PROC. ENTOMOL. SOC. WASH. 85(4), 1983, pp. 633-652 STUDIES ON THE GENUS APHODIUS OF THE UNITED STATES AND CANADA (COLEOPTERA: SCARABAEIDAE). VII. FOOD AND HABITAT; DISTRIBUTION; KEY TO EASTERN SPECIES Robert D. Gordon Systematic Entomology Laboratory, IIBIII, Agricultural Research Service, USDA, % National Museum of Natural History, Washington, D.C. 20560. Abstract.— CuYVQnXXy available information on food and habitat preferences and geographic and temporal distribution is presented, along with a discussion of the factors influencing the distribution of those species associated with deer dung. A key to identify adults of Aphodius species that occur in the eastern forest region is constructed. Although approximately 2 1 species of Aphodius are described from North America (north of Mexico), very little has been published concerning food and habitat preferences. Most published works on the genus deal only with taxonomy, and, in most cases, biological data have not been available. Over the past 12 years, I have collected biological data for species of Aphodius, and these, along with data from similarly inclined colleagues, constitute the basis for this report. There is still relatively little known, and one of the reasons for preparing this publication is to stimulate the collection of additional data. The eastern forest geographic area covered here is not a well defined unit, being based in part on the prevailing distribution patterns of the eastern Aphodius. The eastern forest region as considered here is illustrated in map 1. All species of Aphodius having primary distributions within these limits are treated even though some extend into the Midwest and West. Those species whose primary distri- butions lie west of the limits but extend into the eastern forest region are not treated except in the key to species. All of the imported or holarctic species in the region are included even though the distribution patterns often extend across the continent. Most of the data presented here are previously unpublished and are the result of personal field work and information supplied by fellow workers such as the late O. L. Cartwright (Department of Entomology, Smithsonian Institution, Wash- ington, D.C), Henry Howden (Carleton University, Ottawa, Canada), and Robert Woodruff' (Florida Department of Agriculture, Gainesville, Florida). Published information has been obtained from Cartwright (1957), Robinson (1948), Sim (1930), Brown (1927), Hubbard (1894), and Woodruff" (1973). I thank D. R. Whitehead for critical comments and suggestions, and W. Steiner for testing the key to species. 634 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Map L Eastern forest region. Ellipses indicate the three major areas of species distribution. Food and Habitat The eastern species of native Aphodius can be divided into five categories based on food and habitat preference as indicated in Tables 1-5. Category I is the most important in terms of numbers of species and biological significance. Category I: Species associated with deer dung (Table 1).— Seventeen species of eastern Aphodius are associated with deer dung, either in an obligate fashion or in showing a strong preference for deer dung when it is available. Aphodius odo- coilus Robinson and A. robinsoni Cartwright are examples of the former; A. ruricola Melsheimer and A. rubripennis Horn are examples of species that show a preference for deer dung, but which will utilize other dung (horse, cow, etc.) if available. A very thorough study of the ecology of Aphodius species occurring on the island of Faro in the Baltic sea was done by Landin (1961). The factors Landin VOLUME 85, NUMBER 4 635 Table 1 . Category I, species associated with deer dung. Species Geographic distribution Temporal distribution Spring: Aphodius odocoilis Robinson Aphodius brimleyi Cartwright Aphodius abusus Fall Winter: Aphodius crassulus Fall Aphodius floridanus Robinson Aphodius silvamis Cartwright Aphodius windsori Cartwright Aphodius bicolor Say Aphodius terminalis Say Aphodius lodingi Cartwright Aphodius leopardus Horn Fall: Aphodius robinsoni Cartwright Aphodius stupidus Horn Aphodius lutulentus Haldeman Aphodius manitobensis Summer: Aphodius rubripennis Horn Aphodius ruricola Melsheimer New Jersey North Carolina East Texas Virginia to Florida Florida Maryland (Eastern Shore) South Carolina to Florida New York to Florida, west to Kansas and Texas Pennsylvania to Florida, west to Kansas and Texas South Carolina and Florida, west to Texas North Carolina north to Ontario and west to British Columbia and Alaska New Jersey, West Virginia, North Carolina, South Caroli- na New Jersey and Pennsylvania to Florida, west to east Texas Maryland and Virginia south to Florida, west to Texas and Oklahoma Northeastern U.S. and eastern Canada west to Manitoba Southeastern Canada to North Carolina Ontario to Florida, west to Man- itoba and Texas March-May April-July March-April October-February January-May October-April November-March September-April November-April November-April August-June September-November October-December October-February September-December June-October May-August (most months of year) found, that influence the selection of dung by Aphodius species are, in large part, the factors discussed herein. Landin studied only dung of domestic animals, and perhaps because of that, some of his conclusions differ somewhat from my own observations. For example, Landin states that "it is not the kind of dung, but the climatic conditions of the environment (particularly the microclimatic conditions of the droppings) which restrict dung-beetles to a certain habitat." I find that the obligate deer dung species will not utilize horse or bovine dung even when it occurs in appropriately shaded areas. They will utilize sheep dung, but there 636 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Category II, species associated with rodent burrows or nests, or burrows of the gopher tortoise. Species Geographic distribution Temporal distribution Aphodius aegrotus Horn (pocket North Carolina, Florida gopher) Aphodius laevigatus Haldeman (pocket gopher) Aphodius badiceps Fall (squirrel nests) Aphodius troglodytes Hubbard (gopher tortoise) Florida Southeastern Canada to South Carolina, west to Minnesota, Kansas and Arkansas South Carolina, Florida March-September February-November February-November May-December appears to be very little difference between the dung of deer and sheep. Therefore I conclude that, under natural conditions with dung of native animals, it is often the kind of dung that restricts the beetles to a certain habitat. The moisture content and exposure of deer dung are of paramount importance in regard to selection by beetles. Very few species will use dung that is dry, and, obviously, if eggs are laid in pellets without sufficient moisture content, the larvae will not be able to mature before complete drying occurs. Whether the dung is in shade or exposed to the sun is correlated with moisture. Dung exposed to the sun will dry more quickly than if shaded, and an obvious bias is for shaded dung. The fact that deer dung normally is in the form of a pile of small pellets makes rapid drying inevitable, especially when precipitation is infrequent, making the moisture factor much more important than when horse or bovine dung is the resource. Very few species are associated with deer dung in the arid regions of western North America, and lack of reliable precipitation is probably the reason. The fact that most deer dung species occur in fall, winter, and spring is another result of moisture supply; the pellets maintain moisture much longer in cool weather. Only two deer dung species oi Aphodius are classified as summer species (Table 1), and both of these are capable of utilizing a variety of dung resources. In contrast, eight are classified as "winter" species, four as "fall" species, and three as "spring" species. This breakdown is based on available label data for adults. Several species occur in fall, winter, and spring, but the period of peak abundance usually occurs in one season. That is the season used to classify the species treated here. Aphodius leopardus Horn is an extreme example, with adults active through all except the hottest months but with peak activity in the winter. Latitude is, of course, an important factor for those species having a wide north- south distribution. For instance, A. leopardus occurs as an adult from August to June; in the southern portions of the range it is most abundant in winter, but in the northern portions of the range the population peaks in the fall. Southern winters have warm periods during which flight activity, etc., can take place, but northern winters normally remain too cold for such activity; therefore, "winter" active species do not occur in the north. The thermal factor must also be considered. The species that live in deer dung during fall and winter exhibit a "restricted thermal resistance" (Landin, 1961) VOLUME 85, NUMBER 4 637 Table 3. Category III, native generalists. Species Geographic distribution Temporal distribution Aphodius cuniculus Chevrolat Aphodius vittatus Say Aphodius femoralis Say Aphodius campesths Blatchley Aphodius stercorosus Melsheimer Aphodius rubeolus Beauvois Aphodius lentus Horn Aphodius horealis Gyllenhal South Carolina to Texas (primarily a Caribbean species) Southern Holarctic— North Ameri- ca; southern Canada, U.S. from Maine to South Carolina, west to Washington and Mexico Pennsylvania to South Carolina, west to Nebraska and east Texas New Jersey to Florida and Missis- sippi Southeastern Canada to Florida, west to Kansas and Texas Southeastern Canada to Florida, west to Nebraska and Texas Ontario to Georgia, west to North Dakota and Texas Holarctic, in North America pri- marily boreal forest All months of year All months of year April-June All months of year May-October April-August June-July April-July while the summer species and the majority of the competitor species exhibit "intermediate thermal resistance." This may be very important when attempting to explain partitioning of the resource in terms of winter versus summer. "Thermal resistance" refers to the degree of tolerance for high temperature. Wind is another factor since dung beetles search for droppings with their ol- factory organs. A light wind would aid in the location of droppings, while a strong wind would tend to prevent flight activity. A forested area would tend to favor the search for droppings because strong wind would not be a normal occurrence. Vagility may be an important factor, but data on this subject are very limited. Location of the food supply is accomplished by a searching flight. Most species are capable fliers and the search mode consists of an erratic, rapid flight close to the ground, a pattern very similar to that observed in many Diptera. Personal observations indicate that flights usually take place on overcast days, or in late afternoon, not when it is sunny and hot. A conclusion reached by Landin (1961) with which I agree entirely, is that two or more species can occupy the same "niche," in this case the dung heap. In fact this is the rule rather than the exception. Landin summarized this as follows: "Interspecific as well as intraspecific fluctuations occurring in natural populations of dung-beetles in the dung heaps depend on abiotic factors rather than on the competition factor." However, competition possibly is a factor when food re- sources are limited, as is often the case with deer dung. This competition may come from other species of Aphodius or other species of Scarabaeidae that are not Aphodius. Table 7 lists the major competition from non-Aphodius by season. Note that of the ten species on the list, five are in the summer category and three are found all year, leaving only one species in direct competition with winter 638 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 4. Category IV, native detritivores. Species Geographic distribution Temporal distribution Aphodius paleroides Horn Aphodius parens Horn Aphodius pinguis Haldman Aphodius hyperboreus LeConte New York to South Carolina (coastal sand areas) Florida Quebec to New York, west to Alberta and Montana Quebec to New York, west to British Columbia June-July May-September May-September May-September Aphodius and one in direct competition with spring Aphodius. The temporal distribution of these competitors is almost a complete reversal of the temporal distribution of Aphodius species. The first six names on the list are members of the Aphodiinae, the last four are in the Coprinae. Table 8 lists the species of deer dung Aphodius on the left and non-Aphodius competitors on the right, broken down by season. Where groupings can be indicated by brackets this has been done— where this was not possible lines are drawn directly between the species of Aphodius and the competitor. The brackets on the extreme left group those species of Aphodius that may compete with each other. For example, the two summer species of Aphodius compete with each other whenever they occur to- gether, and all of the competitor species can compete with both Aphodius species. Category II: Species associated with rodent burrows or nests, or burrows of gopher tortoise (Table 2). — In the midwestern and western United States, about 60 species, or approximately 43% of the North American species whose habitats are known, are in this category. However, only four eastern species belong here, presumably because the harsh climatic changes during the Miocene in the western part of North America did not cause the same radical vegetation changes in eastern North America. Aphodius badiceps Fall is found in tree squirrel nests, usually those of the flying squirrel; A. troglodytes Hubbard only in burrows of the gopher tortoise; and A. aegrotus Horn and A. laevigatus Haldeman in pocket gopher burrows. The three burrow inhabiting species are restricted to sand areas of the southeastern United States. Category III: Native generalists (Table 3).— The eight species in this category are general surface dung feeders, some of which are habitat restricted, but are not known to have distinct dung preferences except that they rarely utilize deer dung. Two possible exceptions to this statement are Aphodius lentus Horn and A. boreal is Gyllenhal. I have taken A. lentus in sheep dung, and most species that will utilize sheep dung are also capable of living in deer dung. Aphodius borealis is a northern holarctic species which may use deer dung, but probably also occurs in dung of moose, elk, caribou, and possibly in ground squirrel pellets. Category IV: Native detritivores (Table 4).— The eastern forest fauna in this category is analogous to the fauna of the rodent associated category in that the eastern fauna is comparatively depauperate in both. The native detritivores com- prise about 23% of the North American fauna, but only four species occur in the eastern forest region. Aphodius parcus Horn and A. phaleroides Horn occur in sand areas along the Atlantic seaboard, including Florida; A. pinguis Haldeman VOLUME 85, NUMBER 4 639 Table 5. Category V, native species, habits unknown. and A. hyperboreus LeConte occur in marshy, grassy situations in southern Canada and northern United States. Category V: Native species, habits unknown (Table 5).— Obviously little can be said about these species, but probable habits can be predicted from knowledge of the majority of the fauna. Aphodius mcdonaldi Robinson may be an early spring deer dung species; A. guttatus Escholtz probably utilizes the dung of moose, elk, and possibly deer; A. serval Say is probably a detritivore in sandy riverine situ- ations; A. fordi Gordon is an enigma, all specimens having been taken at light, but the species is morphologically most similar to the European A. scrofa (F.) which is a surface dung feeder. It is possible that this is a recent immigrant because of its apparently restricted distribution (coastal Georgia), but this is pure specu- lation at present. Table 6: Imported species o{ Aphodius; mostly generalists.— These species have the same surface dung feeding habits as the native species in Category III, but have been imported from Europe and have become established. Most are wide- spread in North America; but three species, A. scrofa (F.) and A. subterraneus (L.) are restricted to eastern seaboard localities, and A. rufipes (L.) occurs in the forested mountain regions from New York to Virginia. With the exception oi A. rufipes, all of the imports prefer open pastures and bovine dung. Species Identification The standard reference for determining species of North American Aphodius has been Horn (1887), which is still the only publication dealing solely with the Table 6. Imported species oi Aphodius, mostly generalists. Aphodius fimetarius (L.) Aphodius granarius (L.) Aphodius haemorrhoidalis (L.) Aphodius fossor (L.) Aphodius erraticus (L.) Aphodius lividus (Olivier) Aphodius distinctus (Mull.) Aphodius rufipes (L.) Aphodius prodromus (Brahm) Aphodius scrofa (F.) Aphodius subterraneus (L.) 640 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 7. Competitors for deer dung other than species of Aphodius: geographic and temporal distribution. Species Geographic distribution Temporal distribution Dialytellus humeralis (LeConte) Dialytellus dialytoides (Fall) Dialytes ulkei Horn Dialytes striatulus (Say) Dialytes truncatus (Melsheimer) Aphotaenius carolinus (Van Dyke) Onthophagus concinnus Onthophagus subaeneus (P. de B.) Onthophagus tuberculifrons Harold Copris minutus (Drury) Southeastern Canada to North Car- olina Eastern Canada to North Carolina Eastern Canada to South Carolina Eastern Canada to Georgia, west to Manitoba and Iowa Southeastern Canada to South Car- olina, west to Wisconsin and Illi- nois Maryland to Florida, west to In- diana Pennsylvania to Florida, west to Mississippi Pennsylvania to Florida, west to Kansas and Texas Connecticut to Florida, west to Wisconsin and east Texas Vermont to Florida, west to Kansas and Texas October-May (winter) May-October (summer) June-October (summer) June-October (summer) July-October (summer) June-August (summer) April-December (all year) February-June (spring) February-December (all year) February-December (all year) North American fauna as a whole. Schmidt (1922) published descriptions and keys to subgenera and species of the Aphodius fauna for the world, and included several North American species unknown to Horn in 1887. Since 1922, a few papers dealing with segments of North American Aphodius have appeared (Brown, 1927, 1928, 1929;Saylor, 1 940; Cartwright, 1972; Gordon, 1976, 1977a, 1977b), as have two significant regional publications dealing with the fauna of the Pacific Northwest (Hatch, 1972) and Florida (Woodruff, 1973). In addition, several new species descriptions also have appeared since 1922. The following key to eastern North American Aphodius includes those species restricted to, or centered in, the eastern forest region. In addition to those species, several species of primarily midwestern occurrence are included because they penetrate the eastern forest to a significant degree; these species are indicated by an asterisk (*). The key is arranged to facilitate identification, regardless of phy- logenetic relationships. One species, A. rotundiceps Fall (in Fall and Cockerell, 1907) (type locality. Highlands, North Carolina) is not included in the key nor in the foregoing discussion. The species is apparently known only from the holotype which I examined several years ago. I seriously doubt that A. rotundiceps is a member of the American fauna; more likely, the type is a mislabeled Old World specimen. Most species o^ Aphodius have been assigned to various subgenera, most notably by Schmidt ( 1 922). Most of these "subgenera" are of questionable merit. However, VOLUME 85, NUMBER 4 641 Table 8. Competition among species utilizing deer dung as an energy resource. Species Actual and probable competilors Spring species: Aphodiiis odocoilis Robinson Aphodius brimleyi Cailwright Aphodius abusiis Fall Onthophagiis tuberculifrons Harold Onthophagus subaeneus (P. de B.) Onthophagns concinnus LaPorte Copris mimitus (Drury) Dialytellus humeralis (LeConte) Summer species: Aphodius rubhpennis Horn Aphodius ruricola Melsheimer _ DialyteUus dialytoides (Fall) Dialytes ulkei (Horn) Dialytes striatulus (Say) Dialytes truncatus (Melsheimer) Aphotaenius caroUnus (Van Dyke) Onthophagus subaeneus (P. de B.) Onthophagus concinnus LaPorte Onthophagus tuberculifrons Harold Fall species: Aphodius robinsoni Cartwright Aphodius stupidus Horn Aphodius lutulentus Haldeman Aphodius manitobensis Brown Winter species: Aphodius leopardus Horn Aphodius terminalis Say Aphodius bicolor Say Aphodius silvanus Cartwright Aphodius crassulus Fall Aphodius windsori Cartwright Aphodius lodingi Cartwright Aphodius floridanus Robinson Onthophagus tuberculifrons Harold Onthophagus concinnus LaPorte Onthophagus subaeneus (P. de B.) Copris minutus (Drury) Dialytellus humeralis (LeConte) Onthophagus tuberculifrons Harold Onthophagus concinnus LaPorte Copris minutus (Drury) 642 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON those species occurring in North America that have an elongated scutellum have been placed in two subgenera, or genera, depending on author, and these may need to be separated from Aphodius proper. The subgeneric or generic names involved are: Colobopterus Mulsant, including the species fossor (L.), erraticus (L.), subtenaneus (L.), and haemonhoidalis (L.); and Diapterna Horn, including the species hyperboreus LeConte, pinguis Haldeman, and hamatus Say. The first "subgenus" contains species introduced from Europe, and the latter "subgenus" only native North American species. Similarly Aphodius parens Horn has been placed in the subgenus or genus Didactylia Orbigny, depending on the author. This species lacks the oblique, transverse ridges on the hindtibia so characteristic of most Aphodius species. This species probably deserves at least subgeneric status, but I have followed the same conservative course here in key couplet 9, of treating it as '"Aphodius.^' Key to Eastern North American Species of Aphodius 1 . Scutellum large, Vs to '/j as long as elytron (Figs. 2,3) 2 - Scutellum small, '/lo to % as long as elytron (Fig. 4) 8 2. Head with at least one median tubercle, usually with 3 tubercles (Fig. 1); surface of head roughened or densely punctured 3 - Head without trace of tubercles; surface of head smooth 6 3. Small species, less than 6.0 mm long; elytron black except apical Vs always red or yellow, humerus often red or yellow haemonhoidalis (L.) - Species longer than 6.0 mm; elytron not as described above 4 4. Elytron entirely yellow or yellowish brown except sutural margin black; surface of pronotum dull, densely punctured erraticus (L.) - Elytron entirely black or dark brown; surface of pronotum shiny, not densely punctured 5 5. Intervals on elytron flat; length 9.0 mm or more fossor (L.) - Intervals on elytron convex; length 8.0 mm or less subterraneus (L.) 6. Elytron yellow or yellowish brown with obscurely defined brown blotch medially hamatus hamatus Say - Elytron entirely brown or black 7 7. Mid- and hindfemora and median area of mesostemum coarsely punc- tured pinguis Haldeman - Mid- and hindfemora and median area of mesostemum finely punctured, appearing impunctate hyperboreus LeConte 8. Hindtibia without oblique, transverse ridges (Fig. 5) parens Horn - Hindtibia with oblique, transverse ridges (Fig. 7) 9 9. Elytron pubescent throughout; dorsal color uniformly dark brown or uniformly light brown, obscurely defined reddish areas may be pres- ent 10 - Elytron lacking pubescence, or if pubescent, then pubescence not present throughout, or if so, then pronotum and head entirely or partially brown or black, elytron red or yellow, or uniformly dull gray {lutnlentus) .... 13 10. Pronotum lacking pubescence on dorsal surface 11 - Pronotum pubescent on dorsal surface 12 1 1 . Mesostemum carinate between coxae; dorsal surface reddish brown . . lentus Horn - Mesostemum not carinate between coxae; dorsal surface dark brown to VOLUME 85, NUMBER 4 643 Figs. 1-5. 1, Aphodius fossor, head (X44). 2, A. haemorrhoidalis. scutellum (X44). 3, A. enaticus, scutellum (X60). 4, A. lutulentus, scutellum (XI 00). 5, A. parens, hindtibia (XI 20). 644 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 6-10. 6, Aphodius badiceps, head with clypeal teeth (X40). 7, A. distinctus, hindtibia (X70). , A. campestris, foretibia (XI 10). 9, A. prodromus, foretibia (X80). 10, A. rubripennis. clypeus (X35). VOLUME 85, NUMBER 4 645 black, elytron with obscure red areas, particularly on apical declivity . . . stupidus Horn 12. Basal marginal line of pronotum strongly impressed; Georgia fordi Gordon - Basal marginal line of pronotum obsolete; not known from Georgia . . scrofa (F.) 13. Foretibia punctate on front surface (Fig. 8), punctures may be reduced to a single, irregular row (sallei); 1 st segment of foretarsus longer than 2nd segment (except sallei) 14 - Foretibia impunctate, smooth on front surface (Fig. 9); 1st segment of foretarsus not longer than 2nd segment 19 14. Elytron pubescent in apical '/3 near lateral margin 15 - Elytron without trace of pubescence except hairs on lateral margin may be present 16 1 5. First segment of hindtarsus shorter than next 3 segments; color uniformly reddish brown rubeolus Beauvois - First segment of hindtarsus as long as next 3 segments; pronotum reddish brown, elytron pale brownish yellow campestris Blatchley 16. Head tuberculate; elytron with intervals convex, striae deeply impressed; south Texas; punctures on foretibia arranged in irregular row sallei Harold - Head smooth; elytron with intervals flat, striae finely impressed; not known from south Texas (except stercorosus) 17 17. Pronotal punctures very fine, nearly invisible; lateral margin of elytron with widely spaced, long hairs, burrows of gopher tortoise troglodytes Hubbard - Pronotum with dense, coarse punctures in lateral '/?, discal area im- punctate or with fine punctures; lateral margin of elytron without hairs 18 18. Basal margin of pronotum slightly projecting medially, pronotal disc with distinct, fine punctures; in surface dung stercorosus Melsheimer - Basal margin of pronotum evenly rounded, pronotal disc appearing im- punctate; in pocket gopher burrows aegrotus Horn 19. Each anterior angle of clypeus with short, triangular or spiniform tooth (Fig. 6); dorsal color uniformly black, brown, or dull gray 20 - Each anterior angle of clypeus rounded or obtusely prominent (Figs. 10, 19) dorsal color variable, but if anterior angle of clypeus appears strongly angulate, then elytron pale red, or dark with pale maculation 29 20. Length 8.0 mm or more; clypeal teeth somewhat spiniform (Fig. 6); in squirrel nests badiceps Fall - Length less than 8.0 mm; clypeal teeth more or less triangular 21 2 1 . Dorsal surface dull, with grayish cast; elytron roughened with carinae or alutaceous sculpture 22 - Dorsal surface shiny, uniformly black or brown 23 22. Length 5.0 mm or more; intervals on elytron roughened, alutaceous, flat lutulentus Haldeman - Length 4.50 mm or less; intervals on elytron carinate (Fig. 11) robinsoni Cartwright 646 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 11-14. 11, Aphodius robinsoni. elytron (X200). 12, A. odocoilus, elytron (XI 30). 13, A. windsori, foretibia (X130). 14, A. lodingi. foretibia (X85). 23. Intervals on elytron concave (Fig. 1 2); middle of metastemum pubescent in male; New Jersey Pine Barrens odocoilis Robinson - Intervals on elytron flat or convex; metastemum not pubescent in male; New Jersey Pine Barrens and elsewhere 24 24. Last abdominal sternum concave medially, apex of sternum acuminate; mountains of North and South Carolina brimleyi Cartwright - Last abdominal sternum not concave medially, apex of sternum rounded 25 25. Striae on elytron wider over apical declivity than on disc 26 - Striae on elytron not wider over apical declivity than on disc 27 VOLUME 85, NUMBER 4 647 26. Intervals on elytron convex on disc to strongly convex over apical de- clivity; Virginia to Florida crassulus Horn - Intervals on elytron flat on disc, barely convex over apical declivity; Maryland silvanicus Cartwright 27. Entire clypeus strongly rugose-punctate; east Texas ahusus Fall - Clypeus with small, rounded, well-separated tubercles, or wrinkled in apical V2 only; not occurring in east Texas 28 28. Basal tooth of foretibia nearer base than apex (Fig. 13); dorsal surface of clypeus strongly wrinkled; surface of pronotum smooth, polished throughout; South Carolina to Florida windsori Cartwright - Basal tooth of foretibia nearer apex than base (Fig. 14); dorsal surface of clypeus with distinct, dense tubercles; surface of pronotum smooth, polished except lateral Vs weakly alutaceous, somewhat dull lodingi Cartwright 29. Elytron red, head and pronotum black; elytron pubescent in apical Vs; head smooth between punctures; pronotum lacking basal marginal line rubripennis Horn - Combination of characters not as described above 30 30. Head and pronotum smooth, punctation nearly invisible; length 9.0 mm or more; mountains from New York to North Carolina rufipes (L.) - Head usually with at least some visible punctures; pronotum always punctate, at least laterally; length less than 9.0 mm, or if longer, then pronotum distinctly punctate 31 31. Foretibia with apical spur long, abruptly hooked and acuminate (Fig. 15), or broadly expanded, spatulate (Fig. 16) 32 - Foretibia with apical spur variably modified, but never as described above 33 32. Pronotum with lateral margin broadly explanate (Fig. 17); surface of pronotum smooth, shiny, impunctate medially; Florida and Great Plains; in pocket gopher burrows *haldemani Horn - Pronotum with lateral margin slightly explanate; surface of pronotum densely punctate; Iowa and Great Plains; in pocket gopher burrows . . *nisseus Brown 33. Elytron pubescent throughout, or only in apical '/s and lateral Vs; elytron usually yellow or yellowish brown 34 - Elytron without pubescence 36 34. Foretibia with basal tooth reduced, nearly absent (Fig. 1 8); lateral margin of elytron fringed with hairs as long as scutellum *walshi Horn - Foretibia with basal tooth prominent; lateral margin of elytron fringed with hairs much shorter than scutellum 35 35. Pronotum impunctate medially, or with some scattered, coarse punc- tures; elytron yellow with large, yellowish-brown, central cloud not ex- tending to base or apex pwdromus (Brahm) - Pronotum punctate medially, punctures dense, fine; elytron yellowish brown except basal '/j often yellow femoralis Say 36. Surface of head smooth, without trace of tubercles; clypeus nearly trun- cate, anterior clypeal angle broadly rounded; color completely yellow or elytron with some small, brown spots, median '/j of pronotum usually 648 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 15-18. 15, Aphodius haldemani, female, foretibial spur (XlOO). 16, A. haldemani, male, foretibial spur (XlOO). 17, A. haldemani, pronotum (X35). 18, A. walshi, foretibia (XI 10). indistinctly yellowish brown; Atlantic coastal sand areas phaleroides Horn - Description not as above; not restricted to Atlantic coastal sand areas 37 37. Dorsal color uniformly red or reddish brown; length 7.0 mm or more 38 - Dorsal color not uniformly red or reddish brown, if elytron red, then pronotum black or brown; length 6.0 mm or less, if longer than 6.0 mm, then pronotum darker than elytron 39 38. Surface of pronotum smooth, punctures extremely fine, appearing im- punctate; Rorida, Georgia, Alabama; in pocket gopher burrows laevigatus Haldeman i VOLUME 85, NUMBER 4 649 Figs. 19-22. 19, Aphodius fimetarius, head (X35). 20, A. leptotarsis, pronotum (X42). 21, A. cuniculus, pronotum without basal marginal line (X55). 22, A. granarius. pronotum with basal marginal line (X45). - Surface of pronotum densely, coarsely punctured; Indiana, Illinois, Iowa, Great Plains; in rodent burrows *concavus Say 39. Elytron entirely red; pronotum black except lateral margin or anterior angle pale 40 - Elytron never entirely red, often partially red or yellow; pronotum en- tirely black or dark brown, often with lateral margin and/or anterior angle pale 41 40. Length less than 6.0 mm; head without prominent tubercles . . tenellus Say 650 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 23, 24. 23, Aphodius vittatus, head (X70). 24, A. borealis, head (X80). - Length more than 7.0 mm; head with 3 prominent tubercles and strong clypeal ridge (Fig. 19) fimetarius (L.) 41. Lateral margin of pronotum strongly flared outward, explanate (Fig. 20); hindtarsus as long as tibia; elytron entirely brown; in rodent burrows or nests leptotarsis Brown - Lateral margin of pronotum not flared outward, or only feebly so; elytron black or brown, or variably marked; not known to occur in rodent bur- rows or nests 42 42. Elytron entirely black; meso- and metasterna, and abdomen reddish yellow bicolor Say - Elytron entirely black or brown, or variably marked; meso- and meta- sterna, and abdomen brown to black, if appearing reddish yellow, then elytron not entirely black 43 43. Dorsal surface entirely black except apical V3 of elytron red . terminalis Say - Dorsal surface entirely black or brown, or variably marked, but never with only apical V3 of elytron red 44 44. Elytron appearing speckled or mottled, either with pale spots on dark background, or vice versa (see couplet 52, borealis) 45 - Elytron not appearing speckled or mottled 48 45. Propleuron with few short hairs, hairs not visible in dorsal view; clypeus with anterior angle sharp, nearly dentate serval Say - Propleuron with dense, long hairs, usually visible beyond lateral margin of pronotum in dorsal view; anterior clypeal angle rounded 46 46. Elytron yellow with elongate, black spots varying in pattern; legs dark brown distinctus (Muller) - Elytron brownish yellow, mottled with dark brown spots, or dark brown with pale spots, legs red or reddish brown 47 VOLUME 85, NUMBER 4 65 1 47. Surface of elytron dull, strongly alutaceous; median area of metastemum flat to slightly convex, midline distinct leopardus Horn - Surface of elytron shiny, feebly alutaceous; median area of metastemum concave, midline nearly obliterated guttatus Escholtz 48. Pronotum without basal marginal line (Fig. 21) 49 - Pronotum with basal marginal line (Fig. 22) 50 49. Elytron yellow with large, yellowish brown, median cloud; pronotum yellow in lateral Vs, median area brown lividus (Olivier) - Elytron uniformly brown; pronotum piceous except apical and lateral margin brown cuniculus Chevrolat 50. Scutellum depressed below level of elytron; pronotal punctures coarse, widely, erratically scattered granarius (L.) - Scutellum not depressed below level of elytron; pronotal punctures coarse or fine, but not widely scattered, more or less evenly spaced 51 5 1 . Clypeus dull, strongly alutaceous; apex of elytron below declivity dull, strongly alutaceous; elytron usually with at least small, red, basal and apical areas 52 - Clypeus mostly shiny, never extremely dull; apex of elytron below de- clivity as shiny as rest of elytron; elytron uniformly black or brown . . 53 52. Anterior clypeal angle rounded, apex appearing almost semicircular (Fig. 23); elytron usually mostly red except sutural margin, lateral 'A obscurely or distinctly darkened, red area often variably reduced vittatus Say - Anterior clypeal angle obtusely angulate, apex appearing broadly emar- ginate (Fig. 24); elytron usually with small red area on humerus and apical declivity borealis Gyllenhal 53. Basal segment of hindtarsus longer than long tibial spur manitobensis Brown - Basal segment of hindtarsus shorter than long tibial spur 54 54. Head essentially impunctate except for fine punctures across vertex and around inner margin of eye; intervals on elytron nearly flat mcdonaldi Robinson - Head distinctly punctured; intervals on elytron convex 55 55. Elytron with many fine punctures on interval, strial punctures strongly impressed; pronotal punctures separated by 1-2X the diameter of a puncture; Ontario to South Carolina, west to Manitoba and east Texas ruricola Melsheimer - Elytron with some scattered, fine punctures on interval, strial punctures weakly impressed; pronotal punctures separated by 2-4X the diameter of a puncture; Florida floridanus Robinson Literature Cited Brown, W. J. 1927. A revision of the species of Aphodius of Horn's series I-b. Can. Entomol. 59: 162-167. . 1928. The subgenus Platyderides in North America (Coleoptera). Can. Entomol. 60: 10-21, 35-40. . 1929. Revision of the species of Aphodius of the subgenus Diapterna. (Coleoptera). Can. Entomol. 61: llA-lTiX. Cartwright, O. L. 1 957. Three new North American Aphodius with notes and a key to related species (Scarabaeidae). Proc. U.S. Natl. Mus. 104: 413-462. 652 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON . 1 972. A key to the crassulus group of Aphodius with descriptions of new species from Texas and Maryland (Coleoptera: Scarabaeidae: Aphodiinae). Proc. Biol. Soc. Wash. 85: 57-62. Fall, H. C. and Cockerell, T. D. A. 1907. The Coleoptera of New Mexico. Trans. Am. Entomol. Soc. 33: 145-272. Gordon, R. D. 1976. Studies on the genus Aphodius of the United States and Canada (Coleoptera: Scarabaeidae). IV. A taxonomic revision of Horn's group A. Proc. Entomol. Soc. Wash. 78: 458-478. . 1977a. Studies on the genus Aphodius of the United States and Canada (Coleoptera: Scar- abaeidae). V. An evaluation of Horn's group F. Proc. Entomol. Soc. Wash. 79: 157-167. . 1977b. Studies on the genus Aphodius of the United States and Canada (Coleoptera: Scar- abaeidae). VI. Two new sand associated species from Colorado and Utah. Proc. Entomol. Soc. Wash. 79: 275-280. Hatch, M. H. 1972. The beetles of the Pacific Northwest. Part V. Univ. Washington Press, Seattle. 650 pp. Horn, G. H. 1887. A monograph of the Aphodiini inhabiting the United States. Trans. Am. Entomol. Soc. 14: 1-110. Hubbard, H. G. 1894. The insect guests of the Florida land tortoise. Insect Life 6: 302-315. Landin, B. 1961. Ecological studies on dung beetles (Col. Scarabaeidae). Opusc. Entomol. Suppl. 19: 1-228. Robinson, M. 1948. A new species oi Aphodius with notes on others (Coleoptera: Scarabaeidae). Entomol. News 59: 113-117. Saylor, L. W. 1940. Synopsis of the cadaverinus group of the genus Aphodius with descriptions of three new species (Coleoptera: Scarabaeidae). Proc. Biol. Soc. Wash. 53: 99-104. Schmidt, A. 1922. Aphodiinae. Das Tierreich, Liefrung 45. Berlin. 716 pp. Sim, R. J. 1930. Scarabaeidae, Coleoptera; observations on species unrecorded or little known in New Jersey. J. N.Y. Entomol. Soc. 38: 139-147. Woodruff, R. E. 1973. The scarab beetles of Florida (Coleoptera: Scarabaeidae) part 1. Arthropods Fla. Neighboring Land Areas 8: 1-220.