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Coprinae

The scarab beetle subfamily Scarabaeinae consists of species collectively called true dung beetles. Most of the beetles of this subfamily feed exclusively on dung. However, some may feed on decomposing matter including carrion, decaying fruits and fungi. Dung beetles can be placed into three structural guilds based on their method of dung processing namely rollers, dwellers and tunnelers [3] Dung removal and burial by dung beetles result in ecological benefits such as soil aeration and fertilization; improved nutrient cycling and uptake by plants, increase in Pasture quality, biological control of pest flies and intestinal parasites and secondary seed dispersal.[4] Well-known members include the genera Scarabaeus and Sisyphus, and Phanaeus vindex.

Contents

1 Description 2 Guilds 3 Reproduction 4 Ecology 5 Taxonomy

5.1 Tribes with a global distribution 5.2 Predominantly Old World (New World presence likely represents rather recent dispersal) 5.3 Old World 5.4 New World

6 References 7 External links

Description[edit] Adult dung beetles have modified mouth parts which are adapted to feeding on dung. The clypeus is expanded and covers the mouth parts. The elytra, which cover the wings, expose the pygidium. They also have a space between their middle legs to allow for manipulation of the dung. Dung beetles can be large beetles of a few inches in size to small beetles which are only a few millimeters in size [5] Guilds[edit] Dung beetles are classified into groups based on their method of processing the dung. Rollers are beetles that construct balls of dung from the main food source. They roll away this ball from the position of the original food source and use the dung for feeding or for reproduction. Tunnelers are beetles that dig tunnels beneath the food source creating nests. They relocate food into their nest for reproduction and feeding. Lastly, dwellers are beetles that live and reproduce inside the food source rarely creating nests.[3] Reproduction[edit] Pheromones are thought to aid in dung beetle reproduction. Dung beetles copulate after which both parents dig a tunnel to lay the eggs. This tunnel may have different branches leading to varying egg chambers or may not be branched depending on species. Both parents take dung inside the tunnels in the form of brood balls and the females lay the egg inside the dung. When the larva hatches it feeds on the surrounding dung and forms a pupa undergoing several instars. After this stage the pupa hatches and the newly formed adult evades the tunnel and searches for a fresh dung supply for feeding. After approximately 2 weeks the new adult beetle will be able to reproduce.[6] Ecology[edit] Dung beetle communities are dependent on vegetation and presence of mammalian dung resource. In an area with a lot of vegetation, the dung pads are preserved longer for dung beetle utilization. Vegetation provides conditions suitable for vertebrate trafficking allowing more dung presence in the area. In areas with little vegetation, limited vertebrate fauna may be present limiting the types of dung present. Also, with little vegetation the canopy of the forest may be limiting. This causes the dung pads to be exposed to sunlight, higher temperature and wind action. These factors contribute to dung pad desiccation which decreases the period of suitable dung availability. Dung beetles of different groups may respond to changes in vegetation in different ways.[7] A decline in dung beetle diversity associated with the conversion of native forests to open pastures is known to occur. However, some species are able to utilize dung in open pasture conditions. Tunneler species are able to utilize dung in less optimal condition because the tunnel beneath the dung and utilize it from bottom up. As desiccation occurs on the top of the dung pad, the middle and lower regions may still be suitable for the dung beetle. Also, smaller species are able to survive in such conditions as they require less dung. Larger beetles require larger amounts of dung which may not be available due to the fast desiccation of the dung pads in these conditions. Soil hardness and dryness negatively affect the tunnelers. Species able to tolerate open pasture conditions were extremely abundant. The reduction in the number of dung beetles affected the efficiency of their ecological roles and it was found that dung removal and burial declined.Dung removal and burial by dung beetles result in ecological benefits such as soil aeration and fertilization. These benefits lead to improved nutrient cycling and uptake by plants, increase in pasture quality, biological control of pest flies and intestinal parasites and secondary seed dispersal. Therefore, their role in the ecosystem is essential.[8] Taxonomy[edit] The subfamily is split into 12 tribes.[2][9] These can be grouped by their distribution, though the present classification is expected to require major revision based on recent phylogenetic analyses.[10] Tribes with a global distribution[edit]

Ateuchini - about 20 genera including:

Pedaria Sarophorus

Sarophorus costatus

Pedaria nigra

Coprini (incl. Dichotomiini) - about 20 genera including:

Copris Geoffroy, 1762 Dichotomius Heliocopris Metacatharsius

Copris elphenor female

Copris elphenor male

Deltochilini (= Canthonini) - about 120 genera including:

Anachalcos Hope, 1837 Canthon Hoffmannsegg, 1817 Circellium Latreille, 1825 Dicranocara Frolov and Scholtz, 2003 Epirinus Reiche, 1841 Versicorpus Deschodt Davis and Scholtz, 2011 Deltochilum

Anachalcos convexus

Canthon smaragdulus

Circellium bacchus

Onthophagini - about 40 genera including:

Hyalonthophagus Onthophagus Proagoderus

Hyalonthophagus alcyonides

Onthophagus binodis

Onthophagus gazella

Proagoderus imperator

Predominantly Old World (New World presence likely represents rather recent dispersal)[edit]

Oniticellini (incl. Eurysternini) - 16 genera including:

Euoniticellus Janssens, 1953 Oniticellus Serville, 1825 Eurysternus

Sisyphini - three genera:

Nesosisyphus Neosisyphus Sisyphus

Old World[edit]

Gymnopleurini - four genera:

Allogymnopleurus Garreta Janssens, 1940 Gymnopleurus Illiger, 1803 Paragymnopleurus

Scarabaeini - five genera:

Drepanopodus Kheper Pachylomera Pachysoma Scarabaeus

Parachoriini - one genus:

Parachorius

Onitini - 19 genera including:

Bubas Mulsant, 1842 Cheironitis Lansberge, 1875 Onitis Fabricius, 1798

Cheironitis scabrosus

Chironitis indicus

New World[edit]

Eucraniini - four genera:

Anomiopsoides Ennearabdus Eucranium Glyphoderus

Phanaeini - 12 genera including:

Oxysternon Phanaeus

References[edit]

^ Tarasov, Sergei; Vaz-de-Mello, Fernando Z.; Krell, Frank-Thorsten; Dimitrov, Dimitar (2016). "A review and phylogeny of Scarabaeine dung beetle fossils (Coleoptera: Scarabaeidae: Scarabaeinae), with the description of two Canthochilum species from Dominican amber". PeerJ. 4: e1988. doi:10.7717/peerj.1988. PMC 4986599 . PMID 27547512.  ^ a b Bouchard, P.; Bousquet, Y.; Davies, A.; Alonso-Zarazaga, M.; Lawrence, J.; Lyal, C.; Newton, A.; Reid, C.; Schmitt, M.; Ślipiński, A.; Smith, A. (2011). "Family-group names in Coleoptera (Insecta)". ZooKeys. 88 (88): 1–972. doi:10.3897/zookeys.88.807. PMC 3088472 . PMID 21594053.  ^ a b Louzada, J; Lima, A; Matavelli, R; Zambaldi, L; Barlow, J (2010). "Community structure of dung beetles in Amazonian savannas: role of fire disturbance, vegetation and landscape structure". Landscape Ecology. 25 (4): 631–641. doi:10.1007/s10980-010-9448-3. Retrieved 10 October 2017.  ^ Braga, R; Korasaki, V; Andresen, E; Louzada, J (2013). "Dung Beetle Community and Functions along a Habitat-Disturbance Gradient in the Amazon: A Rapid Assessment of Ecological Functions Associated to Biodiversity". PLoS ONE. 8 (2). doi:10.1371/journal.pone.0057786.  ^ Gill, Bruce. "Scarabaeinae". Generic Guide to New World Scarab Beetles. UNL State Museum. Retrieved 10 October 2017.  ^ Gaikwad, A; Bhawane, G (2016). "Observation on life cycle and nesting behavior of dung beetle Onthophagus Catta (Fabricius)". International Journal of Zoology Studies. 1 (7): 09–13.  ^ Liberal, C; Farias, A; Meiado, M; Filgueiras, B; Iannuzzi, L (2011). "). How Habitat Change and Rainfall Affect Dung Beetle Diversity in Caatinga, a Brazilian Semi-Arid Ecosystem". Journal of Insect Science. 11 (114): 1–11. doi:10.1673/031.011.11401.  ^ Horgan, F (2008). "Dung beetle assemblages in forests and pastures of El Salvador: a functional comparison". Biodiversity and Conservation. 17 (12): 2961–2978. doi:10.1007/s10531-008-9408-2.  ^ Tarasov, S.I. (2017). "A cybertaxonomic revision of the new dung beetle tribe Parachoriini (Coleoptera: Scarabaeidae: Scarabaeinae) and its phylogenetic assessment using molecular and morphological data". Zootaxa. 4329 (2): 101–149. doi:10.11646/zootaxa.4329.2.1.  ^ Tarasov, Sergei; Génier, François (17 March 2015). "Innovative Bayesian and Parsimony Phylogeny of Dung Beetles (Coleoptera, Scarabaeidae, Scarabaeinae) Enhanced by Ontology-Based Partitioning of Morphological Characters". PLoS ONE. 10 (3): e0116671. doi:10.1371/journal.pone.0116671. 

External links[edit]

Data related to Scarabaeinae at Wikispecies Media related to Scarabaeinae at Wikimedia Commons Subfamily Scarabaeinae: photographs of representatives of the tribes

Taxon identifiers

Wd: Q2412715 BugGuide: 2950 EoL: 12044660 Fauna Europaea: 191928 Fossilworks: 70043 iNaturalist: 125819 ITIS:

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