PREDATION ON DUNG BEETLES (Coleoptera: Scarabaeidae): A...
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TRANSACTIONS AMERICAN ENTOMOLOGICAL SOCIETY RESEARCH ARTICLE Publication date: 12 August 2015 urn:lsid:zoobank.org:pub:71896AE0-BEAA-4F6A-BCFA-B530703FDCC6 TAES 141: 111-155 ISSO 0002-8320 http://taes.entomology-aes.org/ PREDATION ON DUNG BEETLES (Coleoptera: Scarabaeidae): A LITERATURE REVIEW ORREY P. YOUNG Formerly at Department of Zoology, University of Maryland, College Park, MD 20742 Current Address: 9496 Good Lion Rd., Columbia, MD 21045 Email: [email protected] http://www.researchgate.net/profile/Orrey_Young ABSTRACT Several reviews of dung beetle (Coleoptera: Scarabaeidae) behavior and ecology, as well as numerous research reports, have either implicitely or explicitely indicated that predation on dung scarabs is rare or absent and thus of minimal or no importance to the biology of the group. This review presents 610 records of predation on dung beetles by 409 species of birds, mammals, reptiles and amphibians from around the world. The involvement of invertebrates as predators on dung scarabs is also documented. It is concluded that this data establishes predation as a potentially major factor in the evolution and current behavior and ecology of dung beetles. The data presented also represents a considerable underestimation of predation on the group. PART 1 – INTRODUCTION, HIGHLIGHTS, DISCUSSION, LITERATURE CITED INTRODUCTION “Dung beetles” refer to those beetles (Coleoptera) that are present at the fecal deposits of animals. Though members of several beetle families may be present, the majority of beetles are typically scarabs (Scarabaeidae). Within that family, all are members of either the subfamilies Scarabaeinae, Aphodinae, or Geotrupinae. Adult individuals of these subfamilies either consume the dung on site or remove it downward or horizontally for eventual consumption by the adult or by subsequent larval progeny (Halffter & Matthews 1966). Current interest in dung scarabs is primarily in the following areas: (1) Secondary disperser of seeds, initially consumed by herbivores with subsequent elimination of seeds in feces and then subsequent capture and dispersal by dung scarabs. Recent studies in the tropical forests of Southeast Asia (McConkey 2005), Central and South America (Lawson et al. 2012, Santos-Heredia et al. 2010), and in temperate forests (Koike et al. 2012) and grasslands (D’hondt et al. 2008) all document the significant role of dung scarabs as secondary seed dispersers. Their associated subsequent involvement in the effects of forest fragmentation in the tropics also has been considered (Nichols et al. 2007). (2) Dung scarabs as important surrogates for detecting changes in tropical forest biodiversity. Their use in analyses has lead to conservation recommendations (Nichols et al. 2009, Gardner et al. 2008, Kohlmann et al. 2007), as well as being used in a broader role as bioindicators (McGeoch et al. 2002). They also can be used as indicators in savannaagroecosystems of pest control practices (Davis et al. 2004). (3) The role of biotic and abiotic factors in the organization and function of communities and
Transcript of PREDATION ON DUNG BEETLES (Coleoptera: Scarabaeidae): A...
TRANSACTIONS AMERICANENTOMOLOGICALSOCIETY
RESEARCH ARTICLE
Publication date: 12 August 2015 urn:lsid:zoobank.org:pub:71896AE0-BEAA-4F6A-BCFA-B530703FDCC6
TAES 141: 111-155ISSO 0002-8320
http://taes.entomology-aes.org/
PREDATION ON DUNG BEETLES (Coleoptera: Scarabaeidae): A LITERATURE REVIEW
ORREY P. YOUNG
Formerly at Department of Zoology,University of Maryland,
College Park, MD 20742
Current Address: 9496 Good Lion Rd., Columbia, MD 21045Email: [email protected]
http://www.researchgate.net/profile/Orrey_Young
ABSTRACT
Several reviews of dung beetle (Coleoptera: Scarabaeidae) behavior and ecology, as well as numerous research reports, have either implicitely or explicitely indicated that predation on dung scarabs is rare or absent and thus of minimal or no importance to the biology of the group. This review presents 610 records of predation on dung beetles by 409 species of birds, mammals, reptiles and amphibians from around the world. The involvement of invertebrates as predators on dung scarabs is also documented. It is concluded that this data establishes predation as a potentially major factor in the evolution and current behavior and ecology of dung beetles. The data presented also represents a considerable underestimation of predation on the group.
PART 1 – INTRODUCTION, HIGHLIGHTS, DISCUSSION, LITERATURE CITED
INTRODUCTION
“Dung beetles” refer to those beetles (Coleoptera) that are present at the fecal deposits of animals. Though members of several beetle families may be present, the majority of beetles are typically scarabs (Scarabaeidae). Within that family, all are members of either the subfamilies Scarabaeinae, Aphodinae, or Geotrupinae. Adult individuals of these subfamilies either consume the dung on site or remove it downward or horizontally for eventual consumption by the adult or by subsequent larval progeny (Halffter & Matthews 1966). Current interest in dung scarabs is primarily in the following areas:
(1) Secondary disperser of seeds, initially consumed by herbivores with subsequent elimination of seeds in feces and then subsequent
capture and dispersal by dung scarabs. Recent studies in the tropical forests of Southeast Asia (McConkey 2005), Central and South America (Lawson et al. 2012, Santos-Heredia et al. 2010), and in temperate forests (Koike et al. 2012) and grasslands (D’hondt et al. 2008) all document the significant role of dung scarabs as secondary seed dispersers. Their associated subsequent involvement in the effects of forest fragmentation in the tropics also has been considered (Nichols et al. 2007).
(2) Dung scarabs as important surrogates for detecting changes in tropical forest biodiversity. Their use in analyses has lead to conservation recommendations (Nichols et al. 2009, Gardner et al. 2008, Kohlmann et al. 2007), as well as being used in a broader role as bioindicators (McGeoch et al. 2002). They also can be used as indicators in savannaagroecosystems of pest control practices (Davis et al. 2004).
(3) The role of biotic and abiotic factors in the organization and function of communities and
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their evolutionary impact has been illustrated by numerous studies of dung scarab communities in many parts of the world (reviewed in Hanski & Cambefort 1991). Dung scarabs as providers of ecosystem services also has been documented (Nichols et al. 2008).
(4) Allometry of secondary sexual characteristics (i.e., horns, legs) and its relationship to fitness has received much recent attention using various dung scarab species as models (House & Simmons 2012, Snell-Rood et al. 2010, Parzer & Moczek 2008)
(5) Sperm competition and genitalic modifications in some dung scarabs have provided insight into the evolutionary consequences of such characters (Parker et al. 2012, House & Simmons 2005, Simmons & Kotiaho 2002).
(6) Medical studies, to include dung scarab relationship to arbovirus vectors (Bishop et al. 2005), as intermediate hosts for nematode parasites of dogs (Du Toit et al. 2012) and monkeys (Boze et al. 2012), and as carriers of nematodes that will consume the scarabs after death (Herrmann et al. 2006).
Competition, predation, and abiotic factors have long been considered to be the major influences on community organization and biodiversity (Holt 2009, Price 2003). Numerous studies have considered the various parameters, such as climate or competition, that influence a particular portion - Scarabaeidae - of dung beetle populations and communities (reviewed in Halffter & Matthews 1966, Hanski & Cambefort 1991, Scholtz et al. 2009, and Simmons & Ridsdill-Smith 2011). For dung scarabs, numerous studies have documented the importance of competition as factors influencing behavior and ecology, both intraspecific (e.g., Kingston & Coe 1977, Matthews 1963) and interspecific (e.g., Bernon 1981, Young 1978), as well as both (e.g., Heinrich & Bartholomew1979). One potentially important parameter -- predation -- has typically been afforded little attention, however, with the result that published evidence of predation is either anecdotal or non- existence for most dung scarab communities. Major field studies of the dung scarab communities in Scandinavia (Landin 1961), France (Lumaret 1978), Kenya (Kingston 1977), Zaire (Walter 1978), Ivory Coast (Cambefort
1984), and the United States (Mohr 1943) make no mention of predation. Other community studies make passing mention of predation, with no specific information; e.g. South Africa (Davis 1977), Australia (Ridsdill-Smith & Hall 1984), Panama (Gill 1984). Reviews of the biology, ecology, behavior, and/or evolution of dung scarab taxa and communities also have little if anything to say about the incidence or role of predation on dung scarabs (e.g., Ridsdill-Smith 1991, Hanski 1989, Zunino & Halffter 1988, Doube 1987, Halffter & Edmonds 1982, Lumaret 1980, Macqueen 1975, Halffter & Matthews 1966, Howden 1955). One review of the ecology, behavior, and evolution of dung scarabs in 1991 had little to say about predation, other than to mention some anecdotal reports and to indicate that predation is probably more important than such reports indicate (Hanski & Cambefort 1991).
Of two more recent reviews, one makes passing reference to potential predation (Scholtz et al. 2009), and the other does not mention predation on dung scarabs (Simmons & Ridsdill-Smith 2011). It is the intent of this literature review to document that predation on dung scarabs by vertebrates and invertebrates does indeed occur, to comment on the quality of the available data, and to make available to researchers the complete database (Excel files and associated citations) developed during the review.
The taxonomy of the various beetle groups associated with dung has been in a state of flux for at least the 140 years represented by the data presented herein. For the sake of convenience, I have continued what may now be an outdated taxonomy by including the Scarabaeinae, Aphodiinae, and Geotrupinae as subfamilies within the Scarabaeidae family (e.g., Lobo & Martin-Piera 1999), and considered them all to be “dung scarabs”.
The quality of the data presented is quite variable. An author may make a statement about dung scarabs suffering predation, but fail to mention where, or when, or how it occurred, or even what specific animal was the predator. I have included such incomplete data because it still demonstrates that predation has occurred. The source of the data can be from observations of the predator in
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action, from a nest or burrow, from internal analysis (stomach, gut, oesophagus, crop, gizzard) of dead specimens, or from the scat, dung, feces (faeces), droppings, or pellets of predators. When an author merely cited prey as dung beetles, I have placed that designation in quotation marks, to highlight the possibility that the prey may have included beetles other than dung scarabs. More specific prey data may be recorded as --Dung scarabs--, or as one or more of the three scarab subfamilies. When just a genus (and sometimes species) is listed as associated with a particular vertebrate, that taxon is in the Coleoptera: Scarabaeidae: Scarabaeinae, Aphodinae, or Geotrupinae. When more than one species in a genus is listed, the genus name is not repeated for the second and subsequent species.
VERTEBRATESBirds
(Tables 1 and 2)
A review of the available information suggests that birds may be the most important predators of dung arthropods in some situations: e.g. Red-billed Chough, Pyrrhocorax pyrrhocorax, and Red Kite, Milvus milvus, at sheep dung in Great Britain (Gatehouse & Morgan 1973; Davis & Davis 1981); Tawny Owl, Strix acluo, in England (Southern 1954); plovers at cattle dung in Tasmania (Green 1966); Marabou Stork, Leptoptilos crumeniferus, at elephant dung in Africa (Seibt & Wickler 1978); Crowned Guinea Fowl, Guttera edouardi, in South Africa (Paschalides 1974); corvids at sheep and cattle dung in Australia (Rowley & Vestjens 1973; Vestjens & Carrick 1974); corvids at cattle dung in Britain (Lockie 1956); corvids and starlings at cattle dung in Poland (Goljan 1953); and Snipe, Capella gallinago, and Western Meadowlark, Sturnella neglecta, at cattle dung in California (White & Harris 1966; Anderson & Merritt 1977). Most of the above-mentioned studies, however, were oriented toward avian behavior/ecology and provide very little information relative to more specific predation on dung scarabs.
The avian literature does provide considerable information concerning the various foraging techniques utilized by birds to obtain beetles and other arthropods from dung. The Common
Crow, Corvus brachyrhynchos, has been observed turning over and searching under cow dung in California, with the stomachs of some individuals containing as many as 50 Aphodius (Kalmbach 1918). Other species have also been observed performing the same behavior: turning over cattle, horse, or camel droppings with the bill or feet, and searching beneath for prey. These bird species include: Pinyon Jay, Gymnorhinus cyanocephala, in Montana (Cameron 1907); Brewer’s Blackbird, Euphagus cyanocephalus, in western U.S.A. (Bent 1958); Hoopoe, Upupa epops, in Africa (Skead 1950); Magpie, Pica pica, in Europe (Waite 1984); Red-billed Chough in Britain (Meyer 1990); and the Jackdaw, Corvus moredula, Rook, Corvus frugilegus, and Carrion Crow, Corvus corone, in Britain (Lockie 1956).
Because many dung scarabs immediately upon arrival burrow into the animal dropping, potential avian consumers must somehow penetrate the dropping to obtain prey. This can be done by either breaking open the dropping or by probing into the dropping. A variety of bird species have been observed breaking dried animal droppings with their bills and/or feet and consuming the exposed dung beetles. These include: Houbara Bustard, Chlamydotis undulata (Cramp 1980); DuPont’s Lark, Chersophilus duponti (Smith 1965); Shore Lark, Eremophila alpestris (Bent 1942); and the Black-crowned Tchagra, Tchagra senegala (Somerev 1956). Some avian species, such as the Alpine Chough, Pyrrhocorax graculus, in Spain, are even more specialized in this regard, tending to feed only at dung previously broken open by other species (Cramp & Perrins 1994a). The Jackdaw often forages in company with the Rook, moving onto a dung patch after the Rook has broken up and scattered the material (Lockie 1956).
Other bird species obtain dung insects by probing deeply in the material. Sandhill cranes, Grus Canadensis, during the autumn in Michigan may probe in every fresh and old cow pat in a pasture, searching for both seeds and insects (Walkinshaw 1949). Other species that have been observed probing include the Hoopoe (Cramp 1985) and the Curlew, Numenius arquata (Hibbert-Ware & Ruttledge 1944).
Some dung beetles arrive at an animal dropping
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by flying into the vicinity of the prospective source, alighting on the adjacent ground, and walking onto the dropping as it arrives. These beetles are potentially exposed to bird predators that are attracted to animal dropping but forage on the ground in the immediate vicinity. Bird species employing such a technique of dung beetle capture include the Red-necked Nightjar, Caprimulgus ruficollis, and the Woodcock, Scolopax rosticola (Cramp 1985, 1983). Probably the most common foraging technique employed by birds at animal droppings is merely the walking onto the fecal material, picking up beetles from the dung surface or probing shallowly for moving forms: e.g., Wheatear, Oenanthe oenanthe (Roberts 1934), Redwing, Turdus iliacus (Tye 1981), and Nuthatch, Sitta europaea (Niethammer 1937).
Most dung scarabs arrive at animal droppings by flight, and thus may be potential prey of aerial predating birds or by birds that glean beetles from a surface without alighting. For many swallows, the most frequently captured Coleoptera are the small dung beetles (Aphodius) that fly low over cow droppings in the evenings; the stomach of one Tree Swallow, Iridoprocne bicolor, contained over 100 Aphodius, another stomach of a Bank Swallow, Riparia riparia, was recorded with 50 Aphodius inquinatus (Beal 1918). Another species that may focus on Aphodius dung beetles is the Mountain Bluebird, Sialia currucoides. One study in Utah that examined the stomach contents of 194 individuals collected over a 14 year period between April and October demonstrated that Aphodius distinctus was the most frequently consumed insect species and was even more frequently consumed than any other family of insects except the Formicidae (ants) (Knowlton & Harmston 1946). Given the fly-catching and gleaning skills of many warblers, it is not surprising that fifteen species of warblers on a Maryland farm were recorded with Aphodius spp. in their stomach (Judd 1902). Demonstrating a very aggressive form of this foraging technique, the White-breasted Kingfisher, Halcyon smyrnensis, has been observed active for more than an hour after dusk, pursuing and capturing in flight the large dung scarab Scarabaeus sacer (Sage 1960).
Avian predators such as hawks, owls, and
shrikes, while typically consumers of small vertebrates, apparently use the same hunting techniques when dung beetles are available. While flying over a likely source of food (animal droppings) or from an adjacent perch, and a dung beetle is observed, the bird will drop down to capture it in its feet or bill, and either consume it on the spot or fly away to consume it elsewhere or feed to its young; e.g., Red-backed Shrike, Lanius collurio (Mann 1983); Tawny Owl (Kirk 1992), Kestrel, Falco tinnunculus (Davis 1975). Dung beetles may actually constitute a major portion of such predators’ diets, as is the case for juvenile Burrowing Owls, Speotyto cunicularia, in the late summer in Iowa (Errington & Bennett 1935), and Kestrels during winter and spring in Britain (Fairley 1973). Burrowing Owls, Athene cunicularia, in Washington may improve their chances of obtaining dung beetle prey by collecting dried mammal feces and placing it around their nesting burrows as attractants for dung beetles (Smith & Conway 2007). One unusual technique is sometimes employed by the Stone Curlew, Burhinus oedicnemus, which captures Typhaeus typhoeus as the beetle is rolling rabbit feces to its burrow (Green et al. 2000). The Gull-billed Tern, Sterna nilotica, in Tanzania can also capture “huge scarabs” as they roll their balls of Wildebeest dung (Peterson 2006). There is even evidence that some bird species may dig in the soil for dung beetles. The Tawny Owl appears to be a specialist on beetles, with pellet and stomach analyses amply demonstrating consumption of dung scarabs (Cramp 1985). Pellets of this bird species containing soil, plant roots, and remains of Typhaeus typhoeus have been obtained near excavated burrows of this dung scarab species, suggesting that the bird will dig for such prey (Burton 1950).
Other specialized foraging techniques employed by birds include the waiting at dung heaps for arthropods to arrive: e.g., Bee-eater, Merops apiaster (Korelov 1948); House Bunting, Emberiza striolata (Cramp & Perrins 1994b); and the following of animals waiting for defecation to occur and the subsequent dung arthropods to arrive; e.g., White-breasted Kingfisher, (Ali & Ripley 1970); Dotterel, Charadrius morinellus (Glutz et al. 1975); Spotless Starling, Sturnus unicolor (Peris 1980).
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In Russia, a variety of bird species have been routinely observed foraging (no specific technique described) in the droppings of domestic animals, particularly horses and camels, for undigested seeds and for dung beetles that have been attracted. These species include: Turnstone, Arenaria interpres; Manchurian Pheasant, Phasianus colchicus; Pander’s Ground Chough, Podoces panderi; Henderson’s Ground Chough, Podocles hendersoni; Red-billed Chough; Snow Finch, Montifringilla nivalis; Tree Sparrow, Passer montanus; and Crested Lark, Galerida cristata (Dementiev & Gladkov 1954a, 1952, 1951c).
In the broader geographic area of the Western Palearctic, many bird species have been observed foraging in the droppings of cattle, horses, and camels, presumably for seeds and dung beetles, though stomach content analyses or specific prey observations supporting this presumption are not available. These bird species include, besides those already noted: Caspian Plover, Charadrius asiaticus; Pin-tailed Sandgrouse, Pterocles alchata; White-crowned Black Wheatear, Oenanthe leucopyga; Rock Thrush, Monticola saxatilis; Blackbird, Turdus merula; Arabian Babbler, Turdoides squamiceps; Sombre Tit, Parus lugubris; Rock Nuthatch, Sitta neumayer; Alpine Chough; Brown-necked Raven, Corvus ruficollis; Fan-tailed Raven, Corvus rhipidurus; Pale Rock Sparrow, Carpospiza brachydactyla; Rock Sparrow, Petronia petronia; Pine Grosbeak, Pinicola enucleator; Snow Bunting, Plectrophenax nivalis; and Corn Bunting, Miliaria calandra (Cramp 1988, 1985, 1983; Cramp & Perrins 1994a, 1994b, 1993).
Tables 1 and 2 present 518 records of 332 bird species as predators of dung scarabs from 48 countries and 14 states within the U.S.A.
MAMMALS(Table 3)
Bats represent the mammal group with the most records of dung scarab consumption. The Greater Horseshoe Bat, Rhinolophus ferrumequinum, in England has captured dung scarabs in captivity (Coward 1907), on the ground (Ransome 1990), and in flight, some identified as Aphodius sp., Typhaeus sp., Geotrupes sterrcorarius, and G.
spiniger (Flanders & Jones 2009, Poulton 1929). The Serotine Bat, Eptesicus serotinus, in England captured dung beetles in grazed pastures (Catto et al. 1994), in some cases Geotrupes stercorarius, Aphodius rufipes and A.erraticus (Robinson & Stebbings 1993), where A. rufipes may constitute the major part of the diet (LaBee & Voute 1983). The congener Eptesicus nilssoni in Sweden also consumes dung scarabs (Rydell 1986). Leisler’s Bat, Nyctalus leisleri, in English pasture areas has captured small dung scarabs (Shiel et al. 1998, Poulton 1929), as has in England the congenera Nyctalus noctula, some identified as Aphodius spp. and Geotrupes stercorarius (Jones 1995, Mackenzie & Oxford 1995, Poulton 1929). In Africa, both the Giant Leaf-nosed Bat, Hipposideros commersoni, (Vaughan 1977) and Nycteris grandis (Fenton et al. 1981) capture large dung beetles, in some cases very large Onitis spp. In Germany various bats are predators on nocturnal dung scarab beetles on the ground (Kolb 1958).
Some species of skunks, badgers, hedgehogs, and bears are predators on dung scarabs. The Spotted Skunk, Spilogale interrupta, in Iowa was found with Canthon sp. and Aphodius sp. in scats (Crabb 1941). The Striped Skunk, Mephitis mephitis, in Illinois (Verts 1967) and the Common Skunk, Mephitis nigra, in Michigan both consumed dung scarabs, the latter with Phanaeus sp. in scats (Kelker 1937). Eurasian Badgers, Meles meles, are known to consume dung scarabs: in Denmark (Andersen 1954) and Sweden (Skoog 1970); in England Geotrupes sp., Aphodius sp. (Neal 1988, 1986); and in Poland Geotrupes sp. (Goszczynski et al. 2000). American Badgers, Taxidea taxus, consume dung scarabs (Neal 1986), while in Iowa Bolboceras sp. has been found in their scats (Errington 1937). Honey Badgers (Ratels), Mellivora capensis, in Africa (Sykes 1964) and Kenya (Kingston & Coe 1977) also consume dung scarabs. Hedgehogs, Erinaceus concolor, in Czechoslovakia had Aphodius spp. remains in feces (Obrtel & Holisova 1981), while Hedgehogs, Erinaceus europaeus, in England (Yalden 1976) and West Germany (Grosshans 1983) both had dung scarabs in stomachs, the former with Geotrupes sp. and Aphodius sp. Black bears, Ursus americanus, have been observed tipping over older bison
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pats to capture various arthropods, which should have included dung scarabs (Murie 1937), and the feces of the Sloth Bear, Melursus ursinus, has contained dung beetles (Onthophagus spp.) (Laurie & Seidensticker 1977). Many rodents are insectivores or omnivores, but only Grasshopper Mice, Onychomys sp., and voles have recorded dung scarabs in their stomachs (Frank 1967, Bailey & Sperry 1929). Of the numerous primates that eat insects (excluding humans), only the Chacma Baboon, Papio ursinus, has been documented as consuming dung scarabs (Paschalidis 1974).
In Africa, several canid and felid species have been recorded preying on dung scarabs. Both Black-backed Jackal, Canis mesomelas, and Golden Jackal, Canis aureus, especially the juveniles, were observed in Tanzania chasing and catching dung scarabs (Wyman 1967). The Black-backed Jackal and the Leopard, Panthera pardus, in the Transvaal of South Africa had Coprinae in their stomachs (Viljoen & Davis 1973). The Striped Hyaena, Hyaena vulgaris, has been observed in Tanzania turning over dung piles with its feet, chasing and jumping at flying dung beetles, and eating fresh dung of herbivores which probably contained dung scarabs (Kruuk 1976). Bat-eared Fox, Otocyon megalotis, and White-tailed Mongoose, Ichneumia albicauda, in Tanzania were observed capturing dung scarabs and the same detected in their feces. Large dung scarabs in other areas of Africa are consumed by: Aardwolf, Proteles cristatus; Aardvark, Orycteropus afer; Genet, Genetta genetta; Civet, Viverra civetta; Ratel, Mellivora capensis; and Wild Cat, Felis libyca (Malcolm 1986, Waser 1980). In England, the Fox, Vulpes vulpes, has been obtained with Aphodius rufipes, Typhaeous typhaeus, Geotrupes stercorarius and G. spiniger in stomachs (Lever 1959), while the Hoary Fox, Pseudalopex vetulus, in Brazil has been observed with the large Pinotus sp. in feces (Courtenay et al. 2005).
A mammal species that seems to be a feeding specialist on dung beetles is the Banded Mongoose, Mungos mungo, of Uganda, Africa (Neal 1970). As a group forager, members of a pack typically would fan out over an area, investigating each piece of dung left by elephant or buffalo, tearing it to pieces with the front paws, and picking out
prey with the mouth. Beetles when disturbed in dung rapidly tried to bury themselves in cracks or the moist soil underneath, but a mongoose would easily dig them out with the long claws of the front feet. Packs would also follow the trails of elephants and buffalo, feeding on the dung scarabs as they went along. Stomach analyses revealed that beetles were the most abundant food item (Neal 1970). Further studies on the Banded Mongoose in Uganda documented up to 15 individuals frequently congregating at a pile of elephant dung, feeding excitedly on dung beetles (Rood 1975). Dung beetle adults and larvae were dug out of burrows, producing excavations to a depth of several cm. Dung balls were also sometimes excavated and then thrown against rocks to crack open and the contained beetle larvae consumed (Rood 1975). The White-tailed Mongoose in the Serengeti also is an avid consumer of dung scarabs, jumping up into the air to capture the beetles as they emerge from the soil at dusk, as well as digging for them several cm below the surface (Waser 1980). Another mongoose species in Hawaii, Herpestes auropunctatus, was found with Onthophagus incensus remains in its scat (Baldwin et al. 1952). Absent are records of feeding on dung beetles from such American opportunistic generalist feeders as peccary, oppossum, armadillo, and coati, as well as subterranean feeders such as moles and shrews.
Table 3 presents 57 records of 43 mammal species as predators of dung scarabs from 17 countries and 5 states within the U. S. A.
AMPHIBIANS and REPTILES(Table 4)
Virtually all terrestrial frogs and toads consume
insects (Wright & Wright 1949). Feeding records, however, that document prey consumption at the taxonomic family level or lower are scarce. The introduced Cane Toad, Bufo marinus, in Queensland Australia feeds voraciously on beetles at dung pads (MacQueen 1975), as it probably does at many other sites around the world. In Ireland, the common frog Rana temporaria is often found with Aphodius spp. in stomachs (Blackith & Speight 1974), as are at least 10 other frog species. Several
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frog species in Pennsylvania and New York also consume dung scarabs (Frost 1924), as do several native frog species in Australia (MacQueen 1975).
Although most lizards are insectivorous (Smith 1946), there are very few records of the specific types of insects consumed. The lizard Cnemidophorus exsanguis was observed in Texas tearing off pieces of dry cow dung with its forelegs, and subsequent examination of stomach contents revealed presence of dung beetle adults and larvae (Scudday & Dixon 1973). Another species in Utah, Sceloporus graciosus, has been captured with dung scarabs in its stomach (Knowlton 1942). A second species in Utah, Uta stansburiana, uses dung piles as perches, waiting for potential prey to arrive (Newbold 2007). Eumeces multivirgatus adults and juveniles have been found under dried cow dung in Nebraska (Gehlbach & Collette 1959). Three species of cordylid lizards in South Africa have been observed with scarab dung beetles in their fecal pellets (Clusell-Trullas & Botes 2008).
There are numerous snakes that probably consume dung beetles, as many species consume Coleoptera (summarized in Ernst and Ernst 2003). Unfortunately, virtually all of the food records these authors list identified stomach material or feeding observations only to the class or ordinal level for insects. Thus there is apparently only one published record (Klinstra 1959) of snakes consuming dung beetles.
A similar situation exists with feeding records for turtles (Ernst, Lovich & Barbour 1994). The Striped Mud Turtle, Kinosternon baurii, sometimes forages in cow dung when on land, probably seeking insects such as dung scarabs. The Ornate Box Turtle, Terrapene ornate, is a frequent disturber of cow dung, and dung beetles constitute the most important staple element of their diet. It has been hypothesized that bison populations on the plains historically must had provided a constant supply of dung beetles for turtle consumption and has lead to the current pattern of beetle utilization in cow dung by the Ornate Box Turtle (Ernst, Lovich & Barbour 1994).
Table 4 presents 33 records of 31 reptile and amphibian species as predators of dung scarabs from 7 countries and 15 states within the U. S. A.
INVERTEBRATES
Several community studies of dung arthropods have documented the presence of various predatory taxa. These taxa include Arachnida, Acari, Hemiptera, Hymenoptera, Diptera, and Coleoptera (e.g., Schoenly 1983, Mohr 1943). Documentation of predation by those arthropods on dung scarab beetles is quite rare, though actual predation may not be. Taxa identified as preying on adult dung scarab beetles include spiders (Macqueen 1975, Robinson & Robinson 1970, Exline & Hatch 1934), vespid wasps (Jones 1984), asilid dipterans (Bernon 1981, Hobby 1931), reduviid hemipterans (Bernon 1981), carabid beetles (Bernon 1981, Young 1980, Shalaby 1979), histerid beetles (Tribe 1976, Paschalidis 1974, Mohr 1943), and staphylinid beetles (see below). One predatory species may even be a specialist on dung scarabs. In an especially thorough study in South Africa, Bernon (1981) was able to document the predation of at least seven species of asilid flies on dung scarabs, with one species, Hoplistomerus nobilis, an apparent specialist on several species of dung scarabs. Another study in southern Africa (Tribe 1976) documented a group of histerid species preying on small-sized adult species of dung scarabs at the interface between a dung pat and the soil, with the temporal and spatial occurrence of the histerids identical to that of the dung scarabs.
Predation on the eggs, larvae, and pupae of dung scarabs may be affected by even a greater variety of organisms. Field observations at dung include: Xylocoris galactinus (Hemiptera) preying on eggs, larvae, and pupae of scarabs in Egyptian cattle dung (Tawfik & Husseini 1971); Hister abbreviatus (Coleoptera: Histeridae) adults predacious on larvae and pupae of Aphodius fimentarius in Tadjikistan (Kryzhanovskii 1944); Sphaeridium spp. (Coleoptera: Hydrophilidae) larvae feeding on Diptera maggots and Aphodius larvae in South Dakota (McDaniel et al. 1971).
The most likely arthropod candidates for having an impact on dung scarab populations would be members of the Staphylinidae (Coleoptera). Staphylinids may constitute up to 42% of the adult insects captured at cattle dung (Peitzmeier et al. 1992). Staphylinid species may exceed by more
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than 4-fold the number of dung scarab species present in an area, with examples of the numbers of staphylinid species at cattle dung including: 3 species in Indiana (Sanders & Dobson 1966), 6 species in Western Australia (Snowball 1944), 8 species in Puerto Rico (Legner & Olton 1970), 9 species in Texas (Pratt 1912), 9 species in Hawaii (Harris et al. 1982), 14 species in Bangladesh (Kabir & Faruquzzaman 1977), 14 species in southern Mexico (Legner & Olton 1970), 15 species in South Dakota (Kessler et al. 1974), 17 species in Egypt (Hafez 1939a), 21 species in Germany (Barth et al. 1994), 21 species in Costa Rica (Legner & Olton 1970), 21 species in northern California (Merritt 1976), 21 species in Texas (Hunter et al. 1991), 26 species in Nebraska (Peitzmeier et al. 1992), 28 species in New York (Valiela 1970), 31 species in Minnesota (Cervenka & Moon 1991), 31 species in northern California (Poorbaugh et al. 1968), 39 species in Uganda (Legner et al. 1981), 100 species in South Africa (Davis et al. 1988), and 134 species in Finland (Hanski & Koskela 1977). In most cited examples, it was not indicated how many species were dung consumers or predators.
Other types of dung and decaying materials where dung scarabs occur also have been examined for presence of Staphylinidae, with similar results: 3 species at poultry manure in northern California (Peck & Anderson 1969), 3 species at yak dung in Tibet (Wu et al. 2011), 4 species at poultry manure in Hawaii (Illingworth 1923), 12 species at gopher tortoise feces in Mississippi (Lago 1991), 12 species at nests of birds of prey in Norway (Hagvar 1975), 17 species at poultry manure in North Carolina (Pfeiffer & Axtell 1980), 21 species at sheep dung in New Jersey (Wilson 1932), 28 species at bear dung in Norway (Mysterud & Wiger 1976), 31 species at compost in Sweden (Palm 1979), 35 species at human dung in the Ivory Coast, Africa (Cambefort 1991), and 54 species at small mammal nests and burrows in the Russian Crimea (Medvedev & Sosnina 1974). Staphylinids also occur at the dung of bats in North and South America, Europe and Asia (Ferreira & Martins 1999), as well as at human feces (Howard 1900).
Many dung scarab species also occur at carrion, where they may also be exposed to predatory
staphylinids. Documentation of these staphylinids at various locations and types of carrion include: 3 species at fish carrion in Ohio (Jaques 1915), 15 species at carrion in New Jersey (Shubeck et al. 1981), 18 species at fish carrion in Tennessee (Walker 1957), 25 species at carrion in Scotland (Hussey & Lane 1956), 27 species at rabbit carrion in northern Italy (Ratti 1980), 27 species at carrion in Maryland (Howden 1950), 28 species at human carrion in England (Easton & Smith 1970), 32 species at carrion in Germany (Topp et al. 1982), 33 species at carrion in England (Kaufmann 1941), 36 species at carrion in Wisconsin (Dodge 1935), 50 species at pig carrion in Georgia (Payne & King 1969), 60 species at fox carrion in England (Easton 1966), 62 species at fish carrion and rotting fruit in New York (Pirone & Sullivan 1980), 71 species at carrion in Poland (Mroczkowski 1978), and 155 species (55 predatory) at carrion in Sarawak, Borneo (Hanski & Hammond 1986).
Not all staphylinids occurring at dung are predators, however, with members of the subfamilies Proteininae and Oxytelinae typically considered to be coprophages, and members of the Omaliinae, Steninae, Paederinae, Staphylininae, Tachyporinae, and Aleocharinae considered to be carnivores (e.g., Koskela & Hanski 1977). Not all predatory staphylinids at dung attack beetles. The majority of predatory staphylinid species at dung probably are primarily consumers of larval dipterans, with relatively few species large enough or with suitable mandibular equipment to penetrate the hard exoskeleton of adult beetles (Tikhomirova 1973). Presence of staphylinid populations in dung in fact have typically thought to be associated with dipteran larvae populations, with numerous studies documenting the relationship (e.g., Davis 1994, Smith et al. 1987, Laurence 1954, Hammer 1941).
The role of staphylinids as predators on dung scarabs is thus not well established. Reviews of the biology of entomophagous insects (Clausen 1940) and entomophagous beetles (Balduf 1935) make no mention of staphylinid predation on beetles. Reviews of staphylinid biology typically make little or no mention of predation by staphylinids on dung beetles (e.g., Newton 1990, Crowson 1981, Evans 1975, Tikhomirova 1973, Voris 1934, Mank 1923). Most reports on the biology of single
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species of predatory staphylinids are also devoid of information on dung scarab predation (e.g., Bauer & Pfeiffer 1991, Aly 1990, Dennis et al. 1990, Ashe & Timm 1987, Miller & Williams 1983, Nield 1976, Drea 1966, Badgley & Fleschner 1956, Hafez 1939b, Abbott 1938). A particularly remarkable occurrence is that in 50 years of observing the behavior of dung insects, Fabre (e.g., 1918) not once mentioned staphylinid predation on dung scarabs.
Actual observations of predation by staphylinids on adult dung scarabs, however, have been reported in the literature. Frost (1916) observed Ontholestes cingulatus in Maine preying upon adult Aphodius fimentarius. Laurence (1954) noted Philonthus splendens feeding on Aphodius adults in cattle dung in England. Tikhomirova (1973) described Ontholestes tesslaus in Russia hunting for Aphodius and Onthophagus spp. on the surface of fresh cattle dung and also consuming Aphodius spp. in the laboratory. Young (1982) in Maryland demonstrated laboratory predation by Staphylinus maculosus on Canthon viridis after suspected predation in the field, and in Panama observed predation on small, medium, and large-sized scarabs at monkey dung by Eulissus chalybaeus (Young 2011, 1998,). This same species Janzen (1983) observed preying on scarabs at horse dung in Costa Rica. Cambefort (1991) mentions the staphylinid Platydracus cantharophagus as "slaughtering dung beetles in elephant dung pats", with most prey small tunnelers in the scarab tribes Oniticellini and Onthophagini. Voris (1934) reported the occurrence of Staphylinus maculosus adults in cavities under dung pats with the remains of many dung scarabs. Besides being potential prey, dung scarabs may also be beneficial to the predatory success of staphylinids on fly larvae. Laboratory and field observations indicate that the presence of burrowing scarab beetles in and below dung create burrows/tunnels that are used by staphylinids and which leads to increased mortality of fly larvae due to predation by staphylinid beetles (Valiela 1974). Wu et al.(2011) describe observations and experiments with yak dung in Tibet that demonstrate two staphylinid species, Philonthus rubripennis and Quedius liangshanensis, not only were predators on numerous dung scarabs, but that
the predation on the most common dung scarab, Aphodius erraticus, caused a significant decrease in the rate of decomposition of the yak dung. Future experiments similar to that of Wu et al. (2011) in other dung scarab communities, may indeed show a similar impact of predatory staphylinids on the entire dung ecosystem.
DISCUSSION
The preceding text and tables should establish conclusively that predation on dung scarabs by numerous vertebrates and invertebrates does occur and is a world-wide phenomena. There is another line of evidence that suggests that predation on the group has a long history. Several types of phenomena support a proposed hypothesis that visually hunting diurnal predators such as birds, mammals, lizards, etc., may have been selective agents influencing the evolution of at least some forms of dung scarabs: (1) the existence of "camouflaged" dung scarabs in Africa that look like pieces of dung (Krikken 1983), or pieces of shredded plant fiber in dung (Davis 1977); (2) the behavioral trait of thanatosis (immobility upon disturbance) in many diurnal but not nocturnal forms (Matthews 1974); (3) the presence of visual mimicry complexes among some groups of African species, suggesting that the model may be distasteful to some presumptive predator(s) (Cambefort 1991); (4) secretion of fluids by a Oniticellus species in Africa from anterior lateral segments when disturbed, fluids that are aromatic and may be distasteful to predator(s) (Davis 1977); (5) digging of breeding chambers at great depths by a Heliocopris species in Africa to possibly avoid predation by a digging mammal, the Ratel (Kingston & Coe 1977); (6) the ability to “jump” vertically in an African species, as much as 60 cm, as a possible escape or frightening mechanism (Davis 1977). Some diurnally active species may also have evolved into crepuscular forms to coincide with the time of maximum dung production by large herbivorous mammals (Tribe 1976), and coincidentally avoid some diurnal predators.
Based on the information presented in this review, one may conclude that of the various
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predatory groups discussed, birds may be the primary predators on dung scarabs and staphylinid beetles the least significant predators. That conclusion may be premature, due to the lack of a significant volume of data from presumptive major predators. That absence is due to a number of factors. Most studies that document the vertebrate consumption of arthropods, either by fecal or stomach analysis or by observation, do not identify prey further than the class or ordinal level. The common omnivorous mammals of the United States, such as raccoons, opossums, skunks, and foxes, all occur in the vicinity of dung but of the literally hundreds of studies that have examined their diets, only 5 identified prey to the family-genus-species level. Of the at least 750 species of insectivorous bats (Fenton 1992), only 8 have been identified as consuming dung scarabs, with most studies identifying prey only to order (Coleoptera) (e.g., Johnson et al. 2012, O’Neill & Taylor 1989). The same situation exists with the various insectivorous lizards, toads, and other reptiles and amphibians. This general lack of identifications to the subfamily and below appears to have a historical component, as many of the reports listed in the four tables are at least 50 years old, and the most detailed, to the family-genus-species level, are more than 90 years old. The classic bird studies of the old U.S.D.A. Biological Survey by Beal and Kalmbach and McAtee and Judd at the turn of the last century are unusual, as literally thousands of bird stomachs were examined and a cadre of U.S. National Museum taxonomists provided identifications, usually to the species level. No such level of examination is likely today for mammals or herptiles, but identification of a beetle to the family and subfamily is possible and relatively easy given the taxonomic sources now available to the professional biologist (Arnett et al. 2002, Arnett & Thomas 2000) and the lay person (White 1983). Recent developments in DNA analysis may improve the future identification of prey fragments in the digestive tract and feces of predators (e.g., Long et al. 2013, Pompanon et al. 2012, Juen & Traugott 2007).
Another conclusion that can be drawn from the present survey of predation records is that documentation of the consumption of dung scarabs
as listed herein is a substantial underestimation of predation, particularly by vertebrates, on members of the Scarabaeidae: Scarabaeinae, Aphodinae,and Geotrupinae. It can also be noted that when predation on dung scarabs is looked for, it usually can be found, such as in the studies of Neal (1970) and Rood (1975) with the Banded Mongoose, Horgan & Berrow (2004) with the Hooded Crow, Bernon (1981) with asilid fly species, and Wu et al. (2011) with staphylinid beetle species. Hopefully this review will encourage others to look more carefully at the possibility of predation on dung scarabs, and the potential role of predation in affecting the currently observed behavior and ecology of the group. An encouraging sign -- one recent research team, after identifying to the species level dung beetles as prey of crows, raised the question of predation impact on competitive relations within the dung beetle community and the effect of dung disturbance by foraging birds on the differential survival of the various species (Horgan & Berrow 2004). I am also hopeful that many investigators, realizing that I have surely missed many published predation records, and that there are probably many more records unpublished, will begin compiling addenda. The Excel files and associated bibliography are available to those who wish to create and publish those addenda.
ACKNOWLEDGMENTS
Beginning in 1975, the librarians of numerous institutions have allowed me to sit in the stacks and peruse the books, journals, reports, proceedings, and various ephemerata in their care. In particular the National Agricultural Library in Beltsville, MD and the Patuxent Wildlife Research Refuge Library in Laurel, MD have been most accommodating. Libraries at the Universities of Georgia and Maryland, and at Mississippi State University, have also provided assistance.
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Table 1. Avian (U.S.A.) predators of dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae, Aphodinae, Geotrupinae).
PART 2 – TABULAR PRESENTATION OF DATA SET, WITH LITERATURE CITED
Common Name Scientific Name Location Source Subfamily, Genus, species Citation ANATIDAE Mallard Anas platyrhynchos U.S.A. stomach Aphodius sp.; Ataenius cognatus, strigatus, sp.;
Canthon sp.; Onthophagus hecate, pennsylvanicus, sp.; Phanaeus carnifex
McAtee, 1918
ACCIPITRIDAE Mississippi Kite Ictinia misisippiensis Kansas pellets Canthon sp. Fitch, 1963 " " " " Oklahoma stomach Dung scarabs; Canthon sp.; Copris sp. Sutton, G.M.,
1939 Red-shouldered Hawk Buteo lineatus U.S.A. stomach Dung scarabs Fisher, 1893 Broad-winged Hawk Buteo platypterus U.S.A. stomach Dung scarabs Fisher, 1893 FALCONIDAE Sparrow Hawk Falco sparverius U.S.A. stomach Dung scarabs Fisher, 1893 TETRAONIDAE Ruffed Grouse Bonasa umbellus Tennessee stomach Phanaeus sp. Stafford, 1975 Greater Prairie Chicken
Tympanuchus cupido Missouri stomach Aphodius fimentarius Schwartz,1945
" " " " " Wisconsin stomach Aphodius fimentarius, distinctus Gross, 1930 Lesser Prairie Chicken Tympanuchus
pallidicinctus Texas stomach Aphodius sp. Lehman, 1941
PHASIANIDAE Bobwhite Colinus virginianus U.S.A. stomach Aphodius inquinatus; Onthophagus pennsylvanicus Judd, 1903 " " " U.S.A. stomach Aphodius sp.; Ataenius cognatus, imbricatus, sp.;
Onthophagus pennsylvanicus, tuberculifrons, sp.; Geotrupes sp.; Bolboceras sp.
Handley & Cottam, 1931
Gambel's Quail Lophortyx gambelii New Mexico crop Aphodius sp. Campbell, 1957 MELEAGRIDIDAE Turkey Meleagris gallopavo Missouri dropping Dung scarabs Dalke et al.,
1942 RALLIDAE American Coot Fulica americana U.S.A. stomach Aphodius distinctus, sp.; Onthophagus sp. Jones, 1940 CHARADRIIDAE Killdeer Charadrius vociferus Utah stomach Dung scarabs Kalmbach, 1914 SCOLOPACIDAE Long-billed Dowitcher Limnodromus
scolopaceus U.S.A. stomach Aphodius sp. Sperry, 1940
Common Snipe Capella gallinago U.S.A. stomach Aphodius distinctus, fimentarius, granarius, sp. Sperry, 1940 " " " " California stomach Dung scarabs White & Harris,
1966 PHALAROPODIDAE Wilson's Phalarope Steganopus tricolor U.S.A. stomach Aphodius sp. Wetmore, 1925 STRIGIDAE Great Horned Owl Bubo virginianus Maryland stomach Dung scarabs Judd, 1902 " " " " " Iowa pellets Aphodius sp.; Canthon sp.; Onthophagus sp.; Copris
sp.; Geotrupes sp.; Trox Errington et al., 1940
Burrowing Owl Athene cunicularia Oklahoma pellets Canthon laevis; Phanaeus carnifex, Tyler, 1983 " " " " Iowa pellets Canthon sp.; Copris sp.; Geotrupes sp.; Bolboceras
sp. Errington & Bennett, 1935
" " " " Iowa pellets Bolboceras farctum Scott, 1940 Spotted Owl Strix occidentalis Oregon pellets Bolboceras obesus Forsman et al.,
1984 CAPRIMULGIDAE Common Nighthawk Chordeiles minor SE U.S.A stomach Dung scarabs Beal et al,, 1916 ALCEDINIDAE Belted Kingfisher Ceryle alcyon U.S.A. nestling Aphodius sp. Judd, 1900 PICIDAE Yellow-bellied Sapsucker
Sphyrapicus varius Maryland stomach Aphodius sp. Judd, 1902
TYRANNIDAE Scissor-tailed Flycatcher
Muscivora forfic U.S.A. stomach Canthon lecontei, cyanellus, laevis, sp.; Onthophagus pennsylvanicus, sp.; Bolboceras farctus
Beal & McAtee, 1912
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Table 1 continued.
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Common Name Scientific Name Location Source Subfamily, Genus, species Citation Eastern Kingbird Tyrannus tyrannus Maryland stomach Aphodius sp.; Ataenius sp. Judd, 1902 " " " " U.S.A. stomach Aphodius fossor, fimetarius, ruricola, granarius,
inquinatus, runeolus, sp.; Ataenius cognatus, sp.; Canthon laevis, practicola, sp.; Onthophagus hecate, nuchicornis, sp.; Copris sp.; Bolboceras sp.
Beal & McAtee, 1912
Western Kingbird Tyrannus verticalis U.S.A. stomach Aphodius fimetarius; Canthon laevis; Onthophagus hecate, sp.; Geotrupes sp.; Bolboceras farctus
Beal & McAtee, 1912
Crested Flycatcher Myiarchus crinitus U.S.A. stomach Aphodius fimetarius, Onthophagus hecate; Phanaeus carnifex
Beal & McAtee, 1912
Eastern Wood Pewee Contopus virens Maryland stomach Onthophagus pennsylvanicus Judd, 1902 " " " " " U.S.A. stomach Aphodius fimetarius, ruricola, inquinatus, granarius,
bicolor, sp.; Ataenius sp.; Canthon lecontei; Onthophagus hecate, pennsylvanicus, sp.
Beal & McAtee, 1912
Western Wood Pewee Contopus sordidulus U.S.A. stomach Aphodius vittatus, sp.; Canthon sp. Beal & McAtee, 1912
Olive-sided Flycatcher Nuttallornis borealis U.S.A. stomach Aphodius fimetarius, Onthophagus sp. Beal & McAtee, 1912
Phoebe Sayornis phoebe Maryland stomach Aphodius fimetarius; Canthon sp.; Onthophagus pennsylvanicus
Judd, 1902
" " " U.S.A. stomach Aphodius fimetarius, granarius, inquinatus, bicolor, sp.; Ataenius cylindrus, imbricatus; Canthon laevis, viridis; Onthophagus hecate, pennsylvanicus, tuberculifrons, sp.
Beal & McAtee, 1912
Black Phoebe Sayornis nigricans U.S.A. stomach Aphodius fimetarius, granarius, ungulatus, vittatus, sp.; Canthon sp.; Onthophagus sp.
Beal & McAtee, 1912
Say's Phoebe Sayornis saya U.S.A. stomach Aphodius inquinatus, sp. Beal & McAtee, 1912
Yellow-bellied Flycatcher
Epidonax flaviventris U.S.A. stomach Aphodius fimetarius, inquinatus, sp.; Canthon viridis, Onthophagus sp.
Beal & McAtee, 1912
Acadian Flycatcher Epidonax virescens U.S.A. stomach Aphodius fimetarius, inquinatus; Onthophagus sp. Beal & McAtee, 1912
Trail's/Alder Flycatcher
Epidonax traillii U.S.A. stomach Aphodius fimetarius, granarius, inquinatus,; Canthon simplex; Onthophagus sp.
Beal & McAtee, 1912
Least Flycatcher Epidonax minimus U.S.A. stomach Aphodius fimetarius, ruricola, inquinatus, sp.; Ataenius sp.
Beal & McAtee, 1912
ALAUDIDAE Horned Lark Eremophila alpestris U.S.A. stomach Aphodius fimetarius, ruricola, inquinatus;
Onthophagus pennsylvanicus McAtee, 1905
HIRUNDINIDAE Purple Martin Progne subis U.S.A. stomach Aphodius fimetarius, inquinatus, stercorosus;
Canthon sp.; Onthophagus sp. Beal, 1918
" " " " Kansas stomach Geotrupes sp. Johnston, 1967 Tree Swallow Iridoprocne bicolor U.S.A. stomach Aphodius fimetarius, inquinatus, ruricola, vittatus,
sp.; Onthophagus hecate, pennsylvanicus, sp. Beal, 1918
Violet-green Swallow Tachycineta thalassina
U.S.A. stomach Aphodius granarius, vittatus, sp. Beal 1918
Cliff Swallow Petrochelidon pyrrhonota
U.S.A. stomach Aphodius fimetarius, granarius, inquinatus, lividus, vittatus, sp.; Ataenius sp.; Onthophagus pennsylvanicus, tuberculifrons, sp.
Beal, 1918
Barn Swallow Hirundo rustica Maryland stomach Aphodius inquinatus; Onthophagus pennsylvanicus Judd, 1902 " " " " U.S.A. stomach Aphodius fimetarius, inquinatus, prodromus, vittatus,
sp.; Ataenius cognatus; Onthophagus hecate, pennsylvanicus, tuberculifrons, sp.
Beal, 1918
Rough-winged Swallow
Stelgidopteryx ruficollis
Utah stomach Aphodius sp. Kalmbach, 1914
" " " " " U.S.A. stomach Aphodius inquinatus, sp.; Onthophagus pennsylvanicus, sp.
Beal, 1918
Bank Swallow Riparia riparia Utah stomach Aphodius sp. Kalmbach, 1914 " " " " U.S.A. stomach Aphodius fimetarius, granarius, inquinatus,
punctatus, ruricola, stercorosus, vittatus, sp.; Ataenius cognatus, sp.; Onthophagus hecate, pennsylvanicus, sp.
Beal, 1918
" " " " New York stomach Aphodius erraticus, haemorrhoidalis Stoner, 1936
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Table 1 continued.
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Common Name Scientific Name Location Source Subfamily, Genus, species Citation CORVIDAE Blue Jay Cyanocitta cristata Maryland stomach Onthophagus sp. Judd, 1902 " " " " U.S.A. stomach Phanaeus carnifex Beal, 1897 Steller's Jay Cyanocitta stelleri Utah stomach Aphodius sp. Kalmbach, 1914 Black-billed Magpie Pica hudsonia W U.S.A. stomach Dung scarabs Kalmbach, 1927 Common Crow Corvus
brachyrhynchos U.S.A. stomach Aphodius fimetarius, fossor, granarius, inquinatus,
lividus, pardalis, ruricola; Ataenius cognatus, cylindrus; Canthon chalcites, laevis, simplex; Onthophagus hecate, pennsylvanicus; Phanaeus carnifex; Copris minutus, anaglypticus, Geotrupes balyi, blackburnii, egerei, semiopacus, Bolboceras farctum; Ateuchus histeroides; Pinotus carolinus
Kalmbach, 1918
White-necked Raven Corvus cryptoleucus U.S.A. stomach Aphodius sp.; Canthon laevis, praticola, puncticollis, sp.; Phanaeus vindex; Bolboceras sp.
Aldous, 1942
TROGLODYTIDAE House Wren Troglodytes aedon Maryland stomach Aphodius sp. Judd, 1902 Bewick's Wren Thryomanes bewickii California stomach Aphodius rugifrons Beal, 1907 MIMIDAE Catbird Dumetella
carolinensis Maryland stomach Aphodius sp.; Onthophagus pennsylvanicus Judd, 1902
Long-billed Thrasher Toxostoma longirostre
Texas stomach Ataenius sp.; Onthophagus sp.; Pseudocanthon sp.; Choeridium lecontei
Cottam & Knappen, 1939
TURDIDAE Robin Turdus migratorius Utah stomach Dung scarabs Kalmbach, 1914 " " " U.S.A. stomach Aphodius alternatus, anthracinus, fossor, fimetarius,
ruricola, inquinatus, pardalis, serval, sp.; Ataenius cognatus, imbricatus, sp.; Canthon simplex sp.; Onthophagus hecate, sp.; Copris minutus; Geotrupes blackburnii, Bolboceras farctum, filicornis
Beal, 1915a
Varied Thrush Ixoreus naevius U.S.A. stomach Aphodius sp. Beal, 1915a Townsend's Solitaire Myadestes townsendi U.S.A. stomach Aphodius sp. Beal, 1915b Wood Thrush Hylocichla mustelina U.S.A. stomach Aphodius sp.; Ataenius sp.; Onthophagus striatulus,
tuberculifrons, sp. Beal, 1915b
Veery Hylocichla fuscescens U.S.A. stomach Aphodius sp.; Ataenius cognatus; Onthophagus sp. Beal, 1915b Grey-cheeked Thrush Hylocichla minima Maryland stomach Ataenius sp. Judd, 1902 " " " " " U.S.A. stomach Aphodius inquinatus, ruricola, sp.; Ataenius
ovatulus, strigatus, sp.; Onthophagus sp. Beal, 1915
Olive-backed Thrush Hylocichla ustulata U.S.A. stomach Aphodius fimetarius; Onthophagus sp. Beal & McAtee, 1912
" " " " " U.S.A. stomach Aphodius fimetarius, hamatus, inquinatus, sp.; Ataenius abditus; Onthophagus hecate, striatulus, tuberculifrons, sp.; Geotrupes sp.
Beal, 1915b
Hermit Thrush Hylocichla guttata U.S.A. stomach Aphodius fimetarius, granarius, inquinatus, pardalis, prodromus, rugifrons, sp.; Ataenius abditus, cognatus, sp.; Onthophagus tuberculifrons, sp.; Geotrupes semipuctata
Beal, 1915b
Eastern Bluebird Sialia sialis Maryland stomach Aphodius sp. Judd, 1902 " " " " U.S.A. stomach Aphodius fimetarius, fossor, granarius, inquinatus,
ruricola, sp.; Ataenius cognatus; Canthon lecontei; Onthophagus hecate, tuberculifrons, sp.; Geotrupes splendidus; Bolboceras farctus
Beal, 1915a
Western Bluebird Sialia mexicana California stomach Aphodius rugifrons Beal, 1907 " " " " U.S.A. stomach Aphodius granarius, inquinatus, lividus, pardalis,
rubidus, sp. Beal, 1915a
Mountain Bluebird Sialia currucoides Utah stomach Aphodius sp. Kalmbach, 1914 " " " " U.S.A. stomach Aphodius fimetarius, inquinatus; Onthophagus sp. Beal, 1915a " " " " Utah stomach Aphodius distinctus, granarius Knowlton &
Harmston, 1946 SYLVIIDAE Golden-crowned Kinglet
Regulus satrapa California stomach Aphodius rugifrons Beal, 1907
MOTACILLIDAE Water Pipit Anthus spinoletta U.S.A. stomach Aphodius distinctus, fimetarius, fossor, Ataenius
figurator, Onthophagus tuberculifrons, sp. Gabrielson, 1924
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Common Name Scientific Name Location Source Subfamily, Genus, species Citation LANIIDAE Loggerhead Shrike Lanius ludovicianus Maryland observ. Copris carolina Judd, 1902 “ “ “ “ Texas impaled Canthon sp. Schwarz, 1896 " " " " Illinois stomach Dung scarabs Graber et al.,
1973 " " " " Missouri pellets Phanaeus vindex; Geotrupes splendidus Kridelbaugh,
1982 " " " " U.S.A. stomach Phanaeus carnifex Bent, 1950 " " " " U.S.A. stomach Aphodius hyperboreus; Canthon sp.; Phanaeus
carnifex; Geotrupes sp. Judd, 1898
STURNIDAE Starling Sturnus vulgaris U.S.A. stomach Aphodius fimetarius, fossor, granarius, inquinatus,
stercorosus, sp.; Ataenius cognatus, sp.; Canthon laevis, sp.; Onthophagus hecate, nuchicornis, pennsylvanicus, sp.; Copris minutus, tullius, sp.; Geotrupes splendidus, sp.; Bolboceras cornigerus, farctum
Kalmbach & Gabrielson, 1921
VIREONIDAE Red-eyed Vireo Vireo olivaceus U.S.A. stomach Aphodius fimetarius, granarius; Ataenius cognatus,
sp. Chapin, 1925
Philadelphia Vireo Vireo philadelphicus U.S.A. stomach Aphodius femoralis, fimetarius, inquinatus, lividus, wolcotti, sp.
Chapin, 1925
Warbling Vireo Vireo gilvus U.S.A. stomach Aphodius fimetarius, granarius Chapin, 1925 Yellow-throated Vireo Vireo flavifrons U.S.A. stomach Aphodius femoralis, fimetarius, granarius, sp. Chapin, 1925 White-eyed Vireo Vireo griseus U.S.A. stomach Aphodius femoralis, granarius, inquinatus Chapin, 1925 Solitary Vireo Vireo solitarius U.S.A. stomach Aphodius fimetarius, granarius, inquinatus, sp. Chapin, 1925 PARULIDAE Macgillivray's Warbler Oporornis tolmiei Utah stomach Dung scarabs Kalmbach, 1914 PLOCEIDAE English Sparrow Passer domesticus U.S.A. stomach Aphodius sp. Judd, 1901 " " " " U.S.A. stomach Aphodius sp.; Ataenius sp.; Canthon sp.;
Onthophagus sp. Kalmbach, 1940
" " " " Utah stomach Aphodius sp. Kalmbach, 1914 ICTERIDAE Eastern Meadowlark Sturnella magna SE U.S.A stomach Dung scarabs Beal et al., 1916 Western Meadowlark Sturnella neglecta California stomach Aphodius granarius, rugifrons, subaeneus Bryant, 1914 " " " " California stomach Aphodius fimetarius Anderson &
Merritt, 1977 " " " " Utah stomach Aphodius granarius, distinctus Knowlton &
Maddock, 1943 Purple Grackle Quiscalus quiscula U.S.A. stomach Aphodius fimetarius, inquinatus; Ataenius sp.;
Canthon sp.; Onthophagus hecate, pennsylvanicus; Phanaeus carnifex; Onthophagus sp.; Bolboceras farctus
Beal, 1900
Brown-headed Cowbird
Molothrus ater Utah stomach Aphodius sp. Kalmbach, 1914
FRINGILLIDAE Cardinal Richmondena
cardinalis U.S.A. stomach Aphodius lividus, inquinatus; Phanaeus carnifex McAtee, 1908
Rose-breasted Grosbeak
Pheucticus ludovicianus
U.S.A. stomach Aphodius fimetarius, granarius, inquinatus McAtee, 1908
Black-headed Grosbeak
Pheucticus melanocephalus
U.S.A. stomach Aphodius inquinatus McAtee, 1908
Blue Grosbeak Guiraca caerulea U.S.A. stomach Aphodius fimetarius McAtee, 1908 Vesper Sparrow Pooecetes gramineus U.S.A. stomach Aphodius sp.; Ataenius sp. Judd, 1901 " " " " Utah stomach Aphodius sp. Kalmbach, 1914 Ipswich Sparrow Passerculus princeps U.S.A. stomach Aphodius fimetarius Judd, 1901 Savannah Sparrow Passerculus
sandwichensis U.S.A. stomach Aphodius sp. Judd, 1901
Grasshopper Sparrow Ammodramus savannarum
U.S.A. stomach Aphodius sp.; Ataenius sp. Judd, 1901
Sharp-tailed Sparrow Ammospiza caudacuta
U.S.A. stomach Aphodius sp.; Ataenius sp. Judd, 1901
White-throated Sparrow
Zonotrichia albicollis U.S.A. stomach Aphodius sp. Judd, 1901
Table 1 continued.
PREDATION OF DUNG BEETLES132
Table 1 continued.
6/30/15 5
Common Name Scientific Name Location Source Subfamily, Genus, species Citation Tree Sparrow Spizella arborea U.S.A. stomach Aphodius inquinatus, sp. Judd, 1901 Chipping Sparrow Spizella passerina U.S.A. stomach Aphodius sp. Judd, 1901 Song Sparrow Melospiza melodia U.S.A. stomach Aphodius fimetarius, inquinatus Judd, 1901 " " " " Utah stomach Aphodius sp. Kalmbach, 1914 Swamp Sparrow Melospiza georgiana U.S.A. stomach Aphodius sp.; Ataenius sp. Judd, 1901 Dickcissel Spiza americana U.S.A. stomach Aphodius sp.; Ataenius sp. Judd, 1901 McCown's Longspur Rhynchophanes
mccownii U.S.A. stomach Aphodius sp. Gabrielson, 1924
Lapland Longspur Calcarius lapponicus U.S.A. stomach Aphodius inquinatus Gabrielson, 1924 Snow Bunting Plectrophenax nivalis U.S.A. stomach Aphodius distinctus Gabrielson, 1924
Table 2. Avian (non-U.S.A.) predators of dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae, Aphodinae, Geotrupinae).
7/14/15 6
Common Name Scientific Name Location Source Subfamily, Genus, species Citation PODICIPEDIDAE Great Crested Grebe Podiceps cristatus Britain Aphodius Witherby, 1938 Black-necked Grebe Podiceps nigricollis Britain Aphodius Witherby, 1938 ARDEIDAE Common Heron Ardea cinerea Britain Aphodius; Geotrupes Witherby, 1938 " " " " Britain stomach Geotrupes stercorarius Newstead, 1908 Purple Heron Ardea purpurea Britain Geotrupinae Witherby, 1938 Little Bittern Ixobrychus minutus Britain Onthophagus Witherby, 1938 PLATALEIDAE Sacred Ibis Threskiornis aethiopicus Australia stomach Scarabaeinae; Aphodius howitti Barker & Vestjens,
1989 " " " " Australia stomach Onthophagus Vestjens, 1977 " " " " Ethiopia observ. Dung scarabs Urban, 1974 " " " " South Africa observ. Dung scarabs Hancock et al.,
1992 " " " " South Africa stomach Onthophagus; Heliocopris; Copris;
Oniticellus; Euoniticellus; Onitis; Liatongus
Clark, 1979
“ “ “ “ South Africa stomach Euoniticellus intermedius, Onthophagus gazelle, Onitis alexis
Davis, 1977
White Ibis Threskiornis molucca Australia stomach Dung scarabs Carrick, 1959 Wattled Ibis Bostrychia carunculata Ethiopia observ. Dung scarabs Hancock et al.,
1992 Hadada Ibis Hagedashia hagedasa South Africa observ. Dung scarabs Davis, 1977 Bald Ibis Geronticus eremita W. Palearctic Dung scarabs Pala, 1971 Bald Ibis Geronticus calvus South Africa observ. Dung scarabs Hancock et al.,
1992 Black Ibis Pseudibis papillosa India observ. Dung scarabs Hancock et al.,
1992 CICONIIDAE Marabou Stork Leptoptilos cruminiferus Kenya observ. Heliocopris Seibt & Wickler,
1978 ANATIDAE Wigeon Anas penelope France stomach Onthophagus Madon, 1935 Pink-eared Duck Malacorhynchus
membranaceus Australia stomach Aphodiinae Firth, 1959
ACCIPITRIDAE Black-winged Kite Elanus caeruleus Egypt stomach Dung scarabs Koenig, 1936 Goshawk Accipiter gentilis Mid Europe esophagus Geotrupes Waller, 1942 Sparrow Hawk Accipiter nisus Mid Europe Geotrupes Glutz et al., 1971 Red Kite Milvus milvus Wales pellets Geotrupes stercorarius Davies & Davis,
1973 " " " " Wales pellets Aphodius luridus, prodromus, rufipes;
Geotrupes spiniger, stercorarius, sp. Davis & Davis, 1981
Black Kite Milvus migrans Australia stomach Geotrupidae Van Tets et al., 1977
Common Buzzard Buteo buteo Britain "dung beetles" Witherby, 1938 " " " " Britain pellets Geotrupes stercorarius Collinge, 1927 " " " " Italy crop/
gizzard Geotrupes Lovari, 1974
" " " " Spain stomach Geotrupes stercorarius Manosa & Cordero, 1992
" " " " Poland Geotrupes Pinowski & Ryszkowski, 1962
" " " " Mid Europe Geotrupes Fuye & Dumast, 1911
" " " " Germany stomach "dung beetles" Schuster, 1909 Lesser Spotted Eagle Aquila pomarina Moldavia (USSR) stomach "dung beetles" Averin et al., 1971 Short-toed Eagle Circaetus gallicus Mid Europe Aphodius sordidus Tobia, 1898 " " " " " Mid Europe Geotrupes Muller, 1904 Montagu's Harrier Circus pygargus Mid Europe Geotrupes Glutz et al., 1971
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Table 2 continued.
7/21/15 7
Common Name Scientific Name Location Source Subfamily, Genus, species Citation Black Kite Milvus migrans Australia stomach Geotrupidae Van Tets et al.,
1977 Common Buzzard Buteo buteo Britain "dung beetles" Witherby, 1938 " " " " Britain pellets Geotrupes stercorarius Collinge, 1927 " " " " Italy crop/
gizzard Geotrupes Lovari, 1974
" " " " Spain stomach Geotrupes stercorarius Manosa & Cordero, 1992
" " " " Poland Geotrupes Pinowski & Ryszkowski, 1962
" " " " Mid Europe Geotrupes Fuye & Dumast, 1911
" " " " Germany stomach "dung beetles" Schuster, 1909 Lesser Spotted Eagle Aquila pomarina Moldavia (USSR) stomach "dung beetles" Averin et al., 1971 Short-toed Eagle Circaetus gallicus Mid Europe Aphodius sordidus Tobia, 1898 " " " " " Mid Europe Geotrupes Muller, 1904 Montagu's Harrier Circus pygargus Mid Europe Geotrupes Glutz et al., 1971 Hooded Vulture Necrosyrtes monachus Africa observ. Dung scarabs Brown et al., 1982 Vulturine Fish Eagle Gypohierax angolensis Congo observ. “dung beetles” Ruggiero & Eves,
1998 FALCONIDAE Peregrine Falcon Falco peregrinus Britain nest "dung beetles" Witherby, 1938 " " " " Britain pellets Geotrupes Collinge, 1927 Hobby Falco subbuteo Britain stomach Geotrupes Witherby, 1938 " " " USSR "dung beetles" Dementiev &
Gladkov, 1951a " " " England pellets Geotrupes Cramp, 1980 " " " Mid Europe Aphodius fimetarius; Geotrupes Glutz et al., 1971 Brown Falcon Falco berigora Australia stomach Scarabaeinae Barker & Vestjens,
1989 Merlin Falco columbarius Britain "dung beetles" Witherby, 1938 Kestrel Falco tinnunculus Britain stom. &
pellets Geotrupes spinipes, stercorarius Newstead, 1908
" " " Britain "dung beetles" Collinge, 1927 " " " England pellets Aphodius; Geotrupes; Typhoeus Yalden &
Warburton, 1979 " " " Italy crop/
gizzard Geotrupes Lovari, 1974
" " " Norway pellets Geotrupes stercorarius Hagen, 1952 " " " England pellets Geotrupes Davis, 1975 " " " Ireland pellets Geotrupes Fairley, 1973 " " " Mid Europe Geotrupes Bouma, 1931 Lesser Kestrel Falco naumanni Britain Ateuchus Witherby, 1938 " " " " Spain pellets "dung beetles" France & Andrada,
1971 " " " " Austria pellets Geotrupes vernalis Glutz et al., 1971 Australian Kestrel Falco cenchroides Australia stomach Scarabaeinae Van Tets et al.,
1977 Red-footed Falcon Falco vespertinus Britain Geotrupes; Ateuchus Witherby, 1938 " " " " " USSR "dung beetles" Dementiev &
Gladkov,1951a " " " " " W. Palearctic Geotrupidae Cramp, 1980 Taita Falcon Falco fasciinucha Zimbabwe pellets Aphodius Hartley et al, 1993 Pale Chanting Goshawk Melierax canorus Kenya stomach Dung scarabs Brown et al., 1982 Southern Caracara Caracara plancus Argentina pellets Aphodius, Athyreus, Canthon,
Diloboderus, Sulcophanaeus Vargas et al., 2007
PREDATION OF DUNG BEETLES134
Table 2 continued.
7/14/15 8
Common Name Scientific Name Location Source Subfamily, Genus, species Citation TETRAONIDAE Hazel Grouse Tetrastes bonasia Central Asia stomach "dung beetles" Formozov, 1934 Black Grouse Lyrurus tetrix Finland stomach Aphodius fimetarius Helminen &
Viramo, 1962 Capercaillie Tetrao urogallus Finland stomach Aphodius fimetarius Helminen &
Viramo, 1962 Ptarmigan Lagopus mutus Mid Europe stomach Aphodius Girtanner, 1880 Crested Francolin Francolinus sephaena East Africa crop "dung beetles and their larvae" Urban et al., 1986 Natal Francolin Francolinus natalensis S.E. Africa observ. "dung beetles" Urban et al., 1986 Swainson's Francolin Francolinus swainsonii South Africa observ. "dung beetles" Urban et al., 1986 “ “ “ “ South Africa observ. “dung beetles” Davis, 1977 Red-necked Spurfowl Francolinus afer Central Africa observ. "dung beetles" Urban et al., 1986 PHASIANIDAE Stubble Quail Coturnix pectoralis Australia stomach Aphodius ambiguus; Ataenius imparilis Frith et al., 1974 Manchurian Pheasant Phasianus colchicus USSR "dung beetles" Dementiev &
Gladkov, 1952 NUMIDIDAE Yellow-necked Spurfowl Pternistis leucoscepus Kenya observ. Dung scarabs Heinrich &
Bartholomew, 1979
Helmeted Guinefowl Numida mitrata Kenya observ. Dung scarabs Heinrich & Bartholomew, 1979
Crowned Guineafowl Guttera pucherani South Africa observ. Dung scarabs Paschalidis, 1974 GRUIDAE Common Crane Grus grus Wales stomach Aphodius fimetarius; Geotrupes Newstead, 1908 " " " " Mid Europe Aphodius; Geotrupes; Scarabaeus Glutz et al., 1973 Demoiselle Crane Anthropoides virgo Mid Europe Aphodius; Copris; Onthophagus;
Scarabaeus Glutz et al., 1973
RALLIDAE Coot Fulica atra Mid Europe stomach Aphodius Schlegel, 1969 OTIDIDAE Great Bustard Otis tarda Britain Geotrupes Witherby, 1938 " " " " Mid Europe Geotrupes Rorig, 1897 " " " " West Asia Coprid dung beetles Dementiev &
Gladkov, 1951b Ruffed Bustard Otis undulata Kazakhstan "dung beetles" Dementiev &
Gladkov, 1951b Ludwig's Bustard Neotis ludwigii South Africa pellets Scarabaeus Earle et al., 1988 " " " " England observ.,
lab Geotrupes vernalis Pocock, 1911
Houbara Bustard Chlamydotis undulata Canary Islands observ. "dung beetles" Cramp, 1980 Kori Bustard Ardeotis kori E. & S. Africa observ. "dung beetles" Urban et al., 1986 BURHINIDAE Stone Curlew Burhinus oedicnemus Britain Geotrupes; Scarabaeus Witherby, 1938 " " " " Mid Europe stomach Dung scarabs Glutz et al., 1982 " " " " England observ. Geotrupes stercorarius, Typhaeus
typhoeus Green et al., 2000
CHARADRIIDAE Masked Lapwing Vanellus miles Australia stomach Geotrupidae; Onthophagus australis Barker & Vestjens,
1989 " " " " Australia stomach Aphodiinae; Onthophagus Van Tets et al.,
1977 " " " " Australia stomach Aphodius spp.; Onthophagus australis Green, 1966 " " " " Australia stomach Scarabaeinae Van Tets &
Vestjens, 1973 Banded Lapwing Vanellus tricolor Australia stomach Geotrupidae; Scarabaeinae Barker & Vestjens,
1989 Wattled Plover Vanellus senegallus Africa pellets "dung beetles" Urban et al., 1986
YOUNG 135
Table 2 continued.
7/21/15 9
Common Name Scientific Name Location Source Subfamily, Genus, species Citation Lapwing Vanellus vanellus Britain Aphodius; Geotrupes Witherby, 1938 " " " Sweden faecal Dung scarabs Johansson &
Blomqvist, 1996 " " " Mid Europe stomach Aphodius; Coprini Stachowski, 1954 " " " Ukraine stomach Dung scarabs Kistjakiwskij,
1957 Black-winged Lapwing Vanellus lugubris South Africa observ. "dung beetles and their larvae" Urban et al., 1986 Greater Sand Plover Charadrius leschenaulti Australia stomach Aphodiinae; Onthophagus Van Tets et al.,
1977 " " " " " Afghanistan stomach Aphodius Burton, 1971 Lesser Sand Plover Charadrius mongolus Kirgiziya (USSR) stomach Aphodius Yanushevitch et
al., 1959 Ringed Plover Charadrius hiaticula Britain Aphodius Witherby, 1938 Little Ringed Plover Charadrius dubius Afghanistan stomach Aphodius Burton, 1971 " " " " " Ukraine stomach Aphodius; Onthophagus Kistjakiwskij,
1957 Kentish Plover Charadrius alexandrinus Afghanistan stomach Aphodius Burton, 1971 Southern Golden Plover Pluvialis apricaria Britain Aphodius Witherby, 1938 " " " " " Britain stomach Aphodius fimetarius, luridus,
prodromus, punctatosulcatus Campbell, 1946
Grey Plover Squatarola squatarola Britain Aphodius Witherby, 1938 Dotterel Eudromias morinellus Britain Aphodius Witherby, 1938 " " " Mid Europe Aphodius fimetarius Bravder, 1963 African Jacana Actophilornis africana Congo observ. “dung beetles” Ruggiero & Eves,
1998 SCOLOPACIDAE Sanderling Calidris alba Australia stomach Aphodius Lea & Gray, 1935 " " " East Europe Aphodius Cramp, 1983 Purple Sandpiper Calidris maritima Mid Europe Aphodius Madon, 1935 Ruff Philomachus pugnax Britain Aphodius Witherby, 1938 Redshank Tringa totanus Britain Aphodius; Onthophagus Witherby, 1938 " " " W. Palearctic Aphodius Cramp, 1983 " " " Germany stomach Aphodius fimetarius, conspucatus,
consputus Schuster, 1909
Greenshank Tringa nebularia Britain Aphodius Witherby, 1938 " " " W. Palearctic Aphodius Cramp, 1983 Wood Sandpiper Tringa glareola Germany pellets Geotrupes Buchholz, 1961 Lesser Yellowlegs Tringa flavipes Mid Europe Aphodius Glutz et al., 1982 Common Sandpiper Actitis hypoleucos Mid Europe stomach Aphodius Koslowa, 1961 " " " " Poland stomach Aphodius; Onthophagus Pek & Fejanina,
1961 " " " " Ukraine stomach Aphodius Kistjakiwskij,
1957 " " " " Afghanistan stomach Onthophagus Burton, 1971 Common Curlew Numenius arquata Britain Aphodius; Scarabaeus Witherby, 1938 " " " " England stomach Geotrupes stercorarius Cramp, 1983 " " " " Ireland observ. Aphodius; Geotrupes stercorarius;
Ceratophyus typhaeus Hibbert-Ware & Ruttledge, 1944
" " " " Hungary stomach Geotrupes; Typhoeus Beretzk, 1959 " " " " Germany stomach Geotrupes stercorarius, vernalis Schuster, 1909 Woodcock Scolopax rosticola W. Palearctic observ. Geotrupidae Cramp, 1983 " " " Ukraine stomach Dung scarabs Kistjakiwskij,
1957 " " " Germany stomach Aphodius fimetarius, conspucatus,
consputus Schuster, 1909
PREDATION OF DUNG BEETLES136
Table 2 continued.
7/14/15 10
Common Name Scientific Name Location Source Subfamily, Genus, species Citation GLAREOLIDAE Collared Pratincole Glareola pratincola Britain Aphodius; Copris; Geotrupes Witherby, 1938 " " " " Hungary stomach Geotrupes Sterbetz, 1974 " " " " Ukraine stomach Aphodius erratus; Caccobius
schreberi; Gymnopleurus mopsus; Onthophagus amyntas, coenobita; Oniticellus fulvus
Kistjakiwskij, 1957
" " " " Mid Europe Aphodius; Oniticellus fulvus; Onthophagus taurus
Glutz et al., 1973
Black-winged Pratincole Glareola nordmanni Britain "dung beetles" Witherby, 1938 " " " " " USSR "dung beetles" Dementiev &
Gladkov, 1951c Oriental Pratincole Glareola maldivarum Australia stomach Geotrupidae Van Tets et al.,
1969 Australian Pratincole Glareola isabella Australia stomach Scarabaeinae Vestjens, 1977 Cream-colored Courser Cursorius cursor USSR "small dung beetles" Dementiev &
Gladkov, 1951c " " " " " Germany stomach Aphodius fimetarius, conspucatus,
consputus Schuster, 1909
LARIDAE Silver Gull Larus novaehollandiae Australia stomach Aphodius; Scarabaeinae Barker & Vestjens,
1989 Black-headed Gull Larus ridibundus Britain stom. &
pellets Aphodius sp., fimetarius; Geotrupes sp., stercorarius
Newstead, 1908
" " " " " Mid Europe pellets Aphodius fimetarius Bezzel, 1976 Common Gull Larus canus Britain observ. Dung scarabs Cramp, 1983 Black-billed Gull Larus bulleri New Zealand stomach Aphodius Moeed, 1976 Crested Tern Sterna bergii Australia stomach Onthophagus McKeown, 1944 Black Tern Chlidonias niger Britain Aphodius Witherby, 1938 White-winged Black Tern
Chlidonias leucopterus Britain Aphodiinae Witherby, 1938
Gull-billed Tern Gelochelidon nilotica USSR Scarabaeus sacer Shevchenko, 1937 “” “ “ Sterna nilotica Tanzania observ. “huge ball-rolling dung scarabs” Peterson. 2006 CUCULIDAE Horsefield's Bronze-cuckoo
Chrysococcyx basalis Australia stomach Scarabaeinae Barker & Vestjens, 1989
Cuckoo Cuculus canorus Germany stomach Geotrupes Link, 1889 STRIGIDAE Long-eared Owl Asio otus Britain pellets Geotrupes spiniger, stercorarius Newstead, 1908 " " " " " Britain pellets Geotrupidae Garzon Heydt,
1968 Short-eared Owl Asio flammeus Britain Geotrupes Witherby, 1938 " " " " " USSR Geotrupes Demetiev &
Gladkov, 1951a Giant Eagle Owl Bubo lacteus South Africa pellets Circellium bacchus Avery et al., 1985 Ural Owl Strix uralensis East Europe Geotrupidae Cramp, 1985 Tawny Owl Strix aluco Britain Geotrupes spinipes, sp.; Copris
lunaris; Onthophagus ovatus, nuchicornis, sp.
Witherby, 1938
" " " " England pellets Geotrupes stercorarius; Typhaeus typhoeus
Kirk, 1992
" " " " England pellets Typhaeus typhoeus Burton, 1950 " " " " Poland pellets Geotrupes Zalewski, 1994 " " " " Poland pellets Geotrupes Gramsz, 1991 " " " " West Asia "dung beetles" Demetiev &
Gladkov, 1951a " " " " England pellets Geotrupes Southern, 1954,
1970
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Table 2 continued.
7/14/15 11
Common Name Scientific Name Location Source Subfamily, Genus, species Citation Little Owl Athene noctua Britain Aphodius; Geotrupes Collinge, 1927 " " " " Wales observ. Geotrupidae Hosking &
Newberry, 1940 " " " " Italy crop/
gizzard Geotrupes stercorarius; Bubas sp. Lovari, 1974
" " " " Germany Typhaeus; Geotrupes stercorarius Haensel & Walther, 1966
" " " " Ukraine "dung beetles" Demetiev & Gladkov, 1951a
" " " " West Asia "dung beetles" Demetiev & Gladkov, 1951a
Needle-legged Owl Ninox scutalata Manchuria Geotrupes stercorarius Demetiev & Gladkov, 1951a
Scops Owl Otus scops Central Asia Geotrupes stercorarius Kadochnikov, 1963
Tengmalm's Owl Aegolius funereus W. Palearctic Geotrupes Cramp, 1985 Chestnut-backed Owlet Glaucidium sjostedti Central Africa pellets "dung beetles" Fry et al., 1988 Burrowing Owl Pholeoptynx cunicularis Argentina observ. Scarabaeus (2 spp.) Hudson, 1874 TYTONIDAE Grass Owl Tyto capensis S.E. Africa observ. Coprine dung beetles Fry et al., 1988 Barn Owl Tyto alba Britain "dung beetles" Collinge, 1927 " " " " Australia pellets Geotrupidae; Onthophagus mniszechi,
pentacanthus Lea, 1922
" " " " Britain Geotrupes Newstead, 1908 " " " " Madagascar pellets Coprinae Goodman et al.,
1993 MOMOTIDAE Blue-diademed Motmot Momotus momotus Costa Rica observ. "burly dung beetles" Skutch, 1983 PODARGIDAE Tawny Frogmouth Podargus strigoides Australia stomach Onthophagus McKeown, 1944 CAPRIMULGIDAE Nightjar Caprimulgus europaeus Britain Aphodius; Geotrupes Witherby, 1938 " " " Britain Geotrupes stercorarius Newstead, 1908 " " " USSR "dung beetles" Demetiev &
Gladkov, 1951a " " " Germany stomach Aphodius Schlegel, 1967 " " " Hungary stomach Geotrupidae Csiki, 1905 " " " Germany stomach Geotrupes stercorarius, vernalis Schuster, 1909 Red-necked Nightjar Caprimulgus ruficollis Germany "dung beetles" Koenig, 1895 Spotted Nightjar Caprimulgus argus Australia stomach Onthophagus Lea & Gray, 1935 Swamp Nightjar Caprimulgus natalensis C. & S. Africa stomach "dung beetles" Chapin, 1939 APODIDAE Swift Apus apus Britain Aphodius merdarius Newstead, 1908 " " " Britain Aphodius Witherby, 1938 ALCEDINIDAE Laughing Kookaburra Dacelo novarguineae Australia stomach Onthophagus Cleland et al., White-breasted Kingfisher
Halcyon smyrnensis Iraq observ. Scarabaeus sacer Sage, 1960
Brown-headed Kingfisher
Halcyon albiventris S.E. Africa Coprine dung scarabs Fry et al., 1988
" " " " " South Africa observ. Dung scarabs Wickler & Seibt, 1984
Red-backed Kingfisher Todiramphus pyrrhopygius
Australia stomach Scarabaeinae Barker & Vestjens, 1989
CORACIIDAE Roller Coracias garrulus Britain Aphodius; Geotrupes Witherby, 1938 " " " Mid Europe Geotrupes vernalis, stercorosus,
stercorarius; Typhaeus typhoeus Klausnitzer, 1960
" " " Germany nest Geotrupidae Klausnitzer, 1963 " " " West Asia Copris lunaris Dementiev &
Gladkov, 1951a " " " Tadzhikistan "dung beetles" Dementiev &
Gladkov, 1951a " " " France nest box Copris umbilicatus Tidmarsh, 2003
PREDATION OF DUNG BEETLES138
Table 2 continued.
7/14/15 12
Common Name Scientific Name Location Source Subfamily, Genus, species Citation UPUPIDAE Hoopoe Upupa epops Britain Aphodius; Scarabaeus Witherby, 1938 " " " Mid Europe Aphodius; Geotrupes; Onthophagus Csiki, 1905 " " " W. Palearctic Geotrupidae Cramp, 1985 " " " Africa stomach Dung scarabs Skead, 1950 BUCEROTIDAE Red-billed Hornbill Tockus erythrorhynchus Kenya observ. Dung scarabs Heinrich &
Bartholomew, 1979
" " " " " C. & S. Africa "dung beetles" Fry et al., 1988 Yellow-billed Hornbill Tockus flavirostris Kenya observ. “dung beetles” Rasa, 1983 Decken’s Hornbill Tockus deckeni Kenya observ. “dung beetles” Rasa, 1983 CAPITONIDAE White-headed Barbet Lybius leucocephalus Central Africa "dung beetles" Fry et al., 1988 MEROPIDAE Rainbow Bee-eater Merops ornatus Australia stomach Scarabaeinae Barker & Vestjens,
1989 " " " " Australia stomach Onthophagus granulatus Lea & Gray, 1935 Golden Bee-eater Merops apiaster USSR "dung beetles" Dementiev &
Gladkov, 1951a " " " " W. Palearctic Geotrupidae Cramp, 1985 " " " " Mid Europe Onthophagus ovatus, taurus Ursprung, 1979 Little Bee-eater Merops pusillus Africa stomach,
pellet Oniticellus; Onthophagus; Phaelops; Sisyphus
Douthwaite & Fry, 1982
ALAUDIDAE Horned Lark Eremophila alpestris West Asia stomach Aphodius melanostictus, segetum Volchanetskii et
al, 1950 Steppe Lark Melanocorypha calandra East Europe Copris lunaris Dementiev &
Gladkov, 1954a Dupont's Lark Chersophilus duponti Morocco "dung beetles" Smith, 1965 Short-toed Lark Calandrella
brachydactyla U.S.S.R. Aphodius Cramp, 1988
" " " " " Mid Europe Aphodius circumcinctus, lugens Glutz & Bauer, 1985
Lesser Short-toed Lark Calandrella rufescens W. Palearctic Geotrupidae Cramp, 1988 Lesser Skylark Calandrella cinerea East Europe stomach Aphodius circumcinctus, lugens Reshetnik, 1937 Crested Lark Galerida cristata W. Palearctic "dung beetles" Cramp, 1988 " " " " East Europe "dung beetles" Dementiev &
Gladkov, 1954a Skylark Alauda arvensis W. Palearctic Geotrupidae Cramp, 1988 " " " East Europe "dung beetles" Dementiev &
Gladkov, 1954a " " " Siberia stomach Aphodius Dementiev &
Gladkov, 1954a HIRUNDINIDAE Swallow Hirundo rustica Britain Aphodius Witherby, 1938 Welcome Swallow Hirundo neoxena Australia stomach Aphodiinae Vestjens, 1977 South African Cliff Swallow
Hirundo spilodera South Africa stomach Onthophagus Keith et al., 1992
House Martin Martula urbica Britain stomach Aphodius fimetarius Newstead, 1908 Crag Martin Ptyonoprogne rupestris Mongolia Aphodiinae Cramp, 1988 MOTACILLIDAE Rock Pipit Anthus spinoletta Britain Aphodius Witherby, 1938 " " " " Norway stomach Aphodius (adults and larvae) Cramp, 1988 " " " " Germany stomach Aphodius fimentarius Schuster, 1909 Richard's Pipit Anthus novaeseelandiae Australia stomach Aphodiinae Barker & Vestjens,
1990 " " " " Kenya "dung beetles" Keith et al., 1992 Tree Pipit Anthus trivialis W. Palearctic Geotrupidae Cramp, 1988 Yellow Wagtail Motacilla flava USSR Geotrupidae Popov, 1978 Citrine Wagtail Motacilla citreola East Europe stomach Geotrupidae Cramp, 1988 Pied Wagtail Motacilla alba Britain Aphodius Witherby, 1938 " " " " East Europe stomach Geotrupidae Cramp, 1988 Grey Wagtail Motacilla cinerea Mid Europe Aphodius prodromus Pek & Fedjanina,
1961
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Table 2 continued.
7/24/15 13
Common Name Scientific Name Location Source Subfamily, Genus, species Citation CAMPEPHAGIDAE White-bellied Cuckoo-shrike
Coracina papuensis Australia stomach Scarabaeinae Barker & Vestjens, 1990
LANIIDAE Lesser Gray Shrike Lanius minor Britain Aphodius; Geotrupes Witherby, 1938 " " " " " Mid Europe Aphodius Schmidt, 1980 Great Grey Shrike Lanius excubitor Britain Geotrupes spp. Witherby, 1938 Red-backed Shrike Lanius collurio Britain Geotrupes Witherby, 1938 " " " " " Mid Europe observ. Dung scarabs; Geotrupidae Mann, 1983 " " " " " Poland observ. Geotrupes mutator, stercorarius,
vernalis Mielewczyk, 1967
" " " " " Mid Europe Dung scarabs; Geotrupidae Mansfeld, 1958 Woodchat Shrike Lanius senator USSR Aphodius Gusev & Bednyi,
1961 " " " " W. Palearctic Geotrupidae Cramp & Perrins,
1993 " " " " Germany stomach Aphodius fimetarius Schuster, 1909 Isabelline Shrike Lanius isabellinus W. Palearctic "dung beetles" Mauersberger,
1982 " " " " W. Palearctic Geotrupidae Cramp & Perrins,
1993 " " " " Kazakhstan stomach Aphodius Cramp & Perrins,
1993 " " " " Mid Europe stomach Aphodius; Geotrupes stercorarius Pek & Fedjanina,
1961 Black-crowned Tchagra Tchagra senegala Africa observ. "dung beetles" Somerev, 1956 PRUNELLIDAE Black-throated Accentor Prunella atrogularis USSR stomach Aphodius prodromus Pek & Fedyanina,
1961 Alpine Accentor Prunella collaris Spain stomach Aphodius Gil, 1928 MUSCICAPIDAE Spotted Flycatcher Muscicapa striata USSR observ. Coprinae Aleksandrova,
1956 Wheatear Oenanthe oenanthe W. Palearctic Geotrupidae Posnett & Bates,
1960 White's Thrush Zoothera dauma Australia stomach Scarabaeinae Green &
McGarvie, 1971 Southern Scrub-robin Drymodes brunneopygia Australia stomach Onthophagus Lea & Gray, 1935 Flame Robin Petroica phoenicea Australia stomach Scarabaeinae Barker & Vestjens,
1990 Scarlet Robin Petroica multicolor Australia stomach Scarabaeinae Barker & Vestjens,
1990 Dusky Robin Melanodryas vittata Australia stomach Scarabaeinae Green &
McGarvie, 1971 Eastern Yellow Robin Eopsaltria australis Australia stomach Scarabaeinae Barker & Vestjens,
1990 Rufous Whistler Pachycephala rufiventris Australia stomach Onthophagus Lea & Gray, 1935 Blue Whistling Thrush Myophonus coeruleus Central Asia stomach Aphodius prodromus Kozlova, 1949 " " " " " USSR Onthophagus; Lethrus Dementiev &
Gladkov, 1954b Grey Shrike-thrush Colluricincla harmonica Australia stomach Aphodius ambiguus Lea & Gray, 1935 " " " " " Australia stomach Scarabaeinae Green &
McGarvie, 1971 " " " " " Australia stomach Scarabaeinae Rose, 1973 " " " " " Australia stomach Scarabaeinae Barker & Vestjens,
1990 Blackcap Warbler Sylvia atricapilla Britain stomach Aphodius punctato-sulcatus Newstead, 1908 Whitethroat Warbler Sylvia communis Britain stomach Aphodius ater, fimetarius Newstead, 1908 " " " " Britain stomach Aphodius ater, erraticus, rufipes Witherby, 1938 Yellow-browed Warbler Phylloscopus inornatus USSR stomach Aphodius melanostictus Kowschar, 1979 Hebridian Song-thrush Turdus ericetorum Britain Aphodius depressus, prodromus Witherby, 1938 Alpine Ring-ouzel Turdus torquatus Britain Aphodius; Geotrupes Witherby, 1938 Blackbird Turdus merula Britain Aphodius Witherby, 1938 Black-throated Thrush Turdus ruficollis W. Palearctic Geotrupidae Cramp. 1988 Song Thrush Turdus philomelos W. Palearctic Geotrupidae Cramp, 1988
PREDATION OF DUNG BEETLES140
Table 2 continued.
7/14/15 14
Common Name Scientific Name Location Source Subfamily, Genus, species Citation Redwing Turdus iliacus W. Palearctic Geotrupidae Cramp, 1988 Mistle Thrush Turdus viscivorus W. Palearctic Geotrupidae Cramp, 1988 ORTHONYCHIDAE Logrunner Orthonyx temminckii Australia stomach Aphodius viridus; Onthophagus
sydneyensis Lea & Gray, 1935
Eastern Whipbird Psophodes olivaceus Australia stomach Onthophagus Cleland et al., 1918
" " " " Australia stomach Onthophagus Lea & Gray, 1935 TIMALIIDAE Grey-crowned Babbler Pomatostomus
temporalis Australia stomach Scarabaeinae Vestjens, 1977
" " " " " Australia stomach Scarabaeinae Barker & Vestjens, 1990
" " " " " Australia stomach Onthophagus McKeown, 1944 Chestnut-crowned Babbler
Pomatostomus ruficeps Australia stomach Scarabaeinae Barker & Vestjens, 1990
PYCNONOTIDAE Yellow-whiskered Greenbul
Andropadus latirostris Central Africa Geotrupidae Keith et al., 1992
SYLVIIDAE Brown Songlark Cinclorhamphus cruralis Australia stomach Aphodius Lea & Gray, 1935 ACANTHIZIDAE Yellow-throated Scrubwren
Sericornis citreogularis Australia stomach Aphodius granarius; Onthophagus Lea & Gray, 1935
White-browed Scrubwren Sericornis frontalis Australia stomach Aphodius sp., granarius Lea & Gray, 1935 Yellow-rumped Thornbill Acanthiza chrysorrhoa Australia stomach Aphodius Lea & Gray, 1935 MELIPHAGIDAE Red Wattlebird Anthochaera carunculata Australia stomach Scarabaeinae Barker & Vestjens,
1990 Yellow Wattlebird Anthochaera paradoxa Australia stomach Onthophagus; Scarabaeinae Barker & Vestjens,
1990 Silver-crowned Friarbird Philemon argenticeps Australia stomach Scarabaeinae Barker & Vestjens,
1990 Noisy Friarbird Philemon corniculatus Australia stomach Scarabaeinae Barker & Vestjens,
1990 Regent Honeyeater Xanthomyza phrygia Australia stomach Onthophagus Lea & Gray, 1936 Bell Miner Manorina melanophrys Australia stomach Scarabaeinae Barker & Vestjens,
1990 Fuscous Honeyeater Lichenostomus fuscus Australia stomach Scarabaeinae Barker & Vestjens,
1990 White-plumed Honeyeater
Lichenostomus penicillatus
Australia stomach Aphodius sp.; lividus Lea & Gray, 1936
Eastern Spinebill Acanthorhynchus tenuirostris
Australia stomach Onthophagus Cleland et al., 1918
White-fronted Chat Ephthianura albifrons Australia stomach Aphodius Major, 1991 PARDALOTIDAE Striated Pardalote Pardalotus striatus Australia stomach Aphodius Barker & Vestjens, EMBERIZIDAE Gray-headed Bunting Emberiza spodocephala East Asia stomach "dung beetles" Dementiev &
Gladkov, 1954a Red-headed Bunting Emberiza bruniceps W. Palearctic Geotrupidae Cramp & Perrins,
1994b FRINGILLIDAE Snow Finch Montifringilla nivalis USSR "dung beetles" Dementiev &
Gladkov, 1954a Chaffinch Fringilla coelebs Britain stomach Aphodius Theobald &
McGowan, 1916 PLOCEIDAE Tree Sparrow Passer montanus USSR "dung beetles" Dementiev &
Gladkov, 1954a House Sparrow Passer domesticus Germany stomach Aphodius Schleh, 1883
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Table 2 continued.
7/14/15 15
Common Name Scientific Name Location Source Subfamily, Genus, species Citation STURNIDAE Common Starling Sturnus vulgaris Australia stomach Aphodiinae; Onthophagus Barker & Vestjens,
1990 " " " " Australia stomach Aphodius; Onthophagus australis Green, 1966 " " " " New Zealand stomach Aphodius Moeed, 1976 " " " " Britain stomach Geotrupes stercorarius; Aphodius
fimetarius, punctatus, granarius, fossor, contaminatus
Theobald & McGowan, 1916
" " " " Scotland stomach Aphodius spp. Dunnet, 1956 " " " " W. Palearctic Geotrupidae Cramp & Perrins,
1994a Spotless Starling Sturnus unicolor Spain Geotrupidae Peris, 1980 Rose-colored Starling Sturnus roseus Mid Europe Aphodius nigrivittis; Onthophagus;
Lethrus apterus Pek & Fedjanina, 1961
Crested Myna Acridotheres cristatellus British Columbia stomach Dung scarabs Scheffer & Cottam, 1935
PTYLONORHYNCHIDAE Green Catbird Ailuroedus crassirostris Australia stomach Scarabaeinae Jarvis, 1929 CORCORACIDAE White-winged Chough Corcorax
melanorhamphos Australia stomach Geotrupidae; Scarabaeinae;
Blackburnianum Barker & Vestjens, 1990
" " " " " Australia observ. "dung beetles" MacQueen, 1975 Apostlebird Struthidea cinerea Australia stomach Scarabaeinae Barker & Vestjens,
1990 ARTAMIDAE White-browed Woodswallow Artamus superciliosus Australia stomach Aphodius lividus Lea & Gray, 1935 CRACTICIDAE White-backed Magpie Gymnorhina tibicen Australia stomach Scarabaeinae Vestjens &
Carrick, 1974 " " " " Australia stomach Onthophagus Cleland et al.,
1918 " " " " Australia stomach Onthophagus Lea & Gray, 1936 " " " " New Zealand stomach Aphodius Moeed, 1976 Pied Currawong Strepera graculina Australia stomach Scarabaeinae Jarvis, 1929 CORVIDAE Rook Corvus frugilegus W. Palearctic Geotrupidae Cramp & Perrins,
1994a " " " Poland stomach Geotrupes; Copris lunaris;
Onthophagus Gromadzka, 1980
" " " Britain stomach Aphodius ater, erraticus, rufipes Lockie, 1956 " " " Britain stomach Aphodius fimetarius; Geotrupes
stercorarius Collinge, 1910
" " " Britain stomach Geotrupes stercorarius, vernalis; Aphodius fimetarius, contaminatus, prodromus, fossor
Leigh, 1914
Carrion Crow Corvus corone Britain Aphodius fimetarius, fossor Newstead, 1908 " " " " W. Palearctic Geotrupidae Cramp & Perrins,
1994a " " " " Mid Europe Dung scarabs Glutz & Bauer, Hooded Crow Corvus corone cornix Ireland pellets,
gizzards Aphodius (14 spp.), Geotrupes spp. Horgan & Berrow,
2004 Raven Corvus corax Britain pellets Geotrupes Collinge, 1927 " " " Mid Europe Dung scarabs Looft, 1971 Australian Raven Corvus coronoides Australia stomach Scarabaeinae Rowley &
Vestjens, 1973 " " " " Australia stomach Onthophagus australis Green, 1966 Forest Raven Corvus tasmanicus Australia stomach Scarabaeinae Rowley &
Vestjens, 1973 White-naped Raven Corvus albicollis Kenya observ. "dung beetles" Seibt & Wickler,
1978 Pied Crow Corvus albus South Africa observ. Kheper subaeneus Tribe, 1976
PREDATION OF DUNG BEETLES142
Table 2 continued.
7/14/15 16
Common Name Scientific Name Location Source Subfamily, Genus, species Citation Common Jackdaw Corvus monedula West Asia stomach "dung beetles" Dementiev &
Gladkov, 1954a " " " " Britain Geotrupes stercorarius Newstead, 1908 " " " " Britain stomach Aphodius ater, erraticus, rufipes Lockie, 1956 Little Raven Corvus mellori Australia stomach Scarabaeinae Rowley &
Vestjens, 1973 Little Crow Corvus bennetti Australia stomach Scarabaeinae Rowley &
Vestjens, 1973 Torresian Crow Corvus orru Australia stomach Scarabaeinae Rowley &
Vestjens, 1973 Magpie Pica pica Britain Aphodius fimetarius, punctato-
sulcatus; Geotrupes Newstead, 1908
" " " W. Palearctic Geotrupidae Cramp & Perrins, 1994a
Slender-billed Nutcracker Nucifraga caryocatactes Britain Dung scarabs Witherby, 1938 " " " " " East Europe stomach Aphodius; Geotrupidae Kumari, 1960 British Jay Garrulus glandarius Britain Geotrupes stercorarius Newstead, 1908 Siberian Jay Perisoreus infaustus W. Palearctic Geotrupidae Cramp & Perrins,
1994a Red-billed Chough Pyrrhocorax pyrrhocorax Britain "dung beetles" Witherby, 1938 " " " " " England pellets,
faeces Dung scarabs; Geotrupidae Meyer, 1990
" " " " " Armenia stomach "dung beetles" Dementiev & Gladkov, 1954a
" " " " " Britain stomach Aphodius (adults and larvae) Warnes & Stroud, 1988
" " " " " W. Palearctic Geotrupidae Cramp & Perrins, 1994a
" " " " " N. Wales stomach Aphodius Gatehouse & Morgan, 1973
" " " " " Wales faeces Aphodius contaminatus, merdarius; Geotrupes
Roberts, 1982
" " " " " Wales faeces "dung beetles" Kerbiriou & Julliard, 2007
Pander's Ground Chough Podoces panderi Turkmenia "dung beetles" Dementiev & Gladkov, 1954a
Henderson's Ground Chough
Podoces hendersoni USSR stomach "dung beetles" Dementiev & Gladkov, 1954a
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TABLE 3. Mammal predators of dung beetles (Coleoptera:Scarabaeidae, Aphodidae, Geotrupidae).
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Common Name Scientific Name Location Source Subfamily, Genus, species Citation Insectivora: Chiroptera Noctule Bat Nyctalus noctula Britain observ. Geotrupes stercorarius Oldham, 1901 " " " " England feces Aphodius spp. Mackenzie & Oxford, 1995 " " " " England observ. Geotrupes spp. Norton, 1883 " " " " England observ. Geotrupes stercorarius Patterson, 1898 “ “ “ “ England droppings Aphodius sp. Jones, 1995 Leisler's Bat Nyctalus leisleri England feces Dung scarabs Shiel et al, 1998 " " " " England observ. "small dung beetles" Moffat, 1900 Greater Horseshoe Bat Rhinolophus ferrum-
equinum Britain pellets;
observ. Geotrupes spiniger, stercorarius, typhaeus
Coward, 1907
" " " " " England feces Aphodius sp.; Geotrupes sp. Flanders & Jones, 2009 " " " " " England feces Geotrupes spp. Poulton, 1929 “ “ “ “ “ England ground Dung scarabs Ransome, 1990 Serotine Bat Eptesicus serotinus England droppings,
observ. Geotrupes stercorarius; Aphodius rufipes, erraticus
Robinson & Stebbings, 1993
" " " " Germany observ. Geotrupes sp. Poulton, 1929 “ “ “ “ England observ. “dung beetles” Catto et al., 1994 “ “ “ “ England droppings Aphodius rufipes LaBee & Voute, 1983 Northern Bat Eptesicus nilssoni Sweden feces Dung scarabs Rydell, 1986 African Bat Nycteris grandis Zimbabwe scat Onitis sp. Fenton et al., 1981 Giant leaf-nosed Bat Hipposideros
commersoni Africa observ. "large dung beetles" Vaughan, 1977
“various bats” “various bats” Germany ground Dung scarabs Kolb, 1958 Carnivora: Procyonidae Raccoon Procyon lotor USA; Iowa scat Canthon sp.; Copris
anaglypticus Giles, 1940
Carnivora: Hyaenidae Aardwolf Proteles cristatus Africa "dung beetles" Waser, 1980 Carnivora: Ursidae Sloth Bear Melurus ursinus India feces Onthophagus sp. Laurie & Seidensticker, 1977 Black Bear Ursus americanus USA. observ. “dung beetles” Murie, 1937 Carnivora: Viverridae Mongoose Herpestes
auropunctatus Hawaii scat Onthophagus insensus Baldwin et al., 1952
Banded Mongoose Mungos mungo Uganda observ. "dung beetles" Rood, 1975 " " " " Uganda observ. "dung beetles" Neal, 1970 White-tailed Mongoose Ichneumia albicauda Tanzania scat, observ. Dung scarabs Waser, 1980 Mongoose Mongoose Kenya observ. Dung scarabs Heinrich & Bartholomew, 1979 Dwarf Mongoose Helogale undulata Kenya observ. “dung beetles” Rasa, 1983 Meerkat Suricata suricata South Africa observ. Dung scarabs Davis, 1977 Common Genet Genetta genetta Africa "dung beetles" Waser, 1980 Large-spotted Genet " tigrina Africa "dung beetles" Waser, 1980 African Civet Viverra civetta Africa "dung beetles" Waser, 1980 Carnivora: Mustelidae Striped Skunk Mephitis mephitis USA; Illinois stomach Aphodius sp.; Ataenius sp.;
Geotrupes sp.; Bolbocerosoma sp.
Verts, 1967
Skunk Mephitis nigra USA; Michigan scat Phanaeus sp. Kelker, 1937 Spotted Skunk Spilogale interrupta USA; Iowa scat Aphodius spp.; Canthon spp. Crabb, 1941 Marten Martes martes Poland scat Geotrupes Goszczynski, 1976 Mink Mustela vison Ireland stomach Geotrupes stercorarius Sleeman & Hutton, 2005 European Badger Meles meles Italy scat Coprinae; Geotrupes niger,
spiniger, spp. Pigozzi, 1991
" " " " Denmark stomach Geotrupes sp. Andersen, 1954 “ “ “ “ Poland feces Geotrupes sp. Goszczynski et al., 2000 “ “ “ “ England stomach Geotrupes sp., Aphodius sp. Neal, 1986, 1988 “ “ “ “ Sweeden stomach Dung scarabs Skoog, 1970 Badger Taxidea taxus USA; Iowa scat Bolboceras sp. Errington, 1937
PREDATION OF DUNG BEETLES144
Table 3 continued.
7/19/15 18
Common Name Scientific Name Location Source Subfamily, Genus, species Citation Honey Badger (Ratel) Mellivora capensis Kenya observ. Heliocopris sp. Kingston & Coe, 1977 " “ “ " " Africa "dung beetles" Waser, 1980 “ “ “ “ “ Africa observ. Dung scarabs Sykes, 1964 Hedgehog Erinaceus europaeus Germany stomach Geotrupes sp. Grosshans, 1983 Hedgehog Erinaceus europaeus England stomach Aphodius sp.; Geotrupes
stercorarius Yalden, 1976
Hedgehog Erinaceus concolor Czechoslovakia scat Aphodius sp. Obrtel & Holisova, 1981 Carnivora: Canidae Fox Vulpes vulpes Britain scat, stomach Aphodius rufipes; Geotrupes
stercorarius Lever, 1959
Swift Fox Vulpes velox USA; Oklahoma
scat Canthon laevis; Bolbocerosoma bruneri
Kilgore, 1969
Bat-eared Fox Otocyon megalotis Tanzania scat, observ. Dung scarabs Waser, 1980 “ “ “ “ “ “ feces Scarabaeinae Malcolm, 1986 Hoary Fox Pseudalopex vetulus Brazil scat Pinotis sp. Courtenay et al., 2006 Striped Hyaena Hyaena vulgaris Tanzania observ. "dung beetles" Kruuk, 1976 Wolf Canis lupus Italy scat Geotrupes stercorarius Meriggi et al., 1991 Black-backed Jackal Canis mesomelas Tanzania observ. "dung beetles" Wyman, 1967 " " " " " Botswana stomach Coprinae Viljoen & Davis, 1973 Golden Jackal Canis aureus Tanzania observ. "dung beetles" Wyman, 1967 Carnivora: Felidae Leopard Panthera pardus Transvaal stomach Coprinae Viljoen & Davis, 1973 African Wild Cat Felis libyca Africa "dung beetles" Waser, 1980 Serval Felis serval Africa "dung beetles" Waser, 1980 Caracal Felis caracal Africa "dung beetles" Waser, 1980 Rodentia: Cricetidae Grasshopper Mice Onychomys sp. USA stomach Onthophagus sp.; Eucanthus
lazarus Bailey & Sperry, 1929
Vole Vole England stomach Dung scarabs Frank, 1967 Rodentia: Dipodidae Jerboa Jaculus jaculus Morocco observ. Scarabaeus cristatus Sanchez Pinero, 2007 Artiodactyla: Suidae Wild Boar Sus scrofa Poland stomach Aphodius subterraneus Genov, 1981 Primates Chacma Baboon Papio ursinus South Africa feces Scarabaeinae Paschalides 1974
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Table 4. Amphibian and reptile predators of dung beetles (Coleoptera: Scarabaeidae, Aphodidae, Geotrupidae).
7/2/15 18
Common Name Scientific Name Location Source Subfamily, Genus, species Citation Amphibia:Anura Red-legged Frog Rana aurora California stomach Canthon simplex Hayes & Tennant, 1985 Bullfrog Rana catesbeiana Missouri stomach Dung scarabs Korschgen & Moyle, 1955 Green Frog Rana clamitans Pennsylvania stomach Aphodius fikmetarius; Geotrupes
blackburnii Surface, 1913
Spotted Frog Rana pretiosa Wyoming stomach Aphodius opacus Turner, 1959 " " " " Oregon stomach Canthon sp. Schonberger, 1945 Dusky Gopher Frog Rana areolata sevosa Alabama stomach Canthon sp. Netting & Goin, 1942 Common Frog Rana temporaria Ireland stomach Aphodius spp. Blackith & Speight 1974 Swamp Cricket Frog Pseudacris nigrita W. USA stomach Aphodius sp. Stebbins, 1951 Colorado River Toad Bufo alvarius Arizona stomach Copris remotus; Onthophagus
velutinus Cole, 1962
Great Plains Toad Bufo cognatus Oklahoma stomach Aphodius granarius Smith & Bragg, 1949 Northern Sonora Toad Bufo compactilis Oklahoma stomach Aphodius granarius Smith & Bragg, 1949 Giant Cane Toad Bufo marinus Fiji stomach Aphodius sp.; Copris incertus Hinckley, 1962 “ “ “ “ “ Australia stomach Dung scarabs MacQueen 1975 Oak Toad Bufo quercicus Florida stomach Ataenius gracilis Hamilton, 1954 American Toad Bufo terrestris Oklahoma stomach Aphodius granarius Smith & Bragg, 1949 " " " " New York stomach Dung scarabs Hamilton, 1954 Rocky Mountain Toad Bufo w. woodhousii Oklahoma stomach Aphodius granarius Smith & Bragg, 1949 Fowler's Toad Bufo w. fowleri Georgia stomach Ataeniussp; Aphodius sp.;
Ateuchus sp. Bush & Menhinick, 1962
Western Spadefoot Toad Scaphiopus hammondi Texas stomach Ataenius cognatus; Aphodius terminalis
Whitaker et al., 1977
Eastern frogs Six species NY, Penn stomach Ataenius cognatus; Aphodius fimetarius
Frost, 1924
Native frogs Several species Australia stomach Dung scarabs MacQueen 1975 Reptilia: Testudinata Striped Mud Turtle Kinosternon baurii U.S.A. observ. Dung scarabs Ernst et al., 1994 Eastern Box Turtle Terrapene carolina Illinois stomach Aphodiinae; Canthon sp. Klinstra & Newsome,
1960
Ornate Box Turtle Terrapene ornata Kansas stomach Dung scarabs Legler, 1960 “ “ “ “ “ U.S.A. stomach Dung scarabs Ernst et al., 1994 Reptilia: Serpentes Racer Coluber constrictor Illinois stomach Canthon sp.; Geotrupes sp. Klinstra, 1959 Reptilia: Sauria Basilisk Lizard Basiliscus basiliscus Panama stomach Dung scarabs Barden, 1943 Lizard Cnemidophorus exsanguis Texas stomach,
observation Dung scarabs (adult and larvae) Scudday & Dixon 1973
Lizard Amphisbaena ridleyi Brazil observ. Dung scarabs Finley, 1945 Lizard (3 species) Cordylids South Africa feces Dung scarabs Clusell-Trullas & Botes
2008
Sagebrush Swift Sceloporus graciosus Utah stomach Aphodius granarius, distinctus Knowlton, 1942
PREDATION OF DUNG BEETLES146
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