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Paleozoogeography of the Wine Mouse (Akodon oenos) & Late Holocene 
Paleoenvironments in South‐Central Mendoza, Argentina 
Fernando Julián Fernández 

 
Author Address: CONICET. Cátedra de Anatomía Comparada, Facultad de Ciencias Naturales y Museo, Universidad Nacional 
de La Plata, Calle 64 s/n (entre diag. 113 y calle 120), La Plata, Argentina 
fernandezf77@yahoo.com.ar  
 
Received: November 24th 2010                       Volume 1:52‐57 
Published: February 14th 2011                       © 2010 Society of Ethnobiology 
 
Abstract: Cranial remains of the wine mouse (Akodon oenos) are documented from an archaeological site  in south‐central 
Mendoza, Argentina (Agua de La Mula, 35º22' S, 68º15' W), which dates to the end of the late Holocene (1610 ± 60; 1260 ± 
60; 1000 ± 50 C14 yr B.P.). The taxonomic status of this small rodent is currently being assessed, but these remains represent 
the  first  fossil  record  for  the  morphotaxon  A.  oenos.  The  species’  present  distribution  is  restricted  to  a  few  records  from 
Mendoza province. Analysis of the remains supports paleoenvironmental reconstruction using the small mammal assemblage 
recovered from this site. From the late Holocene into modernity temperature decreased and winter precipitation increased, 
resulting in advance of Patagonian steppe grading with altitude into Monte desert. Holocene climatic conditions may explain 
the relatively late human occupation of ecologically marginal environments in this region, which probably favored effective 
human occupation of the Payunia region at sites such as Agua de La Mula between 1600 and 1000 years B.P. 

Key Words: Akodon oenos, paleoenvironmental analysis, Agua de la Mula, Mendoza, Argentina 
 

Introduction 
The wine mouse, Akodon oenos (Braun et al. 2000), is a 
poorly known small rodent of the family Cricetidae 
(subfamily Sigmodontinae, tribe Akondotini) for which 
no fossil remains have been reported. Knowledge 
about the biogeography of the species is restricted to a 
few records from Mendoza, Argentina.  These records 
range from north-central arid environments of the 
Monte desert and Puna in localities that are modified 
by modern human activities (mainly wineyards and 
olive groves) to the southwest (see Figure 1A) in 
wetland habitat in the foothills of the volcanic Payunia 
region, which is in the semi-arid Patagonian steppe 
(Braun et al. 2000; Contreras and Rosi 1980; Pardiñas 
et al. 2011). Akodon oenos lives in sympatry with A. 
molinae, the only akodontines reported within Mendoza 
(Braun et al. 2000; Pardiñas et al. 2011). In general, 
knowledge of small mammal paleobiogeography and of 
Holocene paleoenvironmental conditions from the 
volcanic Payunia region of Mendoza is limited. 

In this paper the first fossil record of the A. oenos 
from the Agua de La Mula archaeological site in south-
central Mendoza, Argentina is presented. In addition, 
the species’ taxonomic status and geographic dis-
tribution are discussed. The paleobiogeography of A. 
oenos is integrated into reconstruction of environmental 
conditions during the late Holocene in the region via 
analysis of the small mammal assemblages from Agua 

de La Mula. This paleogeographic and paleoenviron-
mental study expands ongoing discussion of human-
environment interactions during the very late Holocene 
in southern, central Mendoza. 

Study Area 
The Agua de La Mula site is located near the northern 
boundary of the volcanic Payunia region (Figure 1A). 
This area is within the Monte phytogeographic 
province (Cabrera 1976), which is included in the 
climatic region known as the South American Arid 
Diagonal covering a large part of the subcontinent 
from northern Peru along the Andes to the south of 
Neuquén continuing across Patagonia to the Chubut 
River (Bruniard 1982). The east band of the Payunia 
region is exposed to the action of the Atlantic 
anticyclone. However, the great distance traveled by the 
humid Atlantic winds results in low summer 
precipitation (~200 mm). The vegetation is charac-
terized by xerophytic shrubs, such as Prosopis torcuata, P. 
alpataco, Cercidium praecox, Chuquiraga erinacea, Cassia 
aphylla, Larrea, Bulnesia, and Plectrocarpa, and isolated 
stands of Geophroea decorticans that grow in low organic 
matter sandy or rocky soils (Abrahan et al. 2009; 
Cabrera 1976). 

 

 



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Figure 1. A: Map of Mendoza province (Argentina), recording localities for A. oenos: Triangles (archaeological) 1. Agua de La 
Mula (35º22' S, 68º15' W, 967 m above sea level). Circles (modern) 2. 2 km S of Villavicencio Ruta 32 (32º31' S, 68º59' W); 3. 
La Pega (32º48' S, 68º40' W, type locality); 4. Puesto de Lima (32º54' S, 69º01' W) and 5. Llancanelo Natural Reserve (35º38' 
S, 69º11' W). Phytogeography follows Cabrera (1976). B: Map of NW Argentina, recording localities for A. spegazzinii: star 
(type locality); circles (modern); triangles (fossil samples). 
 
 
Methods 
The Agua de La Mula site is a basaltic cave. 
Excavations directed by Dr. Humberto Lagiglia were 
done by the staff of Museo de Historia Natural de San 
Rafael in 1987 in 10 levels of 10 cm each. 1026 small 
mammal bones and bone fragments were recovered 
from levels 4, 5, 6, 7, 9, and 10, as were other organic 
materials such as remains of domestic plants (Zea mays 
and Cucurbita sp.) (LP-563, charcoal sample, level 10, 
1610 ± 60; LP-620, charcoal sample, level 10, 1260 ± 
60; LP-973, charcoal sample, level 5, 1000 ± 50 C14 yr 
B.P.). Two cranial specimens of A. oenos (a juvenile and 
an adult individual) were identified from levels 5 and 6.1 
Morphological description and craniodental measure-
ments of A. oenos (taken with a digital caliper to the 
nearest 0.01 mm) were made following criteria reported 
by Myers (1989). These data were compared with those 
provided by Braun et al. (2000) and with those from a 
sample of ten specimens of A. iniscatus mucus from the 

Neuquén province.2 Un-fortunately, measurements 
from the juvenile specimen of A. oenos from the site 
could not be taken due to its fragmentary condition. 

Paleoenvironmental analysis using small mammal 
remains is based on modern ecological requirements 
and biogeographic distributions for various taxa 
represented in the Agua de la Mula fauna. For 
comparative purposes an additional study was done on 
the area's modern small mammals. A sample of 22 
pellets of black-chested buzzard-eagle (Geranoaetus 
melanoleucus, Accipitridae) and trapping data were 
analyzed, identifying 28 individual micromammals. 

The Modern Small Mammal Assemblage 
The modern fauna is made up primarily of sig-
modontine rodents, with Graomys griseoflavus (MNI% 
17.9), and Eligmodontia sp. (MNI% 10.7) well re-
presented, followed by low frequencies of Phyllotis 
xanthopygus (MNI% 7.1), Calomys musculinus (MNI%  



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Figure 2. Dorsal, ventral and lateral views of cranium 
of Akodon oenos (MHNSR 15.002). Scale: 5 mm. 

7.1), and Akodon molinae (MNI% 3.6). A single hystri-
cognath rodent Galea leucoblephara (MNI% 28.6) was 
recorded, and one chiropteran Tadarida brasiliensis 
(MNI% 3.6), and one marsupial marmosine Thylamys 
pallidior (MNI% 3.6). The modern small mammal 
assemblage also contained one exotic lagomorph 
(MNI% 7.1). 

Small Mammals at Agua de La Mula 
A sample of 1026 cranial remains was analyzed, and 
represents an MNI of 491. In this assemblage, 
hystricognath rodents dominate, Ctenomys sp. (MNI% 
41.9), Microcavia australis (MNI% 7.7), and G. 
leucoblephara (MNI% 5.3), followed by remains of 
sigmodontine rodents, P. xanthopygus (MNI% 16.1), G. 
griseoflavus (MNI% 14.1), Eligmodontia sp. (MNI% 3.1), 
A. molinae (MNI% 2.2), Reithrodon auritus (MNI% 1), C. 
musculinus (MNI% 0.4), and A. oenos (MNI% 0.4). A 
single chiropteran T. brasiliensis (MNI% 0.4) was 
recorded, as were two marsupial marmosines T. 
pallidior (MNI% 6.3), and Lestodelphys halli (MNI% 1). 

The adult specimen of A. oenos is represented by a 
fragmented cranium that is missing the braincase 
(Figure 2). It is medium in size compared to other 
individuals in the genus with a relatively short and wide 
rostrum. The interorbital region is narrow and 
hourglass shaped with frontals that are rounded in the 
dorsolateral margins. The frontoparietal suture is 
crescent-shaped. The zygomatic notches are moderately 
wide and deep. The zygomatic plate is relatively broad, 
and its anterior margin is straight. The zygomatic 
arches are slender, the palatal bridge is short and 
narrow, and the incisive foramina extend posteriorly to 
the protocone of M1. The posterior palatal foramina are 
sligthly enlarged and are located at level of the M2; 
there is also a second pair of small posteropalatal pits 
situated at the level of M3. The mesopterygoid fossa is 
narrow, with an anterior border tending towards a lyre 
shape. Upper incisors are orthodont, and toothrows are 
parallel. Molars are relatively hipsodont and robust, 
with the major cups about equal in size. 

This description agrees with that provided by 
Braun et al. (2000) for A. oenos and differs from other 
species in Akodon that inhabit the region, such as A. 
molinae and A. iniscatus. Akodon molinae is distinguished 
by its comparatively wide interorbital region, square 
frontals, and parallel dorsolateral margins. The 
zygomatic plate of A. molinae is wide with slightly 
convex edges that are oriented obliquely backwards and 
down; the palatal bridge is long and wide. Akodon 
iniscatus is characterized by a long and wide palatal 
bridge, a narrow zygomatic plate, extension of the 
incisive foramina posteriorly to the hipocone of M1, a 
moderately narrow mesopterygoid fossa, a slightly M-
shaped anterior border, opistodont upper incisors, and 
slender molars. In terms of biometry, the specimen 
from Agua de La Mula falls within the range of A. oenos 
described by Braun et al. (2000) (Table 1). 

Discussion 
The taxonomic status of A. oenos is currently being 
assessed. The species was described by Braun et al. 
(2000) based on individuals labeled by Contreras and 
Rosi as “Akodon minoprioi,” and as that original study 
was not formally published A. minoprioi, it became a 
nomen nudum (Galliari et al. 1996). Recent research 
conducted by Pardiñas et al. (2011) based on extensive 
morphologic and molecular data suggests that A. oenos 
is a junior synonym of A. spegazzinii and that it belongs 
to the A. boliviensis species group of Akodon. The 
geographic distribution of A. spegazzinii extends to La 
Rioja, Catamarca, Tucumán, and Salta provinces, with 
the type locality in Lower Cachi, central Salta 
(25º07'11.93''S, 66º09'47''W, 2341 m above sea level,  



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Table 1. Descriptive statistics of craniodental measurements (in mm) of the Agua de La Mula adult specimen of 
Akodon oenos, the A. oenos holotype, and individuals of A. molinae and A. iniscatus nucus. 

   MHNSR  A. oenos Holotype  A. oenos  A. molinae  A. iniscatus nucus 

  15.002  IADIZA‐CM611*  mean*  mean*  mean 

Diastema length  6.4  6.9  6.7 ± 0.35  6.9 ± 0.47  6.5 ± 0.29 

Palatal bridge  3.2  3.6  3.3 ± 0.20  4.0 ± 0.31  2.6 ± 0.21 
Maxillary toothrow length  5.0  4.9  4.8 ± 0.19  4.8 ± 0.04  4.6 ± 0.24 

Incisive foramina length  5.9  5.3  5.8 ± 0.37  5.7 ± 0.15  6.3 ± 0.21 
Zygomatic plate breadth  2.6  2.5  2.8 ± 0.20  3.1 ± 0.19  2.9 ± 0.16 

Mid rostral width  4.1  3.9  4.0 ± 0.15  4.2 ± 0.23  4.2 ± 0.20 

Interorbital constriction  4.6  4.6  4.6 ± 0.15  4.7 ± 0.16  4.5 ± 0.15 
* Data from Braun et al. (2000: 219). 

Figure 1B). If this taxonomic designation is correct, A. 
spegazzinii extends its geographic distribution and 
ecological domain from Salta province to southern 
Mendoza, ranging from 400 to about 3500 m above sea 
level, inhabiting arid environments of the Patagonian 
steppe, the Monte desert, the Puna, and the high 
Andes, as well as the Yungas forests, and the semi-arid 
forests at the ecotone of Yungas-Chaco (see Jayat 2009; 
Jayat et al. 2010). In addition, many fossil remains 
identified as A. spegazzinii have been recovered from 
two paleontological sites in northwestern Argentina, 
located in deposits dating to the middle-upper 
Pleistocene (26º56' S, 65°42' W, La Angostura, 
Tucumán province [Ortiz and Pardiñas 2001]) and to 
the Pleistocene-Holocene boundary (26º51' S, 65º43' 
W, Tafi del Valle, Tucumán province [Ortiz and Jayat 
2007]). 

The presence of pellets preserved in stratigraphy 
and light digestive corrosion on some teeth and 
postcraneal bones recovered from Agua de La Mula 
site, indicate that the main accumulator agent was 
probably an owl (Fernández et al. 2008). Small 
mammals collected by owl are considered good 
indicators of environmental conditions (Andrews 
1995). The modern association of small mammals from 
Agua de La Mula shows a clear predominance of 
species characteristic of the Monte desert and the 
South American Arid Diagonal (G. leucoblephara, G. 
griseoflavus, C. musculinus, A. molinae, and T. pallidior), with 
the addition of two general Andean Patagonian taxa (P. 
xanthopygus and Eligmodontia sp.). Also, the occurrence 
of P. xanthopygus suggests a landscape dominated by 
open rocky areas. 

Remains of A. oenos from Agua de La Mula were 
recovered together with those of small mammal species 
that inhabit the area today (A. molinae, G. griseoflavus, P. 

xanthopygus, C. musculinus, G. leucoblephara and T. 
pallidior). It is interesting to note the presence at this site 
of L. halli, an endemic marsupial of the Patagonian 
steppe that is rare in the Monte desert of Mendoza 
today (Chacras de Coria, 32º45' S, 69º00' W; 
Huayquerías del Oeste, 33º38' S, 68º26' W; 50 km N 
San Rafael, 34º15' S, 68º40' W). Modern populations of 
L. halli are considered relicts of those that were more 
widely distributed earlier in the Holocene. In addition, 
the occurrence of R. auritus suggests the development 
of open, herbaceous steppe environments associated 
with bodies of water. The presence of both species at 
Agua de La Mula indicates wetter and colder conditions 
in the past, which may have been the result of 
westerlies generated by the Pacific anticyclonic center 
(causing winter rainfall) during the late Holocene. 

Palynological studies on materials from an 
archaeological site in the western plains of Mendoza 
(Gruta del Indio, 34º45' S, 68º22' W), 70 km north of 
Agua de La Mula, reveal that the establishment of 
modern climatic conditions occured at roughly 3000 yr 
B.P. (D’Antoni 1983). However, D’Antoni (1983: 97) 
observed a significant change in the pollen sequence 
between 1600 and 1200 yr B.P. when the Patagonian 
steppe vegetation increased (e.g., Poaceae, Cyperaceae, 
and Adesmia) and Monte vegetation decreased (e.g., 
Larrea). D’Antoni suggested that change may have 
resulted from selective wood exploitation by humans 
that inhabited this area. However, no evidence of wood 
exploitation in Agua de La Mula was found, and based 
on the small mammals analysis presented here, I 
propose that near Agua de La Mula between 1600 and 
1000 years B.P. temperature decreased and winter 
precipitation increased, which produced an advance of 
Patagonian steppe and its associated fauna, developing 



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into a transitional mosaic of Patagonian steppe and 
Monte desert. 

Available zooarchaeological and modern data 
indicate that A. oenos are absent from Mendoza’s high 
Andean environments.3 However, the fossil akodontine 
remains from Agua de La Mula represent an 
extralimital record in the palaeoenvironmental mosaic 
of the Patagonian-Monte during the late Holocene 
(Figure 1A).  This species may currently be present in 
such environments and may be common in the 
unexplored volcanic Payunia region, which may be of 
interest to conservation scientists.  

More comprehensive understanding of climatic 
change during the late Holocene in the region is 
important for the study of the late human occupation 
of ecologically marginal environments, such as the 
Payunia. Less dry and cooler environmental conditions 
observed at Agua de La Mula between 1600 and 1000 
years B.P. probably favored human ocupation in all 
environments of Payunia (i.e., effective human 
occupation, sensu Borrero 1994-1995). Human-
environment interactions have been discussed in 
archaeological literature from Mendoza (e.g., Gil 2006; 
Neme 2007). Available evidence suggests that 
economic intensification and effective occupation of 
Andean environments took place at 2000 yr B.P. 
(Neme 2007). This intensification process has been 
interpreted as the consequence of an imbalance 
between environmental carrying capacity and human 
population growth. Over-exploitation of the 
environment reduced the availability of highly-ranked 
resources (e.g., Lama guanicoe, Rhea americana, Pterocnemia 
pennata), and drove subsistence towards the inclusion of 
foods with lower caloric returns and higher processing 
costs such as some plants and small mammals (Gil 
2006; Neme 2007). However, increased human 
occupation and economic intensification in the volcanic 
Payunia region occurred subsequently (about 1000 yr 
B.P) and may relate to climate amelioration indicated 
by evidence presented here. 

In summary, small mammals remains from the 
Agua de La Mula site in southern Mendoza strengthen 
understanding of the biogeography and ecology of A. 
oenos. The presence of this species in deposits that date 
to the late Holocene just prior to human the 
occupation of the Payunia during a period in which 
climate amelioration occurred provides insights into the 
conditions that promoted human occupation of the 
region and into the conservation biology of the species 
today. 

 

Acknowledgements 
I thank Ulyses Pardiñas, Cesar García-Esponda, 
Gustavo Neme, Pablo Teta, and Steve Wolverton for 
editorial comments and discussion. Gustavo Neme and 
Adolfo Gil provided access to the Agua de La Mula 
collection through the Museo de Historia Natural de 
San Rafael. Cesar García Esponda, Gonzalo Martinez 
and Fernando Ballejo assisted with preparation of the 
images. Germán Moreira, Fernando Ballejo, and 
Luciano De Santis provided assistance in the field and 
lab.  Three anonymous reviewers provided constructive 
comments. 

References Cited 
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F. Coronato, and R. Godagnone. 2009. Overview of 
Geography of the Monte Desert Biome (Argentina). 
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Andrews, P. 1995. Mammals as Palaeoecological 
Indicators. Acta Zoológica Cracovensia 38:59-72. 

Borrero, L. A. 1994-1995. Arqueología de la Patagonia. 
Palimpsesto 4:9-56. 

Braun, J. K., M. A. Mares and R. A. Ojeda. 2000. A 
New Species of Grass Mouse, Genus Akodon (Muridae: 
Sigmodontinae), from Mendoza Province, Argentina. 
Zeitschrift für Säugetierkunde 65:216-225. 

Bruniard, E. 1982. La Diagonal Árida Argentina: Un 
Límite Climático Real. Revista Geográfica 95:5–20. 

Cabrera, A. L. 1976. Regiones Fitogeográficas 
Argentinas. Enciclopedia Argentina de Agricultura y 
Jardinería 1:1-85. 

Contreras, J. F. and M. I. Rosi. 1980. Comportamiento 
Territorial y Fidelidad al Hábitat en una Población de 
Roedores del Centro de la Provincia de Mendoza. 
Ecología Argentina 5:17-29.  

D´Antoni, H. 1983. Pollen Analysis of Gruta del Indio. 
Quaternary of South America and Antartic Peninsula 1:83-
104. 

Fernández, F. J., G. J. Moreira, and L. J. M. De Santis. 
2008. Análisis Preliminar del Ensamble de 
Micromamíferos del Sitio Arqueológico “Agua de La 
Mula” (Mendoza, Argentina). Paper presented at the 
First Congreso Nacional de Zooarqueología Argentina, 
Malargüe, Mendoza. 

Galliari, C. A., U. F. J. Pardiñas, and F. J. Goin. 1996. 
Lista Comentada de los Mamíferos Argentinos. 
Mastozoología Neotropical 3:39-61. 



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Gil, A. F. 2006. Arqueología de La Payunia (Mendoza, 
Argentina). El poblamiento humano en los márgenes de la 
agricultura. BAR Internacional Series 1591, Oxford. 

Jayat, J. P. 2009. Roedores Sigmodontinos de los 
Pastizales de Neblina de las Yungas de Argentina. 
Ph.D. Dissertation (Paleontology). Universidad 
Nacional de Tucumán, Tucumán. 

Jayat, J. P., P. E. Ortiz, J. Salazar-Bravo, U. F. J. 
Pardiñas, and G. D’Elía. 2010. The Akodon boliviensis 
Species Group (Rodentia: Cricetidae: Sigmodontinae) 
in Argentina: Species Limits and Distribution, With the 
Description of a New Entity. Zootaxa 2409:1-61. 

Myers, P. 1989. A Preliminary Revision of the Varius 
Group of Akodon (A. dayi, dolores, molinae, neocenus, 
simulator, toba and varius). In Advances in Neotropical 
Mammalogy, eds. K. H. Redford and J. F. Eisenberg, pp. 
5-54. Sandhill Crane Press, Florida.  

Neme, G. 2007. Cazadores Recolectores de Altura en los 
Andes Meridionales. BAR Series 1591, Oxford. 

Ortiz, P. E. and U. F. J. Pardiñas. 2001. Sigmodontinos 
(Mammalia, Rodentia) del Pleistoceno Tardío del Valle 
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y Reconstrucción Paleoambiental. Ameghiniana 38:3-26. 

Ortiz, P. E. and P. Jayat. 2007. Sigmodontinos 
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44:641-660.  

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Biosketch 
Fernando Fernández is a professor of Zooarchaeology on 
the Facultad de Ciencias Naturales y Museo (UNLP) in La 
Plata,  Argentina.  He  is  an  advanced  doctoral  student 
funded  by  CONICET  at  the  same  university  and  has 
produced  more  than  30  publications  and  conference 
presentations  on  taphonomy,  zooarchaeology,  and 
zoogeography of microvertebrates. 

 

 

                                                            

1  The remains were accessioned to the archaeological 
collection of Museo de Historia Natural de San Rafael 

                                                                                                  

as MHNSR 15.002 (adult specimen) and MHNSR 
15.003 (juvenile specimen). 
2  The modern specimens of Akodon iniscatus nucus are 
housed in the Colección de Egagrópilas y Afines “Elio 
Massoia” of the Centro Nacional Patagónico (Puerto 
Madryn, Chubut, Argentina) under the number CNP-E 
88 (owl pellet sample). 
3 Fernández, F. J. Microvertebrados del Holoceno de 
sitios arqueológicos en el sur de Mendoza (República 
Argentina): aspectos tafonómicos y sus implicancias en 
la subsistencia humana. Ph.D. Dissertation (in 
preparation). Facultad de Ciencias Naturales y Museo, 
Universidad Nacional de La Plata, La Plata.