SciELO - Scientific Electronic Library Online

vol.66 número3-4Redescripción de la hembra de Oliveiriella almeidai (Chironomidae: Orthocladiinae)Comunidades de Chironomidae (Diptera) en arroyos de montaña del NOA: comparación entre Yungas y Monte índice de autoresíndice de materiabúsqueda de artículos
Home Pagelista alfabética de revistas  

Servicios Personalizados




  • No hay articulos citadosCitado por SciELO

Links relacionados


Revista de la Sociedad Entomológica Argentina

versión impresa ISSN 0373-5680versión On-line ISSN 1851-7471

Rev. Soc. Entomol. Argent. v.66 n.3-4 Mendoza ago./dic. 2007


The Argentinean dung beetle genus Anomiopsoides (Scarabaeidae: Scarabaeinae: Eucraniini): description of a new species, and new synonymies for A. heteroclyta

El género argentino de escarabajos estercoleros Anomiopsoides (Scarabaeidae: Scarabaeinae: Eucraniini): descripcción de una especie nueva y nuevas sinonimias para A. heteroclyta

Ocampo, Federico C.

Laboratorio de Entomología, Instituto Argentino de Investigaciones de las Zonas Áridas (IADIZA, CRICYT, CONICET), Casilla de correo 507, 5500 Mendoza, Argentina; e-mail:

ABSTRACT. The taxonomy of the genus Anomiopsoides Blackwelder is revised. The species A. catamarcae Martínez and A. aurita (Burmeister) are placed in synonymy with A. heteroclyta (Blanchard). Anomiopsoides fedemariai sp. nov. is described from Argentina. The genus Anomiopsoides now consists of four species: A. biloba (Burmeister), A. cavifrons (Burmeister), A. fedemariai sp. nov. and A. heteroclyta (Blanchard). A key is presented for the identification of the species of Anomiopsoides.

KEY WORDS. Dung beetles; Monte biogeographic province; Scarabaeinae; Eucraniini.

RESUMEN. Se hace una revisión de la taxonomía del género Anomiopsoides Blackwelder. Las especies A. catamarcae Martínez y A. aurita (Burmeister) son consideradas como sinónimos de A. heteroclyta (Blanchard). Se describe una especie nueva, A. fedemariai sp. nov., especie nueva de Argentina. El género Anomioposoides consiste ahora en cuatro especies, A. biloba (Burmeister), A. cavifrons (Burmeister), A. fedemariai sp. nov. y A. heteroclyta (Blanchard). Se presenta una clave para la identificación de las especies de Anomiopsoides.

PALABRAS CLAVE. Escarabajos estercoleros; Provincia biogeográfica del Monte; Scarabaeinae; Eucraniini.


Anomiopsoides Blackwelder is a relatively small genus of dung beetles that belongs to the tribe Eucraniini (Scarabaeidae: Scarabaeinae). Eucraniini consists in four genera, Anomiopsoides, Ennearabdus van Lansberge, Eucranium Brullé and Glyphoderus Westwood. This tribe is endemic to the Chaco and Monte biogeographic provinces in Argentina (Zunino et al 1989; Monteresino & Zunino 2003; Ocampo 2004, 2005)
Species of the genus Anomiopsoides show a high degree of intraspecific variation. This variation, expressed particularly in the shape of the clypeal armature has caused the description of several synonyms, most of them within one species, A. heteroclyta (Blanchard). Clypeal armature in species of this genus consists of two central and two lateral processes. Sexual dimorphism in the clypeal armature is expressed in all but one species of Anomiopsoides, A. biloba (Burmeister) (Ocampo, 2005). Anomiopsoides heteroclyta (Blanchard, 1845), the first-described species in the genus, was originally placed in the genus Anomiopsis Westwood 1837. Anomiopsis Westwood is a synonym of Eucranium Brullé 1834. Burmeister (1861: 62) proposed the name Anomiopsis as a «section» of Eucranium and created a homonym, even when in the same publication he considered the name Anomiopsis as a synonym of Eucranium. As indicated above, the name Anomiopsis was preoccupied by Anomiopsis Westwood 1837 (junior synonym of Eucranium) (Scarabaeidae: Scarabaeinae). In the same publication, Burmeister (1861) described four species: Eucranium (Anomiopsis) aurita, Eucranium (Anomiopsis) biloba, Eucranium (Anomiopsis) cavifrons, and Eucranium (Anomiopsis) furciferum. Burmeister (1873) considered Anomiopsis Burmeister as a «subgenus» of Eucranium. Gillet (1911) elevated Anomiopsis to the generic level. Martínez (1944) described one species, Anomiopsis pereirai, and variety, A. biloba var. schaeferi. Blackwelder (1944) acknowledged the mistake (creation of the homonym) made by Burmeister (1861) and fixed it replacing the name Anomiopsis with Anomiopsoides. Martínez (1945a, 1945b) redescribed the genus Anomiopsoides, redescribed the known species, synonymyzed A. furciferum (Burmeister) with A. heteroclyta (Blanchard), and described three new species, A. aberrans, A. catamarcae, and A. xerophila. Barbero & Palestrini (1993) described the larvae of A. cavifrons, the only known larva of the genus. Ocampo (2005) revised the genus Anomiopsoides and synonymized several species and a subspecies: A. aberrans Martínez, A. pereirai Martínez, and A. xerophila Martínez were synonymized with A. heteroclyta (Blanchard) and A. biloba var. schaeferi Martínez was synonymized with A. biloba (Burmeister). Ocampo (2005) also provided a checklist of the tribe Eucraniini.
The biology of this genus was described by Zunino et al. (1989), Monteresino & Zunino (2003), and Ocampo (2005, 2007).
The aim of this paper is to propose new synonymies for A. heteroclyta, describe a new species of Anomiopsoides from the Monte biogeographic province in Argentina, and provide a new key for the identification of the species.


Body measurements, puncture density, puncture size, fovea density, fovea size, and density of setae are based on the following standards. Body length was measured from the apex of the pronotum (at the middle) to the apex of the elytra (head is excluded because of the variable position of the head, and length of clypeal processes render it unpractical to include in the body length). Body width was measured across midpronotum. Puncture density was considered «dense» if punctures were nearly confluent to less than two puncture diameters apart, «moderately dense» if punctures were two to six diameters apart, and «sparse» if punctures were separated by more than six diameters apart. Puncture size was defined as «small» if punctures were 0.02 mm or smaller, «moderate» if 0.02-0.07 mm, and «large» if 0.07 mm or larger. Surface was defined as «sparsely foveate» if there was (on average) a space of more than one diameter between foveae, «moderately dense» if there were 0.5-1 diameters between foveae, and «densely foveate» if foveae were confluent or the space in between them is less than 0.5 diameters. Setae were defined as «sparse» if there were few setae, «moderately dense» if the surface was visible but with many setae, and «dense» if the surface was not visible through the setae. Elytral carinae were counted from the elytral suture. Specimen labels were copied literally using «/» between lines.
The «Phylogenetic Species Concept» (Wheeler & Platnik, 2000) was used to diagnose species: «Species are the smallest aggregation of populations or lineages diagnosable by a unique combination of character states.»
The results of this study were based on specimens from the following institutions, curators, and collections.
ABTS: Andrew B. T. Smith Collection, Ottawa, Canada.
CASC: California Academy of Sciences, San Francisco, California, USA (R. Brett).
CMNC: Canadian Museum of Nature, Ottawa, Canada (R. S. Anderson, F. Génier).
FCOC: Federico C. Ocampo Collection, Mendoza, Argentina.
IADIZA: Instituto Argentino de Investigaciones de las Zonas Áridas, Mendoza, Argentina (S. Roig-Juñent).
IMLA: Fundación e Instituto Miguel Lillo, Universidad Nacional de Tucumán, Tucumán, Argentina (M. V. Colomo).
MACN: Museo Argentino de Ciencias Naturales, Buenos Aires, Argentina (A. Roig).
MLPA: Museo de La Plata, La Plata, Argentina (A. Lanteri).
MLUH: Martin-Luther-Universität, Zoologisches Institut, Halle, Germany (K. Schneider)
UNSM: University of Nebraska State Museum, Lincoln, NE, USA (B. C. Ratcliffe, M. L. Jameson-Russell).
USNM: United States National Museum, Washington D.C. USA (D. Furth).


New synonymies for Anomiopsoides heteroclyta (Blanchard)

Anomiopsoides heteroclyta (Blanchard, 1845: 156).
Anomiopsoides aberrans Martínez, 1945b: 403, syn. by Ocampo (2005).
Anomiopsoides aurita (Burmeister 1861: 63), new synonymy.
Anomiopsoides catamarcae Martínez,1945b: 400, new synonymy.
Anomiopsis furciferum Burmeister, 1861: 64, syn. by Martínez (1945b).
Anomiopsis heteroclytum Burmeister 1873: 406 (nec Blanchard, 1845), syn. by Martínez (1945a).
Anomiopsoides pereirai (Martínez 1944: 39), syn. by Ocampo (2005).
Anomiopsoides xerophila Martínez 1945b: 395, syn. by Ocampo (2005).

In my revision of the genus Anomiopsoides (Ocampo, 2005), I considered Anomiopsoides catamarcae Martínez as a valid species based on the examination of one paratype male deposited at CMNC from Argentina, Catamarca, Tinogasta. At that time, I could not examine the holotype specimen, which was not at the MACN. Recently, Dr. Arturo Roig, made available for my study the holotype specimen of A. catamarcae, which was returned to the MACN. Differences observed between type material of A. catamarcae and of A. heteroclyta are only observed on the shape of the clypeal armature. The shape of the medial clypeal processes of A. heteroclyta shows a high degree of variation (Fig. 1 a-m) and, by itself, it is not a reliable character to separate species of this genus. Based on my observations of the holotype and one paratype of A. catamarcae at CMNC, and long series of A. heteroclyta, I conclude that there are no character-based differences between A. heteroclyta (Blanchard) and A. catamarcae Martínez, and so I place these species in synonymy.

Fig. 1. Morphological variation observed in the pronotal armature and in size of Anomiopsoides heteroclyta (Blanchard), a-g: males, h-m: females, n: dorsal views showing size variation. Specimens pictured here were all collected at the same locality, same day (Argentina, Salta, 4 km N of Cafayate), and at the same time (10:00-12:00 hs).

The species Anomiopsoides aurita (Burmeister) was originally described in the genus Anomiopsis Burmeister 1861 (junior homonym of Anomiopsis Westwood 1837). According to the original description (Burmeister, 1861) male and female of this species do not exhibit sexual dimorphism. I studied Burmeister material, but I was not able to find any male specimen corresponding to the species described by Burmeister as A. aurita. The holotype of Anomiopsoides aurita is a female specimen deposited at MLUH. Martínez (1945b), in his revision of the genus Anomiopsoides, refers to one male specimen of A. aurita in his collection: «…1 de Catamarca, Tinogasta (Dr. Schaefer-leg)…», I was able to study this specimen and found that is actually a female specimen. I collected extensively in the localities that Burmeister (1961) and Martínez (1945a, 1945b) indicated for A. aurita and never found a male specimen that can be identified as such. In my revision of the genus Anomiopsoides (Ocampo 2005) I indicated that both males (according to Burmeister 1961 and Martínez 1945b) and females of A. aurita can be recognized by «the clypeal surface perpendicular in frontal view between clypeal medial processes», although at that time I had no observations of male specimens attributed to A. aurita. The observation of long series of A. heteroclyta from Argentina, Catamarca and Salta provinces, indicate that the clypeal armature is highly variable among female specimens of this species (Fig. 1 hm). Based on current evidence and the absence of male specimens that indicate the contrary, I consider that A. aurita is a female of A. heteroclyta that presents the normal variation of the clypeal shape found among specimens of this species. I conclude that there are no character-based differences between A. heteroclyta (Blanchard) and A. aurita (Burmeister), and so I place these species in synonymy.

Anomiopsoides fedemariai sp. nov. (Figs. 2-4, 8)

Figs. 2, 3. Anomiopsoides fedemariai sp. nov., 2, dorso-lateral view; 3, dorsal view.

Fig. 4. a: Anomiopsoides fedemariai sp. nov., base of elytron and pronotum showing the teeth-like process at the base of the 7th elytral stria, and 7th stria strongly carinated; b: base of elytron and pronotum of A. heteroclyta (Blanchard).

Diagnosis. Males and females of this A. fedemariai are separated from A. heteroclyta by the pseudoepipleura well developed at level of the 7th stria, and the 7th stria strongly carenated, reflexed and with tooth-like process at elytral base (Figs. 2-4). Males are recognized by the clypeal surface perpendicularly angled downwards with respect to the surface of the frons (Figs. 2, 4); the ventral surface absent; the clypeal medial process developed and reflexed, with the inner margin straight or slightly curved, and the external margin straight or slightly curved, and the apex divergent, reflexed; and the clypeal lateral process developed, with one poorly developed lateral tooth (Fig. 3), this character is different in A. biloba, and A. cavifrons (Figs. 5, 6). Females of this species are recognized by the clypeus with the medial processes short (shorter or as long as the length of frons plus length of clypeus), apex pointed; and clypeal surface not perpendicular in frontal view with respect to surface of frons and between clypeal medial processes, this character is different in A. biloba, and A. cavifrons (Figs. 5, 7).

Figs. 5-7. Head of: 5, Anomiopsoides biloba; 6 and 7, A. cavifrons male and female respectively.

Description. Holotype male. Length 17.0 mm. Width 14.5 mm. Color black, surface shiny to matte. Head (Figs. 2, 3): Shape subrectangular, transverse. Frons slightly foveate to punctate. Frontoclypeal and clypeogenal sutures evident. Frontoclypeal suture elevated on sides, slightly sinuate. Clypeogenal surface strongly foveate. Genal posterior angle rounded; lateral margin irregular, setose. Clypeal surface strongly obliquely angled downwards to perpendicular with respect to surface of frons; ventral surface absent; ventral process developed, acute. Clypeal medial process developed, reflexed, parallel; dorsal and external surface strongly foveate, apex smooth; internal surface smooth to slightly foveate, ventral half with fringe of setae; inner margin straight; external margin straight; apex rounded or acute, divergent, reflexed. Clypeal lateral process developed, apex pointed, with 1 lateral tooth. Pronotum (Fig. 3): Surface punctate, punctures moderately dense to sparse, small to moderate in size. Lateral margin with long, dense setae on basal half, and moderately dense, short setae on apical half; margin beaded, denticulate, denticles larger at middle. Elytron (Figs. 2-4): Striae impressed, punctate; punctures small. Intervals without minute tubercles. Pseudoepipleura well developed at level of 7th stria; 7th stria strongly carenated, reflexed and with tooth-like process at elytral base. Venter: Metasternum with small, apical and posterior longitudinal depressions. Legs: Protibial teeth with pointed apex. Protibial spur curved at apex. External mesotibial spur slightly curved on apical third, apex pointed.
Allotype female. Length 17.6 mm. Width 14.7 mm. As male except in the following respects: Clypeus with surface obliquely angled downwards with respect to surface of frons, ventral surface developed; ventral process developed, acute. Clypeal medial process developed; dorsal and external surface strongly foveate, apex smooth; internal surface densely foveate; inner and external margins slightly sinuous; apex rounded, reflexed.

Variation in paratypes. 77 males and 68 females. Length 11.2-17.8 mm. Width 8.3- 14.8 mm. Anomiopsoides fedemariai, like other species of this genus, shows substantial variation in the shape of the medial clypeal processes. These processes vary in length and width at the base and at the apex. Most specimens have their clypeal processes parallel, but different degrees of convergence or divergence toward the apex was observed in some specimens. Variation of the elytral and pronotal puncture density (sparse or moderate), puncture size (small or medium), and elytral striae impression was also observed.

Etymology: I take great pleasure in naming this species after my son Federico Maria, «Fede».

Type material. Holotype male deposited at IADIZA labeled: «ARGENTINA: San Juan / Salinas de Mascasín, 423 m / S 31°25'39" W 67°04'05" / XII-1-2003. F. C. Ocampo»; Anomiopsoides/ fedemariai / HOLOTYPE / F. C. Ocampo» (red label). Allotype with same label as holotype except: «Anomiopsoides/ fedemariai / ALLOTYPE / F. C. Ocampo» (red label). Eleven male paratypes (in 95° ethanol) and 10 female paratypes (in 95° ethanol) at FCOC with same label as holotype. Thirty five male (14 in 95% ethanol) and 42 female paratypes (five in 95% ethanol) labeled: ARGENTINA: San Juan / RP 20, 12 km E Encón / S 32°12'55" W 68°40'68" / Jan-4- 2003, 524 m / F. C. Ocampo, A. B. T. Smith»; distributed: 20 male (eight in 95% ethanol) and 32 female paratypes (four in 95% ethanol) at FCOC, five males and five females paratypes at UNSM, four male and four female paratypes at ABTS. Nine males (five in 95% ethanol) and four female (three in ethanol) paratypes at FCOC labeled: «ARGENTINA: Mendoza / RP 142 and Rio Mendoza, / S32°35'38" W68°17'27" / Jan-4- 2003, 595 m / F.C. Ocampo, A.B.T. Smith». Six male and five female paratypes at IADIZA labeled: «ARGENTINA: Lavalle / Puente Rio Mendoza / 21 Feb. 2006. E. Ruiz». Three males and one female paratypes (in 95% ethanol) at FCOC labeled: «ARGENTINA: San Juan / Salinas de Mascasín, 423 m / S 31°25'39" W67°04'05" / XII-1-2003. F. C. Ocampo.» Two males and two females at IADIZA labeled: «ARGENTINA: Mendoza / 18 km N Costa de Araujo / 32°25'32" S 68°18'58" W / 29 Jan 2004, 582 m / G. Debandi». Three males and three females paratypes at IMLA (two males and two females), and IADIZA (one male and one female) labeled: «R. ARGENTINA / SAN JUAN / Mirayes / 12.1.1982 / A. Willink y otros.» One male and one female paratypes at USNM labeled: «Pie de Palo San Juan / 11 Mar.1920 Argentina / Cornell U. expedition». Two male and one female paratypes at IADIZA labeled: «Mendoza, 50 km de / Las Catitas, Puesto / La Josefa. 11-XII-79 / col. A. Roig / Sergio Roig». One female paratype at IADIZA labeled: «Rca Arg Mendoza / El Borbollón / I-79 col. A. Roig / Sergio Roig». One paratype male at IADIZA labeled: «R.A. Mza. Lavalle / 25-2-82 / col. Claver / IADIZA». One female paratype at IADIZA labeled: «Bermejo, San Juan / 12-III-1949 / Biraben col.». One male and one female paratypes at IADIZA labeled: «Mendoza: Lavalle / Ruta 142 km 45 / 6/XII/97. Roig». One male paratype at IADIZA labeled: «Mendoza: Lavalle / Ruta 142 / 6/XII/97. Flores». One paratype male at IADIZA labeled: «Mendoza Lavalle / El Quince 16/II/ 98 / S. Roig G. Dabandi». One paratype male at IADIZA labeled: «R. A. San Luis / Dto Ayacucho / Bella Vista (11 hs.) / 14-4-95. G. Flores / IADIZA». One paratype female at CASC labeled: «Mendoza / Argentina»; «Anomiopsoides pereirai / P. Gutierrez 1959». One paratype male at MLPA labeled: «San / Juan»; «Anomiopsoides pereirai / A. Martínez 1958». All paratypes with my yellow paratype label: «Anomiopsoides/ fedemariai / PARATYPE / F. C. Ocampo».

Distribution (Fig. 8). ARGENTINA (147): Mendoza: Costa de Araujo (4); El Borbollón (1); El Quince (1); Lavalle (1); Las Catitas, Puesto La Josefa (3); Ruta 142, intersection with Rio Mendoza (24); Ruta Provincial 142 (no mora data) (1) Ruta Provincial 142, km 45 (2). San Juan: Bermejo (1); Encón, 12 km E, RP 20 (77); Mirayes (6); Salinas de Mascasín (23); Sierra de Pie de Palo (2). San Luis: Bella Vista (1).

Fig. 8. Distribution map of A. fedemariai sp. nov.

Temporal distribution. January (101); February (14); March (3); April (1); December (29).

Remarks. The biology of A. fedemariai is similar to the other species of Anomiopsoides. Specimens were collected at midmorning and late afternoon to early evening (9:00-12:30 am; 4:30-7:30 pm, [ UTC/GMT -3 hours] ). According to my observations, the food relocation behavior and food preferences of A. fedemariai are as those indicated for A. heteroclyta and described by Zunino et al (1989), Monteresino & Zunino (2003), and Ocampo (2005). The larva and nidification behavior of A. fedemariai are unknown.

Key to species of Anomiopsoides

Anomiopsoides species show considerable intraspecific variation due to particular conditions during an individual's development, environmental conditions of the area in which the specimen lived, and/or age of the specimen at the time it was collected. The above factors make identifications difficult for some Anomiopsoides specimens.

1. Medial processes of clypeus with external margin straight, slightly sinuous, or slightly lobed and medial margin concave or straight (Figs. 1-3, 6, 7) ........................................ 2 1'. Medial processes of clypeus with external margin broadly rounded or lobed (Fig. 5) ................................. A. biloba (Burmeister)
2. Clypeal disc with large fovea before and between medial processes. Clypeus with medial processes long (longer than rest of head), with apex divergent (male) (Fig. 6) or clypeus with medial processes short (shorter or as long as rest of head), with apex pointed (female) (Fig. 7) .... A. cavifrons (Burmeister)
2'. Clypeal disc without foveae before medial processes. Clypeus with medial process not as above ................................................... 3
3. Pseudoepipleura developed at level of 7th stria, 7th stria not carenated or reflexed, without tooth-like process at elytral base ...... ............................ A. heteroclyta (Blanchard)
3'. Pseudoepipleura well developed at level of 7th stria, 7th stria strongly carenated, reflexed, with tooth-like process at elytral base (Figs. 2-4) ........ A. fedemariai sp. nov.


I thank the curators and collection mangers of the institutions listed in the material and methods section for making material available for this research, thank you very much! I thank Sergio Roig-Juñent, Gustavo Flores, and Guillermo Debandi (Instituto Argentino de Investigaciones de las Zonas Áridas, IADIZA, Mendoza) for their help and for providing logistic support during my field trips conducted in Argentina in 2003, 2004, 2005 and 2006. I thank Andrew Smith (Canadian Museum of Nature, Ottawa), Gustavo Flores, Guillermo Debandi, and Eider Ruiz-Manzanos (IADIZA) for collecting specimens for this study. I thank Gualberto Zalazar, Eider Ruiz-Manzanos, and German San-Blas (IADIZA) for their help and assistance during the field trips in 2003, 2005, and 2006. I thank Aura Paucar for taking many of the pictures that illustrate this work. I thank Mary Liz Jameson-Russell for her critical review of an earlier version of the manuscript and valuable comments. This project was supported by an NSF/PEET grant (DEB-0118669) to Mary Liz Jameson and Brett Ratcliffe, by an NSF Biodiversity Surveys and Inventories grant (DEB-0342189) to Andrew Smith and Federico Ocampo, and an NSF-DBI grant (0500767) to Mary Liz Jameson and Federico Ocampo.


1. BARBERO, E. & C. PALESTRINI. 1993. Descrizione del terzo stadio larvale di Anomiopsoides cavifrons (Burmeister, 1861). Bollettino della Società Entomologica Italiana 125: 53-60.
2. BLACKWELDER, R. E. 1944. Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America, Part 2. Bulletin of the United States National Museum 185: 189-341.
3. BLANCHARD, C. E. [ 1841-1847] . Insectes. En: A. d'Orbigny, Voyage dans l'Amerique Méridionale, vol. 6, pt. 2. Paris & Strasbourg, pp. 61-222.
4. BRULLÉ, G. A. 1834. Historie Naturelle des Insectes. Vol. 4, Coléoptères I, Pillot, Paris.
5. BURMEISTER, H. 1861. Die Ateuchiden ohne Fulskrallen, monographisch bearbeitet. Berliner Entomologische Zeitschrift 5: 55-67.
6. BURMEISTER, H. 1873. Lamellicornia Argentina. Stettiner Entomologische Zeitung 34: 403-417.
7. GILLET, J. J. E. 1911. Pars 38. Scarabaeidae: Coprinae. I. In: Schenkling (ed.), Coleopterorum Catalogus. W. Junk, Berlin, pp. 1-100.
8. MARTÍNEZ, A. 1944. Insectos nuevos o poco conocidos II. Revista Argentina de Entomología 2: 34-40.
9. MARTÍNEZ, A. 1945a. Insectos nuevos o poco conocidos III (Col. Scarabaeidae). Revista de la Sociedad Entomológica Argentina 12: 261-279.
10. MARTÍNEZ, A. 1945b. Insectos nuevos o poco conocidos IV (Col. Scarabaeidae). Revista de la Sociedad Entomológica Argentina 12: 394-409.
11. MONTERESINO, E. M. & M. ZUNINO. 2003. Sobre el comportamiento de la alimentación de Eucraniini (Coleoptera: Scarabaeidae: Scarabaeinae). En: Onore, G., P. Reyes Castillo & M. Zunino (eds.). Escarabaeidos de Latinoamérica, Estado Actual del Conocimiento. Sociedad Entomológica Aragonesa. Zaragoza, Spain. pp. 75-80.
12. OCAMPO, F. C. 2005. Revision of the southern South American endemic genus Anomiopsoides Blackwelder 1944 (Coleoptera: Scarabaeidae: Scarabaeinae: Eucraniini) with description of its food relocation behavior. Journal of Natural History 39: 2537-2557.
13. OCAMPO, F. C. 2007. Evolution and conservation biology of the South American Eucraniini dung beetles. /SBSAAL/Eucraniini/project.html. Accessed day: May, 2007.
14. WESTWOOD, J. O. 1837. Illustrations of exotic entomology, containing upwards of six hundred and fifty figures and descriptions of foreign insects, interspersed with remarks and reflections on their nature and properties, Vol. 1, H.G. Bohn, London.
15. WHEELER, Q. D. &, N. I. PLATNICK. 2000) The phylogenetic species concept (sensu Wheeler & Platnick). En: Wheeler Q. D. & R. Meier (eds.). Species Concepts and Phylogenetic Theory. Columbia University Press, NY, USA. pp 55-69.
16. ZUNINO, M., E. BARBERO, & M. LUZZATTO. 1989. Food relocation behavior in Eucraniina beetles (Scarabaeidae) and the constraints of xeric environment. Tropical Zoology 2: 235-240.
        [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]         [ Links ]

Recibido: 29-05-2007;
aceptado: 12-09-2007

Creative Commons License Todo el contenido de esta revista, excepto dónde está identificado, está bajo una Licencia Creative Commons