Coleoptera: Scarabaeidae: Aphodiinae), with Descriptions of Two New Genera from South America

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March 2003

Review of New World Aegialiini (Coleoptera: Scarabaeidae: Aphodiinae), with descriptions of two new genera from South America

Zdzislawa Stebnicka Institute of Systematics and Evolution of Animals, Polish Academy of Sciences

Marco Dellacasa Università di Pisa, Centro interdipartimentale

Paul E. Skelley FDACS-DPI

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Stebnicka, Zdzislawa; Dellacasa, Marco; and Skelley, Paul E., "Review of New World Aegialiini (Coleoptera: Scarabaeidae: Aphodiinae), with descriptions of two new genera from South America" (2003). Papers in Entomology. 110. https://digitalcommons.unl.edu/entomologypapers/110

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Review of New World Aegialiini (Coleoptera: Scarabaeidae: Aphodiinae), with descriptions of two new genera from South America.

Zdzislawa T. Stebnicka Institute of Sytematics and Evolution of Animals Polish Academy of Sciences Slawkowska 17 31-016 Krakow, POLAND

Marco Dellacasa Università di Pisa Centro interdipartimentale Museo di Storia Naturale e del Territorio Via Roma, 79 56011 Calci (PI) ITALY

and

Paul E. Skelley, FDACS-DPI Florida State Collection of Arthropods P.O.Box 147100, 1911 SW 34th St. Gainesville, FL 32614-7100, USA

Abstract. Two new taxa of Aegialiini (Coleoptera: Scarabaeidae: Aphodiinae) are described from South America: Argeremazus neuquen Stebnicka and Dellacasa, n.gen, n.sp, and Amerisaprus valdivia Stebnicka and Skelley, n.gen, n.sp. Both genera are presently monotypic. Discussion on the generic status of Caelius Lewis is presented. A key to genera and checklist of aegialiine taxa occurring in the New World are presented.

Key Words. Scarabaeidae, Aphodiinae, Aegialiini, Argeremazus, Amerisaprus

Introduction FMNH – Field Museum of Natural History, Chicago, IL-USA As part of a larger study on New World aphodiine ISEA – Institute of Systematics and Evolution of beetles, an author (PES) discovered a new genus of Animals, PAS, Krakow, Poland aegialiinescarab.Wishingtogetthistaxondescribed, PESC – P. E. Skelley collection, Gainesville, FL-USA the other authors were contacted for assistance. They had an additional taxon needing description and taxonomic comments to make on the group. It was Key to New World Aegialiini genera and decided to review the Aegialiini occurring in the New subgenera World, describe new genera, and present an updated checklist of taxa. The following key is based on those of Stebnicka (1977) and Gordon and Cartwright (1988). Materials 1. Head with surface not setose, variously sculptured, Materials studied are deposited in the following may have with setal fringe on clypeus ...... 2 collections: – Head with surface distinctly setose (Fig. 2) ...... CMN – Canadian Museum of Nature (Martinez col- .. Argeremazus Stebnicka and Dellacasa, n.gen. lection via H.Howden) 74 Volume 17, No. 1-2, March-June, 2003, INSECTA MUNDI

2. Elytral intervals with short hair; abdominal ster- dilated; metatibial spurs divided by base of hind nites narrowed medially, fifth sternite obliterat- tarsomere (rarely not divided). ed at middle by sixth sternite...... Body medium sized, length 1.8 - 8.8 mm, oblong ...... Micraegialia Brown ovaltoparallel-sidedorglobularmoderatelytostrong- – Elytral intervals lacking short hair; abdominal ly convex, glossy, colour yellowish varying to dark sternites not narrowed medially, fifth sternite not obliterated at middle by sixth sternite ....3 brown; margins of head, pronotum and elytra clearly piliferous. Head moderate in size, clypeus shortened 3. Elytral humerus dentate (Fig. 7); body form paral- exposing mouthparts or anterior of clypeus narrowly lel-sided ...... 4 prolongedtocovermouthparts(Argeremazusn. gen.); – Elytral humerus not dentate; body form elongate to surfacepunctateorgranulate-verrucose;clypeo-fron- globose ...... 5 tal suture marked between eyes by blackish line or fine carina, terminal segment of maxillary palpus 4. Pronotum with posterior angle eroded, bearing cylindrical. Eyes small to moderate in size, invisible several small teeth; apical metatibial spurs not from above. Pronotum subquadrate or transverse, separated at base; Chile ...... Amerisaprus Stebnicka and Skelley, n.gen. evenly convex, sides and base finely margined, sur- – Pronotum with posterior angle not notably eroded, face punctured. Scutellum small, triangular. Elytra lacking distinct teeth; apical metatibial spurs usually with striae, surface smooth or punctate. separated at base by metatarsus; western North Ventral surface glabrous or covered with long, pale America and Japan ...... Caelius Lewis hair; metasternum elevated; abdomen with five or six sternites visible, sternites coalesced or not, sutures 5. Maxillary palpus with terminal segment enlarged, membranous or stiff. Legs slender or robust; meso- widest at basal third, abruptly narrowed at base; and metafemora fusiform; protibia relatively long body red-brown, form notably elongate, almost with three to five widely separated lateral teeth and parallel sided; basal marginal line of pronotum indistinct or lacking ...... Rhysothorax Bedel slender terminal spur; meso- and metatibiae with – Maxillary palpus with terminal segment cylindri- distinct transverse ridges or with traces of ridges, cal; species with body black, form elongate to shortly setaceous or with fringe of thick setae and globose, but not parallel sided; basal marginal surface covered with pale hair; terminal spurs of line of pronotum usually distinct, at least later- metatibia slender, acute, semifoliaceous or folia- ally [Aegialia Latreille] ...... 6 ceous, placed separately on each side of tarsal insertion or located close together below tarsal insertion 6. Pronotum approximately one third as long as (genera SaprusBlackburn and Amerisaprus n. gen.); elytron, surface near posterior angle impunctate metatarsus short or moderate in length, tarsomeres or feebly so; meso- and metatibia robust or slender, spurs slender or foliaceous ...... cylindrical or triangular setaceous. Sexual differenc- ...... Aegialia (Aegialia) Latreille es indicated in the length of penultimate abdominal – Pronotum approximately half as long as elytron, sternite, in the shape and sculpture of pronotum, in surface near posterior angle coarsely punctate; the terminal spur of protibia and/or in the length of meso- and metatibia slender, apical spurs slen- penultimate abdominal sternite. der ...... Aegialia (Psammoporus) Thomson Diversity and Distribution. About 65 species are known worldwide, primarily in northern temperate Taxonomic accounts areas. They are the Holarctic Aegialia Latreille, Rhysothorax Bedel, and Caelius Lewis; the Nearctic TRIBE AEGIALIINI Laporte de Castelnau 1840 Micraegialia Brown; the Himalayan-Sichuanan Sil- luvia Landin; and the Central Asia and North Africa = Silluviini Landin 1949 Eremazus Mulsant. The previously known excep- tions are the Australian and Tasmanian genus Sa- Type genus: Aegialia Latreille 1807. prus Blackburn (Stebnicka and Howden 1995), and Aegialia argentina Martinez et al., in Argentina. Two Diagnostic characters. Head flat or slightly con- new genera and species are added to the South vex, surface granulate or punctate. Mandibles with American fauna in this paper. apices exposed, not covered by the clypeus. Pygidium The main references available for the group are lacking longitudinal groove at base, usually lacking Stebnicka’s (1977) world revision and Gordon and transverse ridge at middle. Tibiae stout, frequently Cartwright’s (1988) Nearctic revision. These refer- INSECTA MUNDI, Vol. 17, No. 1-2, March-June, 2003 75

1 3

Figures 1-3. Argeremazus neuquen Stebnicka and Dellacasa, holotype, 1) dorsal habitus, 2) head, right lateral view, 3) right metatibia. ences have keys to species and other important infor- ation, sides slightly arcuate and excised in front of mation on the Aegialiinae. Readers are urged to gena, genae small, setaceous, only slightly exceeding consult them for taxa not considered further. Below, eyes; clypeal surface (Fig. 2) just above median emar- we also discuss an apparent change in the status of gination with narrow band of minute granules, medi- two subgenera. an area between genae coarsely transversely wrin- kled or granulate with short, erect, hair-like setae; References: Gordon and Cartwright 1988 (revision clypeo-genal suture in form of arcuate carina extends of North America, illustrate most species). Stebnicka around of granulate area, base of head with band of 1977(worldrevision),RustandHanks1982(biology), coarse punctures irregularly crenating carina and Jerath and Ritcher 1959 (biology); Porter and Rust concentrated near eyes, middle of vertex polished, 1996, 1997 (phylogenetic and molecular studies), glabrous. Mouthparts adapted for hard saprophagy, Ritcher 1966 (key to known larvae). Epipharynx as in fiig. 4. Eyes moderate in size, invisible from above; antennal club elongate oval; apicalsegmentofmaxillarypalpuscylindrical.Prono- ARGEREMAZUS Stebnicka and Dellacasa, n. gen. tumtrapezoid,relativelysmall,anterioranglesacutely produced, sides and base finely margined, sides short Type species: Argeremazus neuquen Stebnicka and and broadly rounded to base, margin crenate with Dellacasa sp. n. by present designation. close pale hair. Scutellum small, triangular. Elytra convex, sides and apex with short semierect setae, Diagnosis. Body (Fig. 1) elongate oval, margins striae impressed, strial punctures distinct; epipleura fringed with long pale hair. Head convex, anterior of narrow, outer margin fringed with long pale hair. clypeusnarrowlyprolongedtocovermouthparts,edge Ventralsurfacecoveredwithhair;mesocoxaeapprox- finely dentate on each side of deep median emargin- imate,metasternumstronglyelevated;abdomenwith 76 Volume 17, No. 1-2, March-June, 2003, INSECTA MUNDI

ment). However, after examining character states in many genera of Aphodiinae, there is evidence that may be related to Eremazus, which is substantiated by ephipharyngeal characters, as discussed by Steb- nicka (1985). A detailed phylogenetic analysis is needed to substaniate this before any conclusive statementsaremade.Fornow,itissimplyconsidered a member of the Aegialiini.

Etymology.(masuline)Nameformedfromtheprefix Arg-, from “Argentina”, combined with Eremazus to show their generic affinities.

4 Argeremazus neuquen Stebnicka and Dellacasa, sp.n. Figures 1-6

Material examined. Holotype, male “/ [hand writ- ten] ARGENTINA, NEUQUEN, Covunco, Coll. Mar- tinez, [vertical on left side of label] Feb.-976 / H. and A. HOWDEN COLLECTION, ex A. Martinez coll. / 5 [red paper] HOLOTYPE Argeremazus neuquen g.n. Det. Z. Stebnicka /” (dissected, CMN). Two paratypes same data as holotype (CMN, PESC). One paratype “/ EA. COLLUNCO (900 M.S.N.M.), NEUQUEN-ARG. / 19-XII-73, LG.M.GENTILI / H. and A. HOWDEN COLLECTION,exA.Martinezcoll/”(dissected,CMN). 6 One paratype, same data as holotype only “... NEU- QUEN, Lago Blanca, ...” (female, ISEA). Figures 4-6. Argeremazus neuquen Stebnicka and Dellacasa, 4) epipharynx, 5) left lateral view of male genitalia, 6) dorsal Description. Length 3.0-3.2 mm. Body (Fig. 1) view of male genitalia. elongate oval, widest at apical third of elytra, shining; head, pronotum and scutellum brown, elytra and six sternites visible, sternites not coalesced, sutures ventral surface lighter, yellowish brown, antennal membranous, pygidium lightly sclerotized with long club dark brown. Clypeus strongly converging anteri- hair, lacking both transverse ridge and basal groove. orly, clypeal margin denticulate on each side of nar- Legs moderate in length; profemur narrow, parallel- row median emargination, surface granulate-punc- sided, perimarginal groove lacking; meso- and tate and setigerous (Fig. 2). Pronotum relatively metafemora fusiform, scarcely piliferous; protibia small and short, diverging posteriorly, sides and base with two large, widely separated lateral teeth and finely margined, anterior edge widely membranous; small tooth posteriorly; meso- and metatibiae subcy- anterior angles acute, sides short and continuously lindrical, piliferous, mesotibia slightly expanded api- rounded to base, margin crenate-fimbriate; pronotal cally with fine transverse carina; metatibia (Fig. 3) punctures coarse, irregularly spaced, on disc separat- more enlarged than mesotibia, transverse carina ed by 1-3 times their diameter, slightly confluent oblique with row of short, close setae, apical spurs laterally, area from anterior angles to lateral tumos- thick, nearly equal in length, located close together ity with finer but very dense punctures, 2-4 long setae below tarsal articualtion; metatarsus short, tarsom- in anterior angles. Scutellum with few shallow fine eres triangular, claws horn-like. punctures. Elytra moderately convex, sides and apex with short, scarce, semierect setae; striae distinctly Affinity. The taxonomic placement of Argeremazus impressed from base to apex, strial punctures fine to is a subject of various interpretations. General exter- moderate,shallowlycrenatinginnermarginsofinter- nal characters would place it in the Didactylini vals, somewhat larger apically; intervals weakly (unsculptured pygidium and metatibial spur place- convex, sutural ones narrowed toward apex, each INSECTA MUNDI, Vol. 17, No. 1-2, March-June, 2003 77

7 9

Figures 7-9. Amerisaprus valdivia Stebnicka and Skelley, paratype, 7) dorsal habitus, 8) head and pronotum, lateral-oblique view, 9) male genitalia, left lateral view. with irregular row of deep punctures separated by 2- gether below tarsal articulation, metatarsus short, 3 times their diameter on disc, closer toward apex; the tarsomeres triangular,basitarsomeremarkedlyshort- remaining intervals irregularly punctate on apical er than tibial spurs and subequal in length to follow- declivity of elytra. Ventral sclerites shining, partially ing two tarsomeres combined. Epipharynx as in microreticulate; mesosternum lower than metaster- Figure4.Inmale,discofmetasternumsurroundedby num,metasternaldiscelevated,midlinefeeblymarked; setae-bearingpunctures,penultimateabdominalster- abdomen with six sternites visible, each sternite with nite shorter than in female; aedeagus as in Figures 5- pale hair. Profemur without perimarginal groove, 6. surface punctate-setose; mesofemur a little wider than profemur with few large setigerous punctures at Etymology. The species is named after the province base; metafemur two times as wide as mesofemur it was collected in, Neuquen (noun in apposition). with row of setigerous punctures along posterior margin; protibia tridentate, terminal spur short, Remarks. The character states mentioned in the slender; meso- and metatibiae hairy, apically fringed generic description also distinguish the species, since with unequal in length, thick setae; apical spurs of the genus, at present, is monotypic. metatibia slender, equal in length, placed close to- 78 Volume 17, No. 1-2, March-June, 2003, INSECTA MUNDI

CAELIUS Lewis 1895 inanytribeoftheAphodiinaeanditstribalcharacters were not emphasized in the original description. Caelius Lewis 1895: 382 et Auctt. Type species: Caelius Schmidt (1922: 362, “ex Lewis”), assignedC. denticol- denticollis Lewis 1895 by monotypy. lis to the tribe Aphodiini based on its original descrip- = Leptaegialia Brown 1931:12-13; Hatch 1971:440; Steb- tion. This statement was accepted by Balthasar nicka 1977:429-430 (as subgenus ofAegialia); Gordon (1952, 1956) who described three new species of the and Cartwright 1988:4 (as subgenus of Aegialia); typical Aphodius, placing them in the genus Caelius. Dellacasa 1988:103 (catalogued as synonym of Cae- lius); Porter and Rust 1997:306 (as subgenus of Nomura and Nakane (1951) and subsequently Na- Aegialia). Type species: Aegialia (Leptaegialia) hu- kane (1961) expressed an opinion, thatC. denticollis meralis Brown 1931, by original designation. should be placed within Aegialiini, however, these suggestions were criticized by Balthasar (1952, 1964) Diagnosis. Body parallel-sided, color rusty brown to and rejected with no additional evidence and docu- dark castaneous, antennae and palpi yellow. Head mentation (Landin 1965). This endemic Japanese flattened, punctate and/or granulate, clypeo-frontal species, closely related toC. humeralis (Brown), was suture indicated by line with two darkened spots; redescribed and assigned to the tribe Aegialiini by clypeus shortened, mouthparts exposed, adapted for Stebnicka (1986) on the basis of five individuals hard saprophagy, terminal segment of maxillary compared with the type-specimen. palpus widened in basal third. Eyes invisible, con- Parsimony analysis of morphological characters cealed under pronotal margin. Pronotum subquad- of the Nearctic Aegialiini (Porter and Rust 1997) rate, sides and base margined, crenate, side margin supported the monophyly of the four hitherto known fringed with close yellow hair; anterior angles acutely American species of Caelius(=Leptaegialia). The ge- produced, posterior angles broadly rounded, margin nus constitutes a compact group of species well delim- sometimes distinctly serrate; pronotal punctures ited from Aegialia-Psammoporus complex and its evenly convex with mixed fine and coarse punctures. generic status is justified. As here documented, the Scutellum small, triangular. Elytra parallel, disc generic name Caelius Lewis should be considered as subdepressed,humerifeeblytostronglydentate,stri- valid in the meaning of Art. 23 of the ‘Code’ (ICZN ae with moderate to coarse, crenating punctures. 1999). Metathoracic wings functional. Legs moderate in Dellacasa (1988), in his catalogue, has already length; meso- and metatibiae moderately stout or recognized Caelius and Rhysothorax at the generic slender; apical spurs of metatibia equal in length, rank, and lists the species with the correct generic acutely pointed or rounded apically, never foliaceous, combination. This usage has generally not been fol- located on each side of tarsal inserion; metatarsus as lowed. Because of this, we are creating no new combi- long as tibia or shorter, tarsal segments triangular or nations in our recognizing Caelius and Rhysothorax cylindrical. as a distinct genera of Aegialiini. To be thorough in External sexual differences: Male: Terminal our discussion of Caelius, the following account of C. spur of protibia broad, flattened, hooked inward api- denticollis Lewis is presented. It must be emphasized cally; pronotal punctures usually less close than in that it is not a New World species. female;penultimateabdominalsterniteusuallyequal in length to two preceding sternites. Female: Termi- Caelius denticollis Lewis 1895 nal spur of protibia slender, rounded apically; pronotal punctures usually coarser and closer than in male, Caelius denticollis Lewis 1895:382, Fig. 5 (no tribal place- elytrasomewhatshorter;penultimateabdominalster- ment); Schmidt 1922:362 (Aphodiini); Balthasar nite usually equal in length to three preceding stern- 1952:234 (Aphodiini); Nakane 1961:63 (Aegialiini); Balthasar 1964:479, 480 (Aphodiini). ites or only slightly shorter. Aegialia (Leptaegialia) denticollis (Lewis): Stebnicka 1986:341, figs, 1, 2 (Aegialiini). Taxonomic background. The basic features that Caelius denticollis (Lewis): Dellacasa 1988: 119 (cata- distinguishes the aegialiines from other tribes of logued, Aegialiini). Aphodiinae, chiefly from the Psammodiini and Eu- pariini, are the shortened clypeus exposing anterior Distribution: Japan, Honshu (Miynoshita, Kiga, part of labrum and strongly sclerotized scissorial Nikko, Kanagawa Pref. Oyama, 800 m). teeth of mandibles (Stebnicka 1977, Stebnicka and Howden 1995). Caelius denticollis described from Remarks. Caelius denticollisLewis resembles Cae- Japan by Lewis (1895) has not been originally placed lius humeralis (Brown) very closely. It differs fromC. INSECTA MUNDI, Vol. 17, No. 1-2, March-June, 2003 79 humeralis by having the head with less dense punc- Comparative notes. Amerisaprus is most closely tures, the pronotal punctures closer and the pronotal allied to the genus Saprus (Stebnicka and Howden lateral margin distinctly crenulate near anterior 1995). Most of the characters represented in Saprus angles. Other diagnostic characters are: Body more are shared with Amerisaprus. The main synapomor- slender than in C. humeralis; surface of head uni- phies are: 1/ the shape and sculpture of pronotum; 2/ formly, finely punctate throughout, punctures sepa- the heavily punctate elytral striae; 3/ the widely rated by about one diameter; posterior angles of separated mesocoxae; 4/ the apical spurs of metatibia pronotum serrate like in C. humeralis; pronotal placed below tarsal insertion. punctures mixed fine and moderately coarse, the Acombinationofautapomorphiccharacterstates, lattereverywheredistributed,generallyseparatedby distinguishing Amerisaprus from Saprus, are: 1/ their diameter or less; apical spurs of metatibia equal metathoracic wings brachypterous (Saprus is fully in length, slender, acutely pointed, both spurs equal winged); 2/ mesosternum flattened, depressed below in length or only slightly longer than basal tarsomere level of metasternum (in Saprus convex, at same of metatarsus. level); 3/ abdominal sternites fluted along sutures (in Saprus more or less coarsely punctate along sutures); 4/ pygidium carinate with scabrous, rugose disc (in AMERISAPRUS Stebnicka and Skelley, n.gen. Saprus convex, smooth without carina); 5/ mouthparts exposed as usual in Aegialiini (in Saprus la- Type species: Amerisaprus valdiviaStebnicka and brum and bidentate mandibles are more exposed). Skelley, by present designation. Both genera share a number of synapomorphies with Caelius Lewis, that are: 1/ the general shape of Description. Body (Fig. 7) elongate oblong, elytra body; 2/ the head lacking granular surface (like in C. relatively short, metathoracic wings brachypterous. denticollis); 3/ the anterior pronotal angles acutely Head weakly convex, clypeus shortened, exposing produced; 4/ a similar sculpture of the pronotum and mouthparts. Antenna 9-segmented, club ovoid, 3- elytra. segmented. Eye small, can be concealed under prono- In some respects, the abdominal and tibial spur tal margin. Pronotum (Figs. 7-8) subquadrate in characters of Saprus and Amerisaprus are shared shape, sides and base margined, crenate; anterior with some genera of the Eupariini. However, both angles acutely produced, posterior angles broadly genera were apparently isolated in the past from the rounded with two denticles posteriorly, surface close- Holarctic core of the Aegialiini, and may represent a ly punctate. Scutellum small, triangular. Elytra relictualbranchfromacommonancestorbetweenthe suboval, humeral denticles strong, strial punctures more derived Aegialiini and the base of the euparine coarse. Mesosternum deplanate, lower than metast- lineages. ernum, surface punctate; mesocoxae separated; metasternum relatively short, indicating brachyp- Amerisaprus valdivia Stebnicka and Skelley tery; abdominal sternites finely fluted along sutures, n.sp. disc of pygidium transversely carinate, scabrous on Figures 7-9 apical half. Legs moderately long; profemur lacking perimarginal groove, surface punctate; meso- and Materialexamined.Holotypemale:“/Chile,Valdivia metafemora fusiform; protibia with three small later- Prov., 34 km WNW La Union, 700 m, 17.XII.1984 / al teeth; meso- and metatibiae slightly widened to- FMHD#85-921,mixedforest,P#85-36berlese, S.and ward apex with very slight traces of transverse J.Peck/[redpaper]HOLOTYPEAmerisaprusvaldiv- ridges; metatibial apical spurs slender, unequal in ia Stenbicka and Skelley/” (FMNH). Paratypes (3), length, acutely pointed, located close together below same data as holotype, one each in ISEA (male), tarsal insertion. The male genitalia (Fig. 9) of the FMNH, PESC. Aegialiinitype,phallobaseofaedeaguswithindicated dorsal hump. Description. Length 2.8-3.0 mm. Body (Fig. 7) moderately shining, glabrous; color blackish brown, Etymology. (masculine) The generic name was cho- antennae and palpi yellowish; terminal segment of sen to represent apparent relationship of this genus maxillary palpus slender, weakly widened in basal with the Australian genus Saprus Blackburn. Thus, third. Head feebly convex, slightly converging anteri- the name is a combined form of “American - Saprus”. orly with weakly indicated tubercle at middle; genae small, clypeo-frontal suture marked by line; clypeal 80 Volume 17, No. 1-2, March-June, 2003, INSECTA MUNDI surface subopaque, finely closely punctate through- Aegialia clypeata (Say): Dellacasa 1988:359 (nomen out, punctures separated by less than one diameter, dubium) vertical area shining, laterally with slightly larger Aphodius clypeatus Say 1824:278, “North-West puncturesseparatedbyonediameter.Pronotum(Figs. Territory” 7-8) almost quadrate, converging posteriorly; sides [Say’s collection has been almost entirely de- and base narrowly margined and crenate; anterior stroyed (Mawdsley 1993), with no specimen of this angles acutely produced; posterior angles broadly taxon in what remains. LeConte (1859:183-183) rounded toward base with distinct tooth at middle stated the species is probably an Aegialia and Horn that is separated from basaolateral denticle by incon- (1887:98, 110) stated it is probably a Psammodius or spicuous excavation; pronotal surface everywhere Aegialia. However, it appears that neither LeConte closelypunctured,finepuncturesalonganteriormar- nor Horn studied a specimen. The species was not gin become increasingly coarser toward base, on disc mentioned in Cartwright (1955), Gordon and Cart- separated by about one diameter, closer laterally, wright (1988), nor Stebnicka (1977). The identity of nearly contiguous and rugose at anterior angles. this species will remain a mystery.] Elytra suboval, less than 2 times of pronotal length, humeral denticles acute, directed laterad; striae FOSSIL coarsely,closelypunctate,lateralstriae6-9aswideas intervals or wider; intervals flat on disc, narrowly Aegialia rupta Scudder 1890:489, pl. 8, Fig. 19; Green carinate laterally, surface with minute scattered River, USA punctures. Ventral surface shining; mesosternum lower than metasternum, deplanate and cordate in EXTANT TAXA shape, irregularly coarsely punctate; mesocoxae separated, space between mesocoxae slightly smaller AEGIALIA Latreille 1807:96 (28 species in the New than width of mesofemur; metasternum relatively World, 12 in the Palaearctic) short, disc and lateral area coarsely punctate, lateral Type species: Scarabaeus arenarius Fabricius metasternal triangle lacking; abdominal sternites 1787:11, by monotypy finelyflutedalongsuturesandfinelypunctatethrough- out, pygidium with transverse carina, disc rugosely Aegialia(Psammoporus) Thomson 1863:72 (7 spe- sculptured. Legs moderate in length; profemur lack- cies in the New World, 8 in the Palaearctic) ing perimarginal groove, surface closely evenly punc- Type species: Scarabaeus sabuleti Panzer tate; meso- and metafemora smooth, almost impunc- 1797:37, by monotypy tate; protibial lateral teeth small, well separated, = Dimalia Mulsant and Rey 1869:406, Type spe- terminal spur straight and acute in both sexes; meso- cies: Scarabaeus sabuletiPanzer1797:37,bymonoty- and metatibiae externally with weak traces of trans- py verse ridges; metatibia with apical spurs thin, slen- Aegialia (Psammoporus) criddlei Brown 1931:42, der, unequal in length, placed close together below western North America tarsal articulation; metatarsus shorter than tibia, Aegialia (Psammoporus) cylindrica (Eschscholtz segments subtriangular, basal tarsomere slightly 1822:11), western North America shorter than upper tibial spur and subequal to follow- Aegialia (Psammoporus) exarata Mannerheim ing two tarsomeres combined. In male, penultimate 1853:219, ? Nearctic abdominal sternite shorter than and the pygidium [Horn (1887:101, 110) lists as “? syn of A. lacustris”. longer than the female. Male genitalia as in Figure 9. Stebnicka (1977:487) stated it is possibly a mis- identification. It is not treated in Gordon and Etymology. The species is named after the province Cartwright (1988)] in which it was collected, Valdivia (noun in apposi- Aegialia (Psammoporus) lacustrisLeConte1850:225, tion). north and western North America Aegialia (Psammoporus)nanaBrown1931:19,north- Annotated checklist of Aegialiinae eastern North America of New World Aegialia(Psammoporus)opacaBrown1931:17,northwestern North America UNKNOWN TAXA Aegialia (Psammoporus) terminalisBrown 1931:18, Aegialia americana Dejean 1833 [1837:163], north and western North America “Amer.Bor.” (nomen nudum) INSECTA MUNDI, Vol. 17, No. 1-2, March-June, 2003 81

Aegialia (Aegialia) Latreille 1807:96 (21 species in Amerisaprus valdivia Stebnicka and Skelley, South the New World, 4 in the Palearctic) America (Chile) Type species: Scarabaeus arenarius Fabricius 1787:11, by monotypy ARGEREMAZUS Stebnicka and Dellacasa (1 spe- Aegialia (Aegialia) amplipunctata Gordon and Cart- cies in the World) wright 1988:20, northern North America Type species: Argeremazus neuquen Stebnicka Aegialia (Aegialia) argentina Martinez, Pereira, and and Dellacasa, by monotypy Vulcano 1970:336, South America (Argentina) Argeremazus neuguen Stebnicka and Dellacasa, Ar- Aegialia (Aegialia) arenaria (Fabricius 1787:11) Eu- gentina rope, northeastern North America (introduced) = Scarabaeus globosus Kugelann 1792:514 CAELIUS Lewis 1895:381-382 (4 species in the New Aegialia (Aegialia) blanchardi Horn 1887:99, wide- World, 1 species in Japan) spread North America Type species: Caelius denticollis Lewis Aegialia (Aegialia) carri Gordon and Cartwright 1895:382, by monotypy. 1988:15, northwestern North America =LeptaegialiaBrown1931:32,Typespecies: Aegi- Aegialia (Aegialia) cartwrighti Stebnicka 1977:454, alia humeralis Bown 1931, by original designation. eastern North America Caelius browni (Saylor 1934:34), western North Aegialia (Aegialia) concinna Gordon and Cartwright America 1977:48, western North America Caelius humeralis (Brown 1931:13), northeastern Aegialia(Aegialia) conferta Horn 1871:293, northern North America and western North America Caelius montanus (Brown 1931:14), western North Aegialia (Aegialia) convexa Fall 1932:183, western America North America Caelius rufescens (Horn 1887:100), northern North Aegialia (Aegialia) crassa LeConte 1857:42, western America North America = Aegialia rufa LeConte 1878:610 (notScarabae- = Aegialia (Aegialia) insularis Brown 1931:49 us rufus Fabricius 1792:39) Aegialia(Aegialia) crescentaGordon and Cartwright 1977:45, western North America MICRAEGIALIA Brown 1931:11 (1 species in the Aegialia (Aegialia) hardyi Gordon and Cartwright World) 1977:47, western North America Type species: Micraegialia pusilla (Horn Aegialia (Aegialia) kelsoi Gordon and Cartwright 1887:102), by monotypy 1988:24, southwestern North America Micraegialia pusilla (Horn 1887:102), northwestern Aegialia(Aegialia)knightiGordonandRust1997:168, North America western North America Aegialia(Aegialia) latispina LeConte1878:611, west- RHYSOTHORAX Bedel 1911:93 (1 species in the ern North America World) Aegialia(Aegialia) magnifica GordonandCartwright Type species: Scarabaeus rufus Fabricius 1977:43, western North America 1792:39, by monotypy. Aegialia (Aegialia)mcclevei Gordon 1990:271, south- =Anomalaegialia Brown 1931:15, Type species: western North America Aegialia spissipes LeConte 1878:611, by monoty- Aegialia (Aegialia) nigrella Brown 1931:47, western py. North America Aegialia (Rhysothorax) rufa (Fabicius 1792:39), Eu- Aegialia (Aegialia) opifex Horn 1887:104, northeast- rope, northern North America (introduced ?) [see ern North America ICZN 1998] Aegialia (Aegialia) punctata Brown 1931:25, western = Scarabaeus sabuleti var ß Paykull 1798:27 North America = Aegialia spissipes LeConte 1878:611 Aegialia (Aegialia) spinosa Gordon and Cartwright = Aegialia (Rhysothorax) rufina Silfverberg 1988:25, western North America 1977:91

AMERISAPRUS StebnickaandSkelley(1speciesin UNCERTAINPLACEMENT the World) Type species: Amerisaprus valdivia Stebnicka ANNEGIALIA Howden 1971:1466 and Skelley,by monotypy 82 Volume 17, No. 1-2, March-June, 2003, INSECTA MUNDI

Type species: Annegialia ataeniformis Howden Castelnau de Laporte, F. L. 1840. Histoire na- 1971, by monotypy. turelle des Insectes Coleopteres. Paris. 2:1-564. Annegialiaataeniformis Howden1971:1468,western Dejean, P. F. M. A. 1833. Catalogue des Coléoptères North America delacollectiondeM.LeCompteDejean.Paris.pp. [AlthoughpresentlyplacedintheEupariini(Gor- 177-256. don and Cartwright 1988:2), some feel this placement Dellacasa, M. 1987(1988). Contribution to a world- is incorrect and needs to be substantiated by a rigor- wide catalogue of Aegialiidae, Aphodiidae, Au- ous morphological study and cladistic analysis. This lonocnemidae,Termitotrogidae(ColeopteraScar- will be discussed in a separate contribution (ZTS).] abaeoidea). Memorie della Societa Entomologica Italiana 66:1-455. Acknowledgments Eschscholz, J. F. 1822. Descriptions d’Insects nou- veaux. Entomographien, Berlin 1:1-140. For loans of specimens we thank Henry Howden, Fabricius, J. C. l787. Mantissa Insectorum sistens CanadianMuseumofNature,OttawaCanada;andAl eorum Species nuper detectas adiectis Charac- Newton, Field Museum of Natural History, Chicago, teribusgenericis,Differentiisspecificis,Emenda- IL-USA. tionibus, Observationibus, Hafniae. 380 pp. For reviews of the manuscript or critical conver- Fabricius, J. C. 1792. Entomologia systematica sations we thank: W. B. Warner, Chandler, AZ-USA; emendata et aucta. Secundum Classes, Ordines, R. D. Gordon, Willow City, ND-USA; M. C. Thomas Genera, Species adjectis Synonimis, Locis, Ob- andW.Dixon,FloridaDepartmentofAgricultureand servationibus,Descriptionibus.Hafniae.1:1-538. Consumer Services, Gainesville, FL-USA. Fall, H. C.1932. New North American Scarabaeidae, This project was supported, in part, by an NSF/ with remarks on known species. Journal of the PEET grant (DEB-0118669) to M. L. Jameson and B. New York Entomological Society 40:183-204. C. Ratcliffe, University of Nebraska State Museum, Gordon, R. D., and O. L. Cartwright. 1977. Four Lincoln, NE-USA. This is Florida Department of new species of Aegialia (s. str.). from California Agriculture and Consumer Services, Entomology and Nevada sand dunes. Journal of the Washing- Contribution No. 960. ton Academy of Science 67:42-48. Gordon, R. D., and O.L. Cartwright. 1988. North American representatives of the tribe Aegialiini References (Coleoptera:Scarabaeidae).SmithsonianContri- butions to Zoology 461:1-37. Balthasar, V. 1952. Quelques scarabaeides nou- Gordon, R. D., and R. W. Rust. 1997. A new veauxdel’Asiaorientale.CasopisCeskoslovenske southern Nevada species of Aegialia (Aegialia) Spolecnosti Entomologiske 49(4): 222-228. (Coleoptera:Scarabaeidae:Aphodiinae).Pan-Pa- Balthasar, V. 1956. Einige neue laparosticte Scara- cific Entomologist 73:168-171. baeiden. Entomologische Blatter. Zeitschrift fur Gordon, R. D. 1990. A new species of Aegialia Biologie und Systematik der Kafer, Krefeld, 52: (Aegialia)(Coleoptera:Scarabaeidae:Aphodiinae) 66-73. from southern Arizona and new data on North Balthasar, V. 1964. Monographie der Scarabaeidae AmericanAegialia.ColeopteristsBulletin44:271- undAphodiidaederpalaearktischenundoriental- 273. ischenRegion.ColeopteraLamellicornia,Aphodi- Hatch, M. H. 1971. The beetles of the Pacific North- idae - Verlag Tschechoslowakischen Akademie west.Part.V.:Rhypiceroidea,Sternoxi,Phytoph- der Wissenschaften, Praha 3:1-652. aga,RhynchophoraandLamellicornia.Universi- Bedel, L. 1911. Faune des Coleopteres du Bassin de ty of Washington Publications in Biology 16:1- la Seine. I. Scarabaeidae. Société Entomologique 662. de France 4:1-164. Horn, G. H.1871.SynopsisofAphodiinioftheUnited Brown, W. J. 1931. Revision of the North American States. Transactions of American Entomological Aegialiinae(Coleoptera).CanadianEntomologist Society 4:284-297. 63:9-19, 42-49. Horn, G. H. 1887. A monograph of the Aphodiini Cartwright, O. L. 1955. Scarab beetles of the genus inhabiting the United States. Transactions of Psammodius in the Western Hemisphere. Pro- American Entomological Society, Philadelphia ceedings of the United States National Museum 14:1-110. 104(3344):413-462. INSECTA MUNDI, Vol. 17, No. 1-2, March-June, 2003 83

Howden, H. F. 1971. Five unusual genera of New Lewis, G. 1895. On the Lamellicorn Coleoptera of WorldScarabaeidae(Coleoptera).CanadianEnto- Japan,andnoticeofothers.AnnalsandMagazineof mologist103:1463-1471. NaturalHistory16:374-408. ICZN. 1998. Opinion 1890. Scarabaeus rufus Moll, Mannerheim,C.G.1853.DritterNachtragzurKäfer- 1782(currentlyAphodiusrufus),Scarabaeusrufus FaunaderNord-AmerikanischenLaenderdesRus- Fabricius,1792(currentlyAegialiarufa)andScar- sischenReiches.BulletindelaSocieteImperialedes abaeus foetidus Herbst, 1783 (currently Aphodius NaturalistesdeMoscou26:95-273. foetidus)(Insecta,Coleoptera):specificnamesCon- Martinez, A., F. S. Periera and M. A.Vulcano. served.BulletinofZoologicalNomenclature55(1): 1970.Aegialiini,nuevaTribudeApodiinaeparala 54-57. regionneotropical.ActaZoologicaLilloana25:335- ICZN.1999.InternationalCodeofZoologicalNomencla- 341. ture.FourthEdition.InternationalCommissionof Mawdsley,J.R. 1993.Theentomologicalcollectionof ZoologicalNomenclature,London.306p. ThomasSay.Psyche100:163-171. Jerath, M. L., and P. O. Ritcher. 1959. Biology of Mulsant, E., and Rey, C. 1869. In Mulsant E.: Aphodiinae with special reference to Oregon (Co- HistoireNaturelledesColéoptèresdeFrance.Tribu leoptera:Scarabaeidae).Pan-PacificEntomologist des Lamellicornes. Annales de la Société 35:169-175. d’AgriculturedeLyon2:241-650. Kugelann, J. G. l792. In Schneider: Verzeichniss der Nakane, T. 1961. Notes on some Aphodiinae from einigen Gegenden Preussens bis jetzt entdeckten Micronesia (Coleoptera: Scarabaeidae). Scientific Käfer-arten,nebstkurzenNachrichtenvondensel- Report,KyotoPrefectureUniversity(NaturalSci- ben.NeuestesMagazinfurdieLiebhaberderEnto- enceandLivingScience),seriesA,3(3):151-152(25- mologie.l:257-306. 26). Landin, B.O. 1949. Entomological results from the Nomura, S. and T. Nakane. 1951. On some new and Swedish expedition 1934 to Burma and British remarkablespeciesofAphodiinaefromJapanand India.ScarabaeidaecollectedbyReneMalaise.Arkiv Formosa. Kontyu, Tokyo, 19:35-437. forZoologi1:3-9. Panzer, G. W. F. 1797.FaunainsectorumGermaniae Landin, B. O. 1965. Dr. V. Balthasar’s Monographie initia oder Deutschlands Insecten. Nurnberg derScarabaeidaeundAphodiidaederpalaearktis- 37:1.2.3. chen und orientalischen Region. 1-3. A critical Paykull, G. 1798. Fauna Suecica. Insecta, Tomus I. review. Opuscula Entomologica, Lund, 30-:175- Upsaliae, x + 360pp. 182. Porter, J. L. and R. W. Rust. 1996. Allozyme varia- Laporte de Castelnau. See Castelnau de Laporte. tion within five species of Aegialia (Coleoptera: Latreille, P. A. 1807. Genera Crustaceorum et Insec- Scarabaeidae).AnnalsoftheEntomologicalSociety torumsecundumOrdinemnaturaleminFamilias ofAmerica89:710-721. disposita,IconibusEsemplisqueplurimisexplicata. Porter, J. L. and R. W. Rust. 1997. Phylogenetic (1806-1809).ParisisetArgentorati,2:1-280. relationships of North American Aegialia (Co- LeConte, J. L. 1850. General remarks upon the Co- leoptera:Scarabaeidae)asindicatedbyAllozymes leoptera of Lake Superior. In L. Agassiz: Lake andmorphology.AnnalsoftheEntomologicalSoci- Superior its physical characters, vegetation and etyofAmerica90(3):302-315. animals,comparedwiththoseofotherandsimilar Ritcher, P. O. 1966. White grubs and their allies. A regions.WithnarrativeofthetourbyJ.ElliotCabot studyofNorthAmericanscarabaeoidlarvae.Stud- and contributions by other scientific gentlemen. ies in Entomology Number Four. Oregon State 2(4):201-245. University Press, Corvallis, OR, 219 p. LeConte,J.L.1857.Reportsofexplorationandsurveys Rust, R. W. and L. M. Hanks. 1982. Notes on the forarailroadroutefromtheMississippiRivertothe biology of Aegialia hardyi Gordon and Cartwright PacificOcean.I.Reportupontheinsectscollectedon (Coleoptera:Scarabaeidae).Pan-PacificEntomolo- theSurvey.ProceedingsoftheAcademyofNatural gist58:319-325. Sciences,Philadelphia12:1-72. Say, T. 1824. Coleoptera Appendix to: Narrative of an LeConte,J.L.1859.ThecompletewritingsofThomas expeditiontothesourceofSaintPeter’sRiver,etc., Say on the Entomology of North America. Volume under the command of Stephen H. Long, Major 1. Baillière Brothers, New York, 412 p. U.S.T.E.Philadelphia2:268-378.[seeLeConte1859] LeConte, J. L 1878. Coleoptera found in the Lake Superiorregion.ProceedingsoftheAmericanPhilo- sophicalSociety17:610-611. 84 Volume 17, No. 1-2, March-June, 2003, INSECTA MUNDI