Corresponding author: Andrey Frolov (
Academic editor:
The field sampling of the
The tribe
We have summarized the data of multi-year survey in Astrakhan Province and provided a check-list of the tribe
The study area is situated оn the Akhtuba River (fork of Volga River) about 90 km upstream of Volga estuary, Astrakhan Province, European Russia. We were based in Dosang Railway Station (
Four main biotopes were sampled: fixed sands (Fig.
The material used in this work was collected by the authors from 2006 to 2012 in different seasons but primarily in spring and fall. Additional specimens were received from colleagues listed in the acknowledgement section. All the material mentioned is deposited in the collection of Zoological Institute of Russian Academy of Sciences, Sankt-Petersburg (ZIN).
The beetles were collected by the following methods: washing of fresh horse and cow dung, hand collecting from fresh and old dung, mercury lamp traps, soil sifting with a sieve, hand collecting from marmot holes. The larvae were collected from old dung and soil and some specimens from each lot were reared to adults in the laboratory.
Classification and nomenclature of taxa follow
Cattle dung, mostly cow dung.
Southern Palearctic region up to Saur-Tarbagataj in the north-east.
Cattle dung.
Middle Asian deserts, Caspian lowland desert.
Adults and larvae feed on horse dung (Fig.
Central and Eastern Europe, Western Asia up to East Kazakhstan in the east. The species is mostly occur in forest-steppe and steppe zones.
Cattle dung.
Steppe zone from Caucasus to West Siberia.
Adults and larvae feed on horse dung.
The species was recorded from Caucasus, Turkey, Iran, and Middle Asia (
Cattle and wild herbivore dung.
Europe (except for northernmost part), Asia Minor.
Adults were found in sand near plant roots.
Caspian lowland desert.
Middle Asian deserts, Caspian lowland desert.
Adults and larvae feed on cattle dung.
The species is distributed throughout Europe, Transcaucasia, Central Asia, Northern Africa; it was introduced to North America.
Adults were found in horse and cow dung.
Caspian lowland desert.
Cattle and wild herbivore dung.
Transpalearctic species, introduced to North America.
Cattle and wild herbivore dung.
Transpalearctic species, introduced in North America and Australia.
Adults feed on dry cattle dung.
Western Palearctic region South Europe the south-west to West Siberia in the north-east.
Coprophagous generalist.
West Palearctic region (except for northernmost part), introduced in North America.
Cattle dung.
Steppe and semidesert zones from Lower Volga in the west to Mongolia in the east.
Adults and larvae feed on cattle dung.
Volga Region up to Vladimir Province in the north. Kazakh steppe up to Sultankeldy Lake in the East.
Middle Asian deserts.
Horse dung.
West Palearctic region (except for northernmost part).
Cattle dung.
Distributed throughout Europe (except for northernmost part), in North Africa, Asia Minor, Iran, up to Irtysh River in the east.
Cattle dung
Western Palearctic region from North Africa in the south-west to Baikal Lake in the north-east.
Horse dung.
Caspian lowland desert, Middle Asian deserts.
Cattle dung.
Caspian lowland desert, Middle Asian deserts.
Apparently saprophagous species.
Distributed mostly in steppe and desert zones in South Europe, Caucasus, Asia Minor, Middle Asia.
Cattle dung.
West Palearctic region (except for northernmost part).
Cattle dung.
Western Palearctic region from North Africa in the south-west to Saur-Tarbagataj in the north-east.
Cattle dung.
Western Palearctic region up to South Siberia in the north-east.
Cattle dung.
Steppe and semidesert zones from Ciscaucasia in the west to Mongolia in the east.
Horse dung (Fig.
Caspian lowland desert, Middle Asian deserts
Cattle dung
Southern Central Europe, Transcaucasus, Caspian lowland and Central Asian Deserts up to Dzhungar Alatau in the east
Cattle and wild herbivore dung.
Western Palearctic region from North Africa in the south-west to Altai Mountains the north-east.
Cattle and wild herbivore dung.
Transpalearctic species.
Cattle dung.
Western Palearctic region from North Africa in the south-west to southern Urals in the north-east. Mostly in steppe zones.
Cattle and wild herbivore dung.
Transpalearctic species occurring mostly in steppe zone.
Adults were found in horse and cow dung, larvae in horse dung.
Widely distributed in Europe and Asia up to Baikal Lake in the east.
Cattle dung, mostly cow dung.
Transpalearctic species, mostly in steppe and semidesert zone.
Cattle dung.
Distributed throughout Palaearctic region (except for northernmost part), introduced in North America.
Adults occur in large numbers in horse dung in the fall. Larvae were collected among grass roots at the depth of 10–15 cm in riverine secondary forest (
Middle and southern Europe, Minor Asia, North Africa, northern Iran, Central Asia up to South Siberia.
Adults occur in dry cattle dung.
Distributed mostly in steppe zone from South Europe to Central Kazakhstan.
Adults feed on cattle dung.
Widely distributed in western Palaearctic region up to Zaissan Lake in the east.
Adults feed on horse dung (Fig.
Caspian lowland desert.
Rodent nest dweller, adults and apparently larvae feed on rodent excrement (Fig.
Distributed mostly in the steppe and semidesert zones from Lower Volga in the west to West Siberia in the east.
Cattle and wild herbivore dung.
This species is thought to be widely distributed in the Holarctic Region. However,
Caspian lowland desert.
To date, 44 species of the
Long vegetative season with high effective heat sum. The beetles were found active from March to November and it is possible that some species may be active during winter months, especially during mild winters. The climate in the area is arid in general but the high level of groundwater in sands, even in the driest months, provides suitable conditions for psammophilous taxa.
Large livestock including horses and cows provide abundant food resources throughout the year. In the hottest months however, much of the fresh dung is consumed by the
The collecting area is situated in the transition belt between two floristic districts: Volga-Akhtuba Floodplain (intrazonal) and Desert (zonal) of Caspian Subprovince of Turanian Province (
Apparently due to the last factor, the fauna consists of the two components of different origin. Intrazonal habitats are suitable for a large number of mesophilous species which constitute the core Dosang
Another part of the fauna comprises species with the ranges mostly limited to the steppe, semidesert, and desert zones. These are the species with sethian (
We thank V. P. Bulychev (Dosang Antiplague Station, Dosang) for logistics assistance over many years. We also thank A. V. Ivanov (Institute of Plant and Animal Ecology, Ekaterinburg) and K. V. Makarov and A. V. Matalin (Moscow State Pedagogical University, Moscow) for providing additional specimens. Alberto Ballerio (Brescia), Bruce Gill (Ottawa), and Jan Krikken (Leiden) helped to correct and improve the draft manuscript. This study was supported by the Russian Foundation for Basic Research (grants 07-04-00482, 10-04-00539, and 13-04-010020).
Supplementary file with occurences data for cheklist of
Fixed sands near Dosang, Astrakhan Province, Russia.
Barchan sands North-East of Dosang, Astrakhan Province, Russia.
Right side of Akhtuba River, floodplain, Dosang environs, Astrakhan Province, Russia.
Left side of Akhtuba River, riverine secondary forest near Dosang, Astrakhan Province, Russia.