Scatimina, Vaz-De-Mello, 2008

Vaz-De-Mello, Fernando Z., 2008, Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of twelve new genera and review of Genieridium, new genus, Zootaxa 1955, pp. 1-75 : 10-16

Scatimina , new subtribe

Diagnosis: The subtribe Scatimina can be separated readily from all remaining Scarabaeinae by the following combination of characters: (1) hypomeron anteriorly deeply excavated, excavation bordered posteriorly by transverse carina; (2) trochantofemoral pit present; and (3) pseudoepipleuron present lateral to 8th elytral stria, delimited by folding or convexity of 9th interstria, and separated from epipleuron by stria distinctly visible at least basally ( Figs. 5–11).

Description: Clypeus centrally with an emargination, two or more teeth, or both, never simply rounded; external genal angle clearly obtuse; clypeal ventral process forming transverse carina, sometimes angled or tuberculated in middle, united to transversal supraepipharyngeal ridge by angle or short longitudinal carina. Separation between submentum and gula simply curved to obtusely V-shaped, sometimes indistinct, gula laterally glabrous, and anteriorly slightly pointed. Antennae with 9 antennomeres, first antennomere of club completely tomentose. Mentum apically not emarginated, or only slightly so, with medial dorsal, pointed prolongation (serving as "base" for membranous dorsally-positioned ligulae). Labial palpi with 3 palpomeres; third much narrower, smaller than second. Side of pronotal disc with indistinct to distinct callosity, without longitudinal sulcus or distinct fovea; centrally without elevations, sulci, or tubercles, simply convex. Pronotal osterior border unbeaded or very feebly beaded. Hypomeron anteriorly deeply excavated, excavation separated from posterior part by transverse carina. Mesosternum anteriorly with transverse sulcus. Metasternum anteriorly with transverse sulcus parallel and very close to (often coincident with) middle of mesometasternal suture, prolonged parallel to mesocoxa laterally.

Protibial apices mesally truncated, straight (except for mesal apical tooth in some males); protibiae apical border straight, continuous (i. e., without angle) with anterior border of apical tooth, such that two margins (protibial body and apical tooth) lie in straight line ( Figs. 1–4). Protibiae with three teeth or less, with tarsal insertion near the mesal apical border. Protrochantofemoral pit present. Mesocoxal axis very slightly angled, almost parallel to longitudinal body axis. Mesotibial apex obliquely truncated (mesal angle acute). Metatibial apex entire below tarsal insertion. Mesotibiae and metatibiae with longitudinal external carina indicated on disc, and dorso-external longitudinal carina well defined, while ventro-external carina is absent or indicated by row of setae. Mesotibial and metatibial apical fimbriae alternate in size and thickness, with short, thick setae co-mingled with long, narrow setae (sparse short, thick setae also present on the external surface of mesotibiae and metatibiae). Elytra with nine striae. Pseudoepipleura present lateral to 8th elytral stria, delimited by fold or convexity of 9th interstria and separated from epipleuron by epipleural stria well defined at least in basal half ( Figs. 5–11).

Sixth abdominal ventrite never shortened medially. Basal pygidial sulcus present. Male genitalia with genital segment transverse, conformed of a central rounded plate and a lateral rectangular elongated plate at each side. Parameres almost symmetrical (at least in size and shape of sides). Transverse parietal lamella of internal sac (magnifying glass-shaped lamella or circular sclerite of authors) with basal arc very reduced, so that one side of the circle is obliterated or completely absent. Female genitalia with coxites present, sometimes fused ventrally. Spermathecal base simply rounded or pointed, never bilobed or expanded; apex simply pointed.

Type genus: Scatimus Erichson, 1847

Remarks. This new subtribe includes the following Neotropical genera: Besourenga , new genus; Bradypodidium , new genus; Degallieridium , new genus; Eutrichillum Martínez , new status; Feeridium , new genus; Genieridium , new genus; Leotrichillum , new genus; Martinezidium , new genus; Nunoidium , new genus; Onoreidium , new genus; Pedaridium Harold ; Pereiraidium , new genus; Scatimus Erichson ; Scatrichus Génier & Kohlmann ; Silvinha , new genus; and Trichillidium , new genus; Trichillum Harold.

Besides the characters mentioned above, the following are putative apomorphies supporting Scatimina : the presence of a pit in the external face of first and second antennal lamellae (these pits appear to have been secondarily lost in Scatrichus ), the presence of an anterior transverse carina on mesoepimeron (lost in the group formed by Trichillum , Besourenga , Degallieridium , Feeridium , Eutrichillum , and in most species of Scatimus ), the spatulate form of the protibial spur in males (modified in one species of Trichillum and at least some species of Scatimus ), the sinuate form of mesotibial spurs, the absence of a basal raspula and the presence of an elongated flagelliform sclerite in the internal sac.

The Scatimina are distributed in the entire Neotropical region as defined by Morrone (2006) (except for the Antillean Dominion), and in the entire Mexican Transition Zone, in a province of the Nearctic Region (Sonora), and in part of the South American Transition Zone (Monte and perhaps Prepuna provinces) .

The trochanterofemoral pit is not unique to the new subtribe and is also present in the following four genera of Ateuchini : Bdelyropsis Pereira, Vulcano & Martínez , Demarziella Balthasar , Pedaria Laporte , and Uroxys Westwood , and several genera of Canthonini . These genera were previously considered related to members of the Scatimina , but are herein viewed as Ateuchini incertae sedis or transferred to Canthonini (see below).

Although Pedaria and Demarziella have been regarded as relatives of Trichillum and Pedaridium (based on the presence of dorsal setae and fused abdominal sternites), they show the following important differences: (1) elytra with ten striae (as opposed to nine in the Scatimina – the ninth and tenth striae are often partially fused in the middle, but well differentiated both anteriorly and posteriorly); (2) parameres distinctly asymmetrical; and (3) presence of well-developed magnifying glass shaped lamella in the internal sac. These characters of Pedaria and Demarziella are shared with Coptodactyla Burmeister , Thyregis Blackburn , and Microcopris Balthasar , formally belonging to the tribe Coprini , which lack fused abdominal sternites. These genera could be regarded as a separate tribe (Coptodactylini Paulian), but that is beyond the scope of the present paper.

It is tempting to include the genus Uroxys in Scatimina , but the heterogeneity, presumed polyphyletic origin of the genus ( Halffter & Matthews 1966, Halffter & Edmonds 1982, Martínez 1988), and various taxonomic and nomenclatorial difficulties make it inadvisable at the present time. One difficulty is the uncertain position of the type species of Uroxys . Some species presently included in Uroxys are here excluded from Scatimina because of the presence of lateral longitudinal sulci on each side of the pronotum and the complete magnifying glass shaped sclerite in the internal sac. However, it seems that at least one of the species groups now placed in Uroxys belongs or is very close to Scatimina .

Finally, Bdelyropsis has the distinct pseudoepipleuron originating from a fold in the 8th interstria, and appearing related to a group of genera currently in the Canthonini that includes Zonocopris Arrow and Cryptocanthon Balthasar , among others (Vaz-de-Mello 2007, see below on Bdelyrus ).

A new diagnosis for the tribe Ateuchini greatly depends on the reassignment of the genera here considered as incertae sedis to other tribes and will be presented in a subsequent paper. The proposal of a new subtribe in Ateuchini results in the recognition of the subtribe Ateuchina , which is provisionally diagnosed as follows: (1) medial part of clypeal surface concave; (2) clypeal ventral process in the form of a transverse carina; (3) pronotum simply convex, at most with discal longitudinal sulcus; (4) lateral pronotal pit well-defined; (5) hypomeron deeply excavated anteriorly and with a transverse carina (as in Scatimina ); (6) fore legs with trochantofemoral pit absent; (7) protibia with anterior border completely straight (as in Scatimina ), except for anteroapical tooth in male; (8) nine elytral striae (with pseudoepipleuron delimited by a fold or convexity in the 9 th interstria); (9) parameres symmetrical; (10) internal sac sclerites represented by few small flagelliform lamellae, all apical in position.

As defined herein, the subtribe Ateuchina is Neotropical and Nearctic and includes the following genera: Ateuchus Weber (with the exclusion of some species to be assigned to other genera in the near future), Deltorhinum Harold , Aphengium Harold , and Sinapisoma Boucomont (formerly placed in Canthonini ), as well as taxa yet to be described.

Most other ateuchine genera (all lacking trochantofemoral pit) are herein considered to be Ateuchini incertae sedis.

The genera Canthidium Erichson (based on examination of the type species, Canthidium lentum Erichson ) and Parachorius Harold have 10 elytral striae, the 9 th being easily discernible from the 8 th only at elytral apex, and both have hypomera very feebly excavated anteriorly, although the transverse carina is present; they also show an angle between the anterior borders of the apical protibial teeth and the mesal apical margin of tibia (i.e., entire anterior tibial border is not straight). Those characters point to a close relationship between these genera and Holocanthon Martínez & Pereira (presently placed in Canthonini ), and justify their inclusion in the tribe Coprini . Within the Coprini , the most closely related group would be the one including Dichotomius Hope and its allies. The subgenus Eucanthidium Martínez & Halffter (based on examination of the type species, Canthidium cupreum Blanchard ), shares all those characters except that only nine elytral striae are present; Eucanthidium may later turn out to be a taxonomic isolate.

Bdelyrus Harold , has a double pseudoepipleuron originated from folds in both 8 th and 9 th interstriae, lacks the developed trochantofemoral pit, and has a very peculiar assemblage of internal sac sclerites. However, its unusual abdominal shape is similar to that of Paracryptocanthon , and the supraunguicular spines (found otherwise only in Zonocopris ) appears to relate Bdelyrus to the yet unnamed group that includes those genera (Vaz-de-Mello 2007, see above also on Bdelyropsis ). Furthermore, internal sac characters appear to relate Bdelyrus to Onychothecus Boucomont and Paraphytus Harold , both possessing ten elytral striae.

Scatonomus Erichson and Anomiopus Westwood (and the related Hypocanthidium Balthasar ) appear to be true Canthonini , in fact closely related to Canthon . This relationship was suggested by Canhedo (2006), who did not make any formal classification changes, and I here assign Scatonomus and Anomiopus to the tribe Canthonini .

Finally, Sarophorus Erichson , Coptorhina Hope , Frankenbergerius Balthasar , and Delopleurus Erichson have a unique lateral sinuosity on the elytra; a longitudinal groove on the gula; a very simple internal sac, in some cases lacking sclerites; and a very distinctive spermatheca with flat basal paired lobes found otherwise only in some Canthonini . These genera should be considered as a group distinct from Ateuchini ( Zunino 1983, Frolov & Scholtz 2003, Frolov & Scholtz 2005).

A summary of the present considerations on subtribal or tribal position of above-mentioned is given on Table 1.

proposed here. An * indicates genera not examined.

Ateuchini

Ateuchina Scatimina Incertae sedis

Aphengium Harold, 1868 Besourenga , new genus Bdelyropsis Pereira, Vulcano & Martínez, 1960

Ateuchus Weber, 1801 Bradypodidium , new genus Bdelyrus Harold, 1869

Deltorhinum Harold, 1867 Degallieridium , new genus Coptorhina Hope, 1830

Sinapisoma Boucomont, 1928 Eutrichillum Martínez, 1969 , new status Delopleurus Erichson, 1847 Feeridium , new genus Demarziella Balthasar, 1961 Genieridium , new genus Onychothecus Boucomont, 1912 Leotrichillum , new genus Paraphytus Harold, 1877 Martinezidium , new genus Pedaria Laporte, 1832

Nunoidium , new genus * Pleronyx Lansberge, 1874 Onoreidium , new genus * Pseuduroxys Balthasar, 1938 Pedaridium Harold, 1868 Sarophorus Erichson, 1847 Pereiraidium , new genus Uroxys Westwood, 1842

Scatimus Erichson, 1847

Scatrichus Génier & Kohlmann, 2003

Transferred to Canthonini Silvinha , new genus Transferred to Coprini

Anomiopus Westwood, 1842 Trichillidium , new genus Canthidium Erichson, 1847

Hypocanthidium Balthasar, 1938 Trichillum Harold, 1868 Holocanthon Martínez & Pereira, 1956

Scatonomus Erichson, 1835 Parachorius Harold, 1875

The present necessity to split into 15 two previously very heterogenous genera, Trichillum and Pedaridium , is yet another consequence of the tremendous increase in data and material during the remarkable surge in field and museum work on dung beetles during the past 30 or so years. The number of new species known to me in these two groups alone has increased by about 80 (200%) in the last decade; while some are described here, many will be described in subsequent papers. Likewise, in this paper and elsewhere, the recent increase in the number of new taxonomic characters has permitted taxonomic and cladistic studies of higher resolution. Other recent cases of extensive splitting of previously heterogeneous genera are those of Panelus Lewis ( Scholtz & Howden 1987; Ochi et al. 1998; Paulian 1976, 1985; Matthews 1974), Epilissus Reiche ( Olsoufieff 1935, 1947; Paulian 1939), and the “group” Stiptopodius Harold ( Branco 1989, 1990, 1991, 1992a, 1992b, 1995). My personal opinion is that, if these genera (including those treated here) were larger in size and more abundant in collections (it must be said that most collections are very poor in representatives of the diversity of those groups, most having a very small number of species of few subgroups of it), then they would have been split (or independently described one by one) much earlier.

Key to the genera of Scatimina (partly taken from Génier & Kohlmann [2003])

1. Length of last abdominal sternite (measured along midline) about equal to lengths of other sternites or not longer than two of them combined................................................................................................... 2

-. Last abdominal sternite elongate (measured along midline), extending from pygidium to metacoxae, remaining sternites only discernible laterally......................................................................................... 3

2(1). Head with two parallel transverse carinae; lateral edge of pronotum crenulate, with setose punctures; apical declivity of elytron with setiferous punctures on interstriae; trochantofemoral pit of anterior leg oval, transverse; lateral declivity of pronotum with accessory setiferous punctures. Southeastern and central Brazil..........................................................................14. Scatrichus Génier & Kohlmann, 2003

-. Head with one transverse carina, or with one conical tubercle; lateral edge of pronotum simple, lacking setose punctures; apical declivity of elytron lacking setiferous punctures on interstriae; trochantofemoral pit of anterior leg rounded; lateral declivity of pronotum lacking setiferous punctures. Northwestern Mexico to northern South America ........................................................... 13. Scatimus Erichson, 1847

3(1). Elytral pseudoepipleuron forming two sinuosities, the posterior one (near metacoxa) partially covering true epipleuron, and sometimes angled ( Figs. 5, 9) .............................................................................. 4

- Elytral pseudoepipleuron at most forming one long sinuosity on anterior half that does not fold over true epipleuron (sometimes true epipleuron with excavation near metacoxa) ( Figs. 6–8, 10–11) ........ 5

4(3). Clypeal teeth clearly arising below dorsal clypeal margin ( Fig. 91); pseudoepipleuron without anterior longitudinal carina on basal half; posterior pseudoepipleuron sinuosity bent over anteriorly, posterior half of pseudoepipleuron not distinguishable from true epipleuron (i.e., epipleural stria indistinct on posterior half) ( Fig. 9); male with apical tarsomere of anterior tarsi excavated dorsally to receive claws; parameres strongly divergent in the middle and convergent apically ( Fig. 92). Southeastern Brazil....................................................................................................................... 15. Silvinha new genus

-. Clypeal teeth continuous with dorsal clypeal margin (Figs. 98, 100, 102); pseudoepipleuron with a sharp anterior longitudinal carina extending up to posterior pseudoepipleuron sinuosity, which is bent over true epipleuron and angled, posterior half of pseudoepipleuron completely distinguishable posteriorly from true epipleura by epipleural stria ( Fig. 5); male with apical tarsomere of anterior tarsi not modified; parameres flattened, truncated apically, without external lobes. South America east of the Andes excluding southern Argentina......................................................... 17. Trichillum Harold, 1868

5(3). Pseudoepipleuron posteriorly abruptly narrowed near metacoxa, margin forming an angle ( Fig. 10). 6

-. Pseudoepipleuron gradually reduced in width, margin straight near metacoxa ( Figs. 6, 8, 11)............ 7

6(5). Clypeo-genal suture clearly visible, extending from frontoclypeal suture to clypeogenal border; clypeo-genal border incised, with clypeus and gena separately rounded, clypeus evenly rounded between clypeal teeth and clypeo-genal suture ( Fig. 49). Northern Argentina, central and southeastern Brazil, Paraguay, eastern Bolivia, Amazonia (except north), one isolated species in Costa Rica....................... ................................................................................................................ 4. Eutrichillum Martínez, 1969

-. Clypeogenal and frontoclypeal sutures indistinct; clypeal margin lateral to clypeal teeth strongly curved or angled, becoming straight anteriorly to clypeogenal border, itself straight or slightly sinuate ( Figs. 39, 41). Paraguay, eastern Bolivia, central Brazil, and western Amazonia ................................... ........................................................................................................................ 1. Besourenga new genus

7(5). Head with two horns (males) ( Fig. 88) or two feeble separate convexities (females) ( Fig. 90) on frontoclypeal region; anterior margin of pronotum beaded. Southern Brazil... 12. Pereiraidium , new genus

-. Head without horns or tubercles on frontoclypeal region; anterior margin of pronotum unbeaded .... 8

8(7). Protibia with two or three lateral teeth, distributed along one-half or less of tibial length; mesotibia not strongly widened apically, apicolateral area ventrally covered by very long setae............................... 9

-. Protibia with three lateral teeth, distributed along at least apical three-fifths of tibial length, if occupying less than three-fifths of tibial length, then, mesotibia with strong lateroventral tooth and apex abruptly extended laterally, and with sparse apical ventral setae ....................................................... 10

9(8). Head flat to very feebly evenly convex, lacking evident shallow concavities in front of eyes; clypeus laterally straight to slightly curved outward; gena not protruding beyond clypeus ( Figs. 43, 45); body elongated ( Fig. 17); elytral striae non-moniliform (strial punctures small and well separated); claws and apical tarsomere of protarsi of males strongly modified ( Fig. 46). Costa Rica to southeastern Brazil..................................................................................................................2. Bradypodidium , new genus

-. Head evenly convex mesally, with distinct shallow concavities in front of eyes; clypeal margin laterally curved mesad; gena protruding beyond clypeus ( Figs. 93, 95, 97); body strongly rounded ( Figs. 35–36); elytral striae moniliform at least on apical half (strial punctures twice the width of stria, punctures weakly separated or contiguous in posterior half); claws and tarsomeres of male not modified. Nicaragua to Ecuador; Brazil, Argentina, Paraguay, Bolivia................... 16. Trichillidium , new genus

10(8). Sides of pronotum, in dorsal view, sinuous ( Figs. 29, 87); elytra distinctly tectiform, sutural (first) interstriae distinctly elevated, elytral surface flat ( Fig. 29). Southern Brazil........................................... .................................................................................................................. 11. Pedaridium Harold, 1868

-. Sides of pronotum, in dorsal view, approximately straight to uniformly curved; elytra at most slightly tectiform; sutural (first) interstria not distinctly elevated; elytral surface evenly convex ................... 11

11(10). Eyes dorsally as wide as long; dorsal interocular distance less than two times width of one eye ( Figs. 20, 52); pronotum separated from hypomeron by row of punctures, not by carina. Amazon River bor- der and French Guyana................................................................................... 5. Feeridium , new genus

-. Eyes dorsally longer than wide; dorsal interocular distance at least five times width of a single eye; pronotum separated of hypomeron by sharp longitudinal carina or at least weak carina between punctures (Fig. 59) ...................................................................................................................................... 12

12(11). Head disc strongly convex, sometimes with a transverse carina; clypeal emargination indistinct; clypeal teeth upturned and widely separated, area between teeth shallow, very widely U-shaped (Figs. 79, 82–83); head central surface strongly convex, sometimes with a transverse carina; mesotibiae (and metatibiae to lesser degree), in ventral view, with strong tooth near mid-length of lateral border, tooth bearing a very thick and short seta (Fig. 81); mesotibiae with ventral transverse ventral apical carina of the same tibia)with stout setae similar to one on ventral lateral tooth. Ecuador, Panama, and Costa Rica.................................................................................................................... 10. Onoreidium , new genus

-. Head disc flat or anterior margin in the middle concave or clypeal emargination distinct; if clypeal teeth absent, then emargination widely V-shaped (narrowly angled in the middle); mesotibiae and metatibiae, in ventral view, with very reduced external tooth at mid-length; mesotibiae and metatibiae transverse apical ventral carina bearing long setae ............................................................................. 13

13(12). Pseudoepipleuron with longitudinal angle making it approximately vertical on anterior half and horizontal on posterior half; glabrous; width of eye (dorsal view) greater than one-half length ( Figs. 47, 72); body length under 2.8 mm ............................................................................................................ 14

-. Pseudoepipleuron entirely at the same plane, with distinct row of setae along most of length (rarely lacking on anterior third only); width of eye (dorsal view) equal to or less than one half length; body length over 2.8 mm .............................................................................................................................. 15

14(13). Hypomeron with posterior longitudinal carina; elytral interstriae uniformly flat, not distinctly more convex apically; discal interstriae with single row of setose punctures. Central Brazil, Paraguay, Argentina................................................................................................... 7. Leotrichillum , new genus

-. Hypomeron without posterior longitudinal carina; elytral interstriae more convex apically; discal interstriae with two rows of punctures and one row of setae. Central Brazil................................................. ................................................................................................................. 3. Degallieridium , new genus

15(13). Mesotibiae and metatibiae slender, length at least four times apical width. Mexico, Guatemala, Argentina, western Paraguay............................................................................... 8. Martinezidium , new genus

-. Mesotibiae and metatibiae strongly widened posteriorly, length at most three times apical width (in some cases metatibiae strongly modified laterally) ............................................................................. 16

16(15). Eyes dorsally minute or absent, when present separated by more than 10 times their width (Figs. 55, 57, 60, 62, 64, 68, 70); pronotum unbeaded posteriorly, separated from hypomeron by weak lateral carina between punctures; males with modified claws (Figs. 60, 62), length of mesotibiae and metatibiae more than three times apical width; pygidium approximately 45 degrees from vertical plane, in ventral view, its horizontal projection spanning at least 2/5 of abdominal length. Colombian Andes, southern Venezuela, Bolivia, Paraguay, Brazil and Argentina.................................. 6. Genieridium , new genus

-. Eyes dorsally oval and well developed, separated by seven to eight times their width ( Fig. 77); pronotum beaded posteriorly, separated from hypomeron by strong and smooth carina; males with claws unmodified, mesotibiae and metatibiae less than three times as long as wide at apex; pygidium almost vertical, in ventral view, its horizontal projection spanning less than 1/5 of abdominal length. Central Argentina....................................................................................................... 9. Nunoidium , new genus