Dung-beetles (Coleoptera, Scarabaeidae, Aphodiinae, Scarabaeinae) feeding on faeces of steppe marmots Marmotabobak (Rodentia, Sciuridae) in Middle Volga territory.

Background: In open terrestrial biomes of Holarctic realm, ground squirrels are recognised as keystone species inhabiting steppes. They shape the plant species composition and diversity and support a fauna of species associated with their burrows. Ground squirrels and associated dung-beetles are important elements of the steppe food webs, yet the trophic associations between species are still poorly studied. New information: The area in the northern outskirts of Obshchy Syrt plateau, on the border of Samara and Orenburg Provinces of Russia was surveyed and scarab beetles (Scarabaeidae) feeding on steppe marmot (Marmotabobak (Müller, 1776)) faeces were collected from six localities. Twenty eight species of two subfamilies - Aphodiinae and Scarabaeinae, - were identified with the majority of species belonging the genus Aphodius Hellwig, 1798. Seven species are recorded as consumers of marmot faeces for the first time. Only two nidicolous specialist species were found which suggests that the studied population of steppe marmots is as result of the recent secondary colonisation and not all the associated scarab beetle faunas were re-established.

New data on trophic associations of dung beetles (Coleoptera, Scarabaeidae, Scarabaeinae) and lemurs (Primates, Lemuroidea) in Madagascar revealed by metabarcoding.

Resource use and diet specialisation of Madagascan dung beetles have been little studied especially concerning the possible associations between specific dung beetle and lemur species. Pilot studies have demonstrated that amplicon sequencing is a promising tool for the lemur inventories. In the present contribution, we report the results of the gut content analysis of three endemic Madagascan dung beetles species: Helictopleurusclouei (Harold), Epilissusapotolamproides (Lebis) and Nanosdubitatus (Lebis). Amplicon metagenomics revealed trophic associations of these species with Eulemursanfordi (Archbold), Eu.fulvus (É.Geoffroy Saint-Hilaire) and Cheirogaleuscrossleyi (Grandidier), respectively. The reads of other mammal species, revealed by the analysis, including putative contaminations, are discussed.

Phylogenetic analysis of the Neotropical scarab beetle tribe Aegidiini (Coleoptera, Scarabaeidae, Orphninae) with description of new taxa.

In the Neotropics, orphnine scarab beetles are represented by the endemic tribe Aegidiini Paulian, 1984 with five genera and over 50 species. Phylogenetic analysis based on morphological characters of all supraspecific taxa of Orphninae showed that Aegidiini is comprised of two lineages. New subtribes, Aegidiina subtr. nov. (Aegidium Westwood, 1845, Paraegidium Vulcano et al., 1966, Aegidiellus Paulian, 1984, and Onorius Frolov & Vaz-de-Mello, 2015, and Aegidininasubtr. nov. (Aegidinus Arrow, 1904) are proposed to better reflect this phylogeny. Two new species of Aegidinus are described: A.alexanderisp. nov., from the Yungas in Peru and A.elbaesp. nov. from the Caqueta moist forests ecoregion in Colombia. A diagnostic key to Aegidinus species is given.

Amplicon metagenomics of dung beetles (Coleoptera, Scarabaeidae, Scarabaeinae) as a proxy for lemur (Primates, Lemuroidea) studies in Madagascar.

Dung beetles (Scarabaeidae, Scarabaeinae) are among the most cost-effective and informative biodiversity indicator groups, conveying rich information about the status of habitats and faunas of an area. Yet their use for monitoring the mammal species, that are the main providers of the food for the dung beetles, has only recently been recognized. In the present work, we studied the diet of four endemic Madagascan dung beetles (Helictopleurusfissicollis (Fairmaire), H.giganteus (Harold), Nanosagaboides (Boucomont), and Epilissussplendidus Fairmaire) using high-throughput sequencing and amplicon metagenomics. For all beetle species, the ⅔-¾ of reads belonged to humans, suggesting that human feces are the main source of food for the beetles in the examined areas. The second most abundant were the reads of the cattle (Bostaurus Linnaeus). We also found lower but significant number of reads of six lemur species belonging to three genera. Our sampling localities agree well with the known ranges of these lemur species. The amplicon metagenomics method proved a promising tool for the lemur inventories in Madagascar.

New taxa of Athyreacaridae (Acari: Heterostigmata) from Neotropical and Afrotropical realms.

A new genus and three new species of Athyreacaridae (Acari: Heterostigmata) are described associated with beetles of subfamily Bolboceratinae (Coleoptera: Geotrupidae) in Neotropical and Afrotropical realms: Neoathyreacarus pygmephoroides gen., sp. nov. collected on Bolborhinum tubericeps in Chile; Athyreacarus staturosus sp. nov. collected on Zefevazia cantisani  in Argentina; and Athyreacarus camerikae sp. nov. collected on Namibiobolbus helgae in South Africa. An updated key to genera and species of Athyreacaridae is provided. Athyreacarus staturosus sp. nov. is the largest ever described non-physogastric female among Heterostigmata.

A new species of Aegidium Arrow (Coleoptera: Scarabaeidae: Orphninae) from Colombia.

In the Neotropical Region, Orphninae (Coleoptera: Scarabaeidae) are represented by the endemic tribe Aegidiini, which comprises five genera and 49 species (Paulian 1948; Colby 2009; Frolov Vaz-de-Mello 2015; Frolov et al. 2017a, 2017b, 2017c, 2019). Aegidium Westwood is the largest genus of the tribe and it comprises 24 valid species known from the southern Mexico in the north to Central Bolivia in the south (Frolov et al. 2015; Rojkoff Frolov 2017; Frolov Akhmetova 2020).

Identification of larvae of two Aphodius Helwig, 1798 (Coleoptera: Scarabaeidae: Aphodiinae) species using morphology and DNA barcode.

The third instar larvae of Aphodius (Alocoderus) hydrochaeris (Fabricius, 1798) and A. (Bodilus) ictericus (Laicharting, 1781) are described based on scanning electron microscopy and COI sequences. COI barcode sequence for A. (A.) hydrohaeris is provided for the first time. Two haplotypes are discovered in A. (B.) ictericus.

First record of Athyreacaridae (Acari: Heterostigmata) from Asia with description of two new species of Athyreacarus from India and Sri Lanka.

Mites of the family Athyreacaridae are recorded from Asia for the first time. Two new species of Athyreacarus are described, A. indicus sp. nov., collected on the beetles Bolbohamatum laevicolle and B. marginale in India, and A. brevisetosus sp. nov., collected on a beetle Bolboceras sp. in Sri Lanka. An updated key to species of the genus Athyreacarus is provided.

Description of a new brachypterous scarab species, Orphnus brevialatus (Coleoptera: Scarabaeidae: Orphninae) from East Africa, with notes on flightlessness in the orphnines.

A new brachypterous species of the scarab beetle genus Orphnus Macleay, 1819, O. (O.) brevialatus Frolov Akhmetova, new species, is described from Kipengere Range (Livingstone Mountains), Tanzania. The new species differs from other flightless Orphnus species occurring in East Africa in having a well-developed pronotal armature with two long, acute lateral processes. In the shape of the head, pronotum, and male genitalia it is similar to O. compactilis Quedenfeldt, 1882, but differs from it, in addition to the reduced wings and shorter body, in the shape of the endophallic armature. Flightlessness is rather common among the orphnines: 56 orphnine species out of 235 have the wings reduced to different degree making them incapable of flight, although this is a preliminary minimum figure. Flightless species are found in Hybalus Dejean, 1833, Chaetonyx Schaum, 1862, Orphnus Macleay, Hybaloides Quedenfeldt, 1884, Aegidium Westwood, 1845 and Stenosternus Karsch, 1881. All regional orphnine faunas except for Indo-Malayan and Madagascan include flightless species, although flightlessness is more common in African taxa.

The taxonomic status of Aphodius insperatus Petrovitz, A. longipennis Rakovic and A. inylchekensis Frolov (Coleoptera: Scarabaeidae: Aphodiinae).

The subgenus Bodilus Mulsant Rey, 1870 of the genus Aphodius Helwig, 1798 sensu lato comprises 30 species in the Palearctic fauna (Frolov 2001, 2002, Dellacasa et al. 2016 [treated as genus Bodilus]). Frolov (2001) reviewed the species of this subgenus from Russia and adjacent countries and suggested that the monotypic subgenus Paramelinopterus Rakovic, 1984 shares the main diagnostic characters of Bodilus. Frolov (2001) was unable, however, to examine the type species of Paramelinopterus, A. longipennis Rakovic, 1984, at that time. Král (2016) suggested that A. (Bodilus) inylchekensis Frolov, 2001 is a junior synonym of A. longipennis. Recently we have re-examined all the types as well as the additional material accumulated in the museums and can confirm that A. inylchekensis is a distinct species, but A. insperatus Petrovitz, 1967 and A. longipennis cannot be reliably separated by the morphological characters. In the present contribution we illustrate the major diagnostic characters of the species based on the type specimens and also provide a distribution map of the known localities of the species.

A new species of the endemic Madagascan scarab beetle genus, Madecorphnus grebennikovi (Coleoptera: Scarabaeidae: Orphninae): morphological description and DNA barcode.

A new species of the orphnine scarab beetle genus Madecorphnus Paulian, 1992, Madecorphnus grebennikovi Frolov, Akhmetova Vishnevskaya, new species, is described from the Marojejy National Park, Sava Region, northeastern Madagascar. The new species can be distinguished from the congeners by the parameres narrowly rounded in lateral view and having a small but distinct lateral teeth, and by the endophallic armature consisting of 1) a long straight sclerite with attached to its end a 2/3 shorted, somewhat curved sclerite, 2) separate smaller, elongate sclerite, and 3) a rather large area of microspinules. The 811 bp long fragment of the mitochondrial gene COI (DNA barcode) is provided as a part of the diagnosis of the new species. An updated key to the Madecorphnus species is given.

A review of the Malagasy species of the genus Trigonochilus Brenske (Coleoptera: Scarabaeidae: Rutelinae: Adoretini) with description of a new species.

Africa and Madagascar have a large and diverse fauna of the chafers of the ruteline beetle tribe Adoretini (Ohaus 1912, 1918). Many nominal taxa from Madagascar are still poorly known and many new ones were described recently (Akhmetova Montreuil 2010; Montreuil 2010, 2013; Montreuil Frolov 2014; Frolov Montreuil 2018). One of the poorly known Adoretini taxa is the genus Trigonochilus Brenske, 1896. Trigonochilus was proposed to accommodate the East-African species T. coriaceus Brenske, 1896, based on the enlarged and curved metatibiae and a short, spur-like process of the labrum (Brenske 1896). Arrow (1901) noted that the enlarged metatibiae of this species (described from a single male) is a secondary sexual character and added another species to this genus, T. politus Arrow, 1901, from Angola. These species were revised by Krikken (1979). Fairmaire (1903) described Adoretus oedipus from Madagascar (without any precise locality) and noted its enlarged metatibia and the shape of the labrum similar to that in Trigonostonum Burmeister, 1844. Fairmaire was apparently unaware of the Brenske's and Arrow's works as he did not discuss the possible affinities of A. oedipus with Trigonochilus species. Ohaus (1912) moved A. oedipus to Trigonochilus after examination of the type series but he did not discuss the characters of the species in detail as well as its distribution on the island. Therefore, the genus Trigonochilus Brenske comprised three species described so far, two from Africa and one from Madagascar.

A review of the Madagascan species of the genus Cestradoretus Ohaus (Coleoptera: Scarabaeidae: Rutelinae: Adoretini).

The genus Cestradoretus Ohaus, 1912 was erected by Ohaus (1912) to accommodate a Madagascan Adoretini species, C. tenuirostris Ohaus, 1912. This genus is characterized by the adults having the apex of the labrum with a long, fine, acute process curved downwards. In the addition to the above mentioned paper, Ohaus (1912) also described C. acomys Ohaus, 1912 from Tanzania, and later moved Adoretus tarsatus Klug, 1855, described from Mozambique, to Cestradoretus (Ohaus 1916).  Later, Machatschke (1965) described two species from Africa, C. freyi Machatschke, 1965 from Botswana and C. minusculus Machatschke, 1965 from Somalia, and Frey (1976) described C. clypealis Frey, 1976 from Somalia. Thus, Cestradoretus included six species, one from Madagascar and five from Africa.

New and little known species of the Orphninae (Coleoptera: Scarabaeidae) from Madagascar.

Two new species of the orphnine scarab beetle genus Madecorphnus Paulian, 1992 are described: M. heidiae Frolov Akhmetova, new species, and M. ranomafanensis Frolov Akhmetova, new species, from the Ranomafana National Park, Fianarantsoa Province, Eastern Madagascar. The new species can be easily distinguished by the endophallic armature consisting of more than 10 small and similar, conical tubercles in the former species and 3 long sclerites and 2 smaller indistinctly separated clusters of robust spinules in the later. Exact locality data are provided for the first time for two Triodontus Westwood, 1845, species: T. hildebrandtii (Fairmaire, 1883) was found in the northern Madagascar and is probably endemic to the Amber Mountain Range, T. fairmairei Frolov, Montreuil Akhmetova, 2016, was also found in the northern Madagascar: Antsiranana Province, Djangoa. The updated key to the Madecorphnus species is given.

Revision of the genus Athyreacarus (Acari: Athyreacaridae).

Mites of the genus Athyreacarus (Acari: Athyreacaridae) associated with bolboceratine beetles (Coleoptera: Geotrupidae: Bolboceratine) of New World are revised, a new generic concept provided, and the family concept modified. Nine new species are described: A. grandis sp. nov. on Neoathyreus lanuginosus in Brazil; A. magnificus sp. nov. on Athyreus tuberifer in Brazil; A. variabilis sp. nov. on Neoathyreus cf. anthracinus in Brazil; A. similis sp. nov. on Neoathyreus centromaculatus in Bolivia; A. vazdemelloi sp. nov. on Parathyreus rectus in Brazil; A. pusillus sp. nov. on Athyreus brasilicus in Brazil; A. latus sp. nov. on Athyreus cf. bifurcatus from unknown locality; A. angustus sp. nov. on Bolbelasmus horni in USA; and A. ovalis sp. nov. on Bolbelasmus variabilis in Mexico. Bolbapium striatopunctatum, B. minutum, and B. sculpturatum are recorded as new phoretic host beetles for Athyreacarus primitivus. A key to genera and species of the family Athyreacaridae is provided.

A new species, new genus and new records of heterostigmatic mites (Acari: Heterostigmata) phoretic on scarab beetles of the subfamily Orphninae (Coleoptera: Scarabaeidae).

During the study of heterostigmatic mites phoretic on beetles of the subfamily Orphninae (Coleoptera: Scarabaeidae), ten species from five families were found. One species, Premicrodispus hajiqanbari sp. nov. (Acari: Microdispidae) phoretic on Orphnus luminosus Benderitter, found in Central African Republic is described as new for science. A new monobasic genus Scarabadania gen. nov. with type species Bakerdania tenuispina Sevastianov, 1974 is created in the family Neopygmephoridae. Troxodania problematica (Mahunka, 1975) comb. nov. (from Bakerdania) and Pseudopygmephorus peritrematus (Mahunka, 1979) (Acari: Neopygmephoridae) are redescribed. Moreover, Trochometridium tribulatum Cross, 1965 (Acari: Trochometridiidae), Troxodania problematica (Mahunka, 1975) comb. nov. and Aegyptophorus shibinensis Sevastianov and Abo-Korah, 1984 (Acari: Neopygmephoridae) are recorded from Sri Lanka for the first time. Finally, Scarabadania tenuispina (Sevastianov, 1974) comb. nov. and Pygmodispus odontotarsus Sevastianov, 1975 (Acari: Scutacaridae) are recorded from Africa for the first time.

A new species of Formicomotes Sevastianov (Acari: Heterostigmata: Dolichocybidae) associated with termites (Isoptera: Termitidae) from Brazil, with redescription of Formicomotes octipes Sevastianov, 1980.

The new species Formicomotes brasiliensis sp. nov. (Acari: Heterostigmata: Dolichocybidae), phoretic on the termite genus Nasutitermes Dudley (Insecta: Isoptera) from Brazil, is described. The genus Formicomotes Sevastianov, 1980 is recorded from the Neotropical region for the first time. Formicomotes octipes Sevastianov, 1980 is redescribed based on type material collected on the ant species Lasius fuliginosus and new specimens from Crimea collected from forest litter.

New taxa of pygmephoroid mites (Acari: Pygmephoroidea: Pygmephoridae, Neopygmephoridae) phoretic on Enoplotrupes sharpi (Coleoptera: Geotrupidae) from Thailand.

The new species Spatulaphorus enoplotruporum sp. nov. (Acari: Pygmephoridae) and a new monotypic genus and the species Nasutidania orientalis gen., sp. nov. (Acari: Neopygmephoridae) are described phoretic on Enoplotrupes sharpi Rothschild and Jordan (Coleoptera: Geotrupidae) from Thailand. A key to the world species of the genus Spatulaphorus is also provided.

A contribution to the revision of Adorodocia Brenske (Coleoptera: Scarabaeidae: Rutelinae: Adoretini): a new peculiar species from Ankaratra, Madagascar.

The recently revised genus Adorodocia Brenske, 1893 (Coleoptera: Scarabaeidae: Rutelinae: Adoretini), comprises 17 species, one of them represented by two subspecies (Montreuil Frolov 2014). This genus forms, together with Paradorodocia Machatschke, 1957 (four species: Montreuil 2010), and Metadorodocia Machatschke, 1957 (two species: Akhmetova Montreuil 2010), the subtribe Adorodociina Ohaus, 1912, endemic to Madagascar (Ohaus 1912). Representatives of this subtribe are characterized by a strong prominent prosternal apophyse. In addition, all the species of this subtribe have a rather large body and bear a few strong long hairs on the apical hump of the elytra and more or less strong scale-like setae distributed over the elytra.