeces making them vulnerable to declines of mammal populations. Therefore, studying how diverse groups like the Scarabaeinae subfamily utilize alternative food sources, such as decomposing invertebrates, enhances our understanding of their distribution and resilience in defaunated areas. One poorly studied aspect of the feeding habits of dung beetles is the potential differences between the sexes and their predatory behaviour. We evaluated how the sex (male and female) of individuals of Chalconotus convexus affected individual resource attraction to different bait types. Since C. convexus is known to feed on faeces, carrion and dead millipedes, we also test whether it can predate live individuals of millipedes. We used a combination of two field samplings and one laboratory experiment, using pitfall traps baited with faeces, carrion, millipedes and millipede's defensive compounds. While the preference for millipede was not associated with sex, we found a significant relationship between faeces and carrion, where females of C. convexus are significantly less attracted to faeces when compared to males and also males are significantly less attracted to carrion when compared to females. Furthermore, we also show that while actively consuming dead millipedes, C. convexus is unable to predate on live individuals. Our study illustrates that the versatility of food sources may help to explain the high abundance and wide distribution of C. convexus in Africa. [ABSTRACT FROM AUTHOR]

Dysmorphocerinae is a subfamily of Cantharidae erected for a group of genera with a mainly gondwanan distribution whose adult forms could not be reliably assigned to any other subfamily. The systematic position and monophyly of Dysmorphocerinae remains questionable, as recent molecular and morphological studies have produced conflicting results. Despite the importance of immature morphology for characterising lineages of Cantharidae, so far, the larvae of only two dysmorphocerine species had been briefly described: Neoontelus sp., from New Zealand, and Afronycha picta (Wiedemann), from South Africa. Their morphologies considerably differ from one another, and the larvae cannot be readily attributed to any subfamily, as usually occurs with cantharid larvae. Here, we fully describe for the first time the larvae of Asilis Broun (New Zealand) and Plectonotum laterale Pic (Brazil) and redescribe Neoontelus Wittmer (New Zealand). We also diagnose larvae of Heteromastix Boheman (Australia) and A. picta. Dysmorphocerinae cannot be clearly diagnosed because each genus has a unique combination of features, though Neoontelus is the most divergent. We conclude that the Dysmorphocerinae may not be monophyletic with Plectonotum laterale, Asilis, Neoontelus, Heteromastix showing a closer relationship to Malthininae and Afronycha more aligned with Silinae or Cantharinae. The double gland openings present on the body of Neoontelus reported by Crowson (1972) are reinterpreted as a complex character involving a single posterior pore linked to a gland and an anterior sensillum that may serve as a trigger for the release of defensive chemicals. These are also reported in Asilis and Heteromastix and may be a potential synapomorphy for part of the Dysmorphocerinae. Neoontelus has a series of unique features, including a cotyliform glandular pore on abdominal segment IX. [ABSTRACT FROM AUTHOR]

Regional biogeographical patterns result from a complex combination across habitat, climate and environmental variables. Biotic and abiotic variables strongly influence the diversity and spatial distribution patterns. However, very few studies analyse the close interaction and effect of environmental variables on diversity at fine spatiotemporal scales. In this study, we evaluated the influence of soil type, vegetation cover and temperature on species diversity and functional structure of dung beetle assemblages. Dung beetle species were sampled using pitfall traps baited with cattle dung from sandy and clay soils in open woodland and closed canopy areas in Mkhuze Game Reserve, KwaZulu‐Natal, South Africa. We performed a PERMANOVA, a PERMDISP and an NMDS to analyse differences in species composition between soil and vegetation cover types. To test the effect of environmental variables on species richness and abundance, we performed a generalised least squares model. Lastly, we searched for nodes using a bipartite analysis to evaluate network's potential modularity. We collected 1439 specimens comprising 27 species and 13 genera. Species composition differed between environmental variables. Soil type was the most important driver for changes in dung beetle species composition. Species richness and abundance were influenced by the interaction between soil type and vegetation cover. The combination of microenvironmental preferences potentially influenced the dung beetle assemblage diversity and functional structure. Combining all these preferences may act as a dilution mechanism of interspecific competition in saturated habitat conditions, reducing the number of species that potentially interact and generating a modular network structure of the assemblage. [ABSTRACT FROM AUTHOR]

The role of the geological uplift and climatic changes during the late Cenozoic on the species diversification of southern African dung beetles is not fully understood. Therefore, we use a divergence-time-estimated phylogeny, macroevolutionary analyses and ecological niche modelling under different climatic scenarios to investigate diversification of the endemic southern African genus, Epirinus. We predict the ancestral range and vegetation type occupied by Epirinus and how late Cenozoic climatic fluctuations and resulting vegetation changes affected speciation and extinction of Epirinus species. Our results suggest that the genus originated in forest with radiation into three geographical centres: (a) north-east escarpment forest and highland grassland; (b) south-east forest; and (c) south-west lowlands to north-east uplands in open vegetation. Reduced speciation rates in the mid-Miocene and increased extinction rates during the drier and cooler Plio-Pleistocene coincide with the replacement of forest by grassland or savanna in southern Africa. The drier climate in southern Africa may have driven extensive contraction of shaded vegetation, forcing an adaptation of forest inhabitants to upland grassland environments, or driving Epirinus species to extinction. Our study supports hypothesis of climatically driven diversification of Epirinus whereas ecological niche modelling across different geological periods suggest that the south-east and, to a lesser extent, the west coast of South Africa as stable areas. [ABSTRACT FROM AUTHOR]

Aim: The high diversity of species in southern Africa has been attributed to geological and palaeoclimatic factors. The timing of radiations in some groups is held to be linked to these geoclimatic trends. Using the Scarabaeinae dung beetle genus, Sisyphus, as a model system, we investigate how geological uplift and climatic changes in the late Cenozoic affected its diversification patterns in southern Africa. Location: Southern Africa. Taxon: The dung beetle genus, Sisyphus Latreille, 1807 (Coleoptera: Scarabaeidae: Scarabaeinae). Methods: A dated molecular phylogeny of southern African Sisyphus was compared with a factor analysis of species distribution data that statistically defined groups of species according to current climatic distribution. We used these climatic clusters to estimate ancestral ranges using the "BioGeoBEARS" R package. A lineage through time plot was calculated using the R package 'APE'. We used Bayesian diversification models (compound Poisson process on mass extinction times) to test hypotheses on how late Cenozoic uplift and climatic changes affected speciation and extinction of Sisyphus. Furthermore, we implemented ecological niche modelling in MaxEnt to predict the habitat suitability of species under present climatic conditions. Results: Four species groups defined from factor analysis of current climatic distribution data are primarily restricted to the moist summer rainfall region in the northeast. Species diversification occurred from the mid‐Miocene until the Pleistocene, during which the central southern African plateau was uplifted in tandem with the advent of winter rainfall and arid climatic zones to the southwest. The ancestor of Sisyphus was centred in eastern low to mid‐altitude areas. Speciation rates were higher in the Miocene to Pliocene, followed by a dramatic drop during the transition to the Pleistocene. Main conclusions: The taxonomic and eco‐climatic diversification of sisyphines is coincident with geological uplift and changes in climate in east‐central southern Africa. However, phylogenetic relationships of southern African Sisyphus species are not homogenously distributed with respect to geography and climate. [ABSTRACT FROM AUTHOR]

The tribe Sisyphini sensu stricto Mulsant, 1842 comprises only three genera, the widespread Sisyphus Latreille, 1807 and Neosisyphus Müller, 1942 , and the Mauritius endemic, Nesosisyphus Vinson, 1946. In southern Africa, Sisyphus and Neosisyphus are represented by five species groups in each genus. Together, they comprise a total of 33 valid species, of which six are new: Sisyphus auricomus sp. n; Sisyphus australis sp. n ; Sisyphus bicuariensis sp. n ; Sisyphus inconspicuus sp. n ; Sisyphus swazi sp. n ; and Neosisyphus tembyi sp. n. A further Southern African species, Sisyphus crispatus Gory, 1833 , is proposed as a nomen dubium. Sisyphus natalensis Balthasar, 1968 (syn. n), and Sisyphus bornemisszanus Endrödi, 1983 (pars) (syn. n) are made synonyms of Sisyphus sordidus Boheman, 1857. Lectotypes and paralectotypes are designated for Sisyphus costatus (Thunberg, 1818); Sisyphus seminulum Gerstaecker, 1871 ; Sisyphus nanniscus Péringuey, 1901 ; Sisyphus transvaalensis Péringuey 1901 ; Neosisyphus spinipes (Thunberg, 1818) and Neosisyphus barbarossa (Wiedemann, 1823). Diagnoses, photographs of habitus and male genitalia, lists of examined material and distribution maps are presented for all species. An identification key to the southern African sisyphine species is provided. [ABSTRACT FROM AUTHOR]

The tribe Sisyphini Mulsant was recently redefined following the transfer of the endemic southern African genus Epirinus Dejean from the polyphyletic tribe Deltochilini Lacordaire. A molecular phylogeny of the southern African members of Sisyphini supports Epirinus as sister to Sisyphus Latreille and recovered three major clades in Sisyphus classified here as subgenera Sisyphus (Neosisyphus Müller) stat. rev., Sisyphus (Parasisyphus Barbero, Palestrini & Zunino) stat.n. and Sisyphus (Sisyphus) stat.n. A molecular clock analysis suggests that Sisyphus and Epirinus diverged from their last common ancestor during the Lower to Middle Oligocene (c. 29.37 Ma). Biogeographical analysis indicated that southern African Sisyphus species are centred in the east and northeast in Highveld grassland and warmer savannah regions. By contrast, Epirinus species are largely restricted to the southwest and southeast in the cooler winter and bimodal rainfall regions plus arid highland Karoo and Highveld grasslands. Based on morphological and biogeographical differences between Epirinus and Sisyphus, we propose that the monogeneric Epirinus be placed in its own tribe, Epirinini van Lansberge stat. rev. [ABSTRACT FROM AUTHOR]

The tribe Sisyphini sensu stricto Mulsant, 1842 comprises only three genera, the widespread Sisyphus Latreille, 1807 and Neosisyphus Müller, 1942 , and the Mauritius endemic, Nesosisyphus Vinson, 1946. In southern Africa, Sisyphus and Neosisyphus are represented by five species groups in each genus. Together, they comprise a total of 33 valid species, of which six are new: Sisyphus auricomus sp. n; Sisyphus australis sp. n ; Sisyphus bicuariensis sp. n ; Sisyphus inconspicuus sp. n ; Sisyphus swazi sp. n ; and Neosisyphus tembyi sp. n. A further Southern African species, Sisyphus crispatus Gory, 1833 , is proposed as a nomen dubium. Sisyphus natalensis Balthasar, 1968 (syn. n), and Sisyphus bornemisszanus Endrödi, 1983 (pars) (syn. n) are made synonyms of Sisyphus sordidus Boheman, 1857. Lectotypes and paralectotypes are designated for Sisyphus costatus (Thunberg, 1818); Sisyphus seminulum Gerstaecker, 1871 ; Sisyphus nanniscus Péringuey, 1901 ; Sisyphus transvaalensis Péringuey 1901 ; Neosisyphus spinipes (Thunberg, 1818) and Neosisyphus barbarossa (Wiedemann, 1823). Diagnoses, photographs of habitus and male genitalia, lists of examined material and distribution maps are presented for all species. An identification key to the southern African sisyphine species is provided. [ABSTRACT FROM AUTHOR]

The lacewing genus Afroptera Abdalla & Mansell (Neuroptera: Nemopteridae: Nemopterinae) is endemic to southern Africa, predominantly found in the Fynbos and Succulent Karoo biomes. The taxonomy of the genus has been recently resolved. However, the monophyly and evolutionary history of the genus has never been addressed. This study employs an integrative phylogenetic approach, by incorporating three ribosomal genes (16S, 28S and 18S) and two protein‐coding genes (cytochrome oxidase subunit I and carbamoyl‐phosphate synthetase‐aspartate transcarbamoylase‐dihydroorotase), and morphological data to examine the monophyly and historical biogeography of Afroptera. We use Bayesian, parsimony and maximum likelihood phylogenetic methods to assess the monophyly and relatedness of Afroptera within the Nemopterinae. We also use ancestral range reconstruction and diversification analysis to infer the historical biogeography of the genus. Our analyses reveal the genus as a monophyletic lineage. The genus Afroptera originated during the Pliocene (5.24–3.13 Mya) in a desert environment, experiencing rapid speciation during the Pleistocene, primarily within the Fynbos and Succulent biomes; and secondarily dispersed into the Nama Karoo and Savannah (Kalahari) biomes. The current distribution patterns of Afroptera species likely stem from intensified aridification in the southwest during the Plio‐Pleistocene, consistent with the dry‐adapted nature of Afroptera's ancestors. Therefore, our findings suggest a climatically driven diversification model for the genus Afroptera. [ABSTRACT FROM AUTHOR]