Mastozoología Neotropical ISSN: 0327-9383 [email protected] Sociedad Argentina para el Estudio de los Mamíferos Argentina
Ubilla, Martín; Rinderknecht, Andrés A Late Miocene Dolichotinae (Mammalia, Rodentia, Caviidae) from Uruguay, with comments about the relationships of some related fossil species Mastozoología Neotropical, vol. 10, núm. 2, julio-diciembre, 2003, pp. 293-302 Sociedad Argentina para el Estudio de los Mamíferos Tucumán, Argentina
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A LATE MIOCENE DOLICHOTINAE (MAMMALIA, RODENTIA, CAVIIDAE) FROM URUGUAY, WITH COMMENTS ABOUT THE RELATIONSHIPS OF SOME RELATED FOSSIL SPECIES
Martín Ubilla1 and Andrés Rinderknecht2
1 Paleontología, Facultad de Ciencias. Iguá 4225. 11400. Montevideo, Uruguay. ++ 5982 5258618 fax: ++5982 5258617;
ABSTRACT. The oldest dolichotine from Uruguay (late Miocene deposits of southwestern) is described and here assigned to the genus “Prodolichotis” Kraglievich, 1932. It is interme- diate in size between Pediolagus salinicola (Burmeister, 1876) and Dolichotis patagonum (Zimmermann, 1780) and similar to “Prodolichotis” lacunosa (Ameghino, 1888). The material consists in an incomplete skull which has a diastema longer than P4-M3 length, the naso- lacrimal foramen absent in lateral view, the anterior border of mesopterygoid fossa posi- tioned between the anterior and posterior prisms of M2 and the internal posterior fold of M3 has diverging borders forming an angle less than 90º. This M3 morphology shared with the Dolichotinae-Caviinae clade suggests that parallel borders are a derived feature in some Dolichotinae and the condition observed in the Caviinae is a primitive state. Judging by the morphological pattern presented by “Prodolichotis” pridiana Fields (1957)–nasolacrimal fora- men in the maxilla, interorbital width less than the braincase width, and diverging M3 fold– we agree with previous opinions that this species should be included in the Caviinae instead of Dolichotinae. It is discussed a close relationship between Orthomyctera Ameghino, 1889 and Caviinae judging by their similar morphological pattern including interorbital width less than the braincase width, molar series with similar length or longer than the diastema length, nasolacrimal foramen exposed laterally in the maxilla, and similar configuration of the M3 and skull size and overall shape. The occurrence of a dolichotine in the late Miocene of south- western Uruguay suggests open and probably arid or semiarid terrestrial environments.
RESUMEN. Un Dolichotinae (Mammalia, Rodentia, Caviidae) del Mioceno tardío de Uruguay, con comentario sobre las afinidades de especies fósiles relacionadas. Se describe el registro más antiguo de Dolichotinae para el Uruguay proveniente de depósitos del sur-oeste y asignables al Mioceno tardío y se incluye dentro del género “Prodolichotis”. Su tamaño es intermedio entre las especies Pediolagus salinicola y Dolichotis patagonum y similar a “Prodolichotis” lacunosa. El material consiste en un cráneo incompleto con el diastema mas largo que la serie P4-M3; foramen nasolacrimal no apreciable en vista lateral; borde anterior de la fosa mesopterigoidea ubicado entre los prismas del M2; hendidura posterior interna del M3 con bordes divergentes formando un ángulo menor de 90º. Esta configuración del M3 compartida por el clado Dolichotinae-Caviinae sugiere que la condición de poseer bordes paralelos en este molar que presentan los actuales Dolichotinae podría ser una característica derivada y no primitiva, y la configuración existente en el M3 de los cavinos primitiva. Teniendo en cuenta algunas características morfológicas de “Prodolichotis” pridiana –foramen nasolacrimal visible en vista lateral, ancho interorbitario menor que el ancho de la caja craneana, hendidura posterior interna del M3 con bordes divergentes– concordamos con opiniones previas, en que esta especie debe incluirse en Caviinae y no en Dolichotinae. Se discuten evidencias para incluir al género Orthomyctera dentro de la subfamilia Caviinae: ancho interorbitario marcadamente menor que el de la caja craneana;
Recibido 11 diciembre 2002. Aceptación final 23 junio 2003. 294 Mastozoología Neotropical / J. Neotrop. Mammal.; 10(2):293-302 M. Ubilla and A. Rinderknecht
largo de la serie P4-M3 menor que el largo del diastema; foramen nasolacrimal expuesto lateralmente en el maxilar y M3 similar al de los cavinos. La presencia de un dolichotino en el Mioceno tardío del Uruguay sugiere ambientes terrestres abiertos y probablemente áridos o semiáridos.
Key words: Rodentia, Dolichotinae, late Miocene, Uruguay
Palabras clave: Rodentia, Dolichotinae, Mioceno tardío, Uruguay.
INTRODUCTION Walton, 1997; Cione et al., 2000) (Fig. 1). Calcaterra (1972) referred Dolichotis major Despite the fact that there are different opi- (Gervais and Ameghino, 1880) to Pleistocene nions about the monophyletic status of sediments of southwestern Uruguay, the only dolichotines as a subfamily of Caviidae fossil dolichotine known from this country. (Quintana, 1997, 1998), they are recognised as In this paper we describe the oldest record peculiar hystricognath rodents whose age of the subfamily Dolichotinae from Uruguay ranges from the late Tertiary to Recent (Arazatí, San José Department; the Camacho (Vucetich and Verzi, 1995). Living Formation) and discuss the taxonomic value of Dolichotinae are endemic to southern South dental and cranial characters in order to im- America (Redford and Eisenberg, 1992) and prove our knowledge of this poorly known the two living species are usually arranged in group. Comments about the taxonomic assign- one genus Dolichotis or in two different gen- ment of the fossil species “Prodolichotis” era, Dolichotis and Pediolagus, the “mara” and pridiana and fossil genus Orthomyctera to the “conejo del palo” respectively (see Kraglievich, subfamily Caviinae are also provided. 1930; Mares and Ojeda, 1982; Wilson and Reeder, 1993; Quintana, 1998). Dolichotis MATERIAL AND METHODS patagonum is widespread in central and south- ern Argentina and Pediolagus salinicola is Institutional abreviations: MACN-Ma: Museo distributed in NW Argentina and the Chaco of Argentino de Ciencias Naturales, Sección Paraguay and southern Bolivia (Wilson and Mastozoología. MACN-Pv: Museo Argentino de Reeder, 1993) (Fig. 1). These species inhabit Ciencias Naturales-Paleontología de Vertebrados. MLP-DZV: Museo de La Plata, División Zoología arid areas, especially desert, grassland scrub, de Vertebrados. IAVA: Instituto Alfredo Vázquez thorn scrub and dry forest (Mares and Ojeda, Acevedo, Montevideo. MNHN-P-PAM: Muséum 1982). National d’Histoire Naturelle, Paleontologie. Paris, There is also a controversy among authors France about the identity and the subfamilial relation- The following cranial material was used for com- ships of several fossil taxa described as parison: dolichotines or caviines (Quintana, 1997, 1998). Dolichotis patagonum: MACN-Ma: 302; 303; As a complete taxonomic revision of the entire 304; 917; 29879; 30229; 26209; 35406; 49132; subfamily including the fossil taxa is beyond 27152; 14532; 28183; 15533; 29894; 13755; 2665; 28190; 4319; 29193; 15534; 2516; 13756; 4959. the scope of this paper some taxa are included IAVA: 1 specimen: w/n. MLP-DZV: 488, 226, 208, in quotation-marks. 597, 247, 251, 30.X.95.11, 371, 1699, 687, 1783, Fossil dolichotines are known from several 2.VI.60.11, 418, 230, 723, 369, 638. Pediolagus localities in the Neogene and Quaternary ages salinicola: MACN-Ma: 41218, 29872, 41216; in South America, but in general the record is 28188; 4326; 17366; 4286; 17363; 3071; 28213; scarce and not well studied. The oldest record 17365; 29874; 26208; 28143; 30239; 28144; 30154; of the group comes from the Miocene of Co- 33166; 30390; 41217; 3228; 26210; 4279; 30155. lombia and Argentina, whereas the Pliocene MLP-DZV: 672, 673, 1081, 2671. Orthomyctera record is restricted to Argentina (Fields, 1957; rigens (Ameghino, 1889): MACN-Pv: 7319, 1661- 62. Orthomyctera andina (Rovereto, 1914): MACN- A LATE MIOCENE DOLICHOTINAE FROM URUGUAY 295
Fig. 1. A) Geographic ocurrences of Miocene and Pliocene Dolichotinae in South America: 1) Middle Miocene of Colombia, Villavieja Fm. (innominated genus following Walton, 1997), 2) Late Miocene, Catamarca, Argentina, Andalhuala Fm., 3) Late Miocene Entre Ríos, Argentina, Ituzaingó, Fm. 4) Pliocene, Buenos Aires Province. Montehermoso Fm., 5) Pliocene, Chapadmalal, Buenos Aires Province, Argentina. Dotted and shaded areas: recent distribution of Dolichotinae. B-C) geographic locality of MNHN-1633 Dolichotinae fossil from Uruguay. The arrow shows the fossil location.
Pv: 8350, 8351, 8399. “Prodolichotis” prisca GEOLOGICAL SETTING (Rovereto, 1914): MACN: 8346, 8348, 12335. AND BIOSTRATIGRAPHY “Prodolichotis” lacunosa (Ameghino, 1888): MACN-Pv: 7333, 7334. Dolichotis platicephala Three lithostratigraphic units can be recognised Ameghino, 1889: MACN-Pv: 556. Dolichotis sp.: in Arazatí outcrops, these are from the base to MACN-Pv: 10847.”Orthomyctera” chapalmalense Ameghino, 1908: MLP-DP-15-229. Dolichotis the top: Camacho, Raigón (= San José Forma- major: MNHN-P-PAM-267. tion) and Libertad formations (Fig. 2). The Six cranial and dental linear dimensions were outcrops of the Camacho Formation at this measured with a caliper to the nearest 0.1 mm. In locality (considered the Kiyú Formation by Table 1 we provide character abbreviations. Values Francis and Mones, 1965) are the upper facies in Table 2 were calculated by means of Statistic of a Late Miocene transgresive episode, which 4.2-1993 software. Occlusal surfaces of upper mo- includes ostreid biostromes, “patch reefs” and lars were drawn with a Zeiss Stemi SV11 drawing ichnofossils in a greenish-gray clayey silt tube mounted on a stereomicroscope. (Martínez, 1994; Sprechmann et al., 2000; Verde, 2002). The fossil mammal assemblage 296 Mastozoología Neotropical / J. Neotrop. Mammal.; 10(2):293-302 M. Ubilla and A. Rinderknecht
Table 1
Measurements (mm) of the Arazatí material (MNHN-1633) and comparative specimens included in the genus Dolichotis (D), Orthomyctera (O) and “Prodolichotis” (P). 1: constriction interorbital maximum, 2: palatal width (including M3), 3: diastema length, 4: P4-M3 length, 5: M3 length, 6: M3 width.
123456
MNHN-1633 26 25,6 31,6 22,2 7 4,4 O. rigens MACN-Pv-7319 14,2 13,7 16,1 12,7 4 1,4 O. rigens MACN-Pv1661-62 - - - 15,7 4,5 3 O. andina MACN-Pv-8350 12,3 12 13,4 12,5 3,6 2,4 O. andina MACN-Pv-8351 15 15 - 13,4 4,4 3,2 O. andina MACN-Pv-8349 14,5 14 - 13 4,2 2,4 P. prisca MACN-Pv-8348 19 20 21 20,5 6,6 2,6 P. prisca MACN-Pv-8346 22 20 - 17 5,8 2,5 P. lacunosa MACN-Pv-7334 32 28 - 23,4 8 4,4 P. lacunosa MACN-Pv-7333 29 27,4 - 22,5 7,8 4,6 P. prisca MACN-Pv-12335 23 21,6 25 17,3 6,2 3,7 D. platicephala MACN-Pv-556 49 33,4 43,3 27,6 9,4 5,1 O. chapalmalense MLP15-229 29,8 33,5 39,8 27,2 10 6
Table 2
Statistical values for living species Dolichotis patagonum and Pediolagus salinicola. n: sample size; x: mean; min. and max.: range; ds: standard deviation. Measurements explained in Table 1.
Dolichotis patagonum Pediolagus salinicola n x min max ds n x min max ds
1 41 40,9 32,8 46 3,12 27 24,7 20,5 30,5 2,08 2 41 33,6 25,8 38,2 2,35 28 20,4 16,4 22,6 1,47 3 41 41,4 31,8 46,6 3,27 27 24,4 18,8 29,2 2,82 4 41 25,6 20 30,1 2,34 23 16,5 13,1 19,3 1,68 5 37 9 6,9 10,5 0,92 24 5,9 3,9 7 0,93 6 37 5,4 3,9 8,9 0,82 23 3,3 2,1 4,4 0,62
of the Camacho Formation at Arazatí cliffs is Transport processes explain the ocurrence of represented by terrestrial taxa of Huayquerian- terrestrial mammals along with estuarine and “Mesopotamian” affinities (Perea et al., 1994). marine ostreids and ray teeth remains, fresh- It is important to note that Cione et al. (2000) water turtles and flamingoes (Perea et al., 1996; consider the “Mesopotamian” as an invalid unit Ubilla et al., 1990). The Dolichotinae remain correlating the “conglomerado osífero” of the was found a few centimeters above the ostreid Ituzaingó Formation (“Mesopotamian”) with biostromes and close to a tooth ray. The the Late Miocene Huayquerian stage. They taphonomic and sedimentological features of referred Prodolichotis to the Late Miocene this particular lithofacies of the Camacho For- Ituzaingó Formation from Paraná of Argen- mation suggest an estuarine or marginal shal- tina. low-water marine depositional environment A LATE MIOCENE DOLICHOTINAE FROM URUGUAY 297
Fig. 2. Stratigraphic profile of Arazatí exposures. The arrow indicates the location of “Prodolichotis” sp.. c: clay, s: sand, f: fine, m: medium, c: coarse, p: pebbles. Colour codes following Rock Color Chart Committee. 1980. Geological Society of America. 298 Mastozoología Neotropical / J. Neotrop. Mammal.; 10(2):293-302 M. Ubilla and A. Rinderknecht
(Perea et al., 1996), similar to that described DISCUSSION AND CONCLUSION by Behrensmeyer and Hook (1992). Following the taxonomic revision of Quintana SYSTEMATIC PALEONTOLOGY (1998) we consider the skull of Arazatí as a Dolichotinae on the basis of the following Class MAMMALIA Linnaeus, 1758 characters: the diastema is longer than P4-M3 Order RODENTIA Bowdich, 1821 length, the nasolacrimal foramen is absent in Suborder HYSTRICOGNATHI Tullberg, 1899 lateral view and the mesopterygoid fossa posi- Superfamily CAVIOIDEA Kraglievich, tioned at the level of the anterior and posterior 1930. prisms of M2. It is important to note that this Family CAVIIDAE Waterhouse, 1839 Subfamily DOLICHOTINAE Pocock, character assemblage is shared by all taxa in 1922 the subfamily Dolichotinae. With regard to the living species of Genus “Prodolichotis” Kraglievich, Dolichotinae, the Arazatí material differs in size 1932 and in M3 structure. Its size is intermediate “Prodolichotis” sp. (Figs. 3 and 4) between Dolichotis patagonum and Pediolagus salinicola and similar to that “Prodolichotis” Material referred — MNHN-1633: fragmented lacunosa (Figs. 4, 5 and Table 2). Despite the skull with incisors, diastema, palate with left and fact that upper tooth row morphology is quite right P4-M3, part of the infraorbital foramen and similar in the examined genera (Fig. 4), M3 incomplete zygomatic arches. Horizon and locality — in silty sandy and grayish- configuration allows one to differentiate three green sediments of the Camacho Formation (=Kiyú groups which laudably convey its variability: Lithofacies), immediately above an oyster bank (Fig. a) in Dolichotis and Pediolagus the second 2), About 6 km southeast of Sauce Creek, internal fold is penetrating and has cementum; Departament of San José, Puerto Arazatí, Uruguay the posterior border of the second prism and (Fig. 1). the anterior border of the additional posterior Description — incomplete skull of a prism tend to be parallel, b) in Orthomyctera Dolichotinae with some linear dimensions rigens and Orthomyctera andina (which are modified by lateral deformation; intermediate not considered as dolicotines by Quintana, in size between Pediolagus salinicola and 1998) there is approximately 90º between the Dolichotis patagonum and similar to that of borders of the internal folds of the second and “Prodolichotis” lacunosa. Incisors broken at third prisms, c) the “Prodolichotis” group and the alveolar margin; diastema longer than P4- the Arazatí material show diverging borders M3 length; nasals wide and fragmented; in lat- with an angle tending to be less than 90º, an eral view the nasolacrimal foramen is absent; intermediate configuration between the two pre- P4-M2 sub equal with cordiform prisms con- vious groups. nected by an isthmus and covered with an in- Kraglievich (1932) described the genus terrupted layer of enamel; internal fold with Prodolichotis and referred to it several species cementum; external folds delimited by the an- defined on the basis of skull and mandible terior and posterior prisms; M3 with suboval material. Though the genus need revision, the additional posterior prism, smaller than the fossil from Uruguay fit well within its morpho- others; the internal fold between the second logical characteristics. Taking into account the and third prisms slightly penetrating and with- similar size, shape and molar configuration, we out cementum delineated by diverging borders assign MNHN-1633 to “Prodolichotis” sp. and at an angle less than 90º; unexcavated and short we consider the available characters insuffi- palate with posteriorly diverging tooth rows; ciently diagnostic to describe a new species anterior border of mesopterygoid fossa posi- nor assign it to previously described species. tioned between the anterior and posterior prisms In the phylogenetic analysis provided by 3 of M2. Quintana (1998), an M with diverging bor- Measurements — see Table 1. ders is considered a synapomorphy of Caviinae. A LATE MIOCENE DOLICHOTINAE FROM URUGUAY 299
Nevertheless, in our point of view, this should be revised in light of the variability and inter- mediate condition observed in “Prodolichotis”. In fact, diverging borders may have been the ancestral state of the Dolichotinae-Caviinae clade, and the tendency to become parallels a derived feature in some Dolichotinae. So, at this level of resolution the condition observed in the Caviinae could be interpreted as a primi- tive state. However, the Caviinae Microcavia robusta Gervais and Ameghino, 1880 has a deep flexus with parallel borders (Quintana, 1996) which could suggests an independent evolution of this character in each subfamily. The species “Prodolichotis” pridiana de- scribed by Fields (1957) from the Miocene of Colombia differs from the material from Uru- guay by several characters such as the pres- ence of the nasolacrimal foramen in the max- illa, interorbital width less than the braincase width, and clearly diverging M3 flexus. This Fig. 3. MNHN-1633 “Prodolichotis” sp. A: dorsal view, set of characters is shared with the caviines. B: palatal view, C: lateral view. Nevertheless, diastema length is longer than
Fig. 4. Oclusal surface of left upper molars of: A) Dolichotis patagonum, B) “Orthomyctera” chapalmalense, C) Pediolagus salinicola, D) “Prodolichotis” lacunosa (fide Kraglievich, 1930), E) “Prodolichotis” prisca, F) MNHN-1633, G) Orthomyctera rigens and H) Orthomyctera andina. B, D and E are right inverted tooth rows. The arrows indicate the different morphology of the internal fold between the second and third prisms of M3. 300 Mastozoología Neotropical / J. Neotrop. Mammal.; 10(2):293-302 M. Ubilla and A. Rinderknecht
Fig. 5. Bivariant plot of palate width (Y) and P4-M3 length (X) for living species of Dolichotinae, fossil genera and the material from Uruguay (MNHN-1633), along with two species of Orthomyctera. Black bars represent 95% confidence intervals around the mean. cheek teeth length as in dolichotines. Judging with an additional elongation in the anterior by the morphological pattern presented by prism which in general does not become a true “Prodolichotis” pridiana, the generic and prism. In Kerodon and “Prodolichotis” subfamilial relationships of this taxon need to pridiana this additional elongation becomes a be revised. We agree with Quintana (1998) that well defined prism. This also suggests a rela- this species should be included in the Caviinae tionship between these two taxa. instead of the Dolichotinae. It is also interest- There is striking evidence in favour of the ing to note that this suite of characters is also exclusion of Orthomyctera from the found in the peculiar living caviine Kerodon Dolichotinae as was suggested by Quintana
Cuvier, 1825. In caviines P4 has two prisms (1998). Orthomyctera, was described by A LATE MIOCENE DOLICHOTINAE FROM URUGUAY 301
Ameghino (1889) and was represented by four ACKNOWLEDGMENTS species in his original description. In a more restricted sense, considering those species de- We are indebted to L.Castiglioni, who found the fossil fined on skull material (O. andina and O. and allows us to perform this study. J. Bonaparte and A. Kramarz (MACN), G. Vucetich, D. Verzi and M. Merino rigens), this genus shows a similar morpho- (La Plata) and C. De Muizon (Paris) allowed us to study logical pattern to genera of Caviinae. Indeed, comparative material in their collections. To D. Verzi, C. this genus has an interorbital width less than Quintana and an anonymous referee by their invaluable braincase width, molar series with length equal comments. R. Madden and C. Dailey improved the En- glish language of the ms. This is a contribution to the to or longer than diastema length, a well de- project “Cenozoico continental y parálico del Uruguay” veloped nasolacrimal foramen, the third upper (M. Ubilla- CSIC). CSIC facilitated the visit of one of us molar of similar configuration and similar skull (MU) to Buenos Aires, La Plata and Paris Museums. size and overall shape. With reference to size, this genus is clearly smaller than the presumed LITERATURE CITED members of Dolichotinae (Fig. 5). “Orthomyctera” chapalmalense resemble AMEGHINO, F. 1889. Los mamíferos fósiles de la Repú- Dolichotis patagonum in morphology and size blica Argentina. Acta de la Academia Nacional de Ciencias en Córdoba, 6: XXXII+1-1026. (Fig. 5). “Orthomyctera” chapalmalense should BEHRENSMEYER, A.K. and R.W. HOOK. 1992. not be included in this genus as was first sug- Paleoenvironmental contexts and taphonomic modes. gested by Kraglievich (1930), and should in- Pp. 15-135. In: Terrestrial ecosystems through time. stead be considered a dolichotine arguably Evolutionary paleoecology of terrestrial plants and animals (Behrensmeyer, A.K.; J.D. Damuth, W.A. Di Dolichotis. Indeed, “Orthomyctera” Michele, R. Potts, H.D. Sues, and S.L. Wing, eds.). chapalmalense has a diastema longer than P4- Univ. of Chicago Press, Chicago and London, 568 M3 length, the mesopterygoid fossa positioned pp. at the level of the anterior and posterior prisms CALCATERRA, C. 1972. Dos roedores fósiles nuevos 2 para Uruguay y confirmación de otro. Comunicacio- of M , the nasolacrimal foramen is not observed nes Paleontológicas del Museo de Historia Natural de in lateral view, and the posterior border of the Montevideo, 1(2):11-21. second prism and the anterior border of the CIONE, A.L.; M. AZPELICUETA, M. BOND, A.A. additional posterior prism in M3 are parallel. CARLINI, J.R. CASCIOTTA, M.A. COUZZUOL, M. DE LA FUENTE, Z. GASPARINI, F.J. GOIN, J. The continental late Miocene of Uruguay is NORIEGA, G.J. SCILLATO-YANÉ, L. SOIBELZON, not preserved in the landscape of this country. E.P. TONNI, D. VERZI, and G. VUCETICH. 2000. Consequently, the majority of vertebrates in- Miocene vertebrates from Entre Ríos province, eastern cluded in the paralic deposits of the Camacho Argentina. Pp. 191-237. In: El Neógeno de Argenti- na. INSUGEO, Serie Correlación Geológica Formation are our only window into the terres- (Aceñolaza, F.G. and R. Herbst, eds.). Tucumán, 290 trial and fluvial environments of this period. pp. From the ecological requirements of its closest FIELDS, R.W. 1957. Hystricomorph rodents from the Late living relatives among the dolichotines, the Miocene of Colombia, South America. University of California Publication in Geological Sciences, record of “Prodolichotis” in the late Miocene 32(5):273-402. of southwestern Uruguay suggests open and FRANCIS, J.C. and A. MONES. 1965. Contribución a la probably arid or semiarid terrestrial environ- geología y paleontología de las barrancas de San ments. Lagostomopsis Kraglievich, 1926, is Gregorio, Departamento de San José, República Orien- tal del Uruguay. Kraglieviana 1:55-85. also known from these sediments (Francis and GERVAIS, H. and F. AMEGHINO. 1880. Los mamíferos Mones, 1965; Perea et al., 1994). Its most fósiles de la América del Sud. Savy-Igon Hnos. closely-related living genus, Lagostomus, in- XI+1-225. París y Buenos Aires. habits grasslands and thorn scrubs. Even though KRAGLIEVICH, L. 1930. Diagnosis osteológico dental the late Miocene dolichotines may have had de los géneros vivientes de la subfamilia Caviinae. Anales del Museo Nacional de Buenos Aires, 36:59- different ecological requirements than the liv- 96. ing species, these coincident species provide a KRAGLIEVICH, L. 1932. Diagnosis de nuevos géneros y likely environmental setting. especies de roedores cávidos y eumegámidos fósiles de la Argentina. Anales de la Sociedad Científica Argentina, 114:32-49. 302 Mastozoología Neotropical / J. Neotrop. Mammal.; 10(2):293-302 M. Ubilla and A. Rinderknecht
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